ISSN 0006 8241 = Bothalia

Bothalia

A JOURNAL OF BOTANICAL RESEARCH

Vol. 34,1 May 2004

TECHNICAL PUBLICATIONS OF THE NATIONAL BOTANICAL INSTITUTE,

PRETORIA

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

BOTHALIA

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

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

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

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

published annually.

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

STRELITZIA

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

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

MEMOIRS OF THE BOTANICAL SURVEY OF SOUTH AFRICA

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

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

ANNALS OF KIRSTENBOSCH BOTANIC GARDENS

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

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

South African Botany). Discontinued after Vol. 19.

FLOWERING PLANTS OF AFRICA (FPA)

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

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

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

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

or of economic value for illustration.

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

An index to Vols 1-49 is available.

FLORA OF SOUTHERN AFRICA (FSA)

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

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

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

and ecological notes.

Contributions to the FSA also appear in Bothalia.

PALAEOFLORA OF SOUTHERN AFRICA

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

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

available:

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

Anderson.

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

& H.M. Anderson.

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

1 1 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 34,1

Scientific Editor: G. Germishuizen

Technical Editor: B.A. Momberg

NATIONAL

Botanical

INSTITUTE

2 Cussonia Avenue, Brummeria, Pretoria

Private Bag X101, Pretoria 0001

ISSN 0006 8241

May 2004

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, National Botanical Institute, Cape Town, RSA

University of Utrecht, Utrecht, The Netherlands

Acknowledgements to referees

Bandeira, Dr S.O. Universidade Eduardo Mondlane, Maputo, Mozambique.

Bramwell, Dr D. Jardin Botanico Canario Viera y Clavijo, Las Palmas de Gran Canaria, Spain.

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

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

Fellingham, Mrs A.C. 23 Moreson Ave, Valmary Park, Durbanville, RSA.

Fraser-Jenkins, Dr C.R. Student Guest House, Thamel, Kathmandu, Nepal.

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

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

Jordaan, Ms M. National Botanical Institute, Pretoria, RSA.

Kirkwood, D. University of Cape Town, RSA.

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

Lobin, Dr W. Botanische Garten, Bonn, Germany.

Maslin, Dr B. Perth, Western Australia.

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

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

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

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

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

Ross, Dr J. Royal Botanic Gardens, Melbourne, Australia.

Smithies, Mrs S. National Botanical Institute, Pretoria, RSA.

Stuppy, Dr W. Royal Botanic Gardens, Ardingly, West Sussex, UK.

Toelken, Dr H.R. State Herbarium, Adelaide, South Australia.

Van Rooyen, Dr G. University of Pretoria, RSA.

Wigginton, M.J. 36 Big Green, Warmington, Peterborough, UK.

Zmarzty, Dr S. Rotal Botanic Gardens, Kew, UK.

CONTENTS

Volume 34,1

1. The genus Cliffortia (Rosaceae) in KwaZulu-Natal. C.M. WHITEHOUSE 1

2. Two new species of Erica (Ericaceae); one from Western Cape and one from KwaZulu-Natal, South

Africa. E.G.H. OLIVER and I.M. OLIVER 11

3. Two new species of Romulea (Iridaceae: Crocoideae) from the western Karoo, Northern Cape and notes

on infrageneric classification and range extensions. J.C. MANNING and P GOLDBLATT .... 17

4. Studies in the genus Riccia (Marchantiales) from southern Africa. 26. A new species in section Pilifer,

Riccia radiata, is described. S.M. PEROLD 23

5. Notes on African plants:

Boraginaceae. A first record of Echium simplex in South Africa. G.F. SMITH, A.E. VAN WYK,

E.M.A. STEYN and E. RETIEF 44

Crassulaceae. Adromischus schuldtianus subsp. brandbergensis, a new subspecies and a checklist

of the succulent flora of the Brandberg, Namibia. EJ. VAN JAARSVELD, B. NORDENSTAM

and A.E. VAN WYK 35

Ericaceae. Nomenclatural changes in Erica. E.G.H. OLIVER 38

Ericaceae. A new species of indehiscent-fruited Erica from the central Kouebokkeveld, Western

Cape, South Africa. R.C. TURNER and E.G.H.OLIVER 39

Fabaceae. A new species of Acacia (Mimosoideae) from Mpumalanga, South Africa. P.J.H. HURTER

and A.E. VAN WYK 42

Lamiaceae. Plectranthus mzimvubuensis, a new species from Eastern Cape, South Africa. E.J. VAN

JAARSVELD and A.E. VAN WYK 30

Mesembryanthemaceae. A new tribe and adjustments to infrafamilial classification. P. CHESSELET,

A.E. VAN WYK and G.F. SMITH 47

Pteridophyta. Dryopteris fdipaleata (Pteropsida: Dryopteridaceae), a new species from tropical

East Africa. J.P. ROUX 27

Pteridophyta. Dryopteris gorgonea (Pteropsida: Dryopteridaceae), a new species from the Cape

Verde Islands. J.P. ROUX 32

Scrophulariaceae. Type specimens of Selago, Jamesbrittenia and Sutera at Natal University

Herbarium (NU). T.J. EDWARDS 27

6. Functional and taxonomic significance of seed structure in Salix mucronata (Salicaceae). E.M.A. STEYN,

G.F. SMITH and A.E. VAN WYK 53

7. Grass assemblages and diversity of conservation areas on the coastal plain south of Maputo Bay, Mozam-

bique. S.J. SIEBERT, L. FISH, M.M. UIRAS and S.A. IZIDINE 61

8. Obituary: Johannes Jacobus Adriaan van der Walt (1938-2003). P. VORSTER 73

New species, subspecies, name, tribe and varieties in Botbalia 34,1 (2004)

Acacia ebutsiniorum P.J.H.Hurter, sp. nov., 42

Adromischus schuldtianus (Poelln.) Poelln. subsp. brandbergensis B.Nord. & Van Jaarsv., subsp. nov., 35

Cliffortia dracomontana C.M.Whitehouse , sp. nov., 3

Cliffortia paucistaminea Weim. var. australis C.M.Whitehouse , var. nov., 7

Drosanthemeae Chesselet, Gideon F.Sm. & A.E.van Wyk, trib. nov., 49

Dryopteris filipaleata J.P.Roux , sp. nov., 28

Dryopteris gorgonea J.P.Roux, sp. nov., 33

Erica jananthus E.G.H.Oliv. & I.M.OIiv., sp. nov., 11

Erica psittacina E.G.H.Oliv. & I.M.OIiv., sp. nov., 13

Erica salicina E.G.H.Oliv., nom. nov., 38

Erica tragomontana R.C. Turner, sp. nov., 39

Plectranthus mzimvubuensis Van Jaarsv., sp. nov., 30

Riccia radiata Perold, sp. nov., 23

Romulea collina J.C. Manning & Goldblatt, sp. nov., 17

Romulea eburnea J.C. Manning & Goldblatt, sp. nov., 19

iv

Bothalia 34,1 : 1-10 (2004)

The genus Cliffortia (Rosaceae) in KwaZulu-Natal

C.M. WHITEHOUSE*

Keywords: area of endemism, Cliffortia L., Drakensberg, KwaZulu-Natal, Rosaceae

ABSTRACT

The only area of endemism for Cliffortia L. outside of the Cape Floristic Region (CFR) is centred in the northern KwaZulu-

Natal Drakensberg. Eleven species of Cliffortia have been recorded from KwaZulu-Natal and distribution maps are provided

for each. Clarification of the circumscription of C. browniana Burtt Davy is discussed and a new species, C. dracomon-

tana. is described. C. paucistaminea Weim. is subdivided into two varieties: var. australis and var. paucistaminea.

Comments on some of the other species that have frequently been misidentified are also provided along with a key to all

the species in the area.

INTRODUCTION

Cliffortia L. is the largest southern African genus

within the Rosaceae. It was last comprehensively revised

by Weimarck (1934, 1948) and subsequent published

work has focused upon describing new species and clari-

fying taxonomy (Weimarck 1953, 1959; Oliver & Fel-

lingham 1991, 1994; Fellingham 1993a. b. 1994, 1995.

2003). Recent work by Whitehouse (2003) using mor-

phological and molecular techniques has indicated that

some of the species boundaries need revising to reflect

more accurately the diversity found within Cliffortia.

Cliffortia has its centre of diversity in the Fynbos

Biome of the Cape Floristic Region (CFR) and over 80%

of its species are endemic to that region (Goldblatt &

Manning 2000; Whitehouse 2003). Linder (2001 ) stated

several criteria to define areas of endemism: each area

should contain at least two endemic species; the areas

must be mutually exclusive; the areas must be narrower

than the study area and several areas need to be defined

for any discussion about their biogeography to be inter-

esting; and finally, the ranges of the endemic species

should be optimized to be maximally congruent. Using

these criteria, only a single area of endemism exists out-

side of the CFR and is centred on the northern KwaZulu-

Natal Drakensberg (Whitehouse 2003).

Eleven species of Cliffortia have been recorded with-

in KwaZulu-Natal. C. odorata L.f. is recorded from a

single old collection near Port Shepstone (Alexandra

District, Hlokozi, 22 Feb. 1916. Rudatis 2242). The

remaining species are either endemic to the Drakensberg

or widespread, though scattered, through the province.

C. filicauloides Weim. and C. spathulata Weim. are

endemic to the northern KwaZulu-Natal Drakensberg,

whereas the newly described species C. dracomontana

C.M. Whitehouse is also endemic to the main Drakens-

* Bolus Herbarium, University of Cape Town, 7700 Rondebosch, Cape

Town.

MS. received: 2003-02-14.

berg escarpment but is found as far south as Ben

Macdhui in Eastern Cape.

Despite the limited number of species present, the tax-

onomy and identification has often been confused. The

flowers are small and remarkably uniform with few diag-

nostic characters. Therefore, species determination is pri-

marily based upon vegetative characters, especially leaf

form and achenes when present. However, whereas sever-

al species are easily determined from the leaves of their

mature plants, their juvenile foliage (i.e. the first few true

leaves of seedlings or of new shoots resprouting after a

fire) is morphologically very different. Seedlings and

resprouting plants of Cliffortia have frequently been incor-

rectly labelled as unrelated species and this has confound-

ed the identification of otherwise easily discernible spe-

cies. Mature plants also sometimes show ecotypic vari-

ation in the size and shape of their leaves, especially on the

borders of forest or rivers compared with more exposed

slopes. Furthermore, the possibility of hybridization and

subsequent introgression as found in several Cape species

(Weimarck 1934; Fellingham 1993a; Whitehouse 2003)

cannot be dismissed, although no confirmed examples

have yet been demonstrated within KwaZulu-Natal. A key

is presented here that attempts to take account of these

variations and to identify species, where possible, even

when only juvenile foliage is present.

The conservation status of all the KwaZulu-Natal

species of Cliffortia should be regarded as ‘lower risk,

least concern" for IUCN Red Data List assessments

(Golding 2002). The widespread species are common

and often weedy. In particular, C. linearifolia and C.

nitidula subsp. pilosa are sometimes dominant and have

then been used in defining certain vegetation types (e.g.

Killick 1963; Edwards 1967). The endemic species are

more localized but all grow within the protected area of

the uKhahlamba-Drakensberg Park, generally on the

higher slopes and in more inaccessible areas. As a result,

although currently only recorded from a few scattered

localities, they are probably present and common in the

intervening parts of their ranges too.

2

Bothalia 34,1 (2004)

Key to KwaZulu-Natal species of Cliffortia

la Leaves unifoliate, broadly ovate to subcircular, more than 10 mm wide odorata

lb Leaves linear to obovate, less than 10 mm wide:

2a Leaves unifoliate, narrowly oblong, usually more than 20 x 2 mm; growing in or beside water strobilifera

2b Leaves less than 20 mm long, if longer then belonging to a juvenile plant and trifoliate with needle-shaped

leaflets:

3a Leaves with petiole; leaflets broadly obovate and at least middle leaflet toothed:

4a Plants decumbent or with arching stems; outer leaflets narrower than middle leaflet and only middle leaflet

toothed; leaves sparsely hairy, sometimes only on midrib and margins; endemic to northern Drakens-

berg between Loteni River and Cathedral Peak filicauloides

4b Plants erect; middle leaflet similar to outer leaflets, all toothed; leaves hairy throughout

nitidula subsp. pilosa (juvenile foliage)

3b Leaves without petiole, or if present then leaves oblong to linear and not toothed:

5a Leaves unifoliate:

6a Medium to tall shrub with virgate or tangled stems; young stems densely hairy; leaves with midrib and

margins thickened so that they touch each other; sepals 4; stamens 4; stigma white to pinkish . . linearifolia

6b Low shrub with spreading stems that ascend towards apex; young stems glabrous or sparsely and very

shortly hairy; leaves without noticeable thickening on midrib or margins; sepals 3; stamens 6; stigma

red repens

5b Leaves trifoliate:

7a Leaves sparsely hairy especially on young growth; margins usually inrolled beneath nitidula subsp. pilosa

7b Leaves always glabrous; margins not inrolled although sometimes thickened:

8a Very short petiole, <1 mm long, often evident; leaves narrowly oblong, 2-8 x 0. 5-1.0 mm, mucronate to

rounded at apex; sepals 3 ramosissima*

8b Petiole absent; sepals 4, very rarely 3 and then leaves needle-shaped with sharply pointed apex:

9a Leaves linear to needle-shaped:

10a Resprouting plants with leaflets more than 10 mm long, sterile:

11a Stems densely, shortly hairy linearifolia (juvenile foliage)

lib Stems glabrous or almost so repens (juvenile foliage)

10b Mature foliage with leaflets less than 13 mm long:

12a Midrib and margins of leaflets thickened so that they touch each other; apices obtuse to rounded

linearifolia

12b Margins of leaflets not thickened; midrib slightly thickened to form a keel; apices acute to long

acuminate:

1 3a Leaflets curved upwards and towards stem, giving a feathery appearance to branch; George to

Port Elizabeth paucistaminea var. australis

13b Leaflets straight or curved downwards and away from stem; brachyblasts sometimes appearing

star-shaped; Port Elizabeth to KwaZulu-Natal paucistaminea var. paucistaminea

9b Leaves narrowly elliptic to broadly obovate:

14a Leaves glaucous; margins always entire; bracteole margins smooth; achene 3-5 mm long;

2 400-3 100 m a.s.l dracomontana

14b Leaves glaucous or not; at least juvenile leaves toothed; if mature leaves entire then bracteole margins

shortly ciliate and achene less than 3 mm long:

15a Leaves broadly obovate, 1-4 mm wide; mature leaves toothed; achene 3-4 mm long;

1 800-2 800 m a.s.l spatliulata

15b Leaves elliptic to narrowly obovate, 0.5-2 mm wide; mature leaves entire; achene less than 3

mm long:

16a Leaflets narrowly elliptic; apices obtuse to acute; achene darkish brown, indistinctly ribbed;

1 000-2 500 m a.s.l browniana

16b Leaflet narrowly elliptic to obovate; apices rounded to obtuse; achene beige to pale brown,

clearly ribbed; 0-1 600 m a.s.l serpyllifolia

Identity ofC. browniana

One species that has been particularly misunderstood

in the past is C. browniana Burtt Davy. Its type collec-

tion was from the Mpumalanga Drakensberg escarpment

near Lydenburg, from where several similar collections

have subsequently been made. Morphologically, C. browni-

ana has no unique diagnostic character and is very diffi-

cult to separate from some small elliptic-leaved forms of

C. serpyllifolia Cham. & Schltdl., with which it has often

been confused. Within KwaZulu-Natal the two species

need not be confused, as their distribution (see Figure 2)

and altitude ranges do not overlap. However, the C. ser-

pyllifolia grows on the Silotwane Hills of Swaziland,

closer geographically to the type locality populations of

* C. ramosissima has not been recorded from KwaZulu-Natal, but is

included in the key because it is the only species that has also been

recorded from the Free State, Mpumalanga, Limpopo and Lesotho. The

key is therefore made more widely applicable by its inclusion.

C. browniana than those in the KwaZulu-Natal

Drakensberg. Consequently, the exact nature of the rela-

tionship between the Mpumalanga populations and the

specimens also attributed to C. browniana in the

KwaZulu-Natal Drakensberg needs to be clarified by

molecular techniques.

The identity of C. browniana has been further con-

fused by two distinct species being included under the

same name within the KwaZulu-Natal Drakensberg.

Originally Weimarck recognized the two entities as dis-

tinct, describing the specimen Hutchinson 102 separate-

ly in the footnotes to C. browniana. However, when he

later described C. spatliulata he also included a picture

of the latter entity ( Esterhuysen 10183) under the name

C. browniana. Consequently the two species have been

included under the same name (e.g. Hilliard & Burtt

1987). They are in fact easily distinguishable both in the

herbarium and the field and the latter species is here

described as C. dracomontana.

Bothalia 34,1 (2004)

3

Finally, the nature of the juvenile foliage of C. brow-

niana has also caused confusion. Whereas the mature

foliage has small, entire, narrowly elliptic leaves, the

juvenile leaves are toothed. This caused Hilliard & Burtt

(1987) to question whether it could be distinguished

from C. spathulata. However, mature plants are easily

distinguishable, especially if fertile, and the distribution

of true C. spathulata does not actually include Hilliard &

Burtt’s study area of the southern Drakensberg. In con-

trast C. dracomontana has entire juvenile leaves, which

provide another diagnostic character for that species. On

the other hand, seedling leaves of C. setpyllifolia are also

toothed and it is questionable whether juvenile plants of

C. browniana could be distinguished from them on mor-

phology if the locality was not known.

1. C. browniana Burtt Davy , A manual of the

flowering plants and ferns of the Transvaal with Swazi-

land, South Africa 2: xxi, 316 (1932); Weim.: 56, fig.

13A-D (1934); Hilliard & B.L.Burtt: 162 (1987) pro parte.

Type: Eastern Transvaal [Mpumalanga], 2530 (Lyden-

burg): Lydenberg, (-AB), Rogers 22985 (BOL. holo.!; K!).

C. spathulata auctt. non Weim.: Hilliard & B.L.Burtt: 163 (1987).

Erect, medium shrub, up to 0.5 m high; densely divari-

cately branched, forming brachyblasts; young stems

0.8-1. 2 mm wide, reddish tinged, hairy; stem hairs

upwardly adpressed, 0.2-0. 7 mm long. Leaves trifoliate,

chartaceous, 0. 2-0.4 mm thick, midrib not prominent

above lamina, ± held straight or slightly curved upwards

and towards stem, green with two paler stripes on either

side of midrib beneath, glabrous above and beneath;

FIGURE 1. — Individual leaves of Cliffortia species, x 3. A, C. brown-

iana: B, C. dracomontana ; C, C. serpyllifolia ; D, C. spathula-

ta', E, C. fdicauloides\ F, juvenile leaf of C. nitidula subsp.

pilosa : G, mature leaf of C. nitidula subsp. pilosa.

FIGURE 2. — Known distribution of Cliffortia browniana, El; and

Cliffortia serpyllifolia. A, in southern Africa.

sheath 0.9-1. 4 mm long, abaxially glabrous, adaxially

hairy; stipules 0.4-1. 2 mm long, free, margin ciliate;

petiole absent; leaflets elliptic, 1.9^4. 3 x 0.6-1. 3 mm,

apex acute, margins flat, entire. Flowers solitary in axil

of undifferentiated leaves; bracteoles glabrous except for

ciliate margins; sepals 4, glabrous. Male flowers un-

known. Female flowers : bracteoles 1 .3-2. 1 mm long,

longer than immature receptacle; sepals ovate, 0.9-1. 4 x

0.4-0. 7 mm, recurved; carpels 1 or 2; stigma 0.9-1 .3 mm

long, red, feathery, ± hidden by leaves; immature recep-

tacle 1.0-1. 3 x 0.6-0. 8 mm, glabrous, smooth. Achene

2. 1-2.4 x 1.0-1. 3 mm, medium brown, glabrous, faintly

or indistinctly ribbed. Flowering time'. November to

May. Figure 1 A.

Habitat : quartzitic sandstone derived soils in full sun,

often in damp areas or amongst rocks; altitude 1 050-

2 500 m.

Distribution'. KwaZulu-Natal Drakensberg escarpment

from Sehlabathebe in Lesotho to Witsieshoek, with a dis-

junction to the Mpumalanga Drakensberg escarpment

between Graskop and Wolkberg, and an outlying popula-

tion on Blouberg. Figure 2.

Etymology: not explicitly stated but presumably named

after N.E. Brown.

2. C. dracomontana C.M.Whitehouse, sp. nov.,

C. browniana Burtt Davy affinis, sed foliis obovatis

glaucis longioribus, juvenibus foliis integris, bracteolis

glabris, fructibus majoribus differt.

TYPE. — KwaZulu-Natal, 2929 (Underberg): 8-11 km

NNW of Castle View Farm, headwaters of Mlahlangubo

River, (-CB), Hilliard & Burtt 13561 (NU, holo.!; K!,

PRE).

C. browniana auctt. non Burtt Davy: Weim.: 56 (1934) pro parte;

Weim.: t. 5 (1948); Killick: 127 (1963); D.Edwards: 264 (1967) pro

parte; J.H.Ross: 183 (1972); Hilliard & B.L.Burtt: 162 (1987) pro

parte.

Erect, low to medium shrub, up to 0.5 m high; dense-

ly divaricately branched, forming brachyblasts; young

4

Bothalia 34.1 (2004)

FIGURE 3. — Cliffortia dracomontana. A, fruiting branch, x 2.5; B.

achene x 5.

stems 0.7-1. 0 mm wide, often reddish tinged, hairy; stem

hairs upwardly adpressed, 0.2-0. 5 mm long. Leaves trifo-

liate, chartaceous, 0. 1-0.3 mm thick, midrib not promi-

nent above lamina, held ± straight, glaucous, glabrous

above and beneath; sheath 0.9-1. 5 mm long, abaxially

glabrous, adaxially markedly hairy; stipules 0.8- 1.5 mm

long, free, margin smooth; petiole absent; leaflets elliptic

to obovate, 2. 8-6.0 x 0.7-2. 1 mm, apex obtuse to round-

ed, margins flat, entire. Flowers solitary in axil of undif-

ferentiated leaves; bracteoles glabrous, margins smooth;

sepals 4, glabrous. Male flowers : pedicel 0.5-0. 7 mm

long; sepals broadly ovate, 2.5-3. 1 x 0.9- 1.2 mm, acute

to acuminate at apex; stamens 4-6; filaments 1.8-2. 3 mm

long. Female flowers', bracteoles 1 .7-2.5 mm long, longer

than immature receptacle; sepals ovate, 1 .2-1.5 x 0. 5-0.6

mm, recurved; carpel I; immature receptacle glabrous,

smooth. Achene 2.9 — 4.5 x 1.3-1. 8 mm, brown, glabrous,

faintly ribbed and slightly tuberculate to rugose. Flower-

ing lime : October to November. Figures 1 B; 3.

Habitat: basalt-derived, well-drained soils and scree in

full sun; altitude 2 150-3 100 m.

Distribution: Drakensberg escarpment, from Mont-aux-

Sources to Ben Macdhui. Figure 4.

FIGURE 4. — Known distribution of Cliffortia dracomontana, ■; and

Cliffortia strobilifera , ▲, in southern Africa.

Etymology: named after the Drakensberg Mountain Range

in which it is endemic.

Hilliard & Burtt (1987) cite two specimens for

C. browniana, their number 13561 (incorrectly cited as

13651) is C. dracomontana, whereas Guillarmod et al.

218 is a true C. browniana. The two species are easily

distinguished — C. dracomontana has glaucous leaves, ±

3-6 mm long, often obovate with rounded apices and

smooth stipules, whereas C. browniana has green leaves,

often with two pale stripes on either side of the midrib

beneath, usually less than 4 mm long, always elliptic,

with an acute apex and ciliate stipules. Furthermore, the

achenes of C. dracomontana are large, 3. 0-4. 5 mm long,

and have glabrous bracteoles, as opposed to C. browni-

ana., which has achenes 2. 0-2. 5 mm long and bracteoles

with ciliate margins. The juvenile leaves of C. draco-

montana show no evidence of any toothing in contrast to

C. browniana , which are toothed although still narrowly

elliptic.

Preliminary molecular evidence suggests that C. draco-

montana is more closely related to C. spathulata than

C. browniana. C. spathulata shares with C. dracomon-

tana the glaucous leaves, smooth stipules and larger

achenes.

3. C. spathulata Weim. in Botaniska Notiser 90:

1 80, t. 5 ( 1948); J.H.Ross: 1 83 ( 1 972). Type: Natal [Kwa-

Zulu-Natal] 2929 (Underberg): Amawahqua [Mahwaqa]

Mtn, (-DC), Medley Wood 4578 (K, holo.!).

C. browniana auct. non Burtt Davy: Weim.: 56, fig.

13E-H (1934) pro parte.

Erect, medium shrub, up to 1 m high, resprouting after

fire from a single crown; densely divaricately branched,

forming brachyblasts; young stems 0.7-1. 2 mm wide,

reddish tinged, hairy; stem hairs upwardly adpressed,

0. 1-0.6 mm long. Leaves trifoliate, chartaceous, 0.2-0. 3

mm thick, midrib not prominent above lamina, curved

upwards and towards stem, glaucous, glabrous above

and beneath; sheath 0.9-1. 5 mm long, abaxially

glabrous, adaxially markedly hairy; stipule 0.6-1. 1 mm

Bothalia 34,1 (2004)

5

FIGURE 5. — Known distribution of Cliffortia spathulata in southern FIGURE 6. — Known distribution of Cliffortia linearifolia in southern

Africa. Africa.

long, free, margin smooth; petiole absent; leaflets broad-

ly obovate, 2.4-6. 1 x 1. 2-3.4 mm, base contracted

abruptly to form a pseudopetiolule. apex acute to round-

ed, margins flat, markedly toothed or lobed with 3-7

straight teeth, 0.05-0.25 mm long. Flowers solitary in

axil of undifferentiated leaves; bracteoles glabrous, mar-

gins serrate to shortly ciliate; sepals 4, glabrous. Male

flowers : bracteoles 1.6-2. 6 mm long; pedicel 0.6-0. 9

mm long; sepals broadly ovate, 2. 6^1. 2 x 1.3-2. 3 mm,

acute to acuminate at apex; stamens 5 or 6; filaments

2.8—4-. 1 mm long, red; anthers brownish red. Female

flowers : bracteoles 2. 2-2. 8 mm long, longer than imma-

ture receptacle; sepals ovate, 1.9-2. 4 x 0. 7-1.0 mm,

recurved; carpel 1 ; stigma 2. 4-3. 5 mm long, white to red,

feathery; immature receptacle 1 .7-2.2 x 0.6-0. 9 mm,

glabrous, smooth. Achene 3. 2-3. 5 x 1. 0-1.1 mm, medi-

um brown, glabrous, faintly ribbed and slightly tubercu-

late to rugose. Flowering time : November to December.

Habitat : Clarens Formation sandstone or basalt-derived

soils in full sun on well-drained soils; altitude 1 800-

2 750 m.

Distribution : northern Drakensberg between Witsies-

hoek and Monk's Cowl (but see note below). Figure 5.

Etymology, spathulata means spoon- or spatula-shaped,

referring to the leaflets that have a broad apex, then taper

down into a narrow stalk.

The type locality of C. spathulata is recorded as

Mahwaqa Mtn above Bulwer by Medley Wood. How-

ever, this mountain is relatively far from the currently

known distribution of the species on the Drakensberg

escarpment to the north. Furthermore, C. spathulata has

not been recollected from that mountain and was not

reported in a recent survey (Meter et al. 2002). It is there-

fore probable that the type locality has been erroneously

recorded.

4. C. linearifolia Eckl. & Zeyh.

Hilliard & Burtt (1987) note that this species has two

forms in the southern Drakensberg, one forming tall vir-

gate shrubs with unifoliate foliage and the other shorter

and densely branched with trifoliate leaves. The unifoli-

ate form is found in wetter areas, such as along water-

courses, whereas the trifoliate form is found on rocky

outcrops and drier slopes. However, both forms have

similar distribution patterns (Figure 6) and it is not pos-

sible to determine from morphology alone whether they

constitute distinct varieties or just ecotypes. Molecular

work and/or transplantation experiments are needed to

examine these two entities to establish the degree of phe-

notypic plasticity within the species.

5. C. nitidula {Engl.) R.E.Fr. & T.C.E.Fr. subsp. pilosa

Weim. and C. filicauloides Weim.

One of the most marked occurrences of dimorphism

between juvenile and mature leaves is found in C. niti-

dula (Engl.) R.E.Fr. & T.C.E.Fr. subsp. pilosa Weim.

(Figure 7). Whereas the mature foliage has needle-shaped

leaflets typical of many species of Cliffortia , the leaves

of seedlings and resprouting growth after fire are trifoli-

ate with a petiole and broadly toothed leaflets. It has

therefore been frequently confused with the northern

FIGURE 7. — Known distribution of Cliffortia nitidula subsp. pilosa ,

■; and Cliffortia odorata. A, in southern Africa.

6

Bothalia 34,1 (2004)

FIGURE 8. — Known distribution of Clijfortia fiUcauloides, ■: and

Clijfortia ramosissima, A. in southern Africa.

Drakensberg endemic C. fiUcauloides (Figure 8), a

species whose closest relatives are in fact found in the

CFR (Whitehouse 2003). In the field, confusion is un-

likely as the habit of C. nitidula subsp. pilosa is always

erect, whereas C. fiUcauloides forms arching stems that

sprawl across boulders and down slopes, and the juvenile

nature of the shoots is usually evident. Furthermore,

shoots with juvenile leaves appear to be always sterile,

thus flowering material will belong to C. fiUcauloides.

Sterile herbarium material can best be distinguished by

examining the lateral leaflets, which are toothed and simi-

lar in size and shape to the middle leaflet in C. nitidula

subsp. pilosa , but generally untoothed and narrower than

the middle leaflet in C. fiUcauloides.

Subdivision of C. paucistaminea Weim. and typification

of C. galpinii N.E.Br.

C. paucistaminea Weim. is a widespread species from

the Outeniqua Mountains to the KwaZulu-Natal Drakens-

berg. It is easily recognizable over most of its range by

its four-sepalled flowers and needle-shaped leaves that

occur on closely overlapping brachyblasts. However,

two distinct forms can be recognized. Between George

and Port Elizabeth, the leaves are strongly curved up-

wards and towards the stem, accentuating the overlap-

ping nature of the brachyblasts. In addition, the leaflets

are quite fine, 0.3-0. 5 mm wide, and the young stems as

a whole have a feathery appearance, resembling the

Western Cape species C. exilifolia Weim. (which has

only three sepals). To the north and east of Port Elizabeth

the leaves are straighter or sometimes recurved away

from the stem, when the brachyblasts then appear star-

like, and the leaflets are generally broader.

Unlike C. linearifolia Eckl. & Zeyh. above, the two

forms also have a geographical separation and therefore

attributing them a taxonomic rank is appropriate.

However, the variation could be the result of clinal dif-

ferentiation associated with climatic factors: the southern

populations are subject to year-round rainfall, whereas

those further north have an increasingly dominant sum-

mer rainfall pattern. Morphometric studies along with

population-level molecular work are needed across the

range of the species to determine if there is continuous

gene flow between the two varieties or if they would be

better regarded as distinct species. In this case, particular

attention needs to be focused on the border between the

two varieties in the mountains around Port Elizabeth.

Weimarck identified that the type collection of the

name C. galpinii N.E.Br. included two different ele-

ments: C. paucistaminea and C. ramosissima Schltr. He

therefore regarded the name as a nomen confusion.

However, this is an unsatisfactory solution as the name

has precedence over C. paucistaminea but not over

C. ramosissima. To prevent the name C. galpinii being

accidentally resurrected, it is important to typify the

name with the C. ramosissima element and thereby rele-

gate it to synonymy. The Kew collections of Galpin 1607

are ambiguous with regard to their locality, stating

‘Hangklip and Andriesberg Mountains’ for both speci-

mens. However, the Bolus and Grahamstown specimens

state only Andriesberg for the C. ramosissima element,

whereas the BOL and PRE specimens both state

Hangklip Mtn for the C. paucistaminea element. It

would therefore seem logical to deduce that Andriesberg

should now be attributed as the type locality.

6. C. paucistaminea Weim. in Botaniska Notiser

1933: 151, fig. 4c, d (1933); Weim.: 64, fig. 16A-E

(1934); J.H.Ross: 183 (1972); Hilliard & B.L.Burtt: 162,

t. 6C (1987); Pooley: 150 (1998); Fellingham: 615

(2000). Type: Basutoland [Lesotho], Cooper 690 (K,

holo.!; B, BOL!, W, Z).

C. galpinii auct. non N.E.Br. sensu stricto: N.E.Br.: 122 (1901) pro

parte.

C. juniperina auct. non L.f.: Jacot-Guillarmod: 186 (1971).

Erect, medium to tall shrub, up to 2 m high, resprout-

ing after fire and sometimes spreading clonally; densely

divaricately branched, forming brachyblasts, which are

closely overlapping; young stems 0.8-1. 7 mm wide,

hairy; stem hairs upwardly adpressed, 0.2-0. 6 mm long.

Leaves trifoliate, chartaceous, 0. 1-0.4 mm thick, midrib

keeled and prominent, green with two paler stripes on

either side of midrib beneath, glabrous above and be-

neath; sheath 0.6- 1.5 mm long, abaxially glabrous, adax-

ially glabrous to markedly hairy; stipules 1.5-2. 3 mm

long, free, margin smooth; petiole absent; leaflets linear

to needle-shaped, 5.1-10.2 x 0.3-0. 8 mm, apex sharply

acuminate, 0.4-0. 8 mm long, margins flat, minutely ser-

rulate to scabrid. Flowers solitary in axil of undifferenti-

ated leaves; bracteoles hairy, margins serrate to shortly

ciliate; sepals 4, or very rarely 3, glabrous. Male flowers:

bracteoles 0.9- 1.8 mm long; pedicel 0.3-0. 8 mm long;

sepals broadly ovate, 2. 2-3. 6 x 0.9-1. 5 mm, acute to

acuminate at apex; stamens 4(— 6); filaments 1.4-3. 7 mm

long, reddish; anthers yellow to brownish red. Female

flowers: bracteoles 1 .6-2.6 mm long, longer than imma-

ture receptacle; sepals narrowly ovate, 1.1-1. 7 x 0.3-0. 5

mm, erect to recurved; carpel 1 ; stigma 2. 5-3. 7 mm long,

greenish white, feathery, hidden at base of leaves; imma-

ture receptacle 1.0- 1.8 x 0.6-0. 9 mm, glabrous, clearly

ribbed. Achene 2. 1-3.3 x 0.9-1. 2 mm, pale yellowish

brown, glabrous; ribs 8-16, rounded to acute, 0. 1-0.2

mm wide.

Bothalia 34,1 (2004)

7

FIGURE 9. — Example specimen of Clijfortia paucistaminea var.

paucistaminea, Transkei, Engcobo District, hills near Engcobo,

Esterhuysen 29149 (BOL).

Etymology, paucistaminea means few stamens, refer-

ring to the male flowers that only have 4 stamens,

although a number of other species also have that few.

6a. var. paucistaminea

Leaflets 0.4-0. 8 mm wide, held straight or curved

downwards and away from the stem. Flowering time:

predominantly September to February. Figure 9.

Habitat : found in humus-rich soil over sandstone rocks

in full sun; altitude 0-2 350 m.

Distribution : widespread from the Suurberg Mountains

along the Drakensberg escarpment as far as Giant’s

Castle, with outlying populations along the Wild Coast

and KwaZulu-Natal lowlands as far north as Nkandla.

Figure 10.

6b. var. australis C.M.Whitehouse , var. nov., a var.

paucistaminea ramulis plumosis differt.

TYPE. — Eastern Cape, 3424 (Humansdorp): Witte

Els Bosch [Witelsbos], flats, (-AA), Fourcade 2114

(BOL, holo.!; K!, LD, NBG!, PRE).

Leaflets 0.3-0. 5 mm wide, curved upwards and to-

wards stem, giving the branches a feathery appearance.

FIGURE 10. — Known distribution of Clijfortia paucistaminea var.

paucistaminea. HI: and Clijfortia paucistaminea var. australis,

A, in southern Africa.

Flowering time: predominantly November to April.

Figure 11.

Habitat: humus-rich Table Mountain Sandstone derived

soils, in full sun and on well-drained soils, but areas

FIGURE 11. — Holotype of Clijfortia paucistaminea var. australis,

Witte Els Bosch, flats, Fourcade 2114 (BOL).

Bothalia 34,1 (2004)

where it grows are subject to frequent rain and cloud;

altitude 50-1 550 m.

Distribution-, southern Cape mountains and lowlands be-

tween George and Uitenhage. Figure 10.

Etymology: australis means southern, referring to the

distribution of the variety in the southern Cape.

7. C. ramosissima Schltr. in Botanische Jahr-

biicher 24: 444 (1897); Weim.: 73, fig. 18L-R (1934);

Weim.: 419 (1946). Type: Western Cape, 3419 (Cale-

don): Elim, (-DB), Schlechter 7633 (B, holo.; BOL!,

COI, G-DEL, GRA, K!, HBG, L, NH, P, PRE, SAM, W,

WAG, Z).

C. galpinii N.E.Br.: 122 (1901) pro parte. Lectotype, here designat-

ed: Eastern Cape, Queenstown District, Andriesberg, Galpin 1607:

C. ramosissima element (K, lecto. ! ; BOL!. GRA!).

8. C. repens Schltr and C. erectisepala Weim.

Weimarck described C. erectisepala from a single col-

lection from Paarl Mountain. He subsequently comment-

ed on collections from Franschhoek (Weimarck 1940)

and Swellendam (Weimarck 1946). C. repens Schltr. he

applied to unifoliate, needle-leaved collections from Uiten-

hage to Pietersburg. However, morphologically the collec-

tions from Uitenhage to Grahamstown are closer to C.

erectisepala , having shorter, narrower leaves, upright

erect habit, and non-spreading sepals. More recent collec-

tions have extended the range of C. erectisepala from

Gifberg in the north to Rooiberg in the Little Karoo. The

species in general is very inconspicuous and the current

study has revealed that it is in fact much more common

and widespread with many new localities being recorded.

Consequently, the disjunction between Western Cape

populations and the narrow-leaved Eastern Cape popula-

tions of C. repens is not as marked as originally thought.

Therefore, these populations are now included in C. erec-

tisepala, whereas the range of true C. repens is reduced

and extends only as far south as Baziya Mtn near Umtata.

This change is significant biogeographically, as C. repens

is no longer part of the Cape Flora and is better regarded

FIGURE 12.- Known distribution of Cliffortia erectisepala, ■; and

Cliffortia repens, A, in southern Africa.

as a broad Drakensberg endemic, whereas C. erectisepa-

la can be counted as a Cape Ubiquist (sensu Weimarck

1941 ) (see Figure 12).

A single collection is incongruous with this re-organi-

zation. The sheets are labelled as collected by Thomcroft

(in Herb. Rogers 19187 ) from Barberton and are typical

of C. erectisepala , although only male flowers are pre-

sent. However, no other collections have been made in

this well-collected area that resemble C. erectisepala and

it is therefore presumed that the locality is erroneous.

SPECIMENS EXAMINED

Abbott 177 (6a) PRE. Acocks 10217 (1) PRE; 11462 (1) NBG, PRE;

12037 (6a) PRE; 13838. 18690. 20159. 21884 (6a) K, PRE. Acocks &

Hafstrom 571 (4) PRE.

Balkwill, Manning c£ Meyer 790 (5) K, NU; 792 (5) NU; 793 (3) NU.

Balkwill, Manning et al. 1030 (6a) NU. Baur 494 (8) K. Bayer 815 (8)

NU. Bayliss 493, 1347 (6a) PRE. Bean. Viviers & Cloete 2274 (6a)

BOL. Bews 1413 (8) NU. Bolus 250 (6a) BOL. Bond 1210 (6b) BOL,

PRE. Britten 4663 (6a) GRA, PRE; 6408 (6a) GRA. Bruce 495 (8) K,

PRE. Buitendag 827 (8) K, NBG, PRE. H. & J. Burrows 4720 ( 8)

GRA. Burn Davy 1560 (8) BOL, PRE.

Comins 581 (4) NU. Compton 7313 (8) NBG; 10458 (6b) NBG; 19176

(6a) NBG; 19746. 19789. 22377 (8) NBG; 23500 (6b) NBG; 24742 (8)

NBG; 26285. 27606 (8) K. NBG, PRE; 28510. 30531 (8) NBG, PRE.

Cooper 690 (6a) BOL, K. Crewe 24 (5) NU. Curator Pretoria Bot.

Garden 21565 (8) PRE.

Davidson & Mogg 33405 (8) K, PRE. Dealt 2303 (8) PRE. Devenish

1976 (5) NU. Dickin 4 (6b) BOL. Dieterlen 1103. 1203 (6a) NBG,

PRE. Dohne Research Institute 4 (6a) GRA. Dold 815 (6a) GRA.

Downing 104 (4) NU. Drege 5381 (4) K. Dyer & Collett 4700 (8) PRE.

Edwards 244 (4) NU; 596 (2) NU, PRE; 928 (4) NU; 976 (4) NU, PRE;

2013 (3) NU; 2233 (8) K, NU, PRE; 2235, 2248 (5) NU; 2456 (1) K,

NU, PRE; 4054 (8) K, PRE. Esterhuysen 4569 (6b) BOL; 6810 (6b)

BOL, PRE; 7930 (8) BOL; 8843 (3) BOL; 8846 (4) BOL; 10180 (3)

BOL, K; 10181 (5) BOL, K; 10183 (2) BOL; 10184 (8) BOL, K, NBG,

NU; 10816 (6b) BOL; 10854 (6b) BOL, K; 12867 (8) BOL, PRE;

12871 (3) BOL; 12886 (2) BOL; 13222. 13259 (6a) BOL; 13578 (6b)

BOL; 14527 (8) BOL, NBG, PRE; 14528 (8) BOL; 15482 (2) BOL,

NBG, PRE; 15484 juvenile (5) BOL; 15485 (3) BOL; 15489 (8) BOL,

NBG; 15635 (3) BOL; 16272 (6b) BOL; 16770 (6b) BOL, NBG, PRE;

1 7342 juvenile (5) BOL, PRE; 18661 (2) BOL, NBG; 18662 (3) BOL,

PRE; 18684 (5) BOL; 21447(1) BOL, K, PRE; 21687 (2) BOL; 27113

(6b) BOL, PRE; 27374 (6b) BOL; 27541 (6b) BOL, PRE; 27846 (8)

BOL; 29149 (6a) BOL, K. Evans 663 (8) K, PRE.

Fellingham 1631 (6a) NBG, PRE; 1632, 1671, 1672 (6a) NBG.

Feltham 155 (5) NU. Forrester & Gooyer 204 ( 1 ) PRE. Fourcade 2114

(6b) BOL, K, NBG, PRE. Galpin s.n. (8) BOL; 1607 (6a) BOL, K,

PRE; 2255, 8307 (6a) PRE; 9439 (4) K, PRE; 13072 (8) K, PRE.

Germishuizen 5778 (8) PRE. Gerrard 1568 (8) K. Gerstner s.n. (5)

PRE; 3520 (4) PRE; 3934 (5) PRE. Gibbs Russell 3493A (6a) BOL,

GRA, PRE, UFH. Giffen 342 (6a) GRA, PRE. UFH. Gilbert 7593 (2)

PRE. Gillies 60 (4) NU. Glen 380 (6a) NBG; 2444 (4) PRE. Gordon-

Gray 887 (4) NU. Goulimis s.n. (6a) BOL. Grice s.n. (4) NU, (6a) NU.

Grobbelaar 2618 (1) PRE.

Hepburn 125 (6a) GRA. Hilger 7 (6a) PRE. Hilliard 1778A (4) NU;

2648. 3031 (6a) NU; 4708. 8165 (4) NU; 8228 juvenile (5) NU.

Hilliard & Burn 6532 (6a) K, NU, PRE; 9314 (4) NU; 12481, 12581,

13284 (6a) NU; 13285 (6a) K, NU; 13331 (8) NU; 13341 (5) NU;

13342 (1) NU; 13416 (4) NU, PRE; 13428 (5) NU; 13442 (4) K, NU;

13465 (8) NU; 13508, 13513 (4) NU; 13561 (2) K, NU, PRE; 13563

(5) NU; 13592 (4) K, NU; 13653 (4) K, PRE; 13689 (5) NU; 13865

(6a) K, NU; 13866 (6a) K, NU, PRE; 13894 (6a) NU; 14011 (4) NU;

14035 (6a) NU; 14071 (8) K, PRE; 14292 (8) K, NU, PRE; 14323 (1)

NU, PRE; 14383 (8) K, NU, PRE; 14459 (8) K, NU; 14502, 14503 (6a)

NU; 14506 (6a) K, NU; 14510 (6a) K; 14655 (6a) K, NU, PRE; 14753

(6a) K, NU; 14887 ( 4) NU; 14888 (5) NU, PRE; 14925 (6a) NU, PRE;

14965 (4) NU; 15021 (2) K, NU; 15105 (4) K, NU, PRE; 15172 (4)

NU, PRE; 75266(1) K, NU, PRE; 15378 juvenile (5) K, NU; 15397(4)

Bothalia 34. 1 (2004)

9

K, NU, PRE; 15403, 15430 (8) K, NU, PRE; 15447 (5) K, NU. PRE;

15456 (3) K. NU; 15505 (5) NU; 15510 (5) NBG, NU; 15542 (5) NBG;

15591 (6a) NU; 15592 (4) NU; 15634 (8) K, NU, PRE; 15643 (6a) K,

NU, PRE; 15644 (6a) NU, PRE; 15667, 15672 (4) NU; 15673 (4) K,

NU; 15789 (5) K, NU, PRE; 15790 (4) K, NU. PRE; 15815 (4) NU;

15816 (5) NU. PRE; 16273 (8) K. NU, PRE; 16274 (5) NU; 16286 (4)

K. NU. PRE; 16304 (3) K, NU; 16332 (6a) K, NU, PRE; 16333 (6a) K,

NU; 16422 (2) K, NU. PRE; 16489 juvenile (5) NU; 16605 (2) K, NU;

16762 (8) K, PRE; 16870, 16871 (4) K, NU, PRE; 16986 (8) NU;

16987 (4) NU; 17012 (4) K, NU, PRE; 17025 (8) K, NU, PRE; 17030

(4) K, NU. PRE; 17077 (6a) NU, PRE; 17095 (6a) K. NU, PRE; 17110

(4) NU; 17169 (8) K, NU, PRE; 17225 (1) K. NU, PRE; 17667 juve-

nile (5) K, NU, PRE; 17668 (8) K, NU, PRE; 17677, 17681 (1) K, NU.

PRE; 17697(3) K, NU, PRE; 17724 (8) K. NU. PRE; 17761, 17805 (4)

NU; 17955 (6a) NU, PRE; 17956 (4) K, NU; 17994 (1) NU; 18058,

18180 (5) NU; 18284 (5) K. NU, PRE; 18315 juvenile (5) NU; 18329

(6a) K, PRE; 18330 juvenile (5) K. NU. PRE; 18358 (8) K, NU, PRE;

18536 juvenile (5) K. NU. PRE; 18681 juvenile (5) NU; 18837 (6a) K.

NU. PRE. Hilliard, Bunt & Manning 15955 (6a) NU; 15965 (4) NU,

PRE; 16026 (1) K. PRE; 16061, 16062 (1) NU: 16069 (6a) NU; 17261

juvenile (5) NU; 17296 (4) NU. Huntley 409 (5) NU. Hutchinson 4534

(8) K; 4543 (5) K; 4550 (3) K; 4580 (2) K. Hutchinson & Gillen 4314

(8) K. Hutchinson, Forbes & Verdoom 72 (3) PRE.

Jacobsen 22 72 ( 8) PRE. Jacot-Guillannod 3958 (6a) PRE; 7959 (6a)

GRA. Jacot-Guillannod, Getliffe & Mzamane 218(1) GRA, K, PRE.

Johnstone 209 (6a) NU. Jordaan 2910 (1 ) PRE.

Keet STE13384 (6b) NBG, (8) NBG. Kemp 1223 (8) PRE. Kensit &

Copom in NBG. 150/14 (6b) BOL. Kerfoot, Forrester & Gooyer 31 (8)

PRE. Killick 1109 (8) K, PRE; 1932 (3) PRE. Killick & Vahrmeyer

3711 (6a) K, PRE. Kluge 2313 (8) NBG, PRE. Krynauw 310 (8) PRE;

782 (1) PRE. Kuntze s.n. (5) K.

Lambinon & Reekmans 82/460 (8) PRE. Leighton 2697 (6a) BOL;

2998 (6a) BOL, PRE; 3 127 (6a) BOL: 3267 (8) BOL. Levyns 6905 (6a)

BOL; 8240 (4) BOL; 8267, 8273 (8) BOL; 8302 (5) BOL; 8303 juve-

nile (5) BOL; 8308, 9408 (8) BOL; 9470juvenile (5) BOL; 9842, 9856

(6a) BOL; 10482 (6b) BOL.

MacDevette 1574 (6a) PRE. Manin 9067 juvenile (5) GRA. McClean

& Bayer 5 (8) BOL. K, NU, PRE; 103 (5) NU. McDonald 88 (6b) PRE;

240 (8) K, NU, PRE. Medley Wood s.n. (8 ) BOL; 4449 (8) K; 4578 (3)

K; 5987 (8) K; 7903 (6a) BOL. K. Meeuse 9981 (1) PRE. Meyer 58 (8)

PRE. Morris 441 (5) NU. Mudd s.n. (8) BOL, K.

Nel 221 (8) NBG, PRE. Neser NF1969 (4) PRE. Ngwenya 953 (6a) PRE.

Onderstall 170 (1 ) PRE. Ons s.n. (8) NBG.

Palmer 1412 (6a) GRA. Phillips s.n. (6a) NU; s.n. (5) NU; s.n. (8) NU.

Pole Evans 967, 968 (8) K. PRE. Prior 60 (8) K.

Rehmann 6927 (5) K. J. & B. Rennie 168A, 168B (6a) GRA. M. Rennie

120 (4) NU. Roberts 2496 juvenile (5) PRE; 2676 juvenile (5) PRE.

Rogers 21565 (8) K; 21591 (8) BOL, K; 22985 (1) BOL, K. Ross 195

(8) NU; 1755 (8) K, NU. PRE. Ruch 2039 (4) K. Rudatis 2063 (6a)

NBG.

Scharf 13 17 (6b) PRE. Schelpe 440 (4) NU; 673 (5) NU; 1092 (4) NU;

1381 (2) NU. PRE; 7195 (8) BOL, PRE. Schlechter 3318 (4) BOL, K;

5992 (6b) K, PRE; 6494 (8) BOL, PRE. Schmitz 9223 (6a) NU. PRE.

Sidey 2015 (2) PRE. Sim 19430 (6a) NU. PRE; 19431 (4) PRE: 19432

(4) NU, PRE. Skead 23 (4) NU. Smit 631 juvenile (5) PRE. Stayner in

Herb. Bolus 19865 (6a) BOL. Story 3116 (6a) PRE; 3190 (6a) GRA,

PRE; 4003 (8) GRA, K, PRE; 5448 (8) K, PRE. Strey 6933 (6a) NU,

PRE; 7692 (4) K. NU; 11272 (5) NU. Sutherland s.n. (6a) K.

Taylor 4451 (6b) K. NBG. PRE. H. Thode 1598 (8) K. J. Thode 3286,

4330 (8) NBG; 4595 (5) NBG; 6356 (8) NBG. Van Daalen 121 (6b)

NBG, PRE. Van der Schijff 1480 (1) K; 4480 (1) PRE; 5588 (1) K,

PRE. Van der Schut 4475 (8) K. Van der Walt 367 (6a) PRE. Victor 767

(6a) GRA; 1238 (6a) PRE. Von Gadow 472 (6a) GRA.

Wager 181 (8) K. PRE. Werdermann & Oberdieck 2181 (8) K, PRE. C.

& A. Whitehouse 56 (6a) BOL; 286 (2) BOL; 287 (3) BOL; 289(1)

BOL; 290 (8) BOL; 291, 292 (4) BOL; 296 (1) BOL; 297 (6a) BOL;

298 (6b) BOL; 299 (8) BOL; 323 (6a) BOL; 325 (2) BOL; 326 (1)

BOL. Wilkinson s.n. (2) GRA. Williams 342 (6a) PRE. Wilms 595 (8)

K. Wirminghaus 982 (5) NU; 1054 (4) NU. Wylie in Medley Wood

10029 (8) K.

ACKNOWLEDGEMENTS

This study was funded by a Leverhulme Trust, Study

Abroad Studentship, with assistance from a J.W. Jagger

International Scholarship, and was initiated by Prof. H.P.

Linder. I wish to thank the KwaZulu-Natal Parks Board

for granting me a collecting permit and the various sec-

tional managers of the uKhahlamba-Drakensberg Park

for allowing me to collect specimens. I also thank the

curators of BOL, GRA, K, NBG, NU and PRE for per-

mission to examine their collections or in sending loans.

REFERENCES

BROWN, N.E. 1901. Diagnoses Africanae 13. Kew Bulletin 1901:

119-138.

BURTT DAVY, J. 1932. A manual of the flowering plants and ferns of

the Transvaal with Swaziland, South Africa 2. Longmans.

Green. London.

EDWARDS. D. 1967. A plant ecological survey of the Tugela River

Basin. Memoirs of the Botanical Survey of South Africa No. 36.

FELLINGHAM, A.C. 1993a. Observations on Cliffortia micrantha.

Bothalia 23: 65. 66.

FELLINGHAM, A.C. 1993b. Cliffortia fasciculata, a superfluous

name for C. amplexistipula. Bothalia 23: 67, 68.

FELLINGHAM, A.C. 1994. Cliffortia longifolia , a ‘good’ species or

should it be a variety under C. strobiliferal Bothalia 24: 31-34.

FELLINGHAM, A.C. 1995. A new species of Cliffortia from the

Swartberg. Bothalia 25: 104—107.

FELLINGHAM. A.C. 2000. Cliffortia. In P. Goldblatt & J.C. Manning,

Cape plants. A conspectus of the Cape flora of South Africa.

Strelitzia 9: 608-616. National Botanical Institute, Cape Town

and Missouri Botanical Garden.

FELLINGHAM, A.C. 2003. A new serotinous species of Cliffortia L.

(Rosaceae) from Northern Cape. South Africa and section

Arboreae emended. Bothalia 33: 41-47.

GOLDBLATT. P. & MANNING. J.C. 2000. Cape plants. A conspectus

of the Cape flora of South Africa. Strelitzia 9. National Botanic

Institute. Cape Town and Missouri Botanical Garden.

GOLDING, J.S. 2002. Southern African plant Red Data Lists. Southern

African Botanical Diversity Network Report No. 14.

HILLIARD, O.M. & BURTT, B.L. 1987. The botany of the southern

Natal Drakensberg. Annals of Kirstenbosch Botanic Gardens 15.

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

KILLICK. D.J.B. 1963. An account of the plant ecology of the

Cathedral Peak area of the Natal Drakensberg. Memoirs of the

Botanical Survey of South Africa No. 34.

LINDER, H.P. 2001. On areas of endemism, with an example from the

African Restionaceae. Systematic Biology 50: 892-912.

METER. E.B., EDWARDS, T.J., RENNIE, M.A. & GRANGER, J.E.

2002. A checklist of the plants of Mahwaqa Mountain,

KwaZulu-Natal. Bothalia 32: 101-115.

OLIVER, E.G.H. & FELLINGHAM, A.C. 1991. A new species of

Cliffortia from the south-western Cape. Bothalia 21: 60-62.

OLIVER, E.G.H. & FELLINGHAM, A.C. 1994. A new serotinous

species of Cliffortia (Rosaceae) from the southwestern Cape

with notes on Cliffortia arborea. Bothalia 24: 153-162.

POOLEY, E. 1998. Afield guide to wild flowers: KwaZulu-Natal and

the eastern region. Natal Flora Publications Trust, Durban.

ROSS, J.H. 1972. Flora of Natal. Memoirs of the Botanical Survey of

South Africa No. 39.

SCHLECHTER. R. 1897. Plantae Schlechterianae novae vel minus

cognitae describuntur 1 . Botanische Jahrbiicher 24: 434-459.

WEIMARCK, H. 1933. New species in the genus Cliffortia. Botaniska

Notiser 143-164.

WEIMARCK, H. 1934. Monograph of the genus Cliffortia. Hakan

Ohlsson, Lund.

WEIMARCK, H. 1940. Some notes on the genus Cliffortia. Botaniska

Notiser 51: 399-102.

WEIMARCK, H. 1941. Phytogeographical groups, centres and inter-

vals within the Cape Flora. Lunds Universitet Arsskrift 37:

3-143.

WEIMARCK, H. 1946. Further notes on the genus Cliffortia. Botaniska

Notiser 80: 407-420.

WEIMARCK, H. 1948. The genus Cliffortia, a taxonomical survey.

Botaniska Notiser 90: 167-203.

10

Bothalia 34,1 (2004)

WEIMARCK, H. 1953. Two new Cliffortia species from the Cape

Province. Botaniska Notiser 106: 78-80.

WEIMARCK, H. 1959. Four new Cliffortia species. Botaniska Notiser

112: 73-79.

WHITEHOUSE, C.M. 2003. Systematics of the genus Cliffortia L.

(Rosaceae). Ph.D. thesis. University of Cape Town.

Bothalia 34,1: 11-15 (2004)

Two new species of Erica (Ericaceae); one from Western Cape and

one from KwaZulu-Natal, South Africa

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

Keywords: Erica L., KwaZulu-Natal, new species. South Africa, taxonomy. Western Cape

ABSTRACT

Two new species of Erica L. from South Africa are described. E. jananthus E.G.H.Oliv. & I.M.Oliv. is confined to a

single peak in the eastern Groot Swartberg Range in Western Cape and usually forms a small, gnarled, woody, shrublet

growing in rock crevices with sticky white flowers and black subexserted anthers that have obtrullate decurrent appendages.

E. psittacina E.G.H.Oliv. & I.M.Oliv. is from KwaZulu-Natal. It forms large woody shrubs with numerous bright pink

flowers and occurs as a single population on a mountain near Creighton. Both descriptions are accompanied by line draw-

ings and distribution maps.

1. Erica jananthus E.G.H.Oliv. & I.M.Oliv., sp.

nov., fruticulo plerumque parvo lignoso plectile, foliis 3-

natis, foliis bractea bracteolis sepalibusque glandibus

marginalibus sessilibus fulvis ad brunneis, corolla urceo-

lata viscida lobis effusis ad recurvatis, antheris 8

subexsertis calcaribus decurrentibus anguste ad equila-

teriliter obtrullatis et irregulatiter denticulatis, ovario vil-

loso 4-loculare ovulis 6-8, seminibus parietibus anticli-

nalibus percrassis dignoscenda. Figura 1 .

TYPE. — Western Cape, 3322 (Oudtshoom): Great

Swartberg, eastern end, Snyberg. peak just WNW of bea-

con, 1 570 m, (-BD), 16 November 2001, Oliver 11986

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

Shrublet compact to loose, 20 x 30-100 x 150 mm.

usually woody, single-stemmed. Branches : numerous

main branches ± 10 mm long, leafy with occasional leafy

side branchlets 2-5 mm long; stems puberulous to

sparsely strigose with some glands admixed. Leaves 3-

nate, erect-spreading to patent, elliptic to narrowly lance-

olate, 1.5-1. 7 x 0.5-0. 7 mm, rounded to flattened adaxi-

ally and rounded abaxially, sparsely puberulous on both

surfaces to almost glabrous adaxially, sulcus narrow and

open at base, margins rounded with yellowish to pale

brown sessile glands and one apical gland, often com-

pletely glabrous when older; petiole adpressed, 0.4— 0.5

mm long, edged with short hairs and subsessile to sessile

glands. Inflorescence : 1-3 flowers in a single whorl at

ends of leafy, short, main branches and occasional later-

al branchlets. the latter sometimes aggregated at ends of

main branches; pedicel ± 2.4 mm long, sparsely and

shortly strigulose with longer and stouter gland-tipped

hairs or sessile glands admixed; bract partially recaules-

cent in lower quarter of pedicel, elliptic, ± 0.9 x 0.5 mm,

sparsely puberulous ciliate with yellowish to pale brown

sessile glands, white sometimes tinged green; bracteoles

2 in mid position, slightly obovate, otherwise like bract.

Calyx 4-partite; lobes adpressed to corolla, elliptic to

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

7735 Claremont, Cape Town,

t obiit 6-07-2003.

MS. received: 2003-08-08.

broadly so, ± 1.5 x 1.0-1. 2 mm, glabrous, edged with

numerous sessile glands, white with narrowly sulcate

green apex. Corolla 4-lobed, globose urceolate, ± 3.0 x

2.5 mm, glabrous, viscid, white; lobes broadly ovate, ± 1

x 1. 5-2.0 mm, spreading to recurved, obtuse or emar-

ginate, minutely and irregularly toothed. Stamens 8, free;

filaments 1. 8-2.0 x 0.2-0. 3 mm, linear, curved,

glabrous; anthers subexserted, bilobed appendiculate,

thecae ± 0.7 x 0.3 mm and oblong in adaxial view, ± 0.8

x 0.4 mm and elliptic in lateral view, golden brown to

red-brown, pore ± V2 length of theca, appendages decur-

rent on filament, ± 0.3 x 0.2 mm, narrowly to equilater-

ally obtrullate, pendulous, irregularly denticulate, pale

brown to cream-coloured; pollen shed in tetrads. Ovary

4-locular, globose to depressed globose, 0. 5-0.9 x 0.9

mm, obtuse to almost flat apically, villous, white to pale

green, with well-developed, dark nectaries around base;

ovules 6-8 per locule, pendulous from apical placenta;

style exserted, narrowly cylindric, slightly curved, ± 3

mm long, white, glabrous; stigma simple-truncate to

slightly capitellate, dark green. Fruit a dehiscent capsule

± 1.9 x 2.6 mm, valves opening full length, septa equal

on valves and columella. Seeds ± ellipsoid, ± 0.7 x 0.35

mm, sometimes curved, with flattened micropilar end;

testa alveolate, cells ± 100 x 16-20 pm, anticlinal walls

unequally undulate and considerably thickened, pericli-

nal walls with scattered pits. Figure 1.

Diagnostic features: small, woody, gnarled shrublet;

leaves 3-nate; leaves, bract, bracteoles and sepals edged

with yellow to brown sessile glands; corolla urceolate,

viscid white with spreading to recurved lobes; anthers 8

subexserted with appendages decurrent on filament, nar-

rowly to equilaterally obtriangular and irregularly den-

ticulate; ovary 4-locular with 6-8 ovules per locule, vil-

lous; seeds with considerably thickened, unequally undu-

late, anticlinal walls.

Erica jananthus is an isolated species with no clear

indications of its relationships. With the 3-nate leaves,

marginal glands on the bract, bracteoles and sepals pro-

ducing a viscid corolla, it would appear to lie within the

group in §Pachysa that has several species frequenting

the mountains in and around the Little Karoo — E. for-

mosa Thunb., E. spectabilis Klotzsch ex Benth., E. trag-

12

Bothalia 34,1 (2004)

FIGURE 1. — Erica jananthus. A, flowering branch, natural size; B, stem; C, leaf, back and lateral views; D, flower; E, stamen, lateral, front and

back views; F, gynoecium; G, ovary, opened laterally to show placentation; H, capsule with one valve removed; I, seed; J, testa cells. All

drawn from the type, Oliver 11986. Scale bars: B-D, F-H, 2 mm; E, I, 1 mm; J, 100 pm. Artist: Inge M. Oliver.

ulifera Salisb., E. glomiflora Salisb., E. zwartbergensis

Bolus, E. andreaei Compton. In those species, however,

the anthers are very different with their long thick hairs

and narrower appendages, although the latter are often de-

current along the upper part of the filament, and villous

ovary. We recently described E. dolfiana E.G.H.Oliv.

(Oliver & Oliver 2001) from the nearby Blesberg. It has

marginal glands on the above-mentioned organs but these

are stalked and dark red.

This new species was brought to our attention by the

botanist, Jan Vlok of Oudtshoorn, when doing a survey

of that part of the Groot Swartberg. The epithet for this

new species recognizes his considerable contribution to

the recording of plant species and distributions in the

mountains surrounding the Little Karoo, jananthus =

Jan’s flower (Latin), there already being E. vlokii

E.G.H.Oliv. from the Meiringspoort, Kammanassie and

Herold areas (Oliver 2000). We have described several

new species of Erica from the Swartberg in the Meirings-

poort area: E dolfiana E.G.H.Oliv., E. ingeana E.G.H.Oliv.,

E. vlokii E.G.H.Oliv., E. lithophila E.G.H.Oliv. & I.M.Oliv.,

E. jugicola E.G.H.Oliv. & I.M.Oliv. and E. umbratica

E.G.H.Oliv. & I.M.Oliv.

The species is as far as we know confined to the

Snyberg peaks in the eastern part of the Groot Swartberg

Range east of Meiringspoort (Figure 2). There it grows

only in rock crevices on south-facing, small, rocky

ridges and outcrops in shade for most of the day. A sin-

gle plant growing in the ground at the base of one ridge

was more shrubby, thinner stemmed and up to 300 mm

tall. The species does not occur further west on the

peaks around Blesberg but there are several peaks to the

east of Snyberg that have never been explored botani-

cally.

Bothalia 34,1 (2004)

13

FIGURE 3. — Erica psittacina. A, flowering branch, natural size; B, stem with leaves removed; C, leaf; D, flower; E, bracteole; F, sepal; G, anther,

lateral, front and back views; H, gynoecium; I, ovary, L/s showing placenta and ovule positions; J. capsule; K, seed; L, testa cells. All drawn

from the type, Oliver & Turner 12167. Scale bars: B-F, H-J, 2 mm; G, K, 1 mm; L, 100 pm. Artist: Inge Oliver.

No pollinators were noted when the type collection

was made. Pollination is presumed to be carried out by

flying insects because of the presence of nectaries and

the reduced size of the stigma. Crawling insects would

encounter problems with the viscid corolla.

Pciratype material

"WESTERN CAPE. — 3322 (Oudtshoom); Great Swartberg Mtns, Sny-

berg, 1 500 m, (-BD), 22-08-1990, Vlok 2377 (NBG); ibid., 11-03-1991,

fruiting, Vlok 2453 (NBG); ibid., 12-11-1991, Vlok 2533 (NBG, PRE).

2. Erica psittacina E.G.H.OIiv. & I.M.Oliv., sp.

nov., frutice magno lignoso ad 2 m raro 3 m alto unicauli,

foliis 4-natis, inflorescentibus parte inferiore racemosa

superiore umbelliformi, corolla subtiliter breviterque

pubescenti clarorosea, indumento in partibus multis pilis

brevissimis cum pilis paucis crassis plumosis apice fur-

cato dignoscenda. Figura 3.

TYPE. — KwaZulu-Natal, 2929 (Underberg): Creighton,

Hlabeni, steep southeast-facing slopes below summit

ridge, 1 662 m, (-DC), 4 February 2003, Oliver & Turner

12167 (NBG, holo.; BM, CPF, K, NH, NU, NY, PRE).

Shrub woody, bushy, erect, 1 .0— 1 .5(— 2.0)[— 3.0] m tall,

single-stemmed. Branches : main branches up to 400 mm

long, sometimes topped with numerous vegetative sec-

ondary branches or just flowering secondary branches;

secondary branches numerous, 100-160 mm long, with 1

or 2 tertiary branches 30-60 mm long; stem with dense,

short, simple hairs and spreading to reflexed, stouter,

longer hairs admixed, these mostly shortly plumose

mainly towards base of hair and sometimes with longer

14

Bothalia 34.1 (2004)

forked tips; internodes 2-5 mm long. Leaves 4-nate,

suberect to spreading but curved upwards, occasionally

reflexed at base of secondary branches, lanceolate, ± 7 x

1 mm, with rounded margins, partially open-backed with

sulcus open at base, with fine, short, simple hairs and a

few, long, stouter, fork-tipped hairs, these sometimes

shortly plumose at base; petiole ± 1 mm long, with

sparse, short, simple hairs; on margins a few shortly

stalked glands. Inflorescence : flowers 4-nate in 2-5

whorls, terminal on secondary branches, rarely tertiary

branches, lower 1 or 2 whorls with intemodes, upper

whorls umbel-like; pedicel 4-6 mm long, with dense,

fine, simple hairs and sparse, long, stout, spreading,

plumose hairs admixed, red; bract leaf-like and on

branch in lower whorls of inflorescence to partially

recaulescent in mid position in upper whorls; bracteoles

2, lanceolate, ± 3 x 0.4 mm, green, base pinkish, placed

± 2/3 way up pedicel, indumentum like bract. Calyx 4-

partite; segments lanceolate, ± 2. 5-3.0 x 0.5 mm, finely

hairy plus long, stout hairs on margins, the upper termi-

nally forked, the lower also shortly plumose towards

base, acute or attenuated into a long, plumose seta; upper

half of segments green, lower pink, sulcus ± V2 length of

segment. Corolla 4-lobed, globose-urceolate, ± 4 x 3.5

mm. finely puberulous, pink; lobes suberect, ± 0.8 x 1 .5

mm, rounded, entire. Stamens 8, manifest; filaments ± 3

mm long, straight; anthers basifixed, bilobed, oblong in

front view, appendiculate, orange-brown; thecae erect

adpressed, oblong, ± 1.0-1. 3 x 0.5 mm in lateral view,

aculeate, pore ± V2 length of theca, appendages ± 0.7 mm

long, ± ovate-elliptic, flattened, irregularly shallowly and

deeply toothed, yellow; pollen in tetrads. Ovaiy 4-locu-

lar, depressed globose-obovoid, emarginate, ± 1 .2 x 1.5

mm, covered with simple, short hairs, with large nec-

taries around base; ovules ± 50 per locule, spreading

from bulbous placenta on upper 2/3 of columella; style ±

4 mm long, exserted; stigma capitate. Fruit a dehiscent

capsule. ± 1 .8 x 2.3 mm, valves spreading to 45° but

curved-erect, split to base, septa only on valves. Seeds

ellipsoid-ovoid, ± 0.5 x 0.25 mm, sometimes angled,

very shallowly alveolate, yellow-brown to brown; testa

cells oblong, + 50-100 x 25-35 pm, anticlinal walls

undulate to irregularly slightly jigsawed, slightly thick-

ened, periclinal walls finely pitted. Figure 3.

Diagnostic features : large, woody, single-stemmed

shrub, mostly 1.0-1. 5, occasionally up to 2, rarely 3 m

tall; leaves 4-nate; flowers in inflorescences of racemose

lower and umbel-like upper whorls; bract leaf-like and

non-recaulescent in lowest whorls to reduced and par-

tially recaulescent in the middle position in the upper-

most whorls; corolla finely and shortly hairy, bright pink;

indumentum on stems, leaves, pedicel, bract, bracteoles

and sepals finely and densely hairy with simple hairs and

some long, stout hairs admixed, these shortly and some-

times densely plumose, mainly towards the base and

often with longer, forked tips.

Erica psittacina is a very distinct species among the

32 Erica species that occur in KwaZulu-Natal. Its rela-

tionships with these species is not clear. Within the group

of species having 4-nate leaves, it is perhaps most simi-

lar to E. algida Bolus which is widespread along the

whole Drakensberg Range but that differs in being a

multi-stemmed resprouter, and having the flowers in a

single inflorescence arranged in one whorl and with par-

tially recaulescent bract. Similarly most of the remaining

KwaZulu-Natal species with 4-nate leaves have the

wrong type of inflorescence and only a partially re-

caulescent bracts. However, the 4-nate E. revoluta

L.E. Davidson has the racemose inflorescence but with-

out the terminal umbel and the bract varies similarly

from its axial position to partially recaulescent. The

shrub is usually large and woody with a similar indu-

mentum in most parts and the leaves are partially open-

backed, but it differs in the glabrous corolla, and the

reduced, partially fused calyx.

The plants are very vigorous growers. With old flowers

from the previous flowering season often still remaining

on some branches, the annual growth increment is easily

assessable. In many cases it is as much as 200 mm, rarely

to 300 mm. This characteristic is reminiscent of the plants

in the type population of E. oakesiorum E.G.H.Oliv. from

Western Cape (Oliver & Oliver 1996) in which the growth

can be as much as 400 mm in one season.

E. psittacina is the most restricted species in the

province, occurring only in a remarkably small area on

Hlabeni Mountain above Creighton in the KwaZulu-

Natal Midlands (Figure 4). There it grows on a steep

southeast-facing slope in a long, relatively narrow belt of

dense, indigenous, shrubby vegetation up to 4 m tall,

consisting mainly of Tarchonantlius camphoratus and

along the upper limits, species of Passerina , Protea and

Cliffortia. The slope is composed of large boulders and

rocky outcrops. Below that belt there is a pristine stand

of tall indigenous forest above vast tracts of exotic pine

plantations. The summit of Hlabeni is covered in well-

grazed and burnt, short grassland.

It is estimated that there are up to 100 plants of E.

psittacina in this extended population. The shrubs vary in

height from 1 m tall in more exposed open places to almost

2 m tall and rarely taller in the denser stands of vegetation.

They are mostly scattered plants that are easily noted due

to their abundant pink flowers contrasting against the other-

wise drab vegetation. Access to many of the plants is very

difficult due to the nature of the terrain and density of the

vegetation. In those that could be examined closely it was

Bothalia 34.1 (2004)

15

noted that the plants are single-stemmed with a trunk up to

100 mm in diameter. The vegetation appeared to be very

old and not affected by the frequent fires that must sweep

the summit plateau, perhaps being protected by a wind

shadow. From below, the habitat is protected by the dense,

tall belt of indigenous forest.

The flowers were seen to be visited by a few honey

bees. This corresponds with large nectaries around the

base of the ovary, the exserted stigma which with slight-

ly enlarged and the partially exserted anthers, in strong-

ly suggesting an insect pollination syndrome for the

species. A slight scent was noticeable from the fresh

flowers in the wild. It was, however, not possible to see

whether the bees came for nectar or just pollen.

The epithet for this new species honours the very dis-

tinguished and rare inhabitant of Hlabeni, the Cape par-

rot, Poicephalus robustus ( psittacinus = belonging to,

possession of, the parrot). The plants were discovered

there by our daughter and two molecular systematists

from Stellenbosch University who were visiting the

mountain in 2002 to sample the orchids. The indigenous

forests above which the erica grows is inhabited by a rem-

nant population of Cape parrots. It was through the assis-

tance of the local conservationist and parrot enthusiast.

Malcolm Gemmel, that they and ourselves visited the

mountain.

Paratype material

KWAZULU-NATAL. — 2929 (Underberg): Creighton, Hlabeni, ± 1

680 m, (-DC), 24-01-2002, T.A.Oliver 8 (NBG).

ACKNOWLEDGEMENTS

We wish to thank Jan Vlok of Oudtshoom and Malcolm

Gemmel of Creighton for their valued assistance in the

field. The Western Cape Nature Conservation Board is

thanked for granting us permission to collect plants in

Western Cape.

REFERENCES

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

deae): species with indehiscent and partially dehiscent fruits.

Contributions from the Bolus Herbarium 19: 345.

OLIVER, E.G.H. & OLIVER, I.M. 1996. Studies in the Ericeae

(Ericoideae). XX. A rare new species of Erica from South

Africa. Yearbook of the Heather Society 1996: 1-5.

OLIVER, E.G.H. & OLIVER, I.M. 2001. Five new species of Erica

(Ericaceae) from the Swartberg Range, Western Cape, South

Africa and a note on E. esterhuyseniae. Bothalia 31: 155-165.

•i

Bothalia 34,1: 17-22(2004)

Two new species of Romulea (Iridaceae: Crocoideae) from the west-

ern Karoo, Northern Cape and notes on infrageneric classification

and range extensions

J.C. MANNING* and P. GOLDBLATT**

Keywords: Iridaceae, new species, Romulea collina J.C, Manning & Goldblatt, Romulea eburnea J.C. Manning & Goldblatt, South Africa, taxonomy

ABSTRACT

Two new species of Romulea are described from Northern Cape, raising the number of species in southern Africa to 76.

R. collina J.C. Manning & Goldblatt is endemic to the Hantamsberg near Calvinia. It is distinguished in subgenus Spatalanthus

by its clumped habit, yellow flowers with dark markings in the throat, and short papery bracts. A re-examination of rela-

tionships within the subgenus suggests that section Cruciatae is not monophyletic and it is accordingly no longer recog-

nized as separate from section Spatalanthus. R. eburnea J.C. Manning & Goldblatt is a distinctive species of subgenus

Spatalanthus from the Komsberg near Sutherland. It is distinguished by its golden yellow flowers with the apical third of

the tepals coloured pale creamy apricot, bracts with broad, translucent margins and tips, and an unusually long perianth tube,

10-13 mm long.

The genus Romulea (Iridaceae: Crocoideae) compris-

es ± 90 species in sub-Saharan Africa, the Mediterranean

and Near East. The centre of diversity of the genus lies in

the winter rainfall region of southern Africa, where 73

species are currently recognized (Manning & Goldblatt

2001). The most recent revision of the sub-Saharan

species divides the genus into two subgenera and six sec-

tions, based largely on characters of the corm (Manning

& Goldblatt 2001). The fine structure of the corm pro-

vides essential information for accurate identification of

many species, since the flower structure, with few excep-

tions, is conservative (Goldblatt et al. 2002).

Five new species of Romulea have been described

from South Africa in the twenty years since the publica-

tion of De Vos’s (1972) monograph of the genus in south-

ern Africa (De Vos 1983; Manning & Goldblatt 2001).

The subsequent discovery of an undescribed species

from the summit plateau of the Hantamsberg near Cal-

vinia in Northern Cape, described here as Romulea colli-

na, and a second from the Klein Roggeveld south of

Sutherland, described as R. eburnea, now raises the total

number of species in southern Africa to seventy six. Both

new species are members of subgenus Spatalanthus,

defined by the outer corm tunics splitting below into

prominent, recurved or straight teeth without fibrous tips

(Manning & Godblatt 2001). The Roggeveld Escarpment

is well known for the diversity of its endemic geophyte

flora (Manning et al. 2002) and is the centre of diversity

for subgenus Spatalanthus. Over half of the twenty five

known species of the subgenus are endemic to the Rog-

geveld and adjacent Bokkeveld Escarpments (Manning

& Goldblatt 2001) and the description of two new spe-

cies brings the total number of endemic Romulea species

in this centre to fifteen.

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

7735 Claremont, Cape Town.

** B.A. Krukoff Curator of African Botany, Missouri Botanical Garden,

P.O. Box 299, St. Louis, Missouri 63166, USA.

MS. received: 2003-11-27.

The discovery of Romulea collina raises doubts as to

the validity of the distinction between the two sections

Cruciatae and Spatalanthus of subgenus Spatalanthus,

recognized by Manning & Goldblatt (2001) in their

recent account of the genus. This distinction is re-exam-

ined and we conclude that the recognition of section

Cruciatae is no longer justified.

Romulea collina J.C. Manning & Goldblatt, sp. nov.

TYPE. — Northern Cape, 3119 (Calvinia): summit of

Hantamsberg at base of radio mast, (-BD), August 2002,

flowered in cult., July 2003, IBSA 5 ( NBG195425 , holo.).

Plantae 50-100 mm altae caespitosae, caule subter-

raneaneo ad 4-ramoso, ramis ad 10 mm supra terram pro-

ductis, cormo subgloboso asymmetrico base rotundo,

tunicis infra divisis cuspis acuminatis recurvatis supra

fibris grossis 20-40 mm longis, foliis 7-9 laminis ad

50-100 mm longis, floribus solitariis, bracteo extemo

subobtuso pallide papyraceo ad apicem marginibus

apiceque latis translucentibus brunneo vittato 10-12 mm

longo intemo obtuso pallide papyraceo, floribus pro-

funde cupuliformibus pallide flavis cupulo luteo, tubo

perianthii infundibuliforme 4 mm longo parte inferiore ±

1 mm longo, tepalis lanceolatis ± 20 x 7 mm, staminibus

flavis filamentis ± 6 mm longis dense pubescentibus in

dimidio inferiore, antheris 2. 5-3.0 mm longis apiculatis,

ramis styli 1. 5-2.0 mm longis, capsulis subglobosis ± 8

mm oblongo-ovoideis pedicelibus recurvatis.

Plants 50-100 mm high, growing in clumps; stem

subterranean, with up to four branches reaching 10 mm

above-ground, these relatively stout and flushed maroon.

Corm subglobose, asymmetric, base rounded, tunics split

into curved acuminate teeth below, drawn into coarse

fibres above, these 20-40 mm long. Cataphylls 3,

flushed maroon above ground. Leaves 7-9, all basal,

sheaths flushed deep purple, blades two to four times as

long as flowering stems, narrowly 4-grooved, 50-100 x

1.0-1. 5 mm. Inflorescence : up to 4 solitary flowers;

18

Bothalia 34,1 (2004)

FIGURE 1. — Romulea collina. A,

whole plant; B, flower; C,

outer bract; D, inner bract; E,

adaxial markings on outer

tepal; F, adaxial markings on

inner tepal; G, stamens and

style; H, seed; I, t/s leaf.

Scale bars: A-F, 10 mm; G, H,

2 mm. Artist: John Manning.

outer bracts subobtuse, pale and papery throughout with

veins in centre pale green, with broad, translucent mar-

gins and tip flecked pale brown, 10-12 mm long; inner

bracts obtuse, pale and papery throughout with only

main veins pale green, with broad translucent margins

flecked pale brown, about as long as outer. Flowers

deeply cup-shaped, cup ± 1 1 mm deep, pale canary yel-

low with golden yellow cup; outer tepals with dark

median blotch or transverse zone in throat, reverse

marked with dark maroon longitudinal streaks; inner

tepals with narrow brownish maroon median line and

diffuse transverse zone in throat, unscented, ± 30 mm

diam.; perianth tube funnel-shaped, 4 mm long with

lower narrow portion ± 1 mm long, tepals lanceolate, ±

20 x 7 mm. Stamens yellow; filaments inserted at base of

cup, free, densely hairy in lower half, 6 mm long; anthers

apiculate, parallel, 2. 5-3.0 mm long. Style dividing

opposite upper third of anthers, branches 1 .5-2.0 mm

long, divided for about two thirds of their length.

Capsules subglobose, ± 8 mm long, pushed onto soil sur-

face or slightly underground by strongly recurved

pedicels which later become erect and 10-15 mm long.

Seeds subglobose or angled by pressure, reddish brown,

2. 5-3.0 mm diam. Flowering time: late June to mid-July,

possibly to late July. Figure 1.

Distribution and ecology

This clump-forming species is known from a single,

very localized colony on the summit plateau of the

Hantamsberg overlooking Calvinia on the Bokkeveld

Escarpment (Figure 2). The plants grow in seasonally

moist, dolerite clays in open renosterveld, invariably in

small clumps that appear to originate from seeds that are

shed directly at the base of the parent plant. The short

peduncles curve downwards immediately after flowering

but become erect when the fruits mature. They do not,

however, elongate appreciably so that the mature cap-

sules dehisce 10-15 mm above the ground at the base of

the plant. Flowers open around midday and close in mid-

afternoon. They are short-lived, lasting only two days. At

up to 3 mm diam., the seeds of R. collina are among the

largest recorded in the genus, where seeds are mostly 1-2

mm diam. (De Vos 1972).

Romulea collina is yet another of several species of

Iridaceae that are endemic to the Hantamsberg, an isolated,

flat-topped massif representing a northern outlier of the

Roggeveld Escarpment. Iridaceae endemic to the slopes and

summit of the Hantamsberg include Hesperantha hanta-

mensis Schltr. ex Foster, H. oligantha (Diels) Goldblatt,

Bothalia 34,1 (2004)

19

FIGURE 2. — Distribution of Ronnilea collina, O: and Romulea

eburnea. •.

Moraea reflexa Goldblatt and Romulea hantamensis (Diels)

Goldblatt, among others (Van Wyk & Smith 200 1 ).

Diagnosis and relationships

Romulea collim is readily distinguished by its clumped

habit, rounded corm with the outer tunics splitting into acumi-

nate teeth, yellow flowers with dark markings in the throat, and

short bracts, 10-12 mm long, that are entirely pale and papery

at flowering. The corm with tunics splitting into acuminate

teeth at the base are a defining characteristic of Romulea sub-

genus Spatalanthus but within the subgenus, the relationships

of R. collina are more difficult to define. The rounded corm

with curved basal teeth is characteristic of section Spatalanthus

but in other respects, particularly the relatively short, more or

less entirely membranous bracts and the long filaments, the

species closely resembles R. membranacea M.P. de Vos, anoth-

er species from the Roggeveld Escarpment.

Conserx’ation status: Vulnerable ( B 1 + 2c, D).

Apart from the difference in the corm shape, R. collina

is separated from R. membranacea by the longitudinal dark

streaks on the outside of the outer tepals and by the pro-

nounced dark blotches on the inside. Despite their different

corm morphology, the overwhelming similarity between the

two species in other particulars makes it difficult to accept

that they are not immediately related. R. membranacea is

currently placed in section Cruciatae, which is defined by a

corm pointed at base and with straight basal teeth (Manning

& Goldblatt 2001 ) but the relationships of this species have

always been problematic. De Vos (1972) placed it in her

subsection Atrandrae on the basis of the broad membranous

margins of the bracts but pointed out that it was anomalous

here in its pointed corm and lack of marginal veins in the

leaves. Manning & Goldblatt (2001), placing greater

emphasis on corm structure, preferred to ally it with the

other species with straight basal teeth, which it matches in

leaf anatomy. The chromosome number 2n = ± 24 is, how-

ever, inconsistent with this alliance and accords better with

section Atrandrae. The discovery of the new species R. col-

lina, with the rounded corm of section Spatalanthus but the

leaf anatomy and bract morphology of R. membranacea ,

suggests that the earlier placement by De Vos (1972) of R.

membranacea in subsection Atrandrae (= series Atrandrae

of Manning & Goldblatt 2001) is preferable. Within series

Atrandrae , R. collina and R. membranacea are probably

most closely related to R. diversiformis M.P.de Vos, with

which they share yellow flowers, fruiting peduncles that are

curved at maturity rather than coiled, and leaves that lack

marginal veins along the furrows.

The reassessment of the relationships of R. collina and R.

membranacea implies that the pointed corm with straight

basal teeth has arisen more than once within the genus. If

this interpretation is correct then the heterogeneity evident

within the section assumes a new significance. Section

Cruciatae is currently divided into two series. Series

Cruciatae, with the removal of R. membranacea, contains

three species, two of which are clearly sister taxa [77 cruci-

ata (Jacq.) Baker and R. eximia M.P.de Vos] that share the

unusual chromosome number 2 n = 18. On the basis of then-

cytology, R. cruciata and R. eximia are better allied with R.

rosea (L.) Eckl. (section Spatalanthus , series Roseae), the

only other species in the genus with 2 n - 18. The third

species in this series, the poorly known R. vlokii M.P.de Vos

has bracts that more closely resemble those found in series

Atrandrae of section Spatalanthus, which is where De Vos

(1983) placed it. Series Tubiformes contains a single highly

derived species, R. hantamensis (Diels) Goldblatt, with a

chromosome number 2 n = 30, which is unique within sub-

genus Spatalanthus and its relationships remain unclear. If

the pointed corm is interpreted as a convergent character

state, then it is likely to have evolved three times in section

Cruciatae and a fourth time in R. membranacea. Section

Cruciatae as currently construed is probably not mono-

phyletic and it seems more appropriate that the species

within it be removed to section Spatalanthus. Accordingly,

we no longer recognize section Cruciatae, and subgenus

Spatalanthus should no longer be subdivided into sections.

Romulea eburnea J C. Manning & Goldblatt, sp.

nov.

TYPE.— Northern Cape, 3220 (Sutherland): 28 km

along Komsberg Pass road from southern end, foot of

Smoushoogte, alluvial wash along Meintjiesplaasrivier,

(-DC), 28 August 2003, J. Manning 2886 (NBG, holo.;

MO, iso.).

Plantae 100-150 mm altae, caule subterraneaneo 2-

ramoso, ramis ad 30 mm supra terram productis, cormo

subgloboso asymmetrico base rotundo, tunicis infra divi-

sus cuspis acuminatis recurvatis supra fibris grossis ± 2

mm longis, foliis 2-4 laminis ad 150-200 mm longis,

floribus solitariis, bracteo extemo marginibus translucen-

tibus apice membranaceo brunneo vittato 18-22 mm

longo intemo bifido, floribus profunde cupuliformibus

pallide cremeo-armeniacis supra luteis infra ± 30 mm

diam., tubo perianthii infundibuliforme, 10-13 mm longo

parte inferiore 6-8 mm longo, tepalis lanceolatis, 23-27 x

7-9 mm, staminibus flavis filamentis 6-7 mm longis ad

basem pubescentibus, antheris ± 8 mm longis, ramis styli

3 mm longis, capsulis oblongo-ovoideis 10-11 x 5. 5-6.0

mm pedicelibus recurvatis spiralis ubi siccus.

20

Bothalia 34,1 (2004)

Plants 100-150 mm high; stem subten'anean, with up to

two branches reaching 30 mm above ground, these relatively

stout and flushed maroon. Conn subglobose, asymmetric,

base rounded, tunics split into curved acuminate teeth below,

drawn into coarse fibres above, these up to 2 mm long.

Cataphylls 3, flushed maroon above ground. Leaves 2^4,

sheathing portion pale with adhering sand grains, blades of

two longest leaves up to six times as long as flowering stems,

narrowly four-grooved, 150-200 mm long, when more than

two leaves present, then uppermost one or two with blades less

than half as long, 10-100 x 1 mm. Inflorescence : up to 2 soli-

tary flowers; outer bracts pale reddish brown with broad,

translucent margins and broad, membranous tip flecked with

pale brown, 1 8-22 mm long; inner bracts bifid, pale reddish

brown with broad, translucent margins and broad, membra-

nous tip flecked with pale brown, about as long as outer or

slightly longer. Flowers deeply cup-shaped, cup ±12 mm

deep, pale creamy apricot with lower two thirds of tepals and

cup deep yellow; outer tepals with reverse flushed dull olive

in centre in lower half, unscented, ± 30 mm diam.; perianth

tube funnel-shaped, 10-13 mm long with lower narrow por-

tion 6-8 mm long, tepals lanceolate, 23-27 x 7-9 mm.

Stamens yellow; filaments inserted at base of cup, free, pubes-

cent at base, 6-7 mm long; anthers suberect or lightly

incurved, ± 8 mm long. Style dividing opposite tips of anthers,

branches 3 mm long, divided for about half their length.

Capsules oblong-ovoid, 10-11 x 5.5-6.0 mm, pushed onto

soil surface or slightly underground by strongly recurved

pedicels which later coil up when dry. Seeds subglobose or

angled by pressure, reddish brown, 1. 8-2.0 mm diam.

Flowering time: late August to early September. Figure 3.

Distribution and ecology

This distinctively coloured species is known from a single

colony below Smoushoogte Pass south of Sutherland at the

FIGURE 3. — Romulea eburnea. A,

whole plant; B, flower, side

view; C, adaxial markings on

outer tepal; D, adaxial mark-

ings on inner tepal; E, sta-

mens and style; F, dry cap-

sule; G, seed; H, t/s leaf.

Scale bars: A-D, F, 10 mm;

E, 5 mm; G, 1 mm. Artist:

John Manning.

Bothalia 34,1 (2004)

21

foot of the Klein Roggeveldberge (Figure 2). Romulea

ebumea grows in an alluvial wash along the Meintjies-

plaasrivier in seasonally moist, deep sandy loam. The veg-

etation is open scrub dominated by Asparagus capensis ,

Chrysocoma ciliata , Eriocephalus eximius, and Dimorpho-

theca cuneata , and quite distinct from the adjacent renos-

terveld {Elytopappus rhinocerotis) community that occu-

pies fine-grained clays derived from shale.

R. ebumea was first noticed almost a decade ago by local

bulb enthusiasts who reported the occurrence of a pale-flow-

ered species on the Komsberg but in the absence of speci-

mens it was impossible to verify its identity. The possibility

that it represented an outlying population of R. mem-

branacea was suggested by Manning & Goldblatt (2001)

but it is now clear that the plants represent a distinct species.

It is another of several species in series Atrandrae that are

endemic or near-endemic to the Roggeveld Escarpment

around Sutherland, including R. hallii M.P.de Vos, R. koms-

bergensis M.P.de Vos and R. multifida M.P.de Vos.

Diagnosis and relationships

The rounded corms with tunics fragmenting into

curved teeth, place Romulea ebumea firmly in subgenus

Spatalanthus. It is immediately recognized by its largely

golden yellow flowers with the apical third of the tepals

coloured pale creamy apricot, bracts with broad, translu-

cent margins and tips, and the unusually long perianth

tube. This is funnel-shaped and 10-13 mm long with the

lower narrow portion 6-8 mm long. In most other

species of Romulea with a similar-shaped perianth, the

tube is 4- 6(-10) mm long with the lower narrow portion

no more than 2 mm long. Within subgenus Spatalanthus,

the presence of vascular bundles along the margins of

each of the ribs in the leaves, the broad membranous

margins and tips to the bracts, and fruiting peduncles that

coil when dry. are all consistent with series Atrandrae.

Within series Atrandrae, R. ebumea is possibly most

closely allied to R. diver siformis, with which it shares

yellow flowers lacking dark markings in the throat and a

long style that divides at or beyond the tips of the

anthers. In an extraordinary coincidence, both R. diversi-

formis, described just over fifty years ago (De Vos 1952),

and R. ebumea share the same type locality and we have

in the past collected R. diversiformis in flower in late

September within a few dozen metres of the R. ebumea

population, then long past flowering, without being

aware of its existence. R. diversiformis was until recent-

ly thought to be restricted to the Klein Roggeveld area

but is now known to occur also on the Hantamsberg near

Calvinia, a considerable distance away at the northern

end of the Roggeveld Escarpment (Manning & Goldblatt

2001). R. diversiformis is distinguished from R. ebumea

by its greater number of leaves, 6 or more versus 2-4,

uniformly golden yellow flowers with short perianth

tube, 4—6 mm long versus 10-13 mm long and fruiting

peduncles that do not coil up when dry. The two species

differ also in leaf anatomy, R. diversiformis lacking rib

marginal veins but possessing secondary veins in the

ribs. R. diversiformis grows in fine-grained, clay soils

that are seasonally waterlogged or actually shallowly

inundated, whereas R. ebumea is known only from sea-

sonally moist, sandy soils.

Conservation status: Vulnerable (B1 + 2c, D).

Adjustments to distribution records of two Namaqua-

land species

Romulea maculata Manning & Goldblatt in Adansonia

23: 81 (2001).

This species was based on a single collection made

on the summit of the Flaminkberg just southeast of

Nuwerus in southern Namaqualand. The recent discov-

ery of a second collection antedating the type by almost

twenty years, substantially increases the known range of

the species. This collection matches the type in all

respects, including the large white flowers, bracts with

broad, brown-spotted membranous margins and the

white-spotted cataphyll and there is no doubt that it rep-

resents this species. Romulea maculata is now known

from near Nuwerus in the south of Namaqualand to

Komaggas in the north and may be expected to occur

through most of the higher ground of Namaqualand. The

species is restricted to seasonally moist, sandy loam on

granitic slopes.

Additional material examined

NORTHERN CAPE. — 2917 (Springbok): Farm Drierivier, (-DC),

17-08-1980, Van der Westhuizen 109/80 (NBG).

Romulea multisulcata M.P.de Vos in Journal of South

African Botany, Suppl. 9: 139 (1972).

This species was described from seasonal pools on the

Bokkeveld Mountains near Nieuwoudtville in Northern

Cape (De Vos 1972) but later collections from the foot of

the Gifberg-Matsikamma massif nearby (Manning &

Goldblatt 2001), clearly represent a second set of popu-

lations differing from the type only in their white rather

than yellow flowers. However, populations from

Hondeklip Bay in central Namaqualand that were also

assigned to this species on the basis of a single herbari-

um collection (Manning & Goldblatt 2001) prove, on

examination of living plants, to lack the characteristic

multisulcate leaves of R. multisulcata and accord more

closely with R. tabularis Eckl. ex Beg. This species of

waterlogged coastal flats is widely distributed along the

west coast of South Africa from Cape Agulhas in the

south to northern Namaqualand. The Hondeklip Bay

populations thus fall within its recorded range. Romulea

multisulcata thus remains endemic to southern Namaqua-

land, where it is restricted to a few scattered seasonal

pools near Vanrhynsdorp and Nieuwoudtville.

Range extensions for Romulea tabularis

NORTHERN CAPE. — 3017 (Hondeklipbaai): Koingnaas,

Skulpfontein, (-AB), 24-08-1999, Desmet 222 (NBG); Farm

Strandfontein. 15 km S of Wallekraal, (-DA), 25-08-1996, De Villiers

s.n. (NBG 177635); Groen River mouth, slopes above estuary ± 1 km

from ocean, (-DC), 25-8-2002, Goldblatt & Porter 12116 (MO, NBG).

ACKNOWLEDGEMENTS

We thank Roy Gereau for his advice on nomenclature

issues. Support for field work was provided from grants

6704-00 and 7316-02 from the US National Geographic

Society.

22

Bothalia 34,1 (2004)

REFERENCES

DE VOS, M.R 1952. Nuwe en minder bekende Romulea- spesies van

die Roggeveld, Annals of the University of Stellenbosch 28 A,

3: 59-80.

DE VOS, M.P. 1972. The genus Romulea in South Africa. Journal of

South African Botany, Suppl. 9.

DE VOS, M.R 1983. Romulea. In O.A.Leistner, Flora of southern

Africa 7, part 2, fasc. 2: 10-73. Botanical Research Institute,

Pretoria.

GOLDBLATT, P„ BERNHARDT. P. & MANNING. J.C. 2002. Floral

biology of Romulea (Iridaceae: Crocoideae): a progression

from a generalist to a specialist pollination system. Adansonia

24: 243-262.

MANNING, J. & GOLDBLATT, P. 2001 . A synoptic review of Romulea

(Iridaceae: Crocoideae) in sub-Saharan Africa, the Arabian

Peninsula and Socotra including new species, biological notes,

and a new infrageneric classification. Adansonia 23: 59-108.

MANNING, J„ GOLDBLATT, P. & SNIJMAN, D. 2002. The color

encylopedia of Cape bulbs. Timber Press, Oregon.

VAN WYK. A. E. & SMITH, G. F. 2001. Regions of floristic endemism

in southern Africa. Umdaus Press, Pretoria.

Bothalia 34,1: 23-26(2004)

Studies in the genus Riccia (Marchantiales) from southern Africa. 26.

A new species in section Pilifer ; Riccia radiata, is described

S.M. PEROLD*

Keywords: dorsal epithelium, Riccia radiata Perold, section Pilifer O.H.Volk, sporangia, spores, thallus

ABSTRACT

Riccia radiata Perold, a new species endemic to southern Africa, is described. It is referred to section Pilifer O.H.Volk,

which is characterized by the dorsal epithelium of the thalli consisting of short or long, free-standing, hyaline cell pillars.

INTRODUCTION

Including this new species, 18 southern African

species of Riccia have been referred to section Pilifer.

The previous 17 species were treated in Perold (1999).

As I remarked previously in Perold (1990a), the species

assigned to this section, with rare exceptions, have ±

rounded, colourless scales and in several the spores have

radiating ridges on the distal face. The dorsal cell pillars

are composed of two or more hyaline cells: two cells in

R. pulveracea (Perold 1990b) for example, and up to six

cells joined end to end in R. villosa (Volk & Perold

1984). Once the thalli have dried out, the cells in the pil-

lars collapse and it is often not possible to reconstitute

them, especially if the plants have died. Fortunately,

upon keeping some of the thalli of this new species damp

in a closed transparent container for two weeks (16

months after collection), the distal parts of the thalli had

revived sufficiently and could be studied. The ornamen-

tation on both spore faces proved to be quite distinctive

and it was quickly distinguished as a new species, which

is described below.

Riccia radiata Perold , sp. nov.

Thalli gregarii caespitosi, apicem versus laete viridi,

superficies dorsalis columnis cellularum munita mox

collabentibus, caespites lanosos formantibus. Squamae

hyalinae, arete imbricatae. Sporae unice ornatae: superfi-

cies distalis cum cristis pluribus densis e centra radiatis

(itaque nomen), areolis completis vel incompletis inter

cristas; superficies proximalis granulis tenuis, interdum

coalescentibus, confertim tecta.

TYPE. — Northern Cape, 3219 (Wuppertal): Ceder-

berg, foothills of Bloukop along Luiperdskloof 4x4

route, altitude 1 290 m, on mountain slope, on sandy soil,

(-CB), 2002-09-13, M. Koekemoer 2426 (PRE, holo.).

Figure 1.

Thalli perennial, in gregarious patches, sometimes

overlapping, not forming rosettes, dorsal face apically

bright green and glistening, but soon becoming dotted

with scattered white tufts of collapsed cell pillars, proxi-

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

MS. received: 2004-02-26.

mally shaggy, entirely covered with closely crowded,

coarse white woolly tufts (Figure 3A-C); medium-sized,

6-11 mm long, growing from a narrow base, ± 0.5 mm

wide and widening distally to 2.5 mm before bifurcation,

in cross section slightly concave above and rounded

below, 0.9 mm thick, i.e. nearly 3 times wider than thick

(Figure 2C). Branches once or twice symmetrically or

asymmetrically furcate, lobes ligulate, narrowly to mod-

erately divergent, terminal segments 1. 3-5.0 mm long,

up to 2 mm wide, apex ovoid, rounded or wedge-shaped,

once or twice shortly grooved (Figure 3B, C, E, F), mar-

gins acute, sides steep, with overlapping hyaline scales

(Figure 2A). In cross section apically (Figure 2B), thal-

lus branches dorsally V-shaped, only 0.6 mm thick from

bottom of groove to ventral face, at erect sides up to 1 .2

mm thick and here almost as thick as wide; ventrally

rounded, green, hyaline scales barely visible, covered

with rhizoids, 5.0-17.5 pm wide, mostly smooth, rarely

tuberculate. When dry, thallus generally white dorsally,

sometimes tinged with pale green toward apex, slightly

concave, margins erect or partially inflexed, sides white

to pale yellow, covered with appressed scales, occasion-

ally with narrow or broad, deep purple band along length

of side, tailing off proximally. Scales along sides of thal-

lus (Figure 3C, D), extending 200-250 pm above dorsal

face, closely imbricate, ± oblong, somewhat forward

projecting at leading edge (Figure 2F), rounded above,

often sloping at following edge, margins entire, hyaline.

FIGURE 1. — Locality of Riccia radiata in southern Africa.

24

Bothalia 34,1 (2004)

FIGURE 2. — Riccia radiata, M. Koekemoer 2426. A, proximal part of thallus and distal branches after bifurcation; B, c/s branch through api-

cal groove; C, c/s proximal part of thallus before bifurcation, tops of collapsed dorsal pillars indicated by broken line; D, c/s dorsal cell

pillars and top cells of assimilation tissue cell columns; E, horizontal section through basal cells of cell pillars with air pores dotted; F,

scale. Scale bars: A, 2 mm: B, C, 500 pm; D, 100 pm; E, 50 pm; F, 250 pm. Drawn by M. Steyn.

800-950 pm high, 1130-1175 pm across widest part,

cells in body of scale mostly elongated, 5- or 6-sided,

92.5-137.5 x 50.0-67.5 pm, decreasing in size toward

upper margin, where small and brick-shaped, in 1 or 2

rows; rhizoids arising from following edge.

Dorsal epithelium (Figure 2D) consisting of free-

standing 3- or 4-celled hyaline pillars, fragile, 135-250

pm long, top cells ± spherical, conical or elongated with

rounded apex, very variable in size, 50-150 x 30-75 pm,

next cell 50.0-92.5 x 45.0-62.5 pm, basal cell (if 3 cells

iMg

FIGURE 3.- SEM micrographs of Riccia radiata, M. Koekemoer 2426. A, distal branches of thallus after bifurcation; B, apical part of left branch

(in A) with clumps of collapsed dorsal cell pillars; C, apical part of right branch (in A) and scales along sides; D, side view of branch,

showing scales; E, F, groove at branch apex, with intact top cells of dorsal pillars on either side. A, x 8.7, B-D, x 16.5, E, x 60.2; F, x 53.8.

Bothalia 34,1 (2004)

25

FIGURE 4. — Riccia radiata, M. Koekemoer 2426. Spores. A-F, distal face: F, side view. G, H, proximal face; I, apex of triradiate mark. A, H. x

435; B, F. G, x 440; C, x 468; D, x 460; E, x 525; I, x 920.

in pillar) 37.5-72.5 x 35.0-57.5 pm, (if 4 cells in pillar)

37.5-50.0 x 30.0-32.5 pm; air pores small (Figure 2E),

often 4-sided, ± 20 x 20 pm, obscured by cell pillars.

Assimilation tissue 250 x 300 pm thick in section, ± '/3

the thickness of thallus and consisting of vertical

columns of ± 8 cells, 35-50 x 35-40 pm, enclosing nar-

row, 4-sided air canals; storage tissue 35CM-00 pm thick,

up to ± V2 the thickness of thallus, cells crowded togeth-

er, rounded to slightly angular, ± 45 x 50 pm; rhizoids

arising from ventral epidermal cells.

Monoicous? Antheridia not seen; hyaline necks prob-

ably obscured by woolly tufts of collapsed dorsal cells

of thallus. Archegonia with dark red-brown necks, per-

sistent and prominently projecting from bulging sporan-

gia. Sporangia not very common, mostly single,

800-875 pm wide, rarely up to 3 in close proximity near

bifurcation of thallus. Spores 92.5-1 10.0 pm diam., tri-

angular-globular, polar, fairly dark brown, semitranslu-

cent; wing sprinkled with granules, ± 6 pm wide, grad-

ually widening to 15 pm at notched or perforated mar-

ginal angles, margin often minutely crenulate; ornamen-

tation on distal face quite variable (Figure 4A-F), usual-

ly with 3-6 heavy ridges, up to 5 pm wide, radiating

from centre, in between ridges complete or incomplete

areolae, 7.5-15.0 x 7.5-12.5 pm, also thick-walled,

sometimes with a central boss; proximal face (Figure

4G-I) with clearly defined triradiate ridge extending to

margin, all 3 facets densely covered with fine granules,

some coalescing.

This species has only been collected once. It was

found growing on sandy soil on a mountain slope in the

Northern Cape, at the border with Western Cape. This is

a winter rainfall area with vegetation types Northwestern

Mountain Renosterveld and Mountain Fynbos (Low &

Rebelo 1996). The type locality is in a rather inaccessi-

ble place, along a newly opened 4x4 route. It is recog-

nized as a new species mainly by the unique spore orna-

mentation which is characterized by exceptionally heavy

ridges, and is referred to section Pilifer because of the

free-standing cell pillars dorsally on the thalli. The spe-

cific epithet was chosen because of the radiating ridges

on the distal face of many of the spores, quite different

from those previously studied.

ACKNOWLEDGEMENTS

I sincerely thank the referees for their kind advice,

also Dr M. Koekemoer for collecting the type specimen

26

Bothalia 34,1 (2004)

of this new species. Dr H.F. Glen is thanked for suggest-

ing the specific epithet and for translating the diagnosis

into Latin. I also extend my gratitude to Mrs M. Steyn for

the drawings and to Ms D. Maree for typing the manu-

script.

REFEREN-CES

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

and Swaziland. Department of Environmental Affairs & Tourism,

Pretoria.

PEROLD, S.M. 1990a. Taxonomic relevance of the spore-wall ornamen-

tation in the southern African species of Riccia L. (Hepaticae:

Ricciaceae). Unpublished M.Sc. thesis. University of Pretoria.

PEROLD, S.M. 1990b. Studies in the genus Riccia (Marchantiales)

from southern Africa. 19. Two new species: R. pulveracea , sec-

tion Pilifer, and R. bicolorata, section Riccia, group 'Squamatae'.

Bothalia 20: 185-190.

PEROLD, S.M. 1999. Hepatophyta. Part 1: Marchantiopsida, Fascicle

1: Marchantiidae. Flora of southern Africa. National Botanical

Institute, Pretoria.

VOLK, O.H. & PEROLD, S.M. 1984. Studies in the genus Riccia

(Marchantiales) from the southwest Cape. Bothalia 15: 117-124.

Bothalia 34,1: 27-51 (2004)

Notes on African plants

VARIOUS AUTHORS

SCROPHULARIACEAE

TYPE SPECIMENS OF SELAGO, JAMESBR1TTEN1A AND SUTERA AT NATAL UNIVERSITY HERBARIUM (NU)

The recent revisions of the Manuleae and Selagineae

(Hilliard 1994, 1999) provide excellent accounts of these

tribes. Unfortunately there are considerable omissions

regarding type material housed at Natal University

Herbarium (NU). For a number of the taxa. Natal

University Herbarium holds the sole isotype and most of

the holotypes are housed overseas in Edinburgh (E) or

Missouri (MO).

SELAGINEAE

Selago dolosa Hilliard. Type: Eastern Cape, 3225

(Somerset East): Boschberg, (-DA), 30-11-1977, Hilliard

& Burn 10785 (E, holo.; NU).

Selago florifera Hilliard. Type: Cape [Northern

Cape], 3120 (Williston): between Calvinia and Middel-

pos on Bloemfontein road, (-CC), 30-09-1976, Goldblatt

4266 (MO, holo.; E, NU, PRE).

Selago griquana Hilliard. Type: Natal [KwaZulu-

Natal], 3029 (Kokstad): Swartberg, Matatiele on D492,

just past Umzimkulu River, (-AC), 8-01-1986, Hilliard

& Burn 18918 (E, holo.; K, NU, S).

Selago hyssopifolia E.Mey. subsp. retrotricha

Hilliard. Type: Natal [KwaZulu-Natal], 3029 (Kokstad):

near Weza, Zuurberg, (-DA), 26-02-1975, Hilliard &

Burn 8061 (E. holo.; K. NU, S).

Selago inconstans Hilliard. Type: Eastern Cape,

3128 (Umtata): hill above Mhlahlane Forest Station, NW

of Umtata, ± 5000 ft, (-BC), 31-01-1983, Hilliard &

Burn 16343 (E, holo.; K, KEI, NU, PRE, S).

Selago retropilosa Hilliard. Type: Eastern Cape,

3124 (Hanover): S extreme of Renosterberg above Loots-

berg railway halt. Farm Blaauwater. 6000 ft, (-DD), 25-

11-1977, Hilliard & Burn 10651 (E, holo.; MO, NU,

PRE. S).

Selago taraehodes Hilliard. Type: Natal [Kwa-

Zulu-Natal], 2832 (Mtubatuba): St Lucia Estuary,

(-AD), 15-08-1975, Pooley 1749 (E, holo.; K. NU).

Selago variicalyx Hilliard. Type: Eastern Cape,

3326 (Grahamstown): Grahamstown Nature Reserve,

Dassie Krantz, ± 2500 ft, (-AD), 2-12-1977, Hilliard &

Burn 10798 (E, holo.; K, NU, S).

Selago zuluensis Hilliard. Type: Natal [KwaZulu-

Natal], 2831 (Nkandla): Ngoye Forest, (-DC), 15-01-1986,

Hilliard & Burn 19052 (E, holo.; K, NU).

MANULEAE

Jamesbrittenia multisecta Hilliard. Type: East-

ern Cape, 3128 (Umtata): Umtata-Engcobo road near

Umtata, (-DA), 21-11-1977, Hilliard & Burtt 10552 (E,

holo.; NU).

Sutera glandulifera Hilliard. Type: Eastern Cape,

3226 (Fort Beaufort): Katberg Pass, ± 5800 ft, (-DA), 26-

01-1979, Hilliard & Burn 12390 (E, holo.; K, M, NU, S).

REFERENCES

HILLIARD, O.M. 1994. The Manuleae, a tribe of Scrophulariaceae.

Edinburgh University Press, Edinburgh.

HILLIARD. O.M. 1999. The tribe Selagineae ( Scrophulariaceae ). Royal

Botanic Gardens, Kew.

T.J. EDWARDS*

* School of Botany and Zoology, University of Natal, Private Bag X01 ,

3209 Scottsville.

MS. received: 2003-02-28.

PTERIDOPHYTA

DRYOPTERIS FILIPALEATA (PTEROPSIDA: DRY OPTERIDACEAE),

INTRODUCTION

In preparing a taxonomic review of the fern genus Dty-

opteris in Africa as well as for the Flora of tropica! East

Africa , several undescribed species have been identified.

Earlier collectors generally ascribed their Diyopteris collec-

tions to either D. inaequalis (Schltdl.) Kuntze or to D. pentheri

(Krasser) C.Chr., but a critical review of these species by me

showed that they need to be more narrowly defined.

A NEW SPECIES FROM TROPICAL EAST AFRICA

Dryopteris filipaleata J.P.Roux forms part of a group

of species belonging to section Marginatae Fraser-Jenk.

(Fraser-Jenkins 1986), to which both D. inaequalis and

D. pentheri belong. The near similar stoma size in D. fili-

paleata [(40.0— )5(). 1 (—62.0) pm] and D. pentheri

[(34.0-)53.34(-72.0) pm], as well as spore size in D. fili-

paeata [(32.0-)4 1.1 (-54.0) x (18.0-)26.26(-34.0) pm]

and D. pentheri [(38.0-)45.08(-60.0) x (27.0-)31.32

(-40.0) pm], suggest that D. filipaleata is tetraploid.

28

Bothalia 34,1 (2004)

Furthermore, the development of 64 normal spores per

sporangium indicates that the species is sexual.

Dryopteris filipaleata can, however, be separated from

D. pentheri by being exindusiate, eglandular and without

2-celled hairs. In D. pentheri , oblong glands (60. 0-)

1 37.02(— 260.0 ) pm long and 2-celled hairs generally

occur along the lamina axes and veins. The narrow lam-

ina scales also separate it from D. pentheri.

Dryopteris filipaleata appears to be restricted to

Kenya and Tanzania — the Kenyan and Tanzanian moun-

tain forests forming part of the Afromontane Region

(White 1983). Lovett (1988) divided the Tanzanian

forests into a number of subdivisions based on geo-

graphical, edaphic and floristic factors. These include the

coastal forests, the Lake Victoria Basin forests, the

Western mountain forests, also termed the Eastern Arc

mountain forests, and the volcanic mountain forests. The

Eastern Arc mountain forests, which have strong floristic

similarities, show a high percentage of endemism

(Brenan 1978; Lovett 1988). Floristically these forests

differ from those on the adjacent recent volcanic moun-

tains and the basin forests (Lovett 1988). This may be

ascribed to edaphic factors such as the different soil

types.

Dryopteris filipaleata J.P.Roux, sp. nov., laminae

paleis angustis denticulatis, paleis rhizomae stipitisque

majoribus marginibus irregulariter laceratis et gemmis

absentibus differt. Figura 1 & 2.

TYPE. — Tanzania, Eastern Province, Morogoro Dist.

(T6), Uluguru Mountains, Mwere Valley, wet evergreen

forest along stream with abundant Cyathea manniana

and epiphytes, 1 400-1 450 m, 26 Sept. 1970, R.B. Faden,

T. Poes, B.J. Harris, & P. & K. Csontos 70/596 [BOL!,

holo. (2 sheets); K!, iso. (2 sheets)].

Plants terrestrial. Rhizome short-decumbent, up to 12

mm diam., closely set with roots, persistent stipe bases

and scales; scales linear acuminate to narrowly lanceo-

late, up to 15 mm long, up to 5 mm wide, ferrugineous to

castaneous, chartaceous, broadly attached, irregularly

10 mm

FIGURE 1 — Dryopteris filipaleata

J.P.Roux. A, basal pinna; B,

abaxial view of pinnule. Drawn

from R.B. Faden et al. 70/596

(K) by J.P. Roux. Scale bars:

A, 10 mm; B, 5 mm.

Bothalia 34,1 (2004)

29

FIGURE 2 — Dryopteris filipaleata J.P.Roux: vestiture. A, stipe scale; Aa, section showing cellular structure; B, rachis scale; Bb, section showing

cellular structure; C. lamina scale, abaxial surface; Cc, section showing cellular structure; D, moniliform hair from abaxial surface of costa.

Drawn from R.B. Faden et al. 70/596 (K) by J.R Roux. Scale bars: A, B, 1 mm; C, 0.5 mm, Aa-Cc, D, 0.1 mm.

denticulate, irregularly set with scattered capitate glands,

and long, pluricellular. denticulate outgrowths, in larger

scales these outgrowths increase in number towards scale

apex; apex irregularly denticulate, terminating in a short

uniseriate series of cells. Fronds closely spaced, suberect

to arching, up to 1 130 mm long; stipe up to 610 mm

long, up to 7 mm diam.. proximally castaneous and adax-

ially flattened, stramineous higher up and shallowly sul-

cate, proximally densely scaled, sparsely scaled higher

up; scales up to 18 mm long, up to 6 mm wide, ferrugi-

neous, chartaceous, similar to those on the rhizome.

Lamina 2-pinnate-pinnatifid to 3-pinnate, ovate to broad-

ly ovate in outline, up to 590 mm long, up to 430 mm

wide, with up to 13 petiolated pinna pairs; rachis strami-

neous, sulcate adaxially, initially closely scaled, sparsely

scaled later; scales linear-acuminate to filiform, up to 4

mm long, up to 0.5 mm wide, ferrugineous to castaneous,

firmly herbaceous to thinly crustaceous, short-stalked,

cuneate to narrowly cuneate, irregularly denticulate,

apex terminating in a short series of oblong cells. Pinnae

1-pinnate-pinnatifid to 2-pinnate, near opposite to alter-

nate, proximally more widely spaced, overlapping or not,

basal pinnae petiolate, petiole up to 13 mm long, increas-

ingly more broadly attached and basiscopically decurrent

along rachis towards lamina apex, basal pinnae longest,

up to 270 mm long, up to 135 mm wide, basal pair most-

ly conspicuously basiscopically developed, inaequilater-

ally triangular, those higher up mostly near symmetrical.

ovate, oblong-acuminate to lanceolate, with up to 7

stalked pinna pairs; pinna-rachis sulcate adaxially, nar-

rowly winged towards apex, moderately scaled; scales

linear to filiform, up to 3 mm long, up to 0.3 mm wide,

ferrugineous to castaneous, firmly herbaceous to thinly

crustaceous, short-stalked, irregularly denticulate, apex

terminates in a short series of oblong cells. Pinnules

firmly herbaceous, pinnatifid to 1 -pinnate, near opposite

to alternate, spaced to slightly overlapping, basal pin-

nules petiolate; petiolule up to 3 mm long, increasingly

more broadly attached and basiscopically decurrent

along pinna-rachis towards pinna apex; pinnules narrow-

ly lanceolate to oblong-acuminate, up to 78 mm long, up

to 26 mm wide, proximal basiscopic pinnules slightly

basiscopically developed; costa sulcate adaxially, pro-

nounced abaxially, flexuose towards apex, narrowly

winged, sparsely scaled; scales filiform, up to 2.2 mm

long, up to 0.1 mm wide, irregularly denticulate, apex

terminates in a short series of oblong cells; segments and

lobes ovate-obtuse to oblong-obtuse, up to 15 mm long,

up to 7 mm wide, basiscopically decurrent, shallowly

lobed to denticulate, adaxially glabrous or with few hairs

and filiform scales along costa, abaxially sparsely set

with scattered, (4— )6(— 1 8 )-celled moniliform hairs up to

(78.0— )2 1 8.87(— 6 1 5.0) pm long, on and between the

veins. Venation evident, pinnately branched, mostly end-

ing in teeth near margin. Stomata mostly of the polocytic

type, (40.0-)50.1(-62.0) pm long. Sori predominantly 2-

30

Bothalia 34,1 (2004)

seriate along pinnules, 2-seriate on lobes in larger plants,

medial to inframedial on predominantly anadromous vein

branches, exindusiate. Sporangium', stalk simple, glandu-

lar or haired; capsule with ( 1 3—) 1 4(— 19) indurated annu-

lus cells, epistomium 4(-6)-celled, hypostomium (3— )6

(-7)-celled; spores monolete, ellipsoid, with low reticu-

late ridges and bulges, up to (32. 0-) 41.1 (—54.0) x (18.0-)

26.26(-34.0) pm. Figures 1, 2.

Distribution and ecology : Dryopteris filipaleata appears

to be restricted to the mountainous areas of tropical East

Africa occurring at altitudes ranging between 1 350 and

2 000 m. It grows in moist to wet evergreen forests either on

the forest floor or along streambanks with Cyathea manni-

ana. Piper capense, Ensete ventricosa , Symphonia spp.,

Melchiora schliebenii and Allanblackia ulugurensis.

Material examined

KENYA. — South Nyeri Dist. (K4), Kirinyaga Dist., Thiba Fishing

Camp, 31 July 1977, M.G. Gilbert & D. Rankin 4821 (K); Meru Dist.,

Jombeni Range, 1 520 m, H.D. van Someren 438 (K, 2 sheets); Meru,

upper forest, Aug. 1949, H.D. van Someren 493 (K); Kisumu-Londoni

Dist. (K5), Kisumu. bushiand, 2 128 m, Febr. 1915, R.A. Dimmer 1524

& 1727 (K).

TANZANIA. — Morogoro Dist. (T6), Uluguru Mountains, Moming-

side to Bondwa, 1 350-1 900 m, 3-4 July 1970, R.B. Evans et al.

70/351 (K, 2 sheets); Uluguru Mountains, Mwere Valley, 1 400-1 450

m, 26 Sept. 1970. R.B. Faden et al. 70/596 (BOF, 2 sheets, K, 2

sheets); Uluguru Forest Reserve, Lupanga Peak, 2 000 m, 1981, J.B.

Hall s.n. (K); Kanga Mountain, Northern Nguru, 1 800 m, 2 Dec. 1987,

J. Lovett & D.W. Thomas 2800. 2802 & 2802A (MO); Bagamoyo Dist.,

mainland west of Zanzibar, March 1885, J.T. Last s.n. (K).

ACKNOWLEDGEMENTS

My thanks to Dr Ted Oliver for checking the Latin

diagnoses, and the Royal Botanic Gardens, Kew and

Missouri Botanical Garden for making their material

available for study.

REFERENCES

BRENAN, J PM. 1978. Some aspects of the phytogeography of tropical

Africa. Annals of the Missouri Botanical Garden 65: 437^178.

FRASER-JENKINS, C.R. 1986. A classification of the genus Dryop-

teris (Pteridophyta: Dryopteridaceae). Bulletin of the British

Museum (Natural History), Botany 14; 183-218.

LOVETT, J.C. 1988. Endemism and affinities of the Tanzanian montane

forest flora. Monographs in Systematic Botany 25: 591-598.

PARRIS. B.S. 2001. Circum-Antarctic continental distribution patterns

in pteridophyte species. Brittonia 53: 270-283.

ROUX, J.P. 2002. A new species of Dryopteris (Dryopteridaceae;

Pteropsida) from East Africa. Kew Bulletin 57: 735-739.

TRYON, A.F. 1966. Origin of the fern flora of Tristan da Cunha.

British Fern Gazette 9: 269-276.

TRYON, R.M. 1986. The biogeography of species, with special refer-

ence to ferns. Botanical Review 52: 117-156.

WHITE, F. 1983. The vegetation of Africa. UNESCO, Paris.

J.P. ROUX*

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

7735 Claremont, Cape Town.

MS. received: 2003-07-03.

LAMIACEAE

PLECTRANTHUS MZ1MVUBUENSIS, A NEW SPECIES FROM EASTERN CAPE, SOUTH AFRICA

Plectranthus mzimvubuensis Van Jaarsv., sp.

nov., a P. reflexo verticillis brevibus 70-90 mm longis,

tubo corollae usque ad 10 mm longo, lateribus parallelis,

et staminibus tubum corollae aequantibus vel superan-

tibus, differt.

TYPE. — Eastern Cape, 3129 (Port St Johns): near

Ludonga Village, Mzimvubu River, Ecca Group shale

cliffs, (-AD), Van Jaarsvelcl, Xaba, Harrower & Zwide

92 (PRE, holo.).

Perennial, branched, aromatic shrub up to 1 m tall, 3 m

diam., scandent and pendent from cliffs. Roots fibrous to

slightly fleshy, but bearing distinct oblong to rounded

tubers; tubers 25-50 x 14—20 mm, grey, tissue translucent

and slightly yellowish. Stems herbaceous, semisucculent,

4-angled, terete in older branches and with a succulent

basal caudex, 100 mm diam. Bark smooth, grey. Leaves

thin-textured and drying chartaceous, broadly ovate-del-

toid to subrotund, ( 1 5— )25— 50(— 75) x (20-)28-50(-60)

mm, apex acuminate, with a short drip-tip, base truncate

to subcordate, occasionally slightly decurrent on petiole,

adaxial surface sparsely strigose becoming glabrescent,

abaxial surface prominent reticulate-veined, strigose,

becoming less so with age, covered with slightly sunken,

translucent gland dots (sessile glandular trichomes)

becoming yellowish brown in dried specimens, veins

densely strigose and with similar gland dots; margin ser-

rate-dentate with 6-10 pairs of teeth (0.5-)l-2(-4) mm

long, ciliate; petiole reddish purple, 1 0— 20(— 30) mm long,

finely strigose with unbranched, multicellular translucent

hairs, sparsely beset with gland-tipped trichomes.

Inflorescence short, terminal, verticillate, (30— )70— 90(— 120)

mm long, sometimes with a pair of side branches at base;

rachis sparsely strigose, bearing scattered, sessile, yel-

lowish brown gland dots and unbranched, multicellular,

glandular trichomes; bracts broadly ovate, acuminate, 7 x

4 mm. Flowers in sessile, 1-3-flowered cymes forming

2-6-flowered verticillasters, the latter 6— 1 2(— 1 8 ) mm

apart; pedicels 5-8 mm long, finely strigose, bearing few,

multicellular, gland-tipped trichomes. Calyx up to 4 mm

long, accrescent, lengthening to 10-11 mm in fruit,

densely covered with sessile, yellowish brown gland dots

at base, 2-lipped; upper lip erect, broadly ovate, abruptly

acuminate, ± 3 mm long; lower lip 4-toothed, teeth

acuminate; tube ± 8 mm long. Corolla pink; tube straight,

9- 1 0 mm long, laterally compressed, 3 mm wide, slight-

ly deflexed forming a swollen saccate base, sparsely beset

with translucent hairs, 0.2-3. 0 mm long, 2-lipped; upper

lip 4-lobed, 8 mm high, becoming reflexed when stigma

matures; upper lobes bent forward and forming an

ascending, spreading 2-spurred hood, ventral margin of

upper lobes overlapping two upper margins of lateral

lobes and each lobe forming a characteristic spur up to 2

Bothalia 34,1 (2004)

31

mm deep, apex of spur beset with translucent hairs,

0. 5-1.0 mm long; lower lip boat-shaped, 6 mm long,

soon becoming reflexed. Stamens 4, free, fused to tube ±

1 .5-2.0 mm from throat, didynamous, lower pair exposed

for 14-15 mm, upper pair exposed for 10-11 mm, both

pairs becoming reflexed; anthers versatile; pollen cream-

coloured. Style 14 mm long extending up to 20 mm when

mature, exposed for ± 8 mm. Nutlets rounded, 1 .5 x 1.3

mm, dark brown, smooth. Flowering time : autumn

(March and April). Figure 3.

Plectranthus mzimvubuensis is at once distinguished

from P. reflexus Van Jaarsv. & T.J. Edwards by its shorter

parallel-sided corolla tube which is 10 mm long, where-

as that of P. reflexus is longer, 25 mm, and constricted at

the mouth. P mzimvubuensis is a much-branched scram-

bler from a rootstock bearing distinct root tubers, where-

as P. reflexus is an erect shrub with fleshy roots. In both

species the lips and stamens become reflexed and the

mature style is twice the length of the corolla tube.

According to Codd’s (1985) key in his treatment for the

Flora of southern Africa (28,4: 141), the new species

would key out to ‘28’ which includes five species, P

ambiguus, P. ecklonii, P. dolomiticus , P petiolaris and P.

laxiflorus. Of these, the corolla of P. petiolaris, P. laxi-

florus and P. dolomiticus are curved like a ‘Dutchman’s

pipe’. P. mzimvubuensis can be distinguished from P.

ecklonii and P ambiguus by its short corolla of 9-10 mm

with the lobes of the upper lips hooded, their lower mar-

gins overlapping the lateral lobes and forming two short

spurs, each 2 mm long and ending in a translucent hair,

0. 5-2.0 mm long. The spurred upper lobes of the corolla

32

Bothalia 34.1 (2004)

are a unique feature in the genus Plectranthus, possibly

assisting flying insects in effective pollination. The

corolla tube of P. ecklonii is 12-18 mm long and slightly

expanding to the throat, whereas the corolla tube of P.

ambiguus is 20-25 mm long.

Plectranthus mzimvubuehsis is a scrambling shrub

with pendent branches. It is endemic to south-facing

Ecca Group shale cliff faces (Karoo Supergroup)

along the Mzimvubu River, upstream from Port St

Johns in Eastern Cape (Figure 4). P. mzimvubuensis

was first collected during a rubber-boat expedition to

study the cremnophilous flora of the Mzimvubu

River Valley in the autumn of 2002. The Mzimvubu

River, after which this species is named, is the

largest river in the Transkei portion of Eastern Cape.

The Xhosa name Mmzimvubu means 'the home of the

hippopotamus’ ( Hippopotamus amphibius), but these

animals were wiped out in the area more than a cen-

tury ago. The new species was encountered below

Ludonga Village, about 40 km from the river mouth,

at an altitude of ± 600 m. The vegetation consists of

savanna and the rainfall occurs mainly from spring to

autumn. 800-1 000 mm per annum. The climate is

subtropical, with hot summers, dry, sunny, frost-free

winters and cool evenings. The population of P.

mzimvubuensis was encountered on a south-facing

cliff in light shade, together with other succulent

plants such as Adromischus cristatus , Bulbine natal-

ensis, Crassula cordata , C. cultrata, C. multicava

subsp .floribunda, C. orbicularis , Cyanotis speciosus,

Delosperma tradescantioides and Peperomia blanda.

Trees and shrubs in the area include Bauhinia bowk-

eri, Celtis africana , Ficus burkei and Euphorbia tiru-

calli. Cuttings taken from P. mzimvubuensis rooted

rapidly and are grown at Kirstenbosch National

Botanical Garden.

Although only a small population of the new species

was found, no threats seem to exist on or near the cliff

face and the species is expected to be more common in

similar habitats elsewhere in the river valley. Its present

conservation status is classified as Rare, but not threat-

ened.

ACKNOWLEDGEMENTS

We thank Gerrit Germishuizen and Emsie du Plessis

for editing the text and Dr Hugh Glen for translating the

diagnosis into Latin. The Eastern Cape Department of

Tourism & Nature Conservation in Umtata is thanked for

providing plant-collecting permits. The authors also

thank colleagues Phakamani Xaba and Adam Harrower

for their help on the expedition and Godfrey Zwide of the

Eastern Cape Department of Tourism & Nature Con-

servation in Umtata for accompanying the authors and

providing assistance on this expedition.

REFERENCE

CODD, L.E. 1985. Lamiaceae. In O.A. Leistner, Flora of southern

Africa 28,4: 137-172.

E.J. VAN JAARSVELD*f and A.E. VAN WYK**

* National Botanical Institute, Kirstenbosch, Private Bag X7, 7735

Claremont.

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

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

of Pretoria, 0002 Pretoria.

MS. received: 2003-06-17.

PTERIDOPHYTA

DRYOPTERIS GORGONEA (PTEROPSIDA: DRYOPTERIDACEAE), A NEW SPECIES FROM THE CAPE VERDE ISLANDS

INTRODUCTION

The Cape Verde is a group of 10 major islands and sev-

eral smaller islands situated in the Atlantic Ocean

(15-17°N, 23-25°W), ± 620 km west of the coast of

Mauritania. Nine of the islands, which are of volcanic ori-

gin, are inhabited. The Barvalento or Windward Islands

consist of Santo Antao, Sao Vicente, Santa Luzia, Ilheu

Branco, Ilheu Raso, Sao Nicolau, Sal and Boa Vista,

whereas the Sotavento or Leeward group consists of Maio,

Santiago, Fogo and Brava. Santiago, the main island, is

mountainous and like most islands it is arid. Fogo has the

highest peak within Ihe island group with Mt Fogo reach-

ing 2 840 m. Santo Antao, the most northern island has the

highest rainfall. The climate is oceanic with daily highs

ranging between 20-29°C from August to October.

The island group has a depauperate flora as a result of

its arid climate. To date, 35 pteridophyte species have

been recorded for the Cape Verde Islands (Lobin et al.

1998). Dryopteris collections from the islands have been

ascribed to various taxa, of which a summary is provid-

ed by Lobin et al. (1998). Fraser-Jenkins (1982) was the

first to show that two Dryopteris species occur on the

islands, ascribing them to D. oligodonta (Desv.) Pic.Serm.

and D. pentheri (Krasser) C.Chr.

Bothalia 34,1 (2004)

33

During the review of the African Dryopteris species

currently undertaken by me, I had the opportunity to

study most of the relevant types. I therefore can confirm

that D. oligodonta is indeed one of the species occurring

in the Cape Verde Islands. The other species, however,

ascribed to D. pentheri by Fraser- Jenkins (1982), and

embraced by Lobin et al. (1998) is not that species, but

represents a distinct entity. Furthermore, several collec-

tions belonging to this taxon have erroneously been

ascribed to D. oligodonta by Lobin et al. ( 1998).

The earliest available name for this taxon appears to

be Dryopteris elongata (Aiton) Sim var. simplex A.Chev.

However, in describing the variety, three collections

were cited of which Chevalier 45476 is taken as the type,

as he states ‘Covao, 900 m alt. avec le type 45476,’

whereas the other two collections are considered as

paratypes (Chevalier 1935). In spite of Chevalier clearly

having designated his number 45476 as the holotype, it

was lectotypified by Lobin et al. (1998) with the same

collection. This collection, however, is D. oligodonta

whereas Chevalier 45113 and 45499 are not. Dryopteris

elongata var. simplex therefore becomes a synonym of

D. oligodonta. Since no name exists for Chevalier 45113

and Chevalier 45499 , it is here described as D. gorgonea ,

an epithet derived from the Gorgades, an old geographi-

cal name for the island group.

Dryopteris gorgonea J.P.Roux, sp. nov., a D. oligo-

donta stomatibus bene majoribus glandibus laminae differt.

TYPE. — Cape Verde Islands, Fogo, Espia. Mosteiros,

1-08-1934, A. Chevalier 45113 (P!, holo.; COI!, K!, iso.).

Plants terrestrial. Rhizome up to 50 mm long, up to 5

mm in diameter, suberect to erect, closely set with roots,

crowded stipe bases and scales. Fronds up to 1 020 mm

long, suberect to arching; stipe up to 640 mm long, up to

9 mm in diameter, proximally castaneous, adaxially flat-

tened and densely set with glands and pluricellular den-

droid hairs and scales; larger stipe scales up to 23 mm

long, up to 3 mm wide, concolorous or bicolorous, if

bicolorous then centrally castaneous to ferrugineous with

a narrow stamineous margin, firmly herbaceous, broadly

attached, linear-attenuate, margin variously set with long,

reflexed, pluricellular, mostly umseriate hairs and capi-

tate glands; capitate glands also occur on the scale sur-

face; apex flagelliform; stipe higher up stramineous,

shallowly sulcate and sparsely scaled, larger scales up to

5 mm long, up to 4 mm wide, fugaceous, concolorous,

ferrugineous to stramineous, chartaceous, broadly

attached, lanceolate to broadly ovate, variously set with

few short or long marginal outgrowths and glands, sur-

face variously set with capitate glands, apex flagelliform.

Lamina herbaceous, ovate to broadly ovate, up to 650

mm long, up to 31 mm wide, anadromous, catadromous

towards apex, up to 2-pinnate-pinnatifid, with up to 13

petiolated pinna pairs; rachis adaxially shallowly sulcate,

narrowly winged towards apex, variously set with

clavate glands and scales; scales up to 5 mm long, up to

4 mm wide, fugaceous, stramineous to ferrugineous,

chartaceous, broadly attached, margins variously set with

a few short and/or long outgrowths and glands, surface

variously set with capitate glands; apex flagelliform.

Pinnae near opposite to alternate, slightly overlapping or

not, up to I -pinnate-pinnatifid, basal pair mostly longest,

not conspicuously basiscopically developed, ovate,

lanceolate, or oblong-acute distally, up to 193 mm long,

up to 85 mm wide, with up to 6 petiolated pinnule pairs;

petiole up to 8 mm long; pinna-rachis shallowly sulcate

adaxially, narrowly winged for most of the length, close-

ly set with clavate glands, sparsely scaled; scales lanceo-

late to broadly ovate, up to 3.5 mm long, up to 1 mm

wide, ferrugineous to stramineous, chartaceous, sessile,

often somewhat bullate, variously set with short or long,

mostly uniseriate hairs and capitate glands, few capitate

glands also occur on scale surface, apex terminates in a

short or long uniseriate series of oblong cells. Pinnules

near opposite to alternate, slightly overlapping or not,

pinnatifid. ovate to oblong-obtuse, up to 48 mm long, up

to 24 mm wide, adaxially glandular along and between

veins, also with scattered isocytic hairs along costa,

abaxially closely set with clavate glands (50— )68.48(— 98)

pm long and isocytic hairs along veins; hairs often glan-

dular near base; costa adaxially shallowly sulcate, glan-

dular, abaxially variously scaled; scales up to 3 mm long,

up to 1 mm wide, ferrugineous to stramineous, charta-

ceous, sessile to short-stalked, often somewhat bullate,

similar to those on pinna-rachis; petiolule up to 2 mm

long; segments oblong-obtuse, up to 13 mm long, up to

6 mm wide, lobed; lobes dentate, teeth cuneate. Venation

anadromous, catadromous towards apex, pinnately

branched, branches end in teeth near margin. Stomata

mostly of polocytic type, (42— )53.3 1 (—64) pm long. Sori

circular, medial on predominantly anadromous vein

branches, 2-seriate on segments, discrete or touching at

maturity, up to 1.2 mm diam.; indusium brown, firmly

herbaceous, up to 0.5 mm diam., flabellate, entire to

repand, glandular along margin and on surface.

Sporangium: stalk simple, glandular, or haired; capsule

with ( 10—) 1 3(— 1 6) indurated annulus cells, epistomium

4(or 5)-celled, hypostomium 4(or 5)-celled; s pores brown,

ellipsoidal, monolete, perispore folded to form narrow

reticulate ridges, (38-)42.44(-46) x (24-)27.86(-32) pm.

Figure 5A-I.

Diagnostic features and relationships

Dryopteris gorgonea differs from D. pentheri , to

which it was formerly ascribed, in the scales often bear-

ing capitate glands on the surface and the presence of

clavate glands (50-)68.48(-98) pm long along the frond

axes and on the lamina surfaces. In D. pentheri the

glands are oblong, (60—) 1 37.02(— 260) pm long, and

restricted to the frond axes and veins. Also the 2-celled

hairs diagnostic of D. pentheri , have not been recorded

for D. gorgonea. Dryopteris gorgonea is closely related

to D. oligodonta , but differs in the longer clavate glands

[(36— )5 1.31 (—60) vs. (50-)68.48(-98) pm] and larger

stomata [(30— )37.54( — 48) vs. (42— )53.3 1 (—64) pm]. The

larger stomata suggest it being tetraploid, rather than

diploid, as is D. oligodonta (Fraser-Je.nkins 1982).

Distribution and ecology

Dryopteris gorgonea appears to be endemic to the Cape

Verde Islands having been recorded from Santo Antao, Sao

Vicente and the summit of Monte Gordo on Sao Nicolau.

Unfortunately, no ecological information has been record-

34

Bothalia 34,1 (2004)

FIGURE 5. A- I: Dryopteris gorgonea. A, abaxial view of fertile pinnule; B, stipe scale; Bb, section of B showing cellular structure; C, D, rachis

scales; Cc, section of C showing cellular structure; E, secondary rachis scale; Ee, section of E showing cellular structure; F, scale fiom

abaxial surface of costa; Ff, section of F showing cellular structure; G, hair from abaxial surface of lamina; H, clavate gland from abaxial

surface of lamina; I, indusium. Scale bars: A, 10 mm; B, C, E, 1 mm; D. F, I, 0.5 mm; Bb, Cc, Ee, Ff, G, H, 0.1 mm. Drawn from Chevalier

45113 (P) by J.P. Roux.

Bothalia 34, 1 (2004)

35

ed. No recent collections of the species appear to have been

made from any of the islands and it has been suggested to

be extinct (Lobin & Ormonde 1996; Lobin et al. 1998).

Since the species closely resembles D. oligodontci, a care-

ful search for it on the islands should continue.

Other material examined

CAPE VERDE ISLANDS. — Sao Nicolau: in summo monte Gurdo,

1851, C. Bolle s.n. (COI. K); Monte Gordo, 24-02-1864, R.T. Lowe s.n.

(K); Forbes s.n. (K). Santo Antao: Covao, 09-1934. A. Chevalier 45499

(P); Covao, 23. 24-09-1934, A. Chevalier 45532 (COI. K, P). Sine loc.:

Cape Verdes, Cardosa 169 (K); Cap du Vert, Forbes s.n. (K).

ACKNOWLEDGEMENTS

My thanks to Ted Oliver for the Latin diagnosis and

the collection managers of the various herbaria for allow-

ing me access to their holdings.

REFERENCES

CHEVALIER, A. 1935. Les lies du Cap Vert, geographie, biogeogra-

phie, agriculture. Flore de l’Archipel. Revue de Botanique

Appliquee et d' Agriculture Tropicale 15(170-171): 1-336.

FRASER-JENKINS, C.F. 1982. Dryopteris in Spain. Portugal and

Macaronesia. Boletim da Sociedade Broteriana , ser. 2, 55: 175 —

336.

LOBIN, W„ FISCHER, E. & ORMONDE, J. 1998. The ferns and fern-

allies (Pteridophyta) of the Cape Verde Islands, West Africa.

Nova Hedwigia. Beiliefte 115: 1-115.

LOBIN. W. & ORMUNDE. J. 1996. Lista vermelha para os Pterido-

fitos (Pteridophyta). In T. Leyens & W. Lobin, Primeira lista

vermelha do Cabo Verde. Courier Forschungsinstitut Sencken-

berg 193: 37^12.

J.P. ROUX*

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

7735 Claremont, Cape Town.

MS. received: 2003-11-14.

CRASSULACEAE

ADROMISCHUS SCHULDTIANUS SUBSP. BRANDBERGENSIS. A NEW SUBSPECIES AND A CHECKLIST OF THE SUCCULENT

FLORA OF THE BRANDBERG, NAMIBIA

INTRODUCTION

Adromischus schuldtianus (Poelln.) Poelln. subsp.

brandbergensis B.Nord. & Van Jaarsv., a new subspecies

endemic to the Brandberg. northwestern Namibia, is

described. Since its discovery by H ,J. Wiss below Aigub

Peak (Nordenstam 1974), by the second author on

Konigstein and Orabeskop, and later by P.V. Bruyns

(Craven & Craven 2000). this taxon, due to its distinct

leaf characters and isolated distribution, demanded some

form of taxonomic recognition. The new subspecies is at

once distinguished from A. schuldtianus (Poelln.) Poelln.

subsp. schuldtianus by its subfusiform-ellipsoid and

semiterete leaves without a clear margin; they are usual-

ly shallowly concave above, becoming channelled dur-

ing the dry season. Both A. schuldtianus subsp. schuld-

tianus and subsp. juttae have dorsiventrally flattened,

oblanceolate (rarely obovate) leaves.

Adromischus schuldtianus (Poelln.) Poelln.

subsp. brandbergensis B.Nord. & Van Jaarsv ., subsp.

nov.. differ! a A. schuldtiano (Poelln.) Poelln. subsp.

schuldtianus foliis subfusiformibus-ellipsoideis plusmi-

nusve semiteretibus 20-90 mm longis 10-15 mm crassis

supra applanatis vel leviter concavis sine margine dis-

tincto apice acutis-obtusis saepe recurvatis.

TYPE. — Namibia, 2114 (Uis): Orabeskop, 2 300 m,

Brandberg Mountain. (-AA), 06-04-1964, Nordenstam

3677 (ex hort. Bot. Garden, Lund, specimen in WIND,

holo.!; M!, S!, iso.).

Dwarf mat-forming, branched succulent up to 70 mm

high, filling crevices in granite rock fissures. Roots

fibrous. Branches short and succulent, in cultivation up

to 70 x 10 mm. Leaves alternate, spreading, subfusiform-

ellipsoid to ± semiterete, 20-70 x 10-15 mm, without a

distinct margin, flattened to shallowly concave above,

tapering to base and acute-obtuse and often somewhat

recurved at tip, dark green and marbled with white or

dull red areas. Inflorescence a ± one-sided raceme with

2-15 almost patent flowers; peduncle 150-500 mm long

(in cultivation), 1.2-1. 5 mm thick, simple or branching

above middle, terete, glabrous, greenish brown or red-

dish; bracts 1.5 mm long, acute, succulent; bracteoles 2,

basal, subulate, ± 1 mm long, acute; pedicels 5-17 mm

long, somewhat thickened towards apex. Calyx : lobes

narrowly triangular, 1. 8-2.0 x 0. 6-1.0 mm, acute.

Corolla 12-15 mm long, pinkish white or wax-coloured;

tube cylindrical, 2. 5-3.0 mm wide; lobes patent, deltoid,

acute, with somewhat wavy margins; throat bright purple

inside. Styles subulate-filiform, 5-8 mm long, white at

apex, pale green at base. Squamae oblong, bifid, 1 x 0.8

mm, white. Stamens : filaments white or pinkish, five

longer ones adnate for 5 mm, ± 12 mm long, five short-

er ones adnate for 3 mm, ± 10 mm long; anthers oblong,

0.4 mm long. Figure 6.

Adromischus schuldtianus subsp. brandbergensis

appears to be endemic to the Brandberg Mountain in

northwestern Namibia (Figure 7). It occurs in rock

crevices of steep southern and eastern slopes and cliffs of

the mountain from ± I 500 m to ± 2 300 m. It is not com-

mon anywhere but occurs scattered in protected fissures

and crevices in small to dense groups due to vegetative

proliferation. Associated plants in the same habitat

include Aloe dichotoma, A. hereroensis, A. litt oralis,

Cyphostemma currorii and Kalanchoe lanceolata , also

Diospyros acocksii , Euphorbia maitritanica , Ficus ilici-

na, Obetia carruthersiana, Salvia garipensis and

Tetradenia riparia.

In his 1985 revision of the genus Adromischus , Toelken

recognized 27 taxa. A very handy popular version of the

same account was published by Pilbeam et al. in 1998,

36

Bothalia 34,1 (2004)

FIGURE 6. — A. schuldtianus subsp. brandbergensis B.Nord. & Van

Jaarsv. Illustration based on a cultivated plant that flowered in

the Botanical Garden, Lund. Sweden: A, flowering branch, x 1;

B. flower, x 2.5; C, corolla, opened to show styles and stamens,

x 2.5; D, corolla, dorsal view, x 2.5; E, F, leaf outlines in side

view and transect, x 1. Reprinted, with permission, from Nor-

denstam. Drawn by Bertil Nordenstam,

FIGURE 7. — Known distribution of Adromisclius schuldtianus subsp.

brandbergensis.

reflecting the horticultural value of the group. Adro-

mischus schuldtianus subsp. brandbergensis belongs to

section Boreali (Toelken 1978) which includes a few other

taxa confined to the northern parts of South Africa and

Namibia, for example, A. schuldtianus subsp. schuld-

tianus, A. trigynus and A. umbraticola. From these the new

subspecies is at once distinguished by its subfusiform or

almost terete leaves distinctly concave on the upper side.

It is found the furthest north of any Adromischus species.

Of all the members of section Boreali, A. trigynus has

the most southern distribution. It is confined to dolerite

outcrops, growing in shallow soil at altitudes above 1 000

m in the Nama-Karoo Biome in an area that receives pre-

dominantly summer and autumn rain. A. trigynus ranges

from southern Namibia and Pofadder in the west to

Aliwal North and the southern Free State in the east.

Adromischus umbraticola occurs on south-facing

cliffs and in the shallow soil of sandstone and quartzite

outcrops on the Highveld of Gauteng, mountains of the

North-West and further north to the Blouberg and

Chuniespoort (Limpopo). It is common on rocky ridges

of the Witwatersrand and in the Magaliesberg range. The

vegetation of its habitat consists mainly of dry, short

savanna. It has very brittle leaves, and plants often colo-

nize shallow pockets of soil where there is little compe-

tition from mesophytic taxa.

Toelken (1985) recognizes two subspecies of A.

schuldtianus, namely subsp. schuldtianus and subsp. jut-

tae, characterized by their oblanceolate to obovate leaves

but mainly differentiated by their stem and branch

length, 40-80 mm tall, and little branched in subsp. jutta

as opposed to branches 10-30 mm long in subsp. schuld-

tianus, which occurs in arid savanna in central Namibia,

from the Erongo and Auas Mountains in the north to near

Aus Village and the Karas Mountains in the south. It

grows on rock outcrops, usually with a southern aspect.

The second subspecies, A. schuldtianus subsp. juttae is

confined to the Karasberg in southern Namibia and is

differentiated by its longer branches; the plants occur in

the Nama-Karoo. Bruyns (1990) noticed variability in

the leaf shape of A. schuldtianus subsp. schuldtianus on

the Brandberg. He found flat- and fusiform-leaved plants

occurring together. However, in spite of some local vari-

ation, the majority of specimens encountered on the

Brandberg are represented by the subterete-leaved plants

here described as a distinct subspecies.

The Brandberg is an isolated granite inselberg of ± 21

x 25 km and Konigstein (2 573 m) represents the highest

peak in Namibia. It is surrounded by Namib Desert with

typical species such as Welwitschia mirabilis, annual

herbs and grasses, and foothills with woody species

including Acacia montis-usti, Adenolobus garipensis.

Commiphora saxicola, C. virgata, C. wildii and Moringa

ovalifolia. The rainfall on the lower slopes is low (less

then 100 mm per annum) and typical of the Namib.

A total of 480 species was recorded from the moun-

tain by Craven & Craven (2000). The vegetation of the

Brandberg is slowly transformed with altitude (increase

in rainfall, decrease in temperature). At 2 000 m and

above it is reminiscent of renosterveld, not unlike the

Bothalia 34,1 (2004)

37

vegetation found in the winter rainfall Kamiesberg of

Namaqualand, and many genera are shared between the

two regions. The Brandberg itself has a number of

endemics (Nordenstam 1974; Craven & Craven 2000):

Felicia gunillae, Hermannia merxmuelleri , Lithops gra-

cilidelineata subsp. brandbergensis , Nidorella norden-

stamii, Pentzia tomentosa. Plumbago wissii , Ruellia

brandbergensis and the recently described Philyro-

phyllum brandbergense (Herman 2003). Our new Adro-

mischus subspecies brings the total of endemic taxa to

nine. A number of other taxa are near-endemic to the

Brandberg, being known from one or a few localities out-

side the Brandberg. The Brandberg is treated by Van

Wyk & Smith (2001 ) as a local focus of endemism with-

in a larger Kaokoveld Centre of plant endemism.

Other material examined

NAMIBIA. — 2114 (Uis): Brandberg. Konigstein, E slopes, 2 400

m, 31-05-1963. Nordenstam 2837 (cult, in Bot. Garden, Lund, speci-

men in S); Bruyns 3302 (BOL); Van Jaarsveld 17969 (NBG).

ACKNOWLEDGEMENTS

We are grateful to Herta Kolberg from the Ministry of

Environment and Tourism in Nambia for her assistance.

REFERENCES

BRUYNS, P. 1990. New plant records from the Brandberg. Cimbebasia

12: 161-166.

CRAVEN. P. & CRAVEN, D. 2000. The flora of the Brandberg, Namibia.

In A.H. Kirk-Spriggs & E. Marais, Daures — biodiversity of the

Brandberg Massif, Namibia. Cimbebasia Memoir 9: 49-67 .

National Museum of Namibia, Windhoek.

HERMAN, P.P.J. 2003. A new species of PhilyrophyUum (Asteraceae:

Gnaphalieae) from Namibia. Bothalia 33: 118-120.

NORDENSTAM, B. 1974. The flora of the Brandberg. Dinteria 11:1-

67.

PILBEAM. I. RODGERSON, C. & TRIBBLE, D. 1998. Adromischus.

The Cactus File Handbook 3. Cirio Publishing Services,

Southampton.

SMITH. G.F., VAN JAARSVELD, E.J., ARNOLD, T.H., STEFFENS,

F.E.. DIXON, R.D. & RETIEF. J.A. (eds). 1997. List of south-

ern African succulent plants. Umdaus Press, Hatfield.

TOELKEN, H.R. 1978. New taxa and new combinations in Cotyledon

and allied genera. Bothalia 12: 377-393.

TOELKEN. H.R. 1985. Crassulaceae. Flora of southern Africa 14: 1-

244.

VAN WYK. A.E. & SMITH, G.F. 2001. Regions offloristic endemism

in southern Africa: a review with emphasis on succulents.

Umdaus Press, Hatfield.

LIST OF SUCCULENTS RECORDED FROM THE BRANDBERG

To date the most detailed published checklist for the

flora of the Brandberg is that by Craven & Craven

(2000). In the latter list the authors indicate the growth

form for each species/infraspecific entry. Although 'suc-

culent' is one of the growth forms provided for, this state

was applied very inconsistently. For example, no Aloe is

marked as a succulent, whereas this is the prevailing

state in the genus. Hence we here provide an updated list

of those species with a succulent growth form recorded

from the Brandberg. It is hoped that this list will prove

useful to workers interested in comparing the Brandberg

flora with other areas where the proportion of succulent

taxa may differ (e.g. the high Kamiesberg in the Suc-

culent Karoo further south).

Our list was compiled from unpublished field notes

kept by the authors, as well as from a critical assessment

of the published species lists of Nordenstam (1974) and

Craven & Craven (2000). Included in the list are 54

species belonging to 1 7 families, alphabetically arranged

according to family, then according to genus and species.

We have adopted the definition for succulence provided

by Smith et al. (1997), namely that a succulent is a plant

that stores water in its tissues as a mechanism to survive

periods of drought in the growing phase.

Aizoaceae

Sesuvium sesuvioides (Fenzl) Verde, var. sesuvioides

Tetragonia

arbuscula Fenzl

calycina Fenzl

Apocynaceae/Asclepiadaceae

Hoodia gordonii ( Masson ) Sweet ex Decne.

Orbea maculata (N.E. Br.) L.C. Leach subsp. rangeana (Dinter &

A. Berger) Bruyns

Sarcostemma viminale (L). R.Br. subsp. viminale

Stapelia

kwebensis N.E.Br.

longipedicellata (A. Berger) L.C. Leach

Asphodelaceae

Aloe

asperifolia A. Berger (flats surrounding mountain)

dichotoma Masson var. dichotoma

hereroensis Engl. var. hereroensis

littoralis Baker

viridiflora Reynolds

Asteraceae

Othonna brandbergensis B.Nord.

Kleinia longiflora DC.

Burseraceae

Commiphora

glaucescens Engl.

kraeuseliana Heine

pyracanthoides Engl.

saxicola Engl.

tenuipetiolata Engl.

virgata Engl.

wildii Merxm.

Crassulaceae

Adromischus schultdtianus (Poelln.)Poelln. subsp. brandbergensis

B. Nord. & Van Jaarsv.

Cotyledon orbiculata L. var. orbiculata

Crassula

subaphylla [Eckl. & Zeyh.) Harv. subsp. subaphylla

tabularis Dinter

Kalanchoe lanceolata (Forssk.) Pers.

Cucurbitaceae

Corallocarpus

schinzii Cogn.

welwitschii (Naudin) Hook.f.

Euphorbiaceae

Euphorbia

gariepina Boiss. subsp. balsamea (Hiern) L.C. Leach

guerichiana Pax

mauritanica L. var. mauritanica

monteiroi Hook.f. subsp. brandbergensis B.Nord.

virosa Willd. subsp. virosa

Lamiaceae

Aeollanthus neglectus (Dinter) Launert

Tetradenia riparia (Hochst.) Codd

Mesembryanthemaceae

Aptenia geniculiflora (L.) Bittrich ex Gerbaulet

Hereroa puttkameriana (Dinter & A. Berger) Dinter & Schwantes

Lithops gracilidelineata Dinter

subsp. brandbergensis (de Boer) D.T.Cole

subsp. gracilidelineata

Mesembryamhemum guerichianum Pax

Moringaceae

Moringa ovalifolia Dinter & A. Berger

38

Bothalia 34,1 (2004)

Passifloraceae

Adenia

pechuelii (Engl.) Harms

repanda (Burch.) Engl.

Portulacaceae

Avonia albissima (Marloth) G.D. Rowley

Portulaca

kermesina N.E.Br.

*oleracea L.

Talinum sp.

Sterculiaceae

Sterculia

africana (Lour.) Fiori var. africana

quinqueloba (Garcke) K.Schum.

Yitaceae

Cyphostemma

bainesii (Hook.f.) Desc.

congestum (Baker) Desc. ex Willd & R.B.Drumm.

currorii (Hook.f. ) Desc.

Welwitschiaceae

Welwitschia mirabilis Hook.f.

Zygophyllaceae

Zygophyllum simplex L.

E.J. VAN JAARSVELD*t, B. NORDENSTAM**

and A.E. VAN WYK***

* National Botanical Institute, Kirstenbosch, Private Bag X7, 7735

Claremont.

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

0002 Pretoria.

** Department of Phanerogamic Botany, Swedish Museum of Natural

History, PO. Box 50007, SE-104 05 Stockholm, Sweden.

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

University of Pretoria, 0002 Pretoria.

MS. received: 2003-08-19.

ERICACEAE

NOMENCLATURAL CHANGES IN ERICA

Erica salicina E.G.H.Oliv., nom. nov., E. viminalis

E.G.H.Oliv. in Bothalia 31: 5 (2001) non E. viminalis

Salisb.: 298 (1796) [= E. albens L.]. Type: Western Cape,

3319 (Worcester): Hex River Mountains, Milner Peak, on

wet shaded cliffs at base of amphitheatre, E side, 5000 ft

fl 520 m], (-AD), 4 January 1959, Esterhuysen 28100

(BOL, holo.; BM, E, G, K, L, MO, NBG, NY, P, PRE, S, W).

Salisbury’s name (1796) was overlooked when nam-

ing this very localized species with its distinctive, wil-

lowy habit. The new name repeats the etymological ref-

erence to the habit, salicinus - like the genus Salix [ vim-

inalis = willowy]. At the time of description only the

type material was known. Recently the species was re-

collected in the very impressive amphitheatre of the Hex

River Mountains by Nicholas Helme who noted that

there were only two adult plants that had escaped the fire

of three years previously with few signs of any young

plants. The habitat was a permanently wet drip zone

which probably receives direct sunlight for only two

weeks in the year.

Erica pilaarkopensis H. A. Baker in Journal of South

African Botany 41: 128 (1975). Type: 3419 (Caledon):

Pillarkop [Pilaarkop], Riviersondereinde Mtns, (-BB),

near Lindeshof on scree and steep rock slopes, alt. 1 220-

I 524 m, Esterhuysen 33343 (BOL, holo.; NBG).

Baker (1975) gave the specific epithet pillarkopensis

to this species based on the locality of the type collec-

tion, which he cited as ‘Pillarkop’. However the details

given by Esterhuysen in her own hand on the holotype,

refer to the location as ‘Pilaarkop’ which is the spelling

noted on all reference maps. The label details as copied

by Baker onto the isotype in his own herbarium (now in

NBG) give the locality as ‘Pillarkop’ and the specific

epithet as pillarskopensis [sic]. It could be suggested that

Baker deliberately translated the Afrikaans pilaar to the

English pillar. If this is the case then why did he retain

the Afrikaans kop for peak. I contend that being an immi-

grant Englishman unfamiliar with the language, he mis-

copied the name in his manuscript and on the label in

error. I therefore correct the spelling of the specific epi-

thet to reflect the accepted Afrikaans spelling of the

name of the locality as permitted by Article 60. 1 of the

International Code (Greuter et al. 2000).

REFERENCES

BAKER, H.A. 1975. Notes on Erica in South Africa. Journal of South

African Botany 41:1 25-130.

GREUTER, W ., MCNEILL, I, BARRIE, F.R.. BURDET, H.-M.,

DEMOULIN, V., FILGUEIRA, T.S., NICOLSON, D.H.,

SILVA, P.S., SKOG, J.E., TREHANE, P„ TURLAND. N.J. &

HAWKSWORTH. D.L. (eds). 2000. International Code of

Botanical Nomenclature (Saint Louis Code) adopted by the

Sixteenth International Botanical Congress, St. Louis,

Missouri, July-August 1999. Regnum Vegetabile 138. Koeltz

Scientific Books, Konigstein.

OLIVER, E.G.H.& OLIVER, I.M. 2001. Four new species of Erica

(Ericaceae) from Western Cape, South Africa. Bothalia 31: 1-8.

SALISBURY, R.A. 1796. Prodromus stirpium in horto ad Chapel

Allerton vigentimn. London.

E.G.H. OLIVER*

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

7735 Claremont, Cape Town.

MS. received: 2003-08-08.

Bothalia 34,1 (2004)

39

ERICACEAE

A NEW SPECIES OF INDEHISCENT-FRUITED ERICA FROM THE CENTRAL KOUEBOKKEVELD,

WESTERN CAPE, SOUTH AFRICA

INTRODUCTION

The genus Erica L. in southern Africa has recently

been enlarged by the reduction to synonymy of the 18

small, indehiscent-fruited Cape genera, or ‘Minors'

(Oliver 2000). The inclusion of the 84 species resulted in

a total of ± 765 species for the genus. Since then, one

additional indehiscent-fruited species, E. rusticula

E.G.H. Oliv., has been described and this paper adds

another, E. tragomontana.

Erica possesses either a dehiscent, indehiscent or par-

tially dehiscent fruit. Two dehiscence strategies are dis-

played within the partially dehiscent type: 1, partial,

active dehiscence when the drying out of the valves caus-

es an incomplete split along suture lines; and 2, partial,

passive dehiscence when valves do not open indepen-

dently and the swelling of the developing seed forces the

valves to open partially (Oliver 2000: 49, 50). In both

instances seeds are not released from the capsule. The

new species described here has a wholly indehiscent fruit,

no swelling or splitting of the ovary occurring during or

after ripening. Roughly 680 dehiscent-fruited species

occur throughout southern Africa, whereas the 86 inde-

hiscent and partially dehiscent-fruited species are con-

fined to the Cape Floral Kingdom between Niewoudtville

in the northwestern Cape and Port Elizabeth in Eastern

Cape (Oliver 2000). Marked centres of species richness

occur in the Kogelberg/Grabouw area, Riviersonderend

Mountains and Central Kouebokkeveld, 16-18, 15 and 13

species occurring per quarter degree grid (± 14 x 17 km)

respectively (Oliver 2000: 87-95). The discovery of E.

tragomontana therefore raises the number of indehiscent-

fruited species in the central Kouebokkeveld (grid refer-

ence: 3319AB) to 14.

Erica tragomontana R.C. Turner, sp. nov., in

grege specierum olim in generibus minoribus fructibus

indehiscentibus positorum. Ericae bokkeveldiae E.G.H.

Oliv. et Ericae rusticulae E.G.H. Oliv. & I. M. Oliv. affi-

nis, sed ab eis pilis ovarii densis longis lanatis apicalibus,

staminibus 5 vel 6 interdum 4 rare 7, sepalibus brac-

teaque appressis ovato-lanceolatis, foliis latioribus lon-

gioribus, habitu sparsiore erecto differt. Figura 8.

TYPE. — Western Cape, 3319 (Worcester): central Koue-

bokkeveld. Bokberg southwest of Winkelhaak Farm-

stead, southeast-facing slopes below trig, beacon 4,

1 257 m, (-AB), 9 October 2002, Turner 530, (NBG,

holo.; BM, K, NY, PRE).

Plants up to 350 mm tall, erect, single-stemmed reseed-

ers. Branches: several erect, main and numerous short.

FIGURE 8. — Erica tragomontana. A, flowering branch, natural size; B, stem; C, leaf, abaxial view (left), adaxial view (right); D, flower; E, bract;

F, bracteole; G, sepal, abaxial view; H, stamen, back, front and side views; I, gynoecium; J, 1/s ovary; K, testa. All drawn from type. Turner

530. Artist: Traci Polea. Scale bars: B-I, 2 mm; J, 1 mm; K, 100 pin.

40

Bothalia 34,1 (2004)

partially decurved secondary branches; stems, younger

with short stiff hairs, older subglabrous with occasional

short hairs; weak infrafoliar ridges. Leaves 3-nate, nar-

rowly ovate, 1 .0—2.4 x 1.2 mm, narrowly sulcate, abaxial-

ly glabrous, hairy on basal portions adaxially, reddish

green, margins hyaline and ciliate; petiole 0.25-0.4 mm

long, yellow. Inflorescence: flowers 1 to 3-nate, terminal

on short, secondary branchlets; pedicel 0.1 mm long, red,

with short, sparse hairs; bract partially recaulescent,

adpressed, leaf-like, ovate-lanceolate, 1.4—1. 6 mm long,

minutely sulcate, pinkish green, margins ciliate with ses-

sile glands; bracteoles 2, approximate, adpressed, longer

than bract, oblong to lanceolate, 1.6-1 .8 mm long, sub-

acute, keeled, minutely sulcate, pink, margins ciliate with

sessile glands, abaxially and adaxially glabrous. Calyx 4-

lobed; sepals adpressed, ovate-lanceolate, keeled, sub-

acute, 1.6 mm long, glabrous, pink, margins ciliate with

sessile glands. Corolla 4-lobed, broadly funnel-shaped,

2.75-3.3 mm long, glabrous, colliculate, pink, margins

erose, lobes spreading broadly apically, narrow basally.

Stamens (4) 5 or 6 (7), free, manifest; filaments linear with

a slight sigmoid bend, broadening basally, 1.6-1. 8 mm

long, glabrous, pink; anthers dorsally fixed at base, bipar-

tite, erect, scabrous, reddish brown; thecae 0.72-1.0 mm

long; appendages pendulous, dorsally fixed at bases of

thecae, 0.6 mm long, linear, reddish purple, erose,

scabrous, pores 0.2-0.36 mm long; pollen in monads.

Ovary 2-locular, ovoid, 0.8- 1.1 mm long, laterally flat-

tened, dense, long lanate hairs apically and halfway down

lobes, otherwise glabrous, reddish purple, 1 ovule per

locule, placenta apical, no basal nectaries; style truncate,

2. 6-3. 8 mm long, far exserted, glabrous, pink; stigma

minutely capitellate. Fruit indehiscent, ovoid, 0.8-1. 1 mm

long, verrucose, reddish brown with long lanate apical

hairs, otherwise glabrous; pericarp smooth, subcolliculate,

thin, brittle, brown; mesocarp thin, red; endocarp thin,

papery, pale cream. Seeds ellipsoid, ± 0.8 mm long; testa

yellowish brown, cells elongate with unevenly jigsawed

anticlinal walls and numerous small to medium-sized pits.

Flowering time: September to early November. Figure 8.

Diagnostic features and discussion

Superficially resembling E. bokkeveldia and E. rustic-

ula, E. tragomontana differs in having mostly 5 or 6,

sometimes 4 and rarely 7 stamens, with long, woolly,

apical hairs and halfway down the lobes of the ovary;

adpressed ovate-lanceolate sepals, bract and bracteoles;

narrowly ovate leaves of 1.0-2. 4 x 1.0-1. 2 mm; and a

sparser, erect habit. E. bokkeveldia has pendulous, 3-

nate, broadly funnel-shaped flowers with very short

pedicels, 0. 1-0.5 mm long, borne terminally on short,

secondary branches; 3-nate leaves; aculeate anthers with

broad, pendulous appendages; a 2-locular ovary with one

pendulous ovule per locule and an apical placenta; but

differs from E. tragomontana in having scarious, broad-

ly ovate sepals; oblanceolate bract and bracteoles; eight

stamens; and an apically and sparsely pubescent ovary.

The general habit is also compact and more rounded, the

species usually growing on drier, sandy, quartzitic flats in

low proteoid/restioid fynbos at the bases of the sur-

rounding mountains (R.C. Turner pers. obs.).

Erica rusticula, endemic to substrate derived from

quartzitic Witpoort Formations in the northern Koue-

bokkeveld, also has pink, broadly funnel-shaped flowers

with very short pedicels, 0.3 mm long, borne terminally

on short, partially decurved secondary branches; fila-

ments with an apical sigmoid bend, a far exserted, trun-

cate style with a minutely widened stigma; 3-nate leaves;

and a 2-locular indehiscent ovary with one pendulous

ovule per locule. This species differs from E. tragomon-

tana in having branches with short, dense, reflexed hairs;

slightly shorter, imbricate leaves, with entirely hairy

adaxial surfaces; flowers with only four stamens; fila-

ments that broaden apically; non-scabrous anthers with

very short, narrow, pendulous appendages; shorter,

straight, dense hairs covering the top 3/4 of the ovary sur-

face and a subapical placenta; as well as a compact,

woody and more rounded habit, especially in old plants

(Oliver & Oliver 2000; R.C.Turner pers. obs.).

However, E. tragomontana also displays several simi-

lar morphological characters to the dehiscent-fruited E.

argentea Klotzsch ex Benth., particularly the 3-nate

leaves; the branches with weak infrafoliar ridges and fine

hairs; the very short pedicel, 0.16-0.29 mm long, the 3-

nate flowers borne terminally on short, partially

decurved, secondary branchlets; the adpressed bract,

bracteoles and sepals; the filaments with a slight sigmoid

bend; the scabrous thecae with pendulous appendages;

an apically hairy ovary; and an erect, slightly lax habit.

This is not a unique instance, as character similarities

between the indehiscent-fruited E. cetrata E.G.H. Oliv.

and the dehiscent-capsuled E. macrotrema Guthrie &

Bolus var. glabripedicellata Dulfer have been noted

(Oliver 2000: 287). A closer inspection of E. tragomon-

tana reveals an indehiscent, laterally-flattened 2-locular

ovary with 1 ovule per locule and a lack of nectaries;

mostly 5 or 6, sometimes 4 and rarely 7 stamens; and a

broadly funnel-shaped corolla with spreading lobes. E.

argentea has a dehiscent, 4-locular ovary with occasion-

ally 1, usually 2-5 and rarely 7-9 ovules per locule, as

well as basal nectaries; 8 stamens; and a narrowly obcon-

ical to urceolate corolla.

Character adaptations related to the protection of the

fruit differ between the three indehiscent species. In E.

tragomontana the hardened, adpressed sepals continue to

clasp the base of the corolla tightly when dried flowers

fall to the ground, providing added protection for the

slightly sclerified fruit for a further period of time

(Turner 728: pers. obs.). The dried, hardened base of the

corolla tube protects the fruit of E. bokkeveldia after

flowering, whereas in E. rusticula a thicker pericarp and

seed testa perform this function (Oliver & Oliver 2000).

While E. tragomontana usually possesses a 2-locular

ovary with one ovule per locule (G 2/i), an exceptional

situation was observed during dissection of mature fruits

( Turner 728: pers. obs.), in which one locule contained

two developed seeds and the second contained a flat-

tened, unfertilized ovule.

As E. tragomontana possesses an indehiscent fruit,

pollen in monads and a G 2/i ovary, it would appear to be

allied with the major basal polychotomy of 62 taxa with-

in the indehiscent-fruited species (Oliver 2000). Due to

the lack of clear indications of infrageneric relationships

within the now greatly enlarged genus Erica , indehis-

cent-fruited species have been temporarily retained at the

Bothalia 34, 1 (2004)

41

end of the genus (Oliver 2000 in Oliver & Oliver 2000)

and this new species is therefore provisionally placed

after E. rusticula as species number M45.2.

Pollination syndrome

The pollination syndrome of E. tragomontana is unre-

solved. Although the species has no floral nectaries, only

a slight stigmatic enlargement (minutely capitellate) has

occurred, a similar condition also existing with E. rustic-

ula (Oliver 2000) and the question is thus posed as to

what exactly the pollination strategies of these species

are, neither being specifically adapted for entomophily

nor anemophily.

Thrips, common visitors to many Erica species, have

been observed under magnification in the anther pores of

fresh material of E. tragomontana ( Turner 530, 592\

pers. obs.) but these insects are considered too small to

be effective pollinators (Rebelo et al. 1987). The anthers

of E. tragomontana possess relatively large, broad and

pendulous appendages, suggesting some form of ento-

mophily, although no pollinators have been observed

during field study. Even though the flowers of E.

tragomontana have no nectaries and do not emit a

noticeable scent (R.C. Turner pers. obs.), it is very possi-

ble that larger flying insects, such as bees, visit the plant

for its pollen reward alone (Rebelo et al. 1987).

Distribution and habitat

Erica tragomontana is known only from two subpopu-

lations occurring in the central Kouebokkeveld on the

Bokberg and Sandberg (Figure 9), hence the specific epi-

thet tragomontana , referring to the type locality on the

Bokberg: tragus (Latin) = goat = bok (Afrikaans). Re-

markably, the relatively dry Bokberg complex consisting

of the Bokberg, Houdenbeksberg, Sandberg and Vaal-

bokskloofberg, supports at least 14 indehiscent and 38

dehiscent Erica species as well as three subspecies of taxa

occurring within the complex (R.C. Turner pers. obs.).

Upon initial discovery, only 15 plants were found in a

small winter seepage area but it is now known that these

plants represent the fringes of the Bokberg subpopulation

as specimens have since been found to thrive in far greater

numbers on drier, rocky, south- to southeast-facing slopes

in a sandy substrate derived from resistant Witpoort forma-

tions of the Witteberg Group (Spatial Data Management

Unit 1997). This quartzitic sandstone rock comprises both

the Bokberg and Sandberg. At the type locality the species

grows in association with E. eremioicles (MacOwan)

E.G.H.Oliv., E. incamata Thunb., E. melastoma Andrews

variant D (Oliver & Oliver 2002), E. nudiflora L., E.

recurvifolia E.G.H.Oliv., Aulax pallasia Stapf (Proteaceae),

Sorocephalus lanatus Thunb. (Proteaceae) and several

species of Restionaceae. The Sandberg subpopulation

occurs on a similar rocky, sandy, south-facing slope of

identical substrate, growing in association with E. arach-

nocalyx E.G.H.Oliv., E. argentea , E. monsoniana L.f„ E.

nudiflora , E. recurvifolia and E. virginalis Klotzsch ex

Benth., as well as several species of Restionaceae. Both

populations occur between 1 140-1 280 m and may be

snowbound for short periods of time during the winter

months of May to September.

Paratype material

WESTERN CAPE. — 3319 (Worcester): Bokberg, Kouebokkeveld,

upper south-facing slopes S of beacon 4, 1 280 m, (-AB), 06-10-2001,

Turner 381 (NBG); SE-facing slopes of Bokberg S of trig, beacon 4,

fruiting material. 1 257 m, (-AB), 10-03-03, Turner 728 (NBG); rocky

S-facing slopes S of Sandberg, spot height 1 341 m, Peerboomkloof,

Kouebokkeveld. 1 240 m. (-AB), 09-11-2002. Turner 592 (NBG);

Bokberg, Hartebeeskloof 216, southern slopes SE of beacon 4, 1 280

m, (-AB). 10-10-2002, Oliver 12069 (BM, BOL, E, K, MO, NBG, NY,

P, PRE).

ACKNOWLEDGEMENTS

We wish to thank the Western Cape Nature Con-

servation Board for permission to collect plants in the

province and local landowners, in particular the Pohl

family of Winkelhaak, for access to their mountain areas.

REFERENCES

OLIVER, E.G.H. 2000. Systematics of Ericeae (Ericaceae: Ericoi-

deae): species with indehiscent and partially dehiscent fruits.

Contributions from the Bolus Herbarium 19: 1-483.

OLIVER, E.G.H. & OLIVER. I.M. 2000. Three new species of Erica

(Ericaceae) from Western Cape, South Africa. Bothalia 30:

147-153.

OLIVER, E.G.H. & OLIVER, I.M. 2002. The genus Erica (Ericaceae)

in southern Africa: taxonomic notes 1. Bothalia 32: 37-61.

REBELO, A.G. (ed.). 1987. A preliminary synthesis of pollination biol-

ogy in the Cape flora. South African Natural Sciences Pro-

grammes Report No. 141. CSIR, Pretoria.

REBELO, A.G.. SIEGFRIED, W.R. & OLIVER, E.G.H. 1985. Polli-

nation syndromes of Erica species in the southwestern Cape.

South African Journal of Botany 51: 269-280.

SPATIAL DATA MANAGEMENT UNIT. 1997. 1:250 000 Geological

Map, 3319 (Worcester).

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

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

7735 Claremont, Cape Town.

MS. received: 2003-09-01.

42

Bothalia 34,1 (2004)

FABACEAE

A NEW SPECIES OF ACACIA (MIMOSOIDEAE) FROM MPUMALANGA, SOUTH AFRICA

INTRODUCTION

Botanically poorly explored areas of South Africa con-

tinue to yield exciting new plant discoveries and records

(Hurter & Van Wyk 2001). During the course of field-

work aimed at expanding the living plants collection of

the Lowveld National Botanical Garden, Nelspruit, sev-

eral potentially new species of Acacia have been collect-

ed for the first time, one of which is described in the pre-

sent contribution. The new species is a yellow-flowered

member of subgenus Acacia section Acacia subsection

Uniseriae (Vassal 1974; Guinet & Vassal 1978; Chappill

& Maslin 1995; Timberlake et al. 1999).

Acacia ebutsiniorum P.J.H. Hurter, sp. nov., a

speciebus omnibus austro-africanis pedunculis longis-

simus (70-90 mm) robustis, foliolis cum margine hyali-

no differt. A. antunesii similis est sed habitu (suffrutex

vel arbor parva ad 2.5 m alta, non arbor magna ad 6 m

alta), foliis glabris, glandula petiolari magna, crateri-

forme, leguminibus constrictis subfalcatis differt.

TYPE. — Mpumalanga, 2630 (Carolina): Ebutsini tribal

land. Farm Tothiertoe 7 JT, 1 100 m, 8 November 2000,

(-BB), P.J.H. Hurter 133 (PRE, holo.; K, NBG, PRU, iso.).

Small tree up to 2.5 m tall, trunk slender. Bark coarse-

ly flaking or splitting to reveal a reddish or yellowish

brown inner layer. Branches striate, ferruginous; new

growth conspicuously striate, green, glutinous, lenticel-

late. Stipules in pairs, spinescent, 20-30 mm long, white,

glabrous, slender, basally flattened, attenuate, pungent,

antrorse, seldom arcuate, sometimes absent on new

growth. Leaves fastigiate, shiny dark green, glabrous,

glutinous when young, bipinnately compound, pulvinus

vermilion at maturity; petiole sulcate, 5-15 mm long,

with large, raised, crateriform, petiolar nectary gland;

rachis sulcate, 20-90 mm long, with at least one nectary

gland at junction of proximal and distal pinna pair;

rachillae 3-6 pairs, distichous, 30-81 mm long, dotted

with small dark glandular structures, with 8-16 leaflet

pairs; leaflets distichous, narrowly elliptic, 6-15 x

2.4-3.0 mm, entire, eglandular, asymmetrical, apex

mucronulate, base oblique, with conspicuous hyaline

margin, midvein prominent on abaxial surface only, sec-

ondary veins not visible. Inflorescences capitate, glo-

bose, bright yellow, 10-22 mm diam., borne on new

growth, fascicled on axillary peduncles; peduncle 70-90

mm long, glabrous or with few random, small dark

glands, glutinous when young, often with shards of vil-

lose hairs; involucel bracteate, 1 .0-2.2 mm long, 60-70

mm up the peduncle, 2 or 3 prominent, dark, gland-like

tubercles usually present, remnants of a second involu-

cel-like structure sometimes present ± 10 mm below

flowers. Flowers bright yellow, dichlamydeous.

Bracteoles clavate with apices covered by waxy glob-

ules. Calyx campanulate, pentamerous, glabrous, 1 .0-2. 1

mm long, ascending, usually with a few pustular waxy

globules, apex crenulated. Corolla campanulate, pen-

tamerous, membranaceous; lobes ± united, calceiform,

1.4- 1.6 x 0.5-0. 7 mm, ascending, edges and apex

puncticulate, usually with few pustular waxy globules.

Stamens numerous; filaments 2-3 mm long. Ovary ven-

tricose, septate, 1-2 mm long, surface with a few pustic-

ulate waxy globules; style 2-3 mm long. Pods dehiscent,

complanate, eglandular, slightly falcate, 40-1 15 x 7-1 1

mm, 2-6-seeded. edges becoming constricted between

seeds at maturity. Seeds elliptic, 6-9 x 4-7 mm, areole

elliptic, 4-6 x 2-5 mm. Figure 10.

Diagnostic features and affinities'. Acacia ebutsinio-

rum can immediately be distinguished from all other

southern African yellow-flowered acacias, including A.

natalitia E.Mey. and A. karroo Hayne, by its exception-

ally long, 70-90 mm, peduncles and the distinct hyaline

margin of the leaflets. The precise phylogenetic relation-

ships of A. ebutsiniorum are obscure. Its constricted pods

are vaguely reminiscent of those of A. natalitia (at times

included under A. karroo) with which it occurs sympatri-

cally. However, E. ebutsiniorum differs conspicuously

from the latter in that its pods are much broader and more

robust. Moreover, in the field it can readily be distin-

guished from A. natalitia by its much smaller stature and

markedly glutinous new growth. A. natalitia tends to be

a much bigger tree with the new growth never glutinous.

Additional differences between these two species are

provided by floret morphology. In A. natalitia the corol-

la lobes are fused into a short tube above the calyx,

whereas in A. ebutsiniorum the lobes are much shorter

and completely free.

Acacia ebutsiniorum superficially resembles A.

antunesii Harms from the Huila Plateau of southern

Angola in general appearance as well as shape and size

of the leaves (Ross 1979). There are, however, marked

differences in geographical distribution, plant size and

morphology. Salient morphological differences between

the two species are given in Table 1 .

Distribution and habitat', at present A. ebutsiniorum is

known from a single gregarious population at an altitude

of 1 140 m in a mountainous area northeast of Oshoek,

TABLE 1. — Differences between Acacia ebutsiniorum and A. antunesii

Bothalia 34,1 (2004)

43

FIGURE 10. — Acacia ebutsiniorum RJ.H. Hurter. A, flower head; B, individual flower; C, calyx. D, calyx lobe; E, corolla; F, gynoecium; G,-G3,

stipules; H, petiole; I, leaf; J, leaflet; K, pod; L, seed; M, bracteole. Scale bars: A, Gi, I, K, 20 mm; J, 14 mm; G2, G3, L, 10 mm; H, 4 mm;

E, 3 mm; B. 2.2 mm; C, F, 2 mm; D, 1.5 mm, M, 1.8 mm. Drawn from P.J.H. Hurter 133 spirit collection in PRU, by S. Burrows.

44

Bothalia 34,1 (2004)

FIGURE 1 1 . — Known distribution of Acacia ebutsiniorum in South

Africa.

Mpumalanga (Figure 11). The plants grow in exposed,

open grassland on a steep, southeast-facing slope, and

are periodically subjected to fire. A. ebutsiniorum shares

this habitat with several other newly discovered and still

to be described plant taxa, all confined to its immediate

vicinity. No earlier herbarium collections of the new

species are known, which is hardly surprising consider-

ing the remoteness of the locality and the unique and

apparently localized plant community of which it forms

a part. It is possible that other populations might exist in

seemingly similar habitats to the east of the present local-

ity, but this could not yet be confirmed. The known range

of the new species seems to fall just inside a local focus

of plant endemism known as the Barberton Centre (Van

Wyk & Smith 2001). This region is known for its many

edaphic specialists, including taxa endemic to serpenti-

nite-derived soils. Although the precise geological iden-

tity of the rocks at the A. ebutsiniorum locality still needs

to be established, it appears to be a type of serpentinite.

Etymology, the specific epithet commemorates the

people of the Swazi-speaking Ebutsini Tribe living in the

area where the trees occur, in acknowledgement of their

generous hospitality during field work and wealth of

field knowledge they so freely shared with the first

author; without their help this unique plant community

could not have been discovered.

ACKNOWLEDGEMENTS

We are indebted to the curators of GRA, K, NBG,

PRE, PRU, SRGH for access to their herbaria. Dr H.F.

Glen for the Latin diagnosis, Sandie Burrows for the line

drawing, Gerhard Strydom of the Mpumalanga Parks

Board for assistance in the field, SABONET and the

National Botanical Institute for financial assistance.

REFERENCES

CFIAPPILL, J.A. & MASLIN, B.R. 1995. A phylogenetic assessment

of tribe Acacieae. In M. Crisp & J.J. Doyle, Advances in legume

systematics 7: 77-99. Royal Botanic Gardens, Kew.

GUINET. P. & VASSAL, J. 1978. Hypotheses on the differentiation of

the major groups in the genus Acacia (Leguminosae). Kew Bul-

letin 32: 509-527.

HURTER, P.J.H. & VAN WYK, E. 2001. First distribution record for

Brachystegia spicifonnis (Caesalpinioideae) in South Africa.

Bothalia 31 : 43.

ROSS, J.H. 1979. A conspectus of the African Acacia species. Memoirs

of the Botanical Survey of South Africa No. 44. Botanical Re-

search Institute, Pretoria.

TIMBERLAKE, I, FAGG, C. & BARNES, R. 1999. Field guide to the

acacias of Zimbabwe . CBC Publishing, Harare.

VAN WYK, A.E. & SMITH. G.F. 2001 .Regions offloristic endemism

in southern Africa: a review with emphasis on succulents. Um-

daus Press, Pretoria.

VASSAL, J. 1974. Apport des recherches ontogeniques et seminolo-

giques a F etude morphologique, taxonomique et phylogenique

du genre Acacia. Bulletin de la societe d'histoire naturelle de

Toulouse 108: 125-247.

P.J.H. HURTER*f and A.E. VAN WYK**

* Lowveld National Botanical Garden, P.O. Box 1024, 1200 Nelspruit.

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

0002 Pretoria.

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

of Pretoria, 0002 Pretoria.

MS. received: 2003-02-26.

BORAGINACEAE

A FIRST RECORD OF ECHIUM SIMPLEX IN SOUTH AFRICA

INTRODUCTION

The Boraginaceae comprises ±131 genera and 2 500

species of mainly annual or perennial herbs and shrubs,

some trees and a few lianes (Brummitt 1992; Retief &

Van Wyk 1997). The plants are widely distributed in

tropical and especially subtropical and temperate regions,

with major centres of diversity in the Mediterranean

region and western parts of North America (Toelken

1986). In southern Africa the family is represented by 20

genera with 111 species, including 19 alien species re-

garded as naturalized in the flora of southern Africa

(Retief 2003).

The genus Echium L. is one of the non-native genera of

the Boraginaceae in southern Africa. Together with the seg-

regate genus Pontechium Bohle & Hilger, it is the northern

hemisphere counterpart of the closely related genera

Lobostemon Lehm. and Echiostachys Levyns, both endemic

to the Cape Floristic Region of South Africa (Retief & Van

Wyk 1997; Hilger & Bohle 2000; Van Wyk & Smith 2001).

Phylogenetically Echium splits naturally into two distinct

infrageneric groups or clades (Bohle et al. 1996; Hilger &

Bohle 2000). The first group comprises ± 30 species of

mainly herbaceous annuals to perennials. Geographically

this group has a circummediterranean distribution, extending

over most of Europe, reaching western Asia. The second

group of 27 species is endemic to the Canary, Madeira and

Cape Verde Archipelagos of the Macaronesian Islands. All

but two of these island species are woody perennials and

preferential outbreeders. Molecular data and geological evi-

dence suggest that all the contemporary island dwellers are

Bothalia 34.1 (2004)

45

PRETORIA NATIONAL HERBARIUM (PRE)

18.9/2001

- ttf-

ssmmi

FIGURE 12. — Terminal, many-flower-

ed thyrse of Echium simplex

kept as voucher specimen,

Smith & Steyn 14 at PRE.

derived from herbaceous continental antecedents following

a perhaps single founding colonization less than 20 million

years ago (Bohle et al. 1996, but see Perez de Paz 1995 for

evidence from palynology).

ECHIUM SPECIES IN SOUTHERN AFRICA

Two of the herbaceous continental European species of

Echium , namely E. plantagineum L. and E. vulgare L„

were previously discussed in detail (Retief & Van Wyk

1998). Both are naturalized in Western and Eastern Cape,

eastern Free State, Lesotho, the high mountainous areas of

KwaZulu-Natal and in Mpumalanga, where they mainly

occur as annual weeds of roadsides and fallow fields. A

member of the island group of species, E. candicans L.f.

(‘Pride of Madeira'), is widely grown as a garden orna-

mental in South Africa. Based on anecdotal evidence, it

was somewhat doubtfully listed as a potential problem

plant in the summer rainfall regions of South Africa (Wells

et al. 1986). However, there is no evidence that this species

has become naturalized in our region; its occurrence out-

side gardens is rare and accidental. The present paper deals

with a fourth species, Echium simplex DC., another mem-

ber of the group of island dwellers and recently reported

from South Africa for the first time (Willis & Smith 2002).

Echium simplex, the ‘Pride of Tenerife', is a rare

endemic of the Canary Islands. With its basal rosette of

silvery green leaves and massive, erect, up to 2 m tall

inflorescence of several hundred pure white flowers, it is

a most spectacular plant (Figures 12-14). In its native

46

Bothalia 34.1 (2004)

FIGURE 13. — Population of Echium simplex thriving among coastal

scrub near Hermanus in the southwestern Cape. Photograph:

G.F. Smith.

country, it is a coastal cliff species of low altitudes

( 100-400 m), restricted to humid habitats on basalt cliffs

of the Anaga Mountains in the northeastern region of the

island Tenerife (Bramwell 1972; Bohle et al. 1996). In a

taxonomic revision of the Macaronesian members of

Echium , Bramwell (1972) classifies E. simplex together

with E. pininana Webb & Berth, and E. wildpretii

H.H.W.Pearson ex Hook.f. in section Simplicia (H. Christ

ex Sprague & Hutch.) Bramwell. E. simplex has become

naturalized in the flora of South Australia (Toelken 1986)

where it thrives near Robe in the Mediterranean-like cli-

mate of the southeastern parts of the region. In 2001 the

first author found a flourishing population of these alien

plants in the veld among coastal scrub near the coastal

town of Hermanus in the southwestern Cape, where the

climate is also typically Mediterranean. This provenance

is supported by a voucher specimen. Smith & Steyn 14

(Figure 12) kept in the National Herbarium of South

Africa (PRE). This strikingly beautiful plant was most

probably introduced to South Africa as a garden orna-

mental. However, if indeed the case, this must have been

an isolated event as we have no knowledge of the species

being propagated by nurseries in the region and we have

never observed it in a local garden.

In view of the ease with which species of Echium

seem to overcome barriers to produce offspring freely

and to sustain populations over many life cycles in the

FIGURE 14. — Apical part of inflorescence of Echium simplex with

showy, pure white flowers. Photograph: G.F. Smith.

southern African veld, Echium simplex should be regard-

ed as a likely candidate for naturalization (, sensu

Richardson et al. 2000) in the Bora of the southwestern

Cape. On the other hand, this apparent ease of establish-

ment may favour the potential agronomical use of E.

simplex , and perhaps other Macaronesian members of

the genus in the Cape. Seeds of these island species are

one of the richest sources of gamma-linolenic acid

[GLA] found in nature so far. GLA is a commercially

important seed oil for which there is a growing demand

in the food, cosmetic and pharmaceutical industries

(Horrobin 1992; Guil-Guerrero et al. 2000, 2001). A sin-

gle plant of E. simplex produces several thousand propag-

ules, with seed oil comprising 10.04% of seed weight and

containing 19.28% GLA (Guil-Guerrero et al. 2000).

7118000-00065 Echium simplex DC., Catalogus

plantarum horti botanici monspeliensis: 108 (1813);

Bramwell: 75 (1972); Bramwell & Bramwell: 184, t. 228

(1974); Toelken: 1 156 (1986). Type: "E. simplex DC. hab.

in Insula Teneriffa ex Herb. Broussonet’, fide Bramwell

(1972: 75) [MPU, non vidi, fide Bramwell (1972)].

Unbranched, monocarpic rosette plant, lasting 4-5

years. Stem woody, short in vegetative state, lengthening to

± 1 m during flowering. Leaves somewhat leathery, spiral-

ly arranged, crowded at tip of stem in vegetative state, ses-

sile, elliptic-lanceolate, 200-220 mm long, 40-45 mm wide

Bothalia 34.1 (2004)

47

at mid-blade, both surfaces silvery green, densely covered

with silky, short, unicellular, small-based, appressed tri-

chomes, veins on lower surface prominent. Inflorescence a

terminal, dense, cylindrical thyrse up to 600 mm long; lat-

eral cymes in axils of leaf-like bracts, numerous, scoipioid,

1 -5-fid, many-flowered, distinctly stalked; lateral pedun-

cles 15-30 mm long, bracteate. Flowers sessile, actin-

omorphic, nectariferous, unscented, strongly protandrous.

Calyx persistent, 5-partite, divided almost to base; lobes

lanceolate, ± 6 mm long, acute, pubescent. Corolla white,

± 10 mm long, funnel-shaped, sparsely pubescent, 5-lobed;

lobes equal, slightly spreading, rounded. Stamens silvery

white, equal, strongly exserted, arising as single series at ±

3 mm from base of corolla tube; filaments glabrous, cylin-

drical, ± 1 1 mm long; anthers versatile, small, ± 2 mm long

before dehiscence, splitting lengthwise; pollen silvery

white. Ovary deeply 4-lobed; style simple, gynobasic,

exserted beyond anthers; stigma bifid. Fruit a schizocarp

consisting of four greyish brown nutlets; nutlets ±2.7 x 2

mm, tuberculate with tubercles varying in size, convex on

dorsal side, keeled on ventral side, ± triangular at base.

ACKNOWLEDGEMENT

Dr David Bramwell of the Jardin Botanico Canario,

Las Palmas, Gran Canaria, is thanked for positively iden-

tifying the South African specimens of Echium simplex.

REFERENCES

BOHLE, U.-R., HILGER. H.H. & MARTIN. W.F. 1996. Island colo-

nization and evolution of the insular woody habit in Echium L.

(Boraginaceae). Proceedings of the National Academy of

Sciences of the United States of America 93: 11740-11745.

BRAMWELL. D. 1972. A revision of the genus Echium in Maca-

ronesia. Lagascalia 2: 37-115.

BRAMWELL. D. & BRAMWELL. Z.I. 1974. Wild flowers of the

Canary Islands. Excmo. Cabildo Insular de Tenerife in associ-

ation with Thornes, London.

BRUMMITT, R.K. (compiler). 1992. Vascular plant families and gen-

era. Royal Botanic Gardens, Kew.

DE CANDOLLE, A.R 1813. Echium simplex. Catalogus plantation

horti botanici tnonspeliensis. Martel, Paris.

GUIL-GUERRERO, J.L., GOMEZ-MERCADO. F.. GARCIA-MAROTO,

F. & CAMPRA-MADRID. P. 2000. Occurrence and character-

ization of oils rich in (y-linolenic acid. Part I: Echium seeds

from Macaronesia. Phytochemistry 53: 451-456.

GUIL-GUERRERO. J.L., GOMEZ-MERCADO. F.. RODRIGUEZ-

GARCIA, I., CAMPRA-MADRID, P. & GARCIA-MAROTO,

F. 2001. Occurrence and characterization of oils rich in (y-

linolenic acid (III): the taxonomic value of the fatty acids in

Echium (Boraginaceae). Phytochemistry 58: 117-120.

HILGER, H.H. & BOHLE, U.-R. 2000. Pontechium: a new genus dis-

tinct from Echium and Lobostemon (Boraginaceae). Taxon 49:

737-746.

HORROBIN, D.F. 1992. Nutritional and medical importance of gamma-

linolenic acid. Progress in Lipid Research 31: 163-194.

PEREZ DE PAZ, J. 1995. Generalidades, diversidad pollnica y evolucion

de las secciones Macaronesicas: Stricta, Virescentia, Gentia-

noidea y Echium. del genero Echium L. (Boraginaceae). Botanica

Macaronesica 21: 3-28.

RETIEF, E. 2003. Boraginaceae. In G. Germishuizen & N.L. Meyer.

Plants of southern Africa: an annotated checklist. Strelitzia 14:

313-320. National Botanical Institute, Pretoria.

RETIEF, E. & VAN WYK, A.E. 1997. Palynology of southern African

Boraginaceae: the genera Lobostemon , Echiostachys and Echium.

Grana 36: 271-278.

RETIEF, E. & VAN WYK. A.E. 1998. The genus Echium (Boragin-

aceae) in southern Africa. Bothalia 28: 167-177.

RICHARDSON, D.M.. PYSEK, P„ REJMANEK, M„ BARBOUR.

M.G.. PANETTA, F.D. & WEST, C J. 2000. Naturalization and

invasion of alien plants: concepts and definitions. Diversity and

Distributions 6: 93-107.

TOELKEN, H R. 1986. Boraginaceae. In J.P. Jessop & H R. Toelken,

Flora of South Australia 3: 1146-1173.

VAN WYK, A.E. & SMITH, G.F. 2001. Regions offloristic endemism

in southern Africa: a review with emphasis on succulents.

Umdaus Press, Pretoria.

WELLS. M.J., BALSINHAS, A. A., JOFFE, H„ ENGELBRECHT, V.M.,

HARDING, G. & STIRTON, C.H. 1986. A catalogue of prob-

lem plants in southern Africa. Memoirs of the Botanical Survey

of South Africa No. 53. Botanica! Research Institute, Pretoria.

WILLIS, C.H. & SMITH, G.F 2002. Canary Islands: land of the dragon

trees. Aloe 39: 15-22.

G.F SMITH*+, A.E. VAN WYK**, E M. A. STEYN*

and E. RETIEF*f

* National Botanical Institute. Private Bag XI 01, 0001 Pretoria.

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

University of Pretoria, 0002 Pretoria.

+ Affiliation: Acocks Chair, Department of Botany, University of

Pretoria, 0002 Pretoria; corresponding author, e-mail:

smithg@nbi.ac.za

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

0002 Pretoria.

MS. received: 2003-10-31.

MESEMBRYANTHEMACEAE

A NEW TRIBE AND ADJUSTMENTS TO INFRAFAMILIAL CLASSIFICATION

Recently the tribal classification of Mesembryanthe-

maceae Fenzl has been the focus of attention in contri-

butions based on both morphological (Chesselet et al.

2001, 2002) and molecular (Klak et al. 2003) evidence.

Molecular data provides an independent character set for

testing the congruence of morphological characters.

When contrasting these data sets we find that molecular

data largely support the broad infrafamilial classification

based primarily on nectary type proposed by Chesselet et

al. (2002), except in the case of tribe Delospermeae

(Figure 15).

Delosperma N.E.Br. (± 163 species) and Drosanthe-

mum Schwantes (± 120 species) are the largest of sever-

al genera placed by Chesselet et al. (2002) in the Delo-

spermeae. Klak et al. (2003), however, do not maintain

the Delospermeae, but place all taxa assigned to this tribe

in an expanded concept of Ruschieae (Chesselet et al.

2002). Although the molecular phylogeny of Klak et al.

(2003) is not fully resolved, four distinct clades do

emerge for subfamily Ruschioideae, three of which are

formally recognized taxonomically by these authors. The

fourth clade includes all investigated members of

Drosanthemum and a single species of Delosperma ,

namely D. asperulum (Salm-Dyck) L. Bolus (= Dro-

santhemum asperulum (Salm-Dyck) Schwantes). The

latter taxon has since been transferred back to

Drosanthemum by Klak (2003) on the grounds that it

resolves with Drosanthemum in a distinct evolutionary

lineage within the tribe Ruschieae (Klak et al. 2003).

This clade is, however, not given any formal taxonomic

status despite it having 95% bootstrap support in a strict

48

Bothalia 34. 1 (2004)

□ Dot otheanthus bellidiformis

HI Diplosoma retroversum

Disphyma dunsdonii

Q Meyerophytum meyeri

■ o Delosperma echinatum

I o Trichodiadema emarginatum

o Delosperma esterhuyseniae

o Delosperma pageanum

o Malephora lutea

Q Monilaria monolformis

o Gibbaeum pachypodium

I o Oscularia deltoides

1 @ Lampranthus bicolor

o Fai caria felina

— o Mossia intervallaris

o Neohenricia spiculata

o Bijlia tugwelliae

o Orthopterum coeganum

o Cat man thus ringens

o Cerochlamys pachyphylla

o Machairophyllum albidum

i Dinteranthus puberulWs

' Q Lithops julii

Titanopsis luigo-schlechteri

® Conophytum bruynsii

| Namibia cinerea

' IP Ruschia robusta

♦ Nelia pillansii

Antegibbaeum fissoides

. ^ Braunsia gemmata

' 4P Antimima ventricosa

Lampranthus mucronatus

4) Carpobrotus muirii

Enarganthe octonaria

^ Namaquanthus vanheerdei

^ Polymita steenbokensis

^ Lampranthus amphibolnts

^ Lampranthus filicaulis

Q Scopelogena bruynsii

0 Smicrostigma viride

ft Wooleya farinosa

ft Amphibolia laevis

ft Eberlanzia dichotoma

ft Astridia longifolia

ft Ruschia strubeniae

ft Cephalophyllum pillansii

^ Fenestraria rhopalophylla

ft Odontophorus marlothii

ft P/eiospilos simulans

I o Delosperma asperulum

^ o Drosanthemum sp. nov.

■ o Drosanthemum sp.

1 o Drosanthemum speciosum

o Dr os an the m um I ongipes

o Drosanthemum schoenlandianum

o Drosanthemum zygophvlloides

FIGURE 15. - Nectary characters from Chesselet et al. (2002) plotted on molecular phylogeny for mesembs from Klak et til. (2003). The most

striking result is that the Delosperma Group of Hartmann (1991, 1993) is polyphyletic. Note the molecular resolution obtained for mem-

bers of the newly described tribe Drosanthemeae in contrast to polytomy obtained for the Ruschieae, indicating the greater age of the for-

mer group. Members of the Bargeranthus and Stomatium groups of Hartmann (1991. 1993) resolve as a distinct clade within Ruschieae

and are characterized by a lophomorphic meronectary. Key: □, broad flat meronectary; O, lophomorphie meronectary; •, lophomorphic

holonectary.

Bothalia 34.1 (2004)

49

consensus tree of the 5000 most parsimonious trees

based on a four-gene-region analysis. Here we propose

the latter clade as a distinct new tribe, the Drosanthe-

meae. The molecular phylogeny nevertheless shows that

some other Delosperma species as well as several other

genera included in Delospermeae by Chesselet et al.

(2002) are clearly better placed in Ruschieae.

The most unexpected result of the molecular phyloge-

ny provided by Klak et al. (2003) is that the 'Delosperma

group’ of Hartmann (1991, 1993) is polyphyletic. Since

Schwantes (1947), in his System der Mesembrvanthe-

maceen, defined Subtribe Delospermatinae, Delosperma

and Drosanthemum have been uncritically associated

with each other. Despite the simple hygrochastic fruit,

papillate epidermis, often broad, flat mesophytic leaves,

and an exceptionally wide (for mesembs) distribution

range that extends from southern Africa to East Africa

and the Indian Ocean islands of Reunion and Mada-

gascar, the genus Delosperma is not as ‘primitive’ as has

been widely accepted. In contrast, Drosanthemum , with

its characteristic basally-free leaves that bear large,

water-filled bladder cells, emerges as being more basal

than hitherto surmised. The wide distribution range of

Drosanthemum and its strong presence in the Fynbos

Biome supports this hypothesis. Recently, Chesselet et

al. (2003) argued that basal tribes in Ruschioideae have

their origins in the Fynbos Biome, which is considered

the older biome in relation to the Succulent Karoo

(Stuckenberg 1998).

At present the distinct clade named here as tribe

Drosanthemeae contains the single genus Drosanthemum.

Following extensive work on the flowers and fruit of

Drosanthemum (Hartmann & Bruckmann 2000; Rust et

al. 2002) a single possible synapomorphy involving the

fruit stalk was found for the genus. Drosanthemum is a

large and morphologically heterogeneous taxon character-

ized by no less than 1 3 different fruit types and five floral

types. Congruence between these character sets exists in

only three cases. The genus remains one of the most prob-

lematic in terms of identification. The fact that some

Drosanthemum species tend to show free leaves, supports

their more primitive position in the phylogeny. This char-

acteristic is shared with other more primitive ruschioid

taxa in tribes Apatesieae and Dorotheantheae and it can

thus be regarded as a symplesiomorphic feature. Enar-

ganthe N.E.Br. is reported to have basally-free leaves

(Brown 1930), although examination of leaf material does

not confirm this unambiguously. We therefore consider

basally fused leaves as a possible synapomorphy for tribe

Ruschieae (Figure 16). Those species of Drosanthemum

that appear to have leaves that are fused at the base may

indicate convergence with taxa in Ruschieae.

Both Delosperma and Drosanthemum are character-

ized by flowers with a ‘lophomorphic meronectary’ and

were placed by Chesselet et al. (2002) in Delospermeae

together with all other genera with this type of nectary.

The inclusion, into this tribe, of the Bergeranthus and

Stomatium groups of Hartmann (1991, 1993), was

counter-intuitive, though justified according to the prin-

ciple of parsimony, in the absence of other character sets

at the time. The molecular study of Klak et al. (2003) has

shown that taxa belonging in the latter two groups are

better placed in tribe Ruschieae. The presence of a lopho-

morphic meronectary indicates a possible reversal,

switching of major genes or the persistence of the

pleisiomorphic condition in genera allied to Bergeran-

thus and Stomatium.

Following the reassessment of the tribal classification

based on nectary types and together with insights gained

from molecular data, we propose the following nomen-

clatural adjustments. For a comprehensive tribal classifi-

cation refer to Chesselet et al. (2002).

Drosanthemeae Chesselet , Gideon F.Sm. & A.E.van

Wyk , trib. nov. Type: Drosanthemum Schwantes

Plantae perennes. Paria foliorum plerumque basi disc-

reta; folia saepeque caules plerumque cum cellulis vesi-

cariis prominentibus vel pilis. Flores semper cum

meronectariis lophomorphis. Capsulae hygrochasticae,

plerumque 5(6)-carpellatae, interdum cum locellis semi-

nalibus basalibus.

Plants perennial. Leaf pairs mostly basally free,

leaves and often stems, usually with prominent bladder

cells or hairs. Flowers always with crested or lobed

(lophomorphic) separate nectaries (meronectary). Cap-

sules hygrochastic, usually 5(6)-carpellate sometimes

with basal seed chambers.

Includes tribe Delospermeae Chesselet, Gideon F.Sm.

& A.E.van Wyk p.p.

Includes subtribe: Delospermatinae Schwantes p.p.

Comprises the Delosperma Group of Hartmann (1991,

1993) p.p.

Genera: Drosanthemum [at least as to species investi-

gated by Klak et al. (2003)]. Note: the inclusion of addi-

tional species currently placed in Delosperma and/or

Mestoklema is uncertain at this stage, as is the possible

exclusion of some species currently placed in Dro-

santhemum.

Ruschieae Schwantes in Ihlenf., Schwantes & Straka

in Taxon 1 1 : 54 ( 1 962), emend. Klak, Khunou, Reeves &

Hedderson: 1443 (2003). Type: Ruschia Schwantes. Note:

the spelling of ‘Ruschiae’ in Klak et al. (2003) is an ortho-

graphic error.

Plants perennial, sometimes with enlarged rootstock.

Leaves mesomorphic or xeromorphic, always basally

fused, with or without prominent bladder cells and/or

hairs or with homocellular or heterocellular xeromorphic

surfaces. Flowers with crested or lobed (lophomorphic)

separate nectaries (meronectary) or annular nectaries

(holonectary). Capsules hygrochastic, rarely with reduced

expanding keels, and breaking up when dry (schizo-

carpic )', fruit rarely a berry.

Comprises the Bergeranthus, Delosperma p.p., Draco-

philus, Eberlanzia, Lampranthus, Leipoldtia, Mitrophyl-

lum, Ruschia, Stomatium and Titanopsis Groups of

Hartmann (1991, 1993).

50

Bothalia 34,1 (2004)

*Cortical bundles present

‘Stamens and petals forming a tube

Koilomorphic nectary

*n=9

‘semi-epigynous or

epigynous flowers

‘petals of staminodial

origin

Meronectary

‘gynoecium inferior

‘tepal bases free

‘filaments papillate at base

‘placenta basal or parietal

‘expanding keels ±

restricted to valves

Subfamily

Mesembryanthemoideae

Expanding keels reduced or absent

Holonectary

P

Lophomorphic nectary

‘Leaves succulent

‘Wide band tracheids

Tribe Apatesieae

Tribe Dorotheantheae

Tribe Drosanthemeae

Tribe Ruschieae

leaves basally fused

Subfamily

Ruschioideae

FIGURE 16. — Cladogram of the basal groups of Mesembryanthemaceae. The previously named tribe Delospermeae has been replaced with Tribe

Drosanthemeae. Genera previously included in Delospermeae have been redistributed in Drosanthemeae and Ruschieae. From Chesselet

el al. (2002), with modification. Note: characters of the floral nectary indicated as black bars.

Includes subtribes: Carpobrotinae Schwantes, Cono-

phytinae Schwantes, Delospermatinae Schwantes p.p.,

Dracophilinae Schwantes, Erepsiinae Schwantes, Fau-

cariinae Schwantes, Frithiinae Schwantes, Gibbaeinae

Schwantes, Jacobseniinae Schwantes, Jensenobotryinae

Schwantes, Feipoldtiinae Schwantes, Fithopinae Schwan-

tes, Fampranthinae Schwantes, Malephorinae Schwantes,

Mitrophyllinae Schwantes, Nananthinae Schwantes,

Pleiospilinae Schwantes, Psammophorinae Schwantes,

Ruschiinae Schwantes, Stoeberiinae Friedrich, Stoma-

tiinae Schwantes.

Genera: Acrodon, Aloinopsis, Amphibolia, Antegib-

bcieum, Antimima, Arenifera, Argyroderma, Astridia,

Bergeranthus , Bijlia, Braunsia , Carpobrotus, Carruanthus,

Cephalophyllum, Cheiridopsis, Cerochlamys, Chasmato-

phyllum, Circandra, Conophytum, Corpuscularia, Cylin-

drophyllum, Dedanthe, Delosperma, Dicrocaulon, Didy-

maotus, Dinteranthus, Diplosoma, Disphyma, Draco-

philus, Eberlanz.ia, Ebracteola, Ectotropis, Enarganthe ,

Erepsia, Esterhuysenia, Faucaria, Fenestraria, Frithia,

Gibbaeum, Glottiphyllum, Hallianthus, Hammeria, Hart-

manthus, Hereroa, Ihlenfeldtia, Jacobsenia, Jenseno-

botrya, J ordaaniella, Juttadinteria, Khadia, Lampran-

thus, Lapidaria, Leipoldtia, Lithops, Machairophyllum,

Malephora , Marlothistella, Mestoklema, Meyerophytum,

Mitrophyllum, Monilaria, Mossia, Muiria, Namaquan-

thus, Namibia, Nananthus, Nelia, Neohenricia, Octo-

poma, Odontophorus, Oophytum, Orthopterum, Oscula-

ria, Ottosonderia, Pleiospilos, Polymita, Psammophora,

Rabiea, Rhinephyllum, Rhombophyllum , Ruschia, Ruschi-

anthemum, Ruschianthus, Schlechteranthus, Schwante-

sia, Scopelogena, Smicrostigma, Stayneria, Stoeberia,

Stomatium, Tanquana, Titanopsis, Trichodiadema, Vanheer-

dea, Vanzijlia, Vlokia, Wooleya, Zeuktophyllum.

REFERENCES

BROWN, N.E. 1930. Mesembryanthemum. Gardeners' Chronicle 87:

151.

CHESSELET, P„ SMITH, G.F. & VAN WYK, A.E. 2001. A new trib-

al classification for the Mesembryanthemaceae Fenzl 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

flora] nectaries. Taxon 5 1 : 295-308.

CHESSELET, P, VAN WYK, A.E., GRIFFIN, N. & SMITH, G.F.

2003. Patterns of floristic diversity in Mesembryanthemaceae.

Aloe 40: 80-85.

HARTMANN, H.E.K. 1991. Mesembryanthema. Contributions from

the Bolus Herbarium 13: 75-157.

HARTMANN, H.E.K. 1993. Aizoaceae. In K. Kubitzki, J.G. Rohwer

& V. Bittrich, The families and genera of vascular plants 2:

37-69. Springer- Verlag, Berlin.

HARTMANN, H.E.K. & BRUCKMANN, C. 2000. The capsules of

Drosanthemum Schwantes (Ruschioideae, Aizoaceae). Brad-

leya 18: 75-112.

IHLENFELDT, H.D., SCHWANTES, G. & STRAKA, H. 1962. Die

hoheren Taxa der Mesembryanthemaceae. Taxon 1 1 : 52-56.

KLAK, C. 2003. New combinations, a new genus and five new species

in Aizoaceae. Bradleya 21: 107-120.

KLAK, C„ KHUNOU. A., REEVES, G. & HEDDERSON, T. 2003. A

phylogenetic hypothesis for the Aizoaceae (Caryophyllales)

based on four plaslid DNA regions. American Journal of

Botany 90: 1433-1445.

Bothalia 34,1 (2004)

51

RUST, S.. BRUCKMANN, C. & HARTMANN, H.E.K. 2002. The

flowers of Drosanthemum Schwantes (Ruschioideae, Aizoa-

ceae). Bradleya 20: 121-147.

SCHWANTES, G. 1947. System der Mesembryanthemaceen. Sukku-

lentenkunde, Jahrbiicher der Schweizerischen Kakteengesell-

schaft 1 : 34-40.

STUCKENBERG, B.R. 1998. A new Namibian wormlion species, with

an account of the biogeography of Leptynoma Westwood s. str.

and its association with anthophily in the Fynbos and Succulent

Karoo Biomes (Diptera, Vermileonidae). Annals of the Natal

Museum 39: 165-183.

P. CHESSELET*t, A.E. VAN WYK** and G.F. SMITH***

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

7735 Claremont, Cape Town. E-mail: chesselet@nbi.ac.za

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

0002 Pretoria.

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

versity of Pretoria, 0002 Pretoria. E-mail: avanwyk@scientia.up.ac.za

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

Acocks Chair, Department of Botany, University of Pretoria. 0002

Pretoria. E-mail: gfs@nbi.ac.za

'

Bothalia 34,1:53-59 (2004)

Functional and taxonomic significance of seed structure in Salix

mucronata (Salicaceae)

E.M.A. STEYN*f, G.F. SMITH*+and A.E. VAN WYK**

Keywords: anemochory, Flacourtiaceae, hilar aril, hydrochory, plumed seed, rheophyte, Salicaceae, Salix mucronata , seed adaptations, taxonomy

ABSTRACT

The polymorphic African willow, Salix mucronata Thunb., is a widely distributed African tree and a riparian rheophyte.

Ovule-to-seed development is reported for Salix mucronata subsp. woodii (Seemen) Immelman. Contrary to some existing

reports, the tuft of silky hairs enveloping the seed in Salix is derived from the funicle and not the placenta. The micromor-

phological structure of the hilar aril and funicular-placental and arillate hairs is described and illustrated for the first time.

Willow seeds are primarily wind-dispersed, but have additional characters, such as a hydrophobic seed coat and an unwet-

table, hairy, hilar aril as specific adaptations for distribution by water, perhaps even chance dispersal by animal visitors to

the riverine habitat. Seed adaptations linked to different dispersal strategies may account for seemingly marked differences

in ovule/seed structure between Salicaceae s. str. and related, mainly zoochorous flacourtiaceous taxa, recently classified

with the former in a more inclusively circumscribed Salicaeae s.l.

INTRODUCTION

Salicaceae, a classical family recently drastically rede-

fined and classified in the order Malpighiales (Chase et

al. 2002), is now considered a cosmopolitan group com-

prising about 53 genera with ±2415 species of woody

perennials. Apart from Salix L. (willows) and Populus L.

(poplars, cottonwoods), the two genera traditionally mak-

ing up the family (Salicaceae s.str.), the newly proposed

circumscription now also includes most of the genera for-

merly placed in Flacourtiaceae sensu Lemke ( 1988). With

400-450 species included in Salix, this speciose genus of

dioecious, catkin-bearing shrubs and trees is by far the

largest in Salicaceae s.l. sensu Chase et al. (2002).

Salix, commonly known as willows, is the most wide-

ly dispersed genus in the family. Currently, most willow

species occur in cold temperate to arctic regions of the

northern hemisphere, but some grow in warm temperate

to tropical climates (Wilmot-Dear 1985, 1991; Friis

1992; Judd et al. 2002) where the genus possibly origi-

nated (Skvortsov 1968; Dorn 1976). The genus is absent

from Australasia and New Guinea (Wilmot-Dear 1985),

a few species are found in South America (Zuloaga &

Morrone 1999) and one species occurs naturally in

Africa (M. Jordaan pers. comm.). This African species,

Salix mucronata Thunb. (= S. subserrata Willd.) is wide-

ly distributed, extending from southern Arabia (McKean

1996) and Egypt southwestwards to Senegal and south-

wards through Ethiopia to Namibia and South Africa

(Friis 1992). It is a polymorphic species and, in southern

Africa, four (Coates Palgrave 2002; Jordaan 2002) or

five (Immelman 1987) subspecies are recognized.

Characteristically, Salix prefer wet to moist, open habi-

tats (Judd et al. 2002). In almost all African phytochoria.

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

** Department of Botany, University of Pretoria, 0002 Pretoria,

t Corresponding author, e-mail: elsie@nbipre.nbi.ac.za

+ Affiliation: Botany Department, University of Pretoria, 0002 Pretoria.

MS. received: 2003-03-14.

representatives of the genus are riverine in distribution

(Friis 1992). Indeed, many Salix species throughout the

world are riparian rheophytes, i.e. plants growing along

swift-running watercourses and on the banks up to flood

level, but not beyond the reach of regularly occurring flash

floods (Van Steenis 1981; Jordaan 2002). One such species

is the widely dispersed African endemic, S. mucronata

(Van Steenis 1978).

It is generally assumed that the plumed seeds of wil-

lows are wind-dispersed (Ridley 1930; Van der Pijl 1969;

Johri et al. 1992; Judd et al. 2002), but the issue is whether

wind is the principal dispersal agent involved in the wide

geographical distribution of S. mucronata. Our observa-

tions on seed dispersal in Salix mucronata subsp. woodii

(Seemen) Immelman showed that upon shedding, the seed

does not remain airborne for long, despite the cover of

loose wool that initially surrounds the seed and expedites

its release from the capsule. In this contribution we present

data on seed structure, supplemented by a study of capsule

and seed development, suggesting that although willows

are adapted to wind dispersal, the seed has additional modi-

fications for dispersal by water and fortuitously by water-

fowl or other animal visitors to the rheophytic habitat. We

also comment on the taxonomic significance of seed struc-

ture in Salix, considering its recent association with mem-

bers traditionally placed in Flacourtiaceae.

MATERIAL AND METHODS

Catkins in most developmental stages were collected

from a female tree of S. mucronata subsp. woodii (acces-

sion number: GPTA495) in the Pretoria National Botanical

Garden. The plant was grown from seed, collected by Dr

L.E. Codd in 1953 from a tree (Codd 8246 PRE) on the

bank of the Buffalo River, Natal [KwaZulu-Natal], Floral

buds, mature flowers, developing fruit and dehiscing cap-

sules were fixed in a 0.1 M cacodylate-buffered solution

(pH 7.4) containing 2.5% gluteraldehyde and 4% formal-

dehyde. Standard procedures were followed for dehydrat-

ing, infiltrating and imbedding the material in glycol

methacrylate (GMA). For light microscope (EM) study.

54

Bothalia 34,1 (2004)

sagittal sections of ovules and seeds were obtained by sec-

tioning flowers and capsules longitudinally, perpendicular

to the subtending bract. Selected sections were stained with

the periodic acid/Schiff reaction and counterstained with

toluidine blue O (pH 4.4) by using the protocols of O'Brien

& McCully (1981). Seed with mature embryos were

obtained from herbarium sheets {Codd 8246 PRE;

Obermeyer TM31027 PRE) and processed for GMA sec-

tioning and staining as described above.

Tests for flight distance of seed were performed on

windy days in an open part of the Pretoria National

Botanical Garden. Dehisced capsules containing wool-

covered, fertile seed were collected from Codd 8246 and

Obermeyer TM31027, held between tweezers and

exposed to the wind. Sterile ‘seed’ (unfertilized, shriv-

eled ovules covered in wool) produced by Tree No.

GPTA 495 were tested on the same days. Tests for wet-

tability and buoyancy were conducted indoors by blow-

ing loose wool and plumed, fertile seed onto the surface

of muddy or clear, distilled water kept in containers in

the laboratory and noting the flotation period.

For scanning electron microscope (SEM) studies,

seeds from the above-mentioned herbarium specimens

were carefully removed from dehiscing capsules, stuck

onto aluminum stubs, sputter-coated with gold and

viewed in an ISI SX 25 SEM.

RESULTS

Placentation, development and structure of ovules

Catkins on female trees contain a varying number (±

10-50) of flowers. Each flower is borne in the axil of a

bract and consists of an adaxial nectary and a syncarpous,

bicarpellate gynoecium (Figure 1A). The two carpels of

the salicaceous pistil meet in the medium plane of sym-

metry of the flower (Meeuse 1975: 450) so that the pari-

etal placentae, developing on the fused margins of the

carpels, lie in this plane (Figure 1A). The ovuliferous

zones of the placentae are restricted to the lower half of the

unilocular ovary where eight to twelve ovules are formed.

Ovule primordia at the onset of meiosis (Figure IB)

were found inside floral buds still completely covered by

the subtending bracts. In sagittal section (Figure 1A, B),

such primordia consist of a large funicle, a developing

outer integument (Comer 1976: 237) and a nucellus con-

taining an enlarging megaspore mother cell covered by at

least three layers of parietal nucellar cells (Figure 1A).

An inner integument is absent, without any vestiges

remaining. Meiosis results in four megaspores of which

the chalazal one becomes functional (Figure 1C). Finear

tetrads were not found in our material; the three micropy-

lar megaspores lie obliquely to one another and possibly

represent a stage intermediate between linear and T-

shaped tetrads. At the completion of the meiotic process

the integument has reached its full length, it covers the

nucellus and forms the micropyle (Figure 1C).

Mature ovules are small (± 20-25 pm in length, funi-

cle excluded), anatropous and ovoid (Figure 2A). The

lengths of the funicles vary (compare Figure 2A & C);

ovules in the basal-parietal position usually have much

shorter funicles than ovules nearer to the style. The

integument consists of three to four layers of thin-walled

parenchyma. After the elements of the eight-nucleate

embryo sac have been formed, the tip of the sac keeps

intruding into the micropylar nucellus, later breaking

through the nucellus epidermis and reaching the inner

opening of the micropyle canal (Figure 2B).

During the elongation of the embryo sac, a plate-like

intercalary meristem develops in the distal part of the

funicle, forming a narrow band of flattened cells directly

below the ovular body (Figure 2A). This region eventu-

ally forms a small, ring-like structure in the hilar region

of the seed (see further on). At a slightly later stage, the

epidermis cells of the placenta, funicle and hilar meris-

tem form long, unicellular, intra-ovarian hairs, but the

surface of the enlarging ovule remains glabrous (Figure

2C). In developing capsules collected from the female

tree (GPTA No. 495) fertilization did not take place and

the ovules aborted (Figure 2D). In such ovules the cells

of the hilar meristem have disintegrated (Figure 2D).

Seed abscission takes place at the funicular side of the

aborted meristem so that the greater part of the funicle

remains connected to the placenta (Figure 2C, D).

It is interesting that the capsule with its intra-ovarian

hairs keeps developing and later dehisces in a normal

manner, despite the abortion of the ovules. This parthe-

nocarpic development of Salix capsules deserves further

investigation.

Structure of the plumed seed and seed coat

SEM micrographs of fertile seed taken from herbarium

sheets ( Codd 8246 PRE; Obermeyer TM31027 PRE)

show a small, annular structure with numerous long hairs

attached to its perimeter, positioned at the micropylar side

of the seed (Figure 3A, D). These hairs form the plume or

coma on the seed. The free underside of the ring has a

rough surface of broken cells (Figure 3B). When eased

away from the seed (Figure 3C), the upper (micropylar)

side of the ring displays an entire, glabrous exterior

(Figure 3D), suggesting that the ring has enveloped the

base of the seed, without forming an integral part of it. In

structure, origin and position the hairy hilar ring repre-

sents a kind of seed appendage known as a funicular or

hilar aril (Boesewinkel & Bouman 1984: 592). We pro-

pose that this ring has developed from the hilar meristem

in the distal part of the funicle and, during abscission of

the seed, has broken away from the funicle.

Seeds are small (1.3-1. 6 mm long), exalbuminous

and contain straight, chlorophyllous embryos almost fill-

ing the embryo sac, except for an air space underneath

the seed coat (Figure 3E). The seed coat is undifferenti-

ated and the number of layers has not increased during

ovule-to-seed development (compare Figures 2 A, B;

3E). The outer epidermis has unevenly thickened outer

periclinal walls and, in surface view, the uneven thicken-

ings form an irregular micro-morphological pattern

(Figure 4A). Between the thickened parts of the walls the

thinner parts collapse so that numerous, minute depres-

sions are formed on the seed surface (Figure 4A).

The hairs on the aril resemble the placental-funicular

hairs (loose wool) and consists of a mass of seemingly

Bothalia 34,1 (2004)

55

FIGURE 1. — Structure, position and placentation of ovules in Salix mucronata as seen in median sagittal sections. A, young female flower, note

stalked ovule primordia (black arrows); B, ovule primordium in megaspore mother cell stage (white arrowhead) showing parietal nucellar

cells (curved white arrow); C, young, anatropous ovule in megaspore tetrad stage, position of micropyle indicated by black arrowhead, a,

adaxial nectary; b, subtending bract; c, bicarpellate ovary; d. parietal placenta; e, integument; f, funicle; g, functional chalazal megaspore;

i, nucellus. Scale bars: A, 100 pm; B, C, 50 pm.

structureless, cylindrical fibres about 3-4 mm long, with

smooth, transparent, thin walls without pits (Figures

3A-D; 4B). The walls stain blue-green with toluidine blue

and yellow with aniline sulfate, indicating the presence of

lignin. The fibres taper to a point and have a slightly bul-

bous base. When immersed in water, numerous air bub-

bles are trapped in the wide lumen (Figure 4B).

Distance of flight, buoyancy and wettability of seeds and

hairs

Experiments in the garden and laboratory showed that

fertile, wool-covered seeds do not remain airborne for more

than ten metres in a moderately strong wind; after the loose

wool (intra-ovarian hairs supporting the seed) has been

blown apart, the seed with hairy aril (plume) still attached,

slowly descends. The flight distance of sterile ‘seeds' could

not be determined; they flew away so fast that they could not

be retrieved. When the wool-covered seed lands in water, the

loose hairs rapidly float away, but the plume remains

attached to the seed. Loose wool, plumed seed and de-

plumed seed remain buoyant in clear and muddy water until

the water in the containers has evaporated (about five days).

The seed coats of floating seeds are unwettable (hydropho-

bic) and appear translucent white in water because of the

numerous, minute air bubbles trapped in the depressions of

the outer periclinal wall. By the time the muddy water in the

containers has evaporated, the plumes have got stuck in the

mud and the seeds separate easily from the arils.

56

Bothalia 34,1 (2004)

FIGURE 2. — Development of ovule and intra-ovarian hairs in Salix mucronata. A, mature ovule in sagittal section showing hilar meristem (black

arrows) in distal part of funicle; B, I/s micropylar part of ovule with tip of embryo sac breaking through epidermis of nucellus, but stay-

ing inside integument; C, I/s developing capsule showing intra-ovarian hairs originating from placenta and funicle. D, part of sterile cap-

sule with abortive, stalked ovule imbedded in intra-ovarian hairs, note position of disintegrated hilar meristem (small black arrows), black

arrowhead indicates position of micropyle. e, integument; f, funicle, i, nucellus; j, embryo sac; k, intra-ovarian hairs; o, ovule; ol, abortive

ovule; w, ovary wall. Scale bars: A, B, D, 50 pm; C, 100 pm.

Bothalia 34.1 (2004)

57

FIGURE 3. — Seed structure in Salix

mucronata as seen in SEM

micrographs (A-D) and LM

photographs (E). A, plumed

seed with coma of hairs at-

tached to hilar aril; B, micro-

pylar seed region showing

aril and hairs at higher mag-

nification than in A; C, aril

detached from micropylar

part of seed; D. aril as seen

from above at higher magni-

fication than in C; E, 1/s fer-

tile seed showing large chloro-

phyllous embryo surrounded

by air space (black arrows),

note irregular wall thicken-

ings (open black arrow) on

outer periclinal walls of epi-

dermis, curved white arrow,

aril, m, embryo; n, undiffer-

entiated seed coat; s, seed.

All scale bars: 100 pm.

DISCUSSION

The coma of hairs on Salix seed

It has often been alleged that the tuft of silky hairs

enveloping the seed of willows is placental in origin

(Takeda 1936: Van der Pijl 1969; Corner 1976; Johri et

al. 1992). Others (Ridley 1930; Wilmot-Dear 1991;

McKean 1996; Boulos 1999) mention that the hairs are

derived from the funicle. Our observations on ovule-to-

seed development in Salix lend support to the latter view:

sagitally sectioned, developing capsules distinctly show

the presence of stalked ovules with the coma of hairs

resulting from epidermal cells on the most distal part of

the funicle. Takeda's (1936) erroneous conclusion that

Salix ovules have no funicles and that the hairy tuft on

the seed is part of the placenta was based on incorrectly

sectioned capsules. His line drawings (Takeda 1936; 285,

figs 1 & 4) show ovules in longitudinal (not sagittal) sec-

tion in which the funicular connections to the placenta

FIGURE 4. — Seed structure in Salix

mucronata as seen in SEM

micrograph (A) and LM pho-

tograph (B). A, structure of

seed surface; B, hilar aril

(curved white arrow) with air-

filled hairs (black arrows).

Scale bars: A, 20 pm; B, 50

pm.

58

Bothalia 34,1 (2004)

were not visible. Later, well-known embryologists (Van

der Pijl 1969; Corner 1976; Johri et al. 1992), accepted

Takeda's (1936) interpretation, without re-investigating

the matter.

The present report provides the first evidence for the

occurrence of arillate seeds in Salix. The presence of

hairy arils in Salicaceae was suspected, but not substan-

tiated by Corner (1953), or mentioned in subsequent

work (Corner 1976) and was not reconcilable with the

supposed placental origin of the coma (Van der Pijl

1969). Since it is from the narrow distal region of the

funicle that the small, annular and tufted aril arises and

breaks away to form an appendage to the seed during

seed abscission, we have referred to it as a hilar aril.

The micromorphological structure of the funicular-

placental and arillate hairs in Salix is here described and

illustrated for the first time. LM photographs and SEM

micrographs show the mass of hairs consisting of cylin-

drical, structureless, unpitted, lignified and thin-walled

fibres that remain filled with air when immersed in

water. In these characters, Salix fibres closely resemble

those found in the capsules of Bombacaceae, e.g Ceiba

pentandra Gaertn. (kapok tree) and Bombax malabar-

icum DC. (Indian kapok or Semal tree) as described by

Kirby (1963: 389) and Ilvessalo-Pfaffli (1995: 358) and

in Chorisia speciosa A.St.-Hil., the silk-floss tree

(E.M.A. Steyn pers. obs.). Before being largely replaced

by synthetic materials, kapok fibres were used for insu-

lating and stuffing purposes (Kirby 1963). Kapok owes it

use in e.g. life-saving equipment to the fact that the fibres

remain buoyant after long periods in water (owing to its

property of repelling water) and because it can easily be

dried when it has become wet (Kirby 1963). According

to Ilvessalo-Pfaffli (1995) the high buoyancy of kapok is

due to the presence of air bubbles in the lumen of the

fibres which apparently is also the case in Salix fibres.

Dispersal by wind (anemochoiy)

Confined to the rheophytic habitat, i.e. in the beds of

swift-running rivers and streams and on the banks up to

Hood level (Jordaan 2002), Salix mucronata is exposed

to up-and-down-stream breezes all the year round. When

the capsules ripen and dehisce, the placental-funicular

fibres dry out, break loose from the subepidermal tissue,

expand and protrude from the bivalvate fruit. During

tests on dehisced fruit, gusts of wind jerked the seed-

bearing wool from the capsule, carrying it into the air and

keeping fertile seeds airborne for about ten metres. At

that distance the loose wool was blown apart, the hairy

arils alone could not keep fertile seeds airborne and they

descended. We propose that the same occurs in nature.

Wind is therefore the releasing and the primary dispersal

agent of Salix seed. It dislodges the seed from the capsule

and scatters it some distance away from the parent plant.

If the landing surface is not wet, the plumed seed may be

blown further along the ground, but will stick to a wet

substrate, e.g. mud, feathers or hair of wading birds and

animals, bringing the role of wind to an end.

Characters enabling the seed to become airborne in-

clude its relative smallness, the absence of a large, heavy

funicle and low specific weight. The latter is achieved by

the thin, undifferentiated seed coat (testa) with air-filled

cells, an air space below the testa caused by the early dis-

appearance of endosperm and enlargement of the seed

surface by the long, smooth, air-filled hairs of the aril

and seed-bearing wool.

Dispersal by water ( hydrochory )

In its rheophytic habitat, the most important sec-

ondary dispersal agent for Salix seed is water. Our tests

suggest that seed, landing in the bed of the stream or on

the banks when the waters are in spate, will have the abil-

ity to float for days. In this way seeds may spread along

the river edge far away from the parent plant or, caught

in a strong current, they may migrate for many kilome-

tres. Structural modifications for anemochory listed

above, would increase buoyancy (Melcher et al. 2000).

An additional adaptation for hydrochory in Salix

mucronata is the unwettable seed surface. Van der Pijl

(1969: 63) and Rauh et al. (1975: 372) reasoned that air

bubbles, trapped in the numerous depressions in the seed

coat epidermis, would have such an effect. In Salix this

characteristic is aided by the presence of a unique float-

ing apparatus in the form of a small (so as not to unduly

increase seed weight), hilar aril with unwettable, air-

filled hairs. The coma of hairs also ensures that the float-

ing seed is not taken beyond the boundaries of the rheo-

phytic habitat; when floodwaters recede, the fibres stick

to wet mud, retaining the seed (Ridley 1930: 226). In

these watery surroundings, the thin testa does not need to

protect the embryo against desiccation and offers little

resistance to germination. Salix seed germinates easily,

seedlings have strong root systems and rapidly develop

their first leaves (Pax 1894).

Fortuitous dispersal by animals (epizoocliory)

Distribution of seed to upstream locations cannot

occur by means of water and animal visitors to the water

may play an important role here; Salix seeds have been

removed from the coats of animals when the plants were

in fruit (Ridley 1930: 554). It was found that the plumes

attach themselves easily to the hair of animals when the

vegetation is wet. The plumed seed may also become

entangled in the feathers of waterfowl. To be carried in

the mud on the feet of birds and animals, seeds have to

be small and Salix seed certainly falls in this size group.

While the plumes remain stuck in the mud, the small

seeds may be picked up by the feet of waterfowl, hip-

popotamus, elephant and other animals and conveyed to

another suitable rheophytic habitat.

Taxonomic implications

In a report on the phylogenetic relationships of Salix

based on rbcL sequence data, Azuma et al. (2000: 71)

stated that in their analysis 7 desia and Dovyalis of

Flacourtiaceae are the sister groups of the Salicaceae’.

Leskinen & Alstrom-Rapaport (1999) found that Idesia

polycarpa Maxim, shows great sequence similarity with

Salicaceae and that the 5.8S rDNA sequences in their

study suggest that Flacourtiaceae and Salicaceae may

have shared a common ancestor within the order Violates.

Chase et al. (2002) provide evidence that if a wider range

of flacourt genera is included in an analysis, Itoa Hemsl.

and Poliothyrsis Oliv. (Flacourtieae: Flacourtiaceae)

seem to be the closest relatives of Salicaceae s.str.

Bothalia 34,1 (2004)

59

As far as embryological data are concerned, Meeuse

(1975) compared anther and ovule characters of Salicaceae

s.str. and Flacourtiaceae s.l. and found 'many points of

agreement' (Meeuse 1975: table 1). To these shared charac-

ters should be added the presence of arillate seeds. The

presence of an aril on the seed of Salix offers some mor-

phological support for combining on phylogenetic grounds

Salicaceae s.str. with several flacourtiaceous taxa, members

of which are reported to often have arillate seed (Chase et

al. 2002). Several other authors who have previously sug-

gested a close relationship between Salicaceae and some

Flacourtiaceae are listed in Chase et al. (2002). However, if

the putative close relationship between Salicaceae s.str. and

Flacourtiaceae (Meeuse 1975; Chase et al. 1996; Nandi et

al. 1998; Chase et al. 2002) is accepted, a radical change

must have occurred during the evolution of the unitegmic,

salicaceous seed with its undifferentiated seed coat. In

Flacourtiaceae the seed coat develops from a bitegmic ovule

and contains a fibrous exotegmen (Corner 1976; John et al.

1992) that protects the zoochorous seed, often produced in

edible berries (e.g. Idesia , Flacourtia L'Her., Dovyalis)

against animal feeders. No rheophytes have been reported

in Flacourtiaceae (Van Steenis 1981) and anemochorous

seed is extremely rare but may, according to Sleumer ( 1954)

be granted to the winged seed of Itoa. Poliothyrsis also has

winged seed (Judd et al. 2002). It is noteworthy that these

two anemochorous genera are probably the sister groups of

Sailicaceae s.str. (Chase et al. 2002) — both are from China

with Itoa also in tropical Asia. To our knowledge, theiur seed

coat structure has not been reported.

CONCLUSION

We propose, therefore, that the total absence of an

inner integument in Salix ovules and its suppression in

early ovular stages in Populus (Nagaraj 1952) might be

due to an adaptive change from animal to wind and water

dispersal. This integument with its exotegmic protective

layers was no longer necessary and would only increase

seed weight. This could be another example of the well-

known evolutionary tendency for unused structures, or

unnecessary complication of structures, to degenerate

(Cronquist 1988: 232). A study of ovule-to-seed devel-

opment in Dovyalis is in progress and may shed more

light on the possible evolutionary changes that have led

to the Salicaceae s.str. clade.

ACKNOWLEDGEMENTS

The National Botanical Institute, Pretoria is acknowl-

edged for providing the infrastructure to execute this

investigation. Thanks are due to Dr Sarie Perold for

assisting at the scanning electron microscope.

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Bothalia 34,1: 61-71 (2004)

Grass assemblages and diversity of conservation areas on the coastal

plain south of Maputo Bay, Mozambique

S.J. SIEBERT*, L. FISH**, M.M. UIRAS*** and S.A. IZIDINEt

Keywords: dunes, forest, grasses, grassland, Maputaland, Mozambique, Poaceae, woodland

ABSTRACT

A floristic analysis of the grass species assemblages of the Licuati Forest and Maputo Elephant Reserves south of

Maputo Bay, Mozambique, is presented. Sampling of grass data was undertaken in six previously described, major vegeta-

tion types. TWINSPAN divisions distinguished grass assemblages that are characteristic for these major vegetation types of

the study area. The results were supported by an Indirect Gradient Analysis. Further TWINSPAN divisions of a larger

Maputaland data set indicated a floristic relationship between grass assemblages of similar major vegetation types in the

study area and South Africa. This relationship was supported by high similarity values (> 65%), obtained with Sorenson’s

Coefficient. The coefficient also indicated varying degrees of similarity between grass assemblages of different major veg-

etation types within the study area. A rich diversity of 1 15 grass species and infraspecific taxa was recorded for the study

area. The Chloridoideae and Panicoideae dominate the grass diversity and the genera with the most species include

Eragrostis, Panicum and Digitaria. Most grass species in the study area are perennials and have a tufted growth form, but

this varies considerably between vegetation types.

INTRODUCTION

Despite the importance of the Poaceae to both subsis-

tence and commercial agriculture (Myre 1971; Tainton et

al. 1976) and the continued documentation of the rich

diversity of this family in southern Africa (Gibbs Russell

et al. 1990; Kobisi & Kose 2003), some areas still remain

poorly studied and documented. One particular area is

the coastal plain of Mozambique south of Maputo Bay.

Although Myre (1964) and De Boer et al. (2000) provid-

ed a comprehensive account of the vegetation south of

Maputo Bay in Mozambique, the descriptions of the

grass layer contained limited taxonomic and floristic

information.

In this paper the grass diversity of the coastal plain

south of Maputo Bay is revisited to investigate the fol-

lowing hypotheses based on current knowledge: 1, dif-

ferent grass assemblages characterize the major vegeta-

tion types of the study area; 2, a floristic relationship

exists between grass assemblages of different major

vegetation types within the study area; 3, there is a

floristic relationship between grass assemblages of the

study area and similar vegetation types in South Africa;

4, species that form the grass assemblage for a major

vegetation type are characterized by certain life/growth

forms; 5, dominance of different Poaceae subfamilies in

the study area correspond with predictions made previ-

ously; and 6, there is a rich diversity of grasses in the

study area.

* Department of Botany, University of Zululand, Private Bag XI 001,

3886 KwaDlangezwa.

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

*** National Botanical Research Institute, Private Bag XI 3 184, Wind-

hoek, Namibia.

t National Institute for Agronomic Research, P.O. Box 3658, Mavalane,

Maputo, Mozambique.

MS. received: 2003-03-14.

STUDY AREA

The study area comprises the Maputo Elephant Reserve

and Licuati Forest Reserve on the Maputaland coastal plain,

south of Maputo Bay in Mozambique (Figure 1). Ma-

putaland is an important centre of plant endemism and

diversity of Mozambique, South Africa and Swaziland (Van

Wyk & Smith 2001), defined as the biogeographical area

bounded by the Inkomati-Limpopo River in the north,

Indian Ocean in the east, foothills of the Lebombo

Mountains in the west and St Lucia estuary in the south.

The topography comprises high, linear, north-south ori-

ented dune cordons along the inland margin of the coast.

The youngest of these dunes are probably 10 000-30 000

years old, making them in geological terms some of the

youngest formations in southern Africa (Botha 1997).

These high dune cordons mark a succession of marine

regressions that deposited these sediments. Marine silt-

stone underlies these sediment deposits and in turn, the

sediment deposits underlie the dune sand deposits current-

ly defining the surface relief in this area.

Maputaland lies within a transitional zone between

the tropics and subtropical coastal conditions to the south

(Bruton & Cooper 1980), with warm to hot summers

(mean of 27°C in January) and cool to warm winters with

no frost (mean of 16°C in July). Mean relative air humid-

ity is high along the coast, namely 55% in August and

90% in February. Summers are wetter than winters,

although rain is received throughout the year. Mean

annual rainfall is higher along the coast (1 100 mm/year)

and declines progressively inland (600 mm/year).

Morning mist is common in the dry season.

The study area comprises the major terrestrial vegeta-

tion types recognized and defined for this floristic region

by Myre (1964), De Boer et al. (2000) and Matthews et

al. (1999, 2001), namely Coastal Woodland, Dune

Forest, Licuati (Sand) Forest, Primary Dunes, Reed Beds

and Woody Grassland.

62

Bothalia 34,1 (2004)

METHODS

Sampling sites were randomly selected using topograph-

ic maps and aerial photographs. Plots were placed in vege-

tation types conforming to the descriptions of Matthews et

al. (1999, 2001 ), avoiding the edges of these and refraining

from sampling in disturbed areas. Presence/absence of diag-

nostic woody species was used to identify vegetation types

in the field. A vegetation type is a composition of species

that recurs in a region as a result of specific combinations of

environmental factors (Barbour et al. 1999). Reed Beds

could not be sampled adequately due to high water tables.

However, wetlands that feed into the Reed Beds were sam-

pled, and these are referred to as Hygrophilous Grassland

for the purpose of this study.

Twenty-one sites were sampled at the beginning of sum-

mer 2001, the peak flowering season for grasses. At each

site all the grass species in a 20 x 20 m grid were collected

and identified. Scientific names conform to Fish (2003).

Voucher specimens are listed in a checklist (Appendix 1)

and housed at the Maputo Herbarium (LMA), with dupli-

cates in the Luanda Herbarium (LUAI), Natal Herbarium

(NH), Pretoria National Herbarium (PRE) and University

of Zambia Herbarium (UZL). The LMA collection was

consulted to identify sterile specimens and to locate fertile

voucher specimens for these species. If none were found,

sterile specimens were provisionally identified, but listed

without voucher specimens in the checklist.

FIGURE 1 . — Locality of the Maputo

Elephant and Licuati Forest

Reserves in Mozambique and

the Tembe Elephant Park and

Sileza Nature Reserve in South

Africa.

A floristic classification of grass data for the

Licuati Forest and Maputo Elephant Reserves was ob-

tained by the application of Two-Way Indicator Species

Analysis (TWINSPAN) (Hill 1979a). The multivariate

analysis divided the data set, comprising all collected

grass species and their presence/absence data per plot,

into nodes. In MEGATAB (Hennekens 1996) the re-

sultant classification was summarized in a synoptic

table to reflect percentage occurrence of each species

per node (major vegetation type) and refined with

Braun-Blanquet procedures to group species in as-

semblages. Detrended Correspondence Analysis

(DECORANA) (Hill 1979b) was applied to the data

set to illustrate vegetation gradients and floristic rela-

tionships.

A second data set containing grass species pres-

ence/absence data from the study area was merged with

data from studies conducted in Sileza Nature Reserve

(Matthews et al. 1999) and Tembe Elephant Park

(Matthews et al. 2001 ). Vegetation types from these con-

servancies in South Africa were chosen for comparison

because of their similar grass flora, status as pristine

environments, proximity to the study area, and formal

classification and description as representative of north-

ern Maputaland. The TWINSPAN divisions depicting

the floristic relationship are illustrated in a dendro-

gram (Figure 2).

Bothalia 34,1 (2004)

63

FIGURE 2. — Dendrogram of a TWINSPAN division depicting the floristic relationships between the grass assemblages of the major vegetation

types of four protected areas of the Maputaland coastal plain in Mozambique and South Africa. LFR, Licuati Forest Reserve; MER,

Maputo Elephant Reserve; SNR, Sileza Nature Reserve; TEP, Tembe Elephant Park.

The species richness of grasses was calculated as the

number of species per major vegetation type (homoge-

neous unit) in the study area (Whittaker 1978). Rare and

threatened species were identified as either endemic (Van

Wyk 1996) and/or Red Data List (Izidine & Bandeira

2002). Character species were defined as species that are

relatively restricted to specific vegetation types (West-

hoff & Van der Maarel 1978). Recognition as an intro-

duced alien species was based on Fish (2003). Life cycles

(annual or perennial) and growth form (tufted, rhizoma-

tous and/or stoloniferous) of each species followed Gibbs

Russell et al. (1990). Sorenson’s Index (Mueller-Dom-

bois & Ellenberg 1974) was used to determine the beta

diversity between the vegetation types of the study area

and between different reserves in Maputaland.

Floristic diversity of the Poaceae subfamilies/tribes,

and the distribution of these taxonomic units within the

different vegetation types were presented in tables to

interpret and compare current patterns of grass diversity

with what was previously predicted for southern Africa

(Gibbs Russell 1986, 1988).

RESULTS AND DISCUSSION

The resultant hierarchical division of the sample plots

from the TWINSPAN analysis (Table 1) resulted in the

classification of distinct grass assemblages associated

with six major terrestrial vegetation types of the Ma-

putaland coastal plain in Mozambique, namely Coastal

Woodland, Dune Forest, Hygrophilous Grassland, Licuati

Forest, Primary Dunes and Woody Grassland. The ordi-

nation clusters (Figure 3) obtained for the first and second

axes tended to substantiate the groups identified in the

TWINSPAN classification. A TWINSPAN division of the

Mozambican grass data merged with sample plots from

South Africa resulted in a hierarchy of assemblages that

confirms that a floristic link exists between these differ-

ent parts of Maputaland (Figure 2).

Grass assemblages

1. Grass assemblage of Primary Dunes

Locality: Maputo Elephant Reserve

This assemblage is restricted to the upper reaches of

beaches, bordering on the seaward side of primary dunes.

Grasses are mostly pioneers and are not only associated

with the unstable seashore dunes, but also occur further

inland along freshwater and saline marshes, and in dis-

turbed places such as road reserves. Grasses of the

assemblage prefer light shade, but tolerate full sun.

Character species for this grass assemblage are given

in species group A (Table 1). The assemblage is species

poor and although predominantly characterized by

perennials, has the highest proportion of annuals (36%)

in the study area. It also has the highest proportion of

grass species with a stoloniferous growth form (35%).

One naturalized alien grass, Cenchrus brownii, was

recorded. Taxonomically the vegetation type is unique in

that it is not characterized by the Paniceae (Panicoideae)

as in the grass assemblages of the other vegetation types

(Table 2), but is dominated by the Eragrostideae

(Chloridoideae). The community is floristically most

related to Hygrophilous Grassland (Sorenson Coefficient

= 15%; Table 1) (Figure 3).

2. Grass assemblage of Hygrophilous Grassland

Locality: Maputo Elephant Reserve

Hygrophilous Grassland represents a vegetation type

of open grassland on seasonally wet, sandy or clay soils.

These seasonally wet areas may occur as inter-dune

64

Bothalia 34,1 (2004)

FIGURE 3. — Relative positions of sampling plots along the first and second ordination axes. Numbers refer to plots; polygons group the plots into

grass assemblages.

depressions, pans, lake seepages, riverbanks and flood-

plains, and are characterized by high levels of organic

matter and a water table of less than one metre below the

soil surface. Soils are also less sandy and with a higher

clay content. The biomass of the hygrophilous grass

component is the highest due to dense growth under con-

stant wet conditions.

Character species for the grass assemblage are given in

species group B (Table 1). This grass assemblage is

species rich (56 taxa) and tufted perennials dominate. It

also has the highest proportion of rhizoinatous taxa (36%).

There are 15 character grass species, which is the most for

any of the major vegetation types. This is the only grass

assemblage in the study area characterized by representa-

tives from the Arundineae, Aveneae and Oryzeae (Table

2) . Floristically this assemblage is most related to Woody

Grassland and Dune Forest (Sorenson Coefficient = 56%

and 57% respectively; Table 1 ) (Figure 3). It also shows a

high similarity in grass species composition when com-

pared with plots from Maputaland Hygrophilous Grass-

land in South Africa (Sorenson Coefficient = 76%; Table

3) (Figure 2).

3. Grass assemblage of Woody Grassland

Locality: Maputo Elephant Reserve

Extensive grasslands occur along the coast south of

Maputo Bay and is characterized by deep, well-drained

sandy to loam soils and an undulating topography (dunes

and floodplains). This grassland type is endemic to

Maputaland and is dominated by geoxylic suffrutices

which are dwarf woody plants with annual or short-lived

shoots sprouting from woody, perennial underground

axes. Woody Grassland is not as rich in grass species as

the Hygrophilous Grassland.

Character species of the grass assemblage are given in

species group D (Table 1 ). A low number of four charac-

ter species were recorded, because grassland represents a

transition between other major vegetation types. This

assemblage is characterized by tufted, perennial species.

The Andropogoneae (Panicoideae) dominate the assem-

blage (Table 2). One rare grass species, namely the Ma-

putaland endemic Trichoneura schlechteri , was recorded.

Floristically it is most related to Dune Forest, Coastal

Woodland and Hygrophilous Grassland (Sorenson Co-

efficient = 62%, 58% and 56% respectively; Table 1)

(Figure 3). Of all the grass assemblages in the study area,

it has the lowest similarity in grass species composition

when compared with similar vegetation further south in

South Africa (Sorenson Coefficient = 63%; Table 3)

(Figure 2).

4. Grass assemblage of Dime Forest

Locality: Maputo Elephant Reserve

Forests mainly occur on well-established secondary

dunes and further inland. Soils are moist, deep and sandy.

Forest trees have a higher diversity of creepers and under-

storey vegetation than the vegetation of surrounding areas.

Grass species of Dune Forest prefer semi-shade and are

especially common along forest margins, riverbanks, and

partially disturbed and overgrazed areas.

Seven character species for this grass assemblage are

given in species group F (Table 1). This grass assem-

blage is the most species rich (57 taxa). It is character-

ized by tufted perennials, has the highest number of

recorded rare grass species (5) and taxonomically it is

dominated by the Panicoideae, although the Eragrosti-

deae, a tribe of the Chloridoideae, supports the highest

diversity (Table 2). Along the coast, Dune Forest tends to

Bothalia 34,1 (2004)

65

TABLE 1. — Diagnostic grass assemblages of the major vegetation types of the Maputaland coastal plain south of Maputo Bay in Mozambique.

p, perennial; a, annual; s, stoloniferous; t, tufted; r, rhizomatous

66

Bothalia 34,1 (2004)

TABLE 1. — Diagnostic grass assemblages of the major vegetation types of the Maputaland coastal plain south of Maputo Bay in Mozambique,

p, perennial; a, annual; s, stoloniferous; t, tufted; r, rhizomatous (cont.)

Bothalia 34,1 (2004)

67

TABLE 2. — Numbers of species of Poaceae subfamilies/tribes recorded

within major vegetation types of Maputaland, south of Maputo Bay

* CW, Coastal Woodland; DF, Dune Forest; HG. Hygrophilous Grass-

land; LF, Licuati Forest; PD, Primary Dunes; WG, Woody Grassland

be continuous, but inland it forms a mosaic with other

vegetation types. Its grass assemblage therefore shows a

strong floristic link with Woody Grassland, Coastal

Woodland, Hygrophilous Grassland, and to a lesser ex-

tent Licuati Forest (Sorenson Coefficient = 62%, 62%,

57% and 39% respectively; Table 1) (Figure 3). It also

shows a high similarity in grass species composition

when compared with plots from Maputaland Dune Forest

in South Africa (Sorenson Coefficient = 71%; Table 3)

(Figure 2).

5. Grass assemblage of Coastal Woodland

Locality; Maputo Elephant Reserve and Licuati

Forest Reserve

Savanna covers extensive areas along the coast south

of Maputo Bay and stretches westwards to the foothills

of the Lebombo Mountains. The vegetation type occurs

on sandy soils in drier habitats with a deeper water table.

The grass assemblage forms a dense herbaceous layer

and is associated with light shade or full sunlight and

TABLE 3. — Similarity indices of grass assemblages shared between

major Maputaland vegetation types in Mozambique (Moz) and

South Africa (RSA)

Total Total Shared Sorenson

Vegetation species species between Index

type* (Moz) (RSA) Moz and RSA (Ss)%

occurs in a wide range of habitats including floodplains,

dune crests, drainage lines, transition zones and dis-

turbed areas.

Nine character species were recorded for this assem-

blage and are given in species group J (Table 1). Tufted

perennials typically dominate this grass assemblage.

Two rare species were recorded and one naturalized alien

grass, Coix lacryma-jobi, has colonized this assemblage.

This is the only grass assemblage that contains a member

of the Pappophoreae and is dominated by the Pani-

coideae (Table 2). Dune Forest forms localized patches

within Coastal Woodland, and in turn. Coastal Woodland

within Woody Grassland. Hence, its grass assemblage

shows a strong floristic link with Dune Forest and Woody

Grassland, and to a lesser extent with Licuati Forest on

which it borders (Sorenson Coefficient = 62%, 58% and

41% respectively; Table 1) (Figure 3). The assemblage is

similar to that of corresponding Maputaland woodlands

in South Africa (Sorenson Coefficient = 67%; Table 3)

(Figure 2).

6. Grass assemblage of Licuati Forest

Locality: Licuati Forest Reserve

This vegetation type is endemic to Maputaland and

has many rare plant species. It is restricted to the ancient

coastal dunes of Maputaland, and drier (600 mm per

annum) conditions than most of the other coastal forest

types in southern Africa. This grass assemblage prefers

moister habitats in shady places and is common along

forest margins. Based on vegetation structure and species

composition the forests of ancient dunes in Maputaland

can be divided into Licuati Forest and Licuati Thicket

(Izidine et a\. 2003).

Character species of the grass assemblage are given in

species group M (Table 1). Although it is a forest system

and tufted perennials dominate the grass assemblage, this

vegetation type has a high proportion (24%) of annual

species. The assemblage is species poor, but a high pro-

portion of three rare species are recorded for the assem-

blage. This is the only vegetation type in the study area

with representatives from the Arundinelleae, Centotheceae

and Ehrharteae, and the only one without a representative

of the Andropogoneae (Table 2). This grass assemblage is

related to Coastal Woodland and Dune Forest as a result of

similar microhabitats (Sorenson Coefficient = 41% and

39% respectively; Table 1) (Figure 3). It also shows a high

similarity in grass species composition when compared

with plots from Maputaland sand forest in South Africa

(Sorenson Coefficient = 71%; Table 3) (Figure 2).

Floristic analysis

In a provisional checklist compiled from available lit-

erature for the two Maputaland reserves in Mozambique,

the Poaceae numbered 52 species/infraspecific taxa and

36 genera. Currently, with 95% of the collected speci-

mens identified, the updated checklist (Appendix 1) con-

tains 115 species/infraspecific taxa and 56 genera.

However, 15 species previously recorded were not col-

lected again. Most of these taxa are either locally rare

(e.g. Panicum genuflexum and Triraphis andropogo-

noides) or wetland species (e.g. Leersia hexandra and

68

Bothalia 34.1 (2004)

TABLE 4. — Floristic analysis of subfamilies and tribes recorded for Maputo Elephant and Licuati Forest Reserves

*CW, Coastal Woodland; DF. Dune Forest; HG, Hygrophilous Grassland; LF, Licuati Forest; WG, Woody Grassland

Dinebra retroflexa var. condensata), for which high

water tables made their localized habitats inaccessible.

In the study area the most commonly represented sub-

families are the Panicoideae with 66 species/infraspecif-

ic taxa (accounting for 57% of the total checklist) and

Chloridoideae with 39 taxa (34%) (Table 4). Together

they account for 91% of the species in the checklist. The

tribes with the most species are the Paniceae (41 taxa),

Eragrostideae (32 taxa) and Andropogoneae (24 taxa)

(Table 4). The genera with the most species are Era-

grostis (12), Panicum ( 10), Digitaria (8) and Sporobolus

(6). Andropogon, Brachiaria and Dactyloctenium are repre-

sented by four species each.

Interesting species were recorded, such as Panicum

kalaharense (usually associated with much drier areas on

Kalahari sands) and Monocymbium ceresiiforme (usual-

ly associated with much higher altitudes on mountains).

Many grass species of direct significance for conserva-

tion initiatives, reserve management and sustainable uti-

lization were also recorded:

1 , three Maputaland endemic grasses belonging to the

Eragrostidae (Van Wyk 1996). Brachychloa schiemanni-

ana occurs in Dune Forest, Eragrostis moggii var. moggii

is associated with Licuati Forest and Dune Forest, and

Trichoneura schlechteri is found in Coastal Woodlands,

Dune Forest, Licuati Forest and Woody Grassland.

Brachychloa fragilis was not recorded, although it is a typ-

ical endemic associated with deep sands in South Africa;

2, three alien grasses belonging to the Panicoideae. As is

the case in many Centres of Endemism in the world (Stohl-

gren et al. 1999), alien taxa have also invaded Maputaland.

Cenchrus brownii has invaded vegetation of Primary Dunes

and Coix lacryma-jobi and Digitaria didactyla occurs in

Coastal Woodland in the vicinity of villages;

3, two Red Data List grass species: Panicum plei-

anthum from Dune Forest which is assessed as Low Risk

(Izidine & Bandeira 2002), and Alloteropsis papillosa

from Dune Forest, Coastal Woodland and Licuati Forest

which is assessed as Insufficiently Known (Hilton-

Taylor 1996). Both species are locally common and

known to occur as far north as Kenya and Tanzania;

4, eight grasses that are important for rural livelihoods.

Some species are used as important components to build

huts: Phragmites australis for the walls and Cymbopo-

gon excavatus , Imperata cylindrica and Hyperthelia clis-

soluta to thatch the roof (Mangue 1999). Dactyloctenium

giganteum, Eleusine coracana subsp. africcma , Echino-

chloa pyramidalis and Sorghum bicolor subsp. arundi-

naceum are used as indigenous grass cereals (Scudder

1971).

Correlation analyses of the floristic data did not reveal

any meaningful relationships. The only significant posi-

tive correlation (n = 6; r = 0.827; P = 0.04) was obtained

between the number of rare/endemic grass species and

the number of annual grass species per vegetation type.

Forests/woodlands have higher numbers of rare/endemic

and annual grass species than grasslands.

Distribution patterns

All known distributions of the six grass subfamilies

extend into southern Mozambique. The diversity of the

Chloridoideae and the Panicoideae in the study area was

expected and subsequently were the best represented in

the major vegetation types. The dominance of the Pani-

coideae (both C3 and C4 grasses) in the terrestrial vegeta-

tion types coincides with the centre of diversity of the

subfamily in mesic summer rainfall regions (Gibbs Rus-

sell 1986). However, its diversity was much lower than

would be expected (19% instead of the predicted

46-60%) (Table 4). The diversity of species of the Chlori-

doideae (mainly C4 aspartate producers) was within the

expected range of 16-30% (17% of the subfamily’s

species was recorded) (Table 4), as this subfamily’s cen-

tre of diversity is mainly situated further north in arid

summer rainfall regions (Gibbs Russell 1986). Chlori-

doideae dominated the grass assemblage of the Primary

Dunes. The species diversity of the other four subfami-

lies was as predicted by Gibbs Russell (1988). Arundi-

Bothalia 34,1 (2004)

69

noideae is dominant in Hygrophilous Grassland and the

remaining three subfamilies are associated mainly with

Hygrophilous Grassland and Licuati Forest.

The tribes, Eragrostideae (C4 aspartate), Paniceae (both

C3 and C4) and Andropogoneae (C4 malate) dominate the

grass assemblages of the major vegetation types, probably

due to specific regional climatic conditions, such as high

temperatures at the local scale, which favour these C4

groups to successfully colonize specific habitats. Eragrosti-

deae dominate the grass assemblages of Dune Forest and

Primary Dunes, the Paniceae dominate in Coastal Wood-

land, Hygrophilous Grassland and Licuati Forest, and the

Andropogoneae in Woody Grassland (Table 2). The Aristi-

deae and Cynodonteae are found in nearly all the vegetation

types, but are restricted to a maximum of four species per

vegetation type. This is low when compared to the maxi-

mum of 21, 18 and 15 species respectively recorded for the

three dominant tribes. The remaining seven tribes are

restricted to single vegetation types. Three of these tribes

are associated with Licuati Forest, two with Hygrophilous

Grassland and one with Coastal Woodland.

CONCLUSIONS

It is evident from the analysis of the grass diversity on

the coastal plain of southern Mozambique that a specific

grass flora is present and that six species assemblages are

associated with and characteristic of certain major vege-

tation types of the coastal plain.

The grass assemblage of Dune Forest was qualitative-

ly the most similar and central to the grass assemblages

of the other vegetation types south of Maputo Bay.

Primary Dunes have the most floristically unrelated

grass assemblage to other vegetation types.

Similarity in grass assemblage composition was more

than 60% for each vegetation type shared between

reserves in Mozambique and South Africa. This relation-

ship with areas further south links the grass assemblages

to the Maputaland floristic region.

Grasses of the study area are mostly perennial. Woody

Grassland has the highest proportion of perennial species

and Primary Dunes the highest proportion of annuals.

Tufted grasses are the most common growth form with the

highest proportions in Licuati Forest and Coastal Woodland.

Chloridoideae and Panicoideae dominate the grass

composition in the study area. The tribes Paniceae, Era-

grostideae and Andropogoneae are represented most and

Eragrostis , Panicum and Digitaria are the largest genera

of this part of Maputaland.

The coastal plain south of Maputo Bay has a grass

diversity of 115 species and infraspecific taxa. Dune

Forest and Hygrophilous Grassland have the richest

grass diversity. Dune Forest has the most rare and en-

demic grass species and Hygrophilous Grassland the

most character species.

ACKNOWLEDGEMENTS

The authors thank Ms Teresa Martins (LUAI, Angola)

and Ms Florence Nyirenda (UZL, Zambia) for their grass

collections and identifications. Mr Calane da Silva

(LMA, Mozambique) is thanked for hosting the

SABONET Mozambique Expedition. The Global En-

vironment Facility (GEF)/United Nations Development

Programme (UNDP), through the National Botanical

Institute of South Africa, financially supported this

research as part of the capacity building Southern Afri-

can Botanical Diversity Network (SABONET) Project.

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APPENDIX 1 . — Checklist of grasses recorded for the major vegetation types of the Maputo Elephant

and Licuati Forest Reserves

Arrangement and classification follow Clayton & Renvoize (1986) and author citations follow Brummitt &

Powell (1992). Species names follow Fish (2003) and general practice at Maputo Herbarium (LMA). All specimens

are housed at LMA, with duplicates specified for the herbaria of Luanda (LUAI), Natal (NH), Pretoria (PRE) and

University of Zambia (UZL). Naturalized alien species are marked with an asterisk*, Maputaland endemic species

with **, and Red Data List species with ***. Abbreviations for collectors' names: B , Balsinhas; DHN, De Koning,

Hiemstra & Nuvunga; G, Greenwood; H , Hornby; Ma , Martins; MB , Myre & Balsinhas; MC, Myre & Carvalho; MD,

Myre & Duarte; My, Myre; N, Nyirenda; P, Pedro; PP, Pedro & Pedrogao; S, Siebert; U, Uiras; V, Viana; Z, field

observations of sterile specimens.

BAMBUSOIDEAE

ORYZEAE

Leersia hexandra Sw.. Z 14

EHRHARTEAE

Ehrharta erecta Lam. var. natalensis Stapf Z 1

POOIDEAE

AVENEAE

Agrostis lachnantha Nees var. lachnantha, Ma 101 (LUAI), H 133

CENTOTHECOIDEAE

CENTOTHECEAE

Megastachya mucronata (Poir.) P.Beauv., S 2180 (PRE), U 99 (NH)

ARUNDINOIDEAE

ARUNDINEAE

Phragmites

australis (Cav.) Steud., H 123

mauritianus Kunth, DHN 8846

ARISTIDEAE

Aristida congesta Roem. & Schult.

subsp. barbicollis (Trin & Rupr.) De Winter, S 2137 (PRE), U 65

(NH)

subsp. congesta, P 3933

Aristida stipitata Hack.

subsp. graciliflora (Pilg.) Melderis, S 2161 (PRE), U 106 (NH)

subsp. ramifera (Pilg.) Melderis, S 2188 (PRE)

CHLORIDOIDEAE

PAPPOPHOREAE

Enneapogon

cenchroides (Roem. & Schult.) C.E.Hubb ., MB 349

scoparius Stapf, Z 2

ERAGROSTIDEAE

Bewsia biflora (Hack.) Gooss., S 2128 (PRE), N 506 (UZL)

**Brachych)oa schiemanniana (Schweick.) S.M. Phillips, S 2141 (PRE),

U 60 (NH)

Dactyloctenium

aegyptium (L.) Willd., N 350 (UZL)

australe Steud., My 3743

geminatum Hack., S 2158 (PRE), U 79 (NH)

giganteum Fisher & Schweick., U 94 (NH)

Dinebra retroflexa (Vahl) Panz. var. condensata S.M. Phillips, Z 3

Eleusine

coracana (L.) Gaertn. subsp. africana (Kenn. -O'Byrne) Hilu & De

Wet, S 2130 (PRE)

indica (L.) Gaertn., U 98 (PRE)

Eragrostis

capensis (Thunb.) Trin., S 2122 (PRE), U 20 (NH)

ciliaris (L.) R.Br., S 2189 (PRE), U 27 (NH)

gummiflua Nees, U 104 (NH)

heteromera Stapf, S 2185 (PRE), U 97 (NH)

inamoena K.Schum., S 2125 (PRE)

lappula Nees, S 2144 (PRE), U 58 (NH)

**moggii De Winter var. moggii, S 2151 (PRE), N 372 (UZL)

pallens Hack., S 2174 (PRE), U 85 (NH)

sarmentosa (Thunb.) Trin., S 2164 (PRE)

sclerantha Nees subsp. sclerantha, Z 13

sp„ U61 (NH)

superba Peyr., N 362 (UZL), U 101 (NH)

Pogonarthria squarrosa (Roem. & Schult.) Pilg., S 2173 (PRE), U 53

(NH)

Sporobolus

africanus (Poir.) Robyns & Tournay, S 2140 (PRE), U 63 (NH)

fimbriatus (Trin.) Nees, S 2176 (PRE), U 89 (NH)

sanguineus Rendle, S 2186 (PRE), U 100 (NH)

subtilis Kunth, N 354 (UZL), U 81 (NH)

subulatus Hack., S 2123 (PRE)

virginicus (L.) Kunth, S 2106 (PRE), U 59 (NH)

Trichoneura

grandiglumis (Nees) Ekman, S 2136 (PRE), U 67 (NH)

**schlechteri Ekman, S 2170 (PRE), U 86 (NH)

Triraphis

andropogonoides (Steud.) E. Phillips, Z4

schinzii Hack., S 2154 (PRE), U 57 (NH)

CYNODONTEAE

Chloris

gayana Kunth, S 2196 (PRE), U 115 (NH)

virgata Sw., S 2187 (PRE)

Cynodon dactylon (L.) Pers., S 2149 (PRE), U 29 (NH)

Eustachys paspaloides (Vahl) Lanza & Mattei, S 2133 (PRE), U 42

(NH)

Perotis patens Gaud., S 2169 (PRE), U 32 (NH)

PANICOIDEAE

PANICEAE

Acroceras macrum Stapf, B 1 708

***Alloteropsis papillosa Clayton, S 2166 (PRE), U 109 (NH)

Brachiaria

chusqueoides (Hack.) Clayton, S 2147 (PRE), U 55 (NH)

deflexa (Schumach.) C.E.Hubb. ex Robyns, Ma 102 (LUAI), 11 3047

humidicola (Rendle) Schweick., S 2157 (PRE)

nigropedata (Ficalho & Hiern) Stapf, S 2182 (PRE), U 95 (NH)

Bothalia 34,1 (2004)

71

Cenchrus

*brownii Roem. & Schult., S 2129 (PRE)

ciliaris L„ S 2199 (PRE), U 111 (NH)

Digitaria

argyrothricha ( Andersson ) Chiov., S 2163 (PRE), U 50 (NH)

ciliaris (Retz.) Koeler , MB 541

debilis (Desf) Willd ., Ma 103 (LUAI), My 1088

*didactyla Willd., Z 11

eriantha Steud., S 2126 (PRE), U 107 (NH)

gymnostachys Pilg., U 112 (NH)

longiflora {Retz.) Pers., S 2181 (PRE)

natalensis Stent, U 21 (NH)

Echinochloa

colona (L.) Link, S 2195 (PRE), U 118 (NH)

holubii (Stapf) Stapf, Z 12

pyramidalis {Lam.) Hitchc. & Chase. H 3057

Eriochloa meyeriana (Nees) Pilg. subsp. meyeriana, N 382 (UZL), U 116

(NH)

Melinis repens (Willd.) Zizka

subsp. grandiflora (Hochst.) Zizka, U 90 (NH)

subsp. repens, S 2198 (PRE), U 52 (NH)

Oplismenus hirtellus (L.) P.Beauv., Z 5

Panicum

coloratum L. var. coloratum, N 385 (UZL)

deustum Plumb. , S 2139 (PRE), U 108 (NH)

genuflexum Stapf, Z 15

glandulopaniculatum Renvoize, B 1196

heterostachyum Hack., Z 6

infestum Peters, Z 9

kalaharense Mez, S 2184 (PRE)

laticomum Nees, Z 8

maximum Jacq., S 2179 (PRE), U 92 (NH)

*** pleianthum Peters, S 2143 (PRE), U 49 (NH)

Paspalum scrobiculatum L., V 38

Sacciolepis curvata (L.) Chase, S 2150 (PRE), U 64 (NH)

Setaria

incrassata {Hochst.) Hack., S 2194 (PRE), U 114 (NH)

megaphylla (Steud.) T.Durand & Schinz, S 2160 (PRE)

sphacelata (Schumach.) Moss var. sericea (Stapf) Clayton, S 2132

(PRE), U 73 (NH)

Stenotaphrum dimidiatum (L.) Brongn., S 2108 (PRE)

Tricholaena monachne (Trin.) Stapf & C.E.Hubb., Z 10

Urochloa mosambicensis (Hack.) Dandy, S 2138 (PRE), U 96 (NH)

ARUNDINELLEAE

Tristachya nodiglumis K.Schum., S 2193 (PRE), U 105 (NH)

ANDROPOGONEAE

Andropogon

eucomus Nees, S 2155 (PRE), U 72 (NH)

gayanus Kunth var. polycladus (Hack.) Clayton, S 2168 (PRE), U 84

(NH)

huillensis Rendle, Ma 104 (LUAI), G 1

schirensis A. Rich., MC 1138

Bothriochloa insculpta (A. Rich.) A. Camus., Z 7

*Coix lacryma-jobi L., My 1235

Cymbopogon

excavatus (Hochst) Stapf ex Burtt Davy, S 2138 (PRE), U 62 (NH)

nardus (L.) Rendle, S 2197 (PRE), U 117 (NH)

pospischilii (K.Schum.) C.E. Hubb., PP 1068

Diheteropogon amplectens (Nees) Clayton, MD 3942

Elionurus muticus (Spreng.) Kuntze, S 2191 (PRE), U 102 (NH)

Hemarthria altissima (Poir.) Stapf & C.E.Hubb., MC 1163

Heteropogon contortus (L.) Roem. & Schult., MB 348

Hyparrhenia

dichroa (Steud. ) Stapf, MB 632

filipendula (Hochst.) Stapf var. filipendula, 5 2134 (PRE), U 43 (NH)

Hyperthelia dissoluta (Nees ex Steud.) Clayton, S 2118 (PRE), U 41

(NH)

Imperata cylindrica (L.) Raeusch., S 2117 (PRE), U 16 (NH)

Ischaemum fasciculatum Brongn., S 2121 (PRE), U 78 (NH)

Monocymbium ceresiiforme (Nees) Stapf, MC 1161

Sorghastrum stipoides (Kunth) Nash, B 1579

Sorghum bicolor (L.) Moench subsp. arundinaceum (Desv.) De Wet &

Harlan, S 2198 (PRE), U 113 (NH)

Themeda triandra Forssk., S 2135 (PRE), U 39 (NH)

Trachypogon spicatus (L.f) Kuntze, S 2165 (PRE), U 44 (NH)

Urelytrum agropyroides (Hack.) Hack., S 2167 (PRE), U 37 (NH)

Bothalia 34.1: 73-76 (2004)

OBITUARY

JOHANNES JACOBUS ADRIAAN VAN DER WALT ( 1938-2003)

Johannes Jacobus Adriaan (Adri) van der Walt (Figure

1), renowned for his research and books on Pelargonium,

died on 28 November 2003 in Kleinmond, Western Cape.

He was bom on 16 October 1938 in Krugersdorp,

where he grew up and attended school. Though he never

spoke of those days, I presume that his parental home was

not too distant from open veld and that the young Adri

roamed the fields and ridges in his free time, acquiring his

love for plants which served him so well in later years. In

1956 he enrolled at the University of Pretoria, attaining a

B.Sc. with Botany and Zoology in 1958. Having a didac-

tic leaning, he enrolled for a Higher Education Diploma

which he attained in 1961 and a B.Ed. attained in 1966,

and from 1961 to 1967 taught biology for the Transvaal

Education Department. Meanwhile he enrolled for a

M.Sc. at the University of Pretoria, studying the secondary

growth of certain lianes under the tutelage of Professors

H.G. Schweickerdt (Van der Schijff 1977) and H.P. van

der Schijff (Theron 1998), and graduating in 1963. In

1968 he was appointed lecturer of Botany at the Uni-

versity of Zululand near Empangeni. In 1969 he joined

FIGURE 1.— J.J.A. van der Walt (1938-2003).

the Botany Department of the University of Stellenbosch

as lecturer and succeeded P.G. Jordaan as Professor in

1979 (Van der Walt 1987).

He also enrolled at Pretoria University for further

studies on his research project, the taxonomy of Com-

miphora (Burseraceae) in South Africa and Namibia, for

which he was awarded a doctorate in 1971.

At Stellenbosch University he consolidated his position

and soon became known for his emphasis on teaching and

research, aspects dear to the University. He started looking

for a good long-term research project, and the catalyst was

the meeting with botanical artist Ellaphie Ward-Hilhorst

who had the ambition of making a water-colour painting

of every Pelargonium species [see dedication to Ellaphie

Ward-Hilhorst in Flowering Plants of Africa 54 (1995),

and Rourke 1994], This resulted in the first volume of

Pelargoniums of southern Africa (Van der Walt 1977), the

second and third volumes following after I had joined the

team (Van der Walt & Vorster 1981, 1988). This work on

Pelargonium became his main focus for the rest of his life.

He was a great organizer, with a knack for persuading peo-

ple to work for his goals, and at one time a considerable

proportion of the department’s staff was working for him.

He recruited a number of local and international collabo-

rators including Dr Mary Gibby at the Natural History

Museum in London, Prof. Focke Albers of the University

of Munster (both cytologists), and Dr Frederic Demame of

CIRAD-IRAT Reunion (essential oils). His interest

encompassed the whole of the Geraniaceae as having a

bearing on Pelargonium, and his encouragement led to

monographs of Sarcocaulon (Moffett 1979), Monsonia

(Venter 1979), as well as an account of the South African

Erodium species (Venter & Verhoeven 1990). In 1990 he

organized an International Geraniaceae Symposium in

Stellenbosch to which came the world’s experts in this

field (Vorster 1990).

Adri took research seriously. During the late 1970s

and 1980s he travelled extensively throughout South

Africa, collecting Pelargonium both as herbarium speci-

mens and as living plants which were grown in his

department's botanic garden. This living collection

proved to be of great value, enabling several new species

to be recognized, chromosome numbers to be determined,

essential oil compositions to be established, anatomical

studies to be executed, and illustrations to be prepared.

Molecular research currently undertaken overseas is

largely based on cuttings originating from this material.

He preferred to refer to his chosen field as biosystemat-

ics rather than taxonomy, defining it as taxonomy

beyond the alpha level.

He was a popular teacher on all levels. He had a way

of attracting students and I well remember his engrossing

lectures and happy field excursions when I first came to

Stellenbosch. He had no difficulty in recruiting post-

74

Bothalia 34,1 (2004)

FIGURE 2. — J.J.A. van der Walt admiring a Commiphora in Namibia,

early 1980s.

graduate students. Many of his students eventually made

headway in botany, including Jo Beyers, Matt Buys,

Leanne Dreyer, Loretta Hugo, Gillian Maggs, Bettie

Marais, Mike Muller, Koos Roux, Marianna Schonken,

Mary Thompson, Ben-Erik van Wyk, and Alvaro Viljoen.

He also had endless patience with amateur growers and

interested members of the public, and this attribute, no

less than his books and other publications, fostered a

world-wide interest in the collection and cultivation of

Pelargonium species as opposed to cultivars.

Honours and awards followed, including the Compton

Prize for the best article in the Journal of South African

Botany in 1988 and 1993, honorary membership of the

International Geranium Society based in the USA in

1994, and the Havenga Prize for Biological Sciences

awarded by the South African Academy for Science and

the Arts in 2000. He served on the Board of the National

Botanical Gardens from 1989 to 1992, and coined the

name ‘National Botanical Institute’.

Besides Pelargonium, his main interest lay in almost

any team sport, as observer rather than participant, and

invariably his day was started by reading the sport pages

of Die Burger. In his younger days he was fond of fresh-

water angling, sometimes with his pupils. He was also

very fond of the veld and always hugely enjoyed the field

trips and hunting down elusive species. He often spoke

of his wonderful field trips with W. Giess (Kolberg 2001 )

in Namibia (Figure 2). He enjoyed telling about the out-

rageous escapades of his fellow students while at univer-

sity, but it appeal s as if he himself never took part in these

exercises. He was a devoted member of his church, the

Gerefonneerde Kerk , as well as a member of the Broeder-

bond.

Prior to the commencement of his work on Pelargonium ,

he collected at least 306 numbers of herbarium speci-

mens which are housed in PRE and PRU, and some in

WIND. No collecting register of this period seems to be

in existence. Subsequently his numbering system

became somewhat confusing in that the material which

he and his collaborators collected were assigned STEU

numbers which were also applied to plants in the living

collections, and starting at 400. Some of these specimens

bear a Van der Walt collection number as well as an iden-

tical STEU number, whereas others have different Van

der Walt and STEU numbers. It is recommended that

these specimens be referred to as, for example. Van der

Walt 538 sub STEU483 or Van der Walt s.n. sub

STEU2786. Again, no collecting register exists: the data

were entered in little 164 x 87 mm landscape format

pocketbooks, one accession to a page. There are 30 of

these books containing 4 371 accessions collected by

Van der Walt as well as his collaborators. At the time of

writing, these books are still in the Botany Department,

University of Stellenbosch. The main set of the Pelar-

gonium herbarium collection is in PRE, with at least

some duplicates in NBG, STEU, K, MO, and BM in

order of abundance.

Surprisingly, he took early retirement in 1996 at the

age of 58 and made what he called a clean break with

botany. He settled at Kleinmond where he lived out his

days. He also acquired a small farm in the Caledon

District where he pottered about, keeping bees, experi-

menting with growing indigenous plants, and battling

with alien invaders, while delighting in the re-appear-

ance of indigenous species after clearing the land.

Shortly before his retirement he was diagnosed with

cancer. The treatment appeared to be successful, but after

a few years the cancer flared up again and turned out to

be terminal, mercifully rapidly so.

He is survived by his wife Isabel, and three of their

four children. All three of these scions are involved with

plants: Dawid as nurseryman, Riaan as conservation offi-

cer, and Gerda as farmer.

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ERDT, H.G. 1973. Anomalous secondary growth in the stems

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of Pelargonium section Campylia (Geraniaceae). South African

Journal of Botany 54: 145-171.

VAN DER WALT, J.J.A., VENTER, H.J.T., VERHOEVEN, R. &

DREYER, L.L. 1990. The transfer of Erodium incarnation to

the genus Pelargonium (Geraniaceae). South African Journal of

Botany 56: 560-564.

VAN DER WALT, J.J.A. & VORSTER, P.J. 1980a. Application of the

name Pelargonium tabulare (Geraniaceae). Journal of South

African Botany 46: 279-281.

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Botany 46: 283-292.

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aceae). Taxon 30: 307.

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Bothalia 14: 517-523.

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bosch Botanic Gardens 16. National Botanic Gardens, Kirsten-

bosch. Cape Town.

VAN DER WALT, J.J.A., WERKER, E. & FAHN, A. 1987. Wood

anatomy of Pelargonium (Geraniaceae). I AW A Bulletin 8: 95-

108.

VAN WYK, C.M. & VAN DER WALT, J.J.A. 1995. Three new sub-

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133-139.

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SWART, J.P.J. 1995. A study of the variation in the essential oil

and morphology of Pelargonium capitatum (L.) L’Herit.

(Geraniaceae). Part III. Geographical variation in essential oil

composition and floral structure. South African Journal of

Botany 61: 105-113.

VILJOEN, A.M., VAN DER WALT, J.J.A., SWART, J.P.J. &

DEMARNE, F.E. 1995. A study of the variation in the essential

oil and morphology of Pelargonium capitatum (L.) L'Herit.

(Geraniaceae). Part II. The chemotypes of P. capitatum. Journal

of Essential Oil Research 7: 605-611.

VOLSCHENK, B. & VAN DER WALT, J.J.A. 1982. The subspecies of

Pelargonium cucullatum (L.) L’Herit. Bothalia 14: 45-51.

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Pelargonium L’Herit. (Geraniaceae). South African Journal of

Botany 2: 76-81.

P. VORSTER*

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BOTHALIA

Volume 34,1 May 2004

CONTENTS

1. The genus Cliffortia (Rosaceae) in KwaZulu-Natal. C.M. WHITEHOUSE 1

2. Two new species of Erica (Ericaceae); one from Western Cape and one from KwaZulu-Natal, South

Africa. E.G.H. OLIVER and I.M. OLIVER . 11

3. Two new species of Romulea (Iridaceae: Crocoideae) from the western Karoo, Northern Cape and notes

on infrageneric classification and range extensions. J.C. MANNING and P. GOLDBLATT .... 17

4. Studies in the genus Riccia (Marchantiales) from southern Africa. 26. A new species in section Pilifer,

Riccia radiata, is described. S.M. PEROLD 23

5. Notes on African plants:

Boraginaceae. A first record of Echium simplex in South Africa. G.F. SMITH, A.E. VAN WYK,

E.M.A. STEYN and E. RETIEF 44

Crassulaceae. Adromischus schuldtianus subsp. brandbergensis, a new subspecies and a checklist

of the succulent flora of the Brandberg, Namibia. E.J. VAN JAARSVELD, B. NORDENSTAM

and A.E. VAN WYK 35

Ericaceae. Nomenclatural changes in Erica. E.G.H. OLIVER 38

Ericaceae. A new species of indehiscent-fruited Erica from the central Kouebokkeveld, Western

Cape, South Africa. R.C. TURNER and E.G.H.OLIVER 39

Fabaceae. A new species of Acacia (Mimosoideae) from Mpumalanga, South Africa. P.J.H. HURTER

and A.E. VAN WYK 42

Lamiaceae. Plectranthus mzimvubuensis , a new species from Eastern Cape, South Africa. E.J. VAN

JAARSVELD and A.E. VAN WYK 30

Mesembryanthemaceae. A new tribe and adjustments to infrafamilial classification. P. CHESSELET,

A.E. VAN WYK and G.F. SMITH 47

Pteridophyta. Dryopteris filipaleata (Pteropsida: Dryopteridaceae), a new species from tropical

East Africa. J.P ROUX 27

Pteridophyta. Dryopteris gorgonea (Pteropsida: Dryopteridaceae), a new species from the Cape

Verde Islands. J.P. ROUX 32

Scrophulariaceae. Type specimens of Selago, Jamesbrittenia and Sutera at Natal University

Herbarium (NU). T.J. EDWARDS 27

6. Functional and taxonomic significance of seed structure in Salix mucronata (Salicaceae). E.M.A. STEYN,

G.F. SMITH and A.E. VAN WYK 53

7. Grass assemblages and diversity of conservation areas on the coastal plain south of Maputo Bay, Mozam-

bique. S.J. SIEBERT, L. FISH, M.M. UIRAS and S.A. IZIDINE 61

8. Obituary: Johannes Jacobus Adriaan van der Walt (1938-2003). P. VORSTER 73

Abstracted, indexed or listed in • AETFAT Index • AGRICOLA • AGRIS • BIOSIS: Biological Abstracts/RRM • CABS ♦ CABACCESS • CAB

ABSTRACTS • ISI: Current Contents, Scisearch, Research Alert • Kew Record of Taxonomic Literature • Taxon: reviews and notices.

ISSN 006 8241

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