ISSN 0006 8241 = Bothalia

Bothalia

A JOURNAL OF BOTANICAL RESEARCH

Vol. 34,2

Oct. 2004

TECHNICAL PUBLICATIONS OF THE SOUTH AFRICAN NATIONAL

BIODIVERSITY INSTITUTE, PRETORIA

Obtainable from the South African National Biodiversity Institute, Private Bag XI 01, Pretoria

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

BOTHALIA

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

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

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

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

jects are published annually.

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

STRELITZIA

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

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

MEMOIRS OF THE BOTANICAL SURVEY OF SOUTH AFRICA

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

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

ANNALS OF KIRSTENBOSCH BOTANIC GARDENS

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

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

South African Botany). Discontinued after Vol. 19.

FLOWERING PLANTS OF AFRICA (FPA)

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

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

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

Betty Connell, Stella Gower, Rosemary Holcroft, Kathleen Lansdell, Cythna Letty (over 700

plates), Claire Linder-Smilh 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. & 14. 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. &

I I. M. Anderson. Obtainable from: A. A. Balkema Marketing, Box 3 1 7, 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 34,2

Scientific Editor: G. Germishuizen

Technical Editor: B.A. Momberg

national

biodiversity

institute

S A N B I

2 Cussonia Avenue, Brummeria, Pretoria

Private Bag X101, Pretoria 0001

ISSN 0006 8241

Oct. 2004

Editorial Board

D.F. Cutler

B.J. Huntley

P.H. Raven

M.J.A. Werger

Royal Botanic Gardens, Kew, UK

National Botanical Institute, Cape Town, RSA

Missouri Botanical Garden, St Louis, USA

University of Utrecht, Utrecht, The Netherlands

Acknowledgements to referees

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

Campbell, Prof. B.M. Jakarta, Indonesia.

Cleevely, R.J. High Croft, Gunswell Lane, South Molton, Devon, UK.

Cowling, Prof. R.H. University of Port Elizabeth, RSA.

Cutler, Prof. D.F. Royal Botanic Gardens, Kew, UK.

Fellingham, Mrs A.C. 23 Moreson Ave, Valmary Park, 7550 Durbanville, RSA.

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

Glen, Dr H.F. National Botanical Institute, Durban, RSA.

Gregory, Dr M. Royal Botanic Gardens, Kew, UK.

Hurter, P.J.H. Lowveld Garden, National Botanical Institute, Nelspruit, RSA.

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

Kloppers, Ms R. National Botanical Institute, Pretoria, RSA.

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

McCracken, Prof. D.P. University of KwaZulu-Natal, Durban, RSA.

Netnou, Ms N.C. Biology Department, MEDUNSA, RSA.

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

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

Ortiz, Dr S. Universidade de Santiago, Santiago de Compostela, Spain.

Paterson- Jones, Dr D. National Botanical Institute, Pretoria, RSA.

Ross, Dr J. Royal Botanic Gardens, South Yarra, Victoria, Australia.

Steyn, Dr E.M.A. National Botanical Institute, Pretoria, RSA.

Stotler, Prof. R. Southern Illinois University, Carbondale, USA.

Swartz, Ms P. 362 Brook St, Menlo Park, Pretoria, RSA.

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

Tindale, Dr M.D. Royal Botanic Gardens, Sydney. Australia.

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

CONTENTS

Volume 34,2

1. New species in the section Multinerviae of Cliffortia (Rosaceae). C.M. WHITEHOUSE 77

2. Taxonomic notes and new species of the southern African genus Babiana (Iridaceae: Crocoideae).

P. GOLDBLATT and J.C. MANNING 87

3. Studies in the genus Riccia (Marchantiales) from southern Africa. 27. Riccia lanceolata and R. radicosa

now also locally reported. S.M. PEROLD 97

4. Notes on African plants:

Asteraceae. Dicoma picta, the correct name for Cypselodontia eckloniana. P.P.J. HERMAN and

M. KOEKEMOER ' 106

Fabaceae. A new species of Acacia (Mimosoideae) from the province of Limpopo, South Africa.

PJ.H. HURTER and A.E. VAN WYK 109

Hyacinthaceae. Ornithogalum laikipiense, a synonym of Drimia macrocarpa. J.C. MANNING . . 112

Iridaceae. A new species of Thereianthus (Crocoideae) from Western Cape, South Africa, nomen-

clatural notes and a key to the genus. J.C. MANNING and P. GOLDBLATT 103

Pteridophyta. A new combination in Lastreopsis (Tectariaceae) from Madagascar. J.P. ROUX . . 108

Zamiaceae. Typification of Encephalartos. P. VORSTER 112

5. Extrafloral nectaries in Combretaceae: morphology, anatomy and taxonomic significance. P.M. TILNEY

and A.E. VAN WYK 115

6. Cape heaths in European gardens: the early history of South African Erica species in cultivation, their

deliberate hybridization and the orthographic bedlam. E.C. NELSON and E.G.H. OLIVER .... 127

7. Vegetation of high-altitude fens and restio marshlands of the Hottentots Holland Mountains, Western

Cape, South Africa. E.J.J. SIEBEN, C. BOUCHER and L. MUCINA 141

8. National Botanical Institute South Africa: administration and research staff 31 March 2004, publications

1 April 2003-31 March 2004. Compiler: B.A. Momberg 155

9. Guide for authors to Bothalia 175

New genus, species, name, combinations and status in Bothalia 34,2 (2004)

Acacia sekhukhuniensis PJ.H.Hurter , sp. nov., 109

Babiana cuneata J .C .Manning & Goldblatt, sp. nov., 89

Babiana fragrans ( Jacq .) Goldblatt & J .C Manning, comb, nov., 95

Babiana longiflora Goldblatt & J.C .Manning, sp. nov., 94

Babiana praemorsa Goldblatt & J C .Manning, sp. nov., 90

Babiana regia (G.J. Lewis) Goldblatt & J.C. Manning, comb, et stat. nov., 94

Cliffortia acanthophylla C.M.Whitehouse , sp. nov., 79

Cliffortia ceresana C.M.Whitehouse , sp. nov., 80

Cliffortia denticulata (Weim.) C.M.Whitehouse, comb, et stat. nov., 81

Cliffortia oligodonta C.M.Whitehouse, sp. nov., 82

Cliffortia prionota C.M.Whitehouse, sp. nov., 83

Cliffortia recurvata (Weim.) C.M.Whitehouse, comb, et stat. nov., 82

Cliffortia reniformis (Weim.) C.M.Whitehouse, comb, et stat. nov., 77

Cliffortia scandens C.M.Whitehouse, sp. nov., 84

Cliffortia schlechteri (Weim.) C.M.Whitehouse, comb, et stat. nov., 78

Lastreopsis perrieriana (C.Chr.) J.P.Roux, comb, nov., 108

Thereianthus montanus J .C .Manning & Goldblatt, sp. nov., 103

IV

Bothalia 34,2: 77-85 (2004)

New species in the section Multinerviae of Cliffortia (Rosaceae)

C.M. WHITEHOUSE*

Keywords: Cliffortia L., new combinations, new species, new statuses, Rosaceae, section Multinerviae DC., species complexes

ABSTRACT

Three species complexes in section Multinerviae DC. of Cliffortia L. are discussed. Two of the varieties of C. ilicifolia L.

are raised to specific rank: C. reniformis and C. schlechteri, as are both varieties of C. grandifolia Eckl. & Zeyh.: C. den-

ticulata and C. recurvata. Two new species are described that were previously included within the variable species

C. dregeana C. Presl: C. acanthophylla and C. ceresana. Three further new species belonging to section Multinerviae are

also described: C. oligodonta, C. prionota and C. scandens.

INTRODUCTION

The sectional classification of the genus Cliffortia L. is

in need of revision (Whitehouse 2003). However, the cur-

rent circumscription (Weimarck 1934, 1948) of section

Multinerviae DC. can be retained for the most pan.

Species of section Multinerviae are generally medium to

tall shrubs with tough, rigid, unifoliate leaves that are

often sharply pointed and toothed and have several veins

from the base. They also have male flowers with numer-

ous stamens (20+), which are almost always white to yel-

low in colour, and female flowers tucked down at the base

of the leaves and consequently often hidden from sight.

Within section Multinerviae there are three species com-

plexes, which have previously been regarded as three vari-

able species each with a number of varieties. The C. ilici-

folia L. complex has a southern Cape distribution, between

Cape Town and Port Elizabeth, whereas C. dregeana C.

Presl has a distribution along the western Cape mountains,

from the Riviersonderend Mountains northwards to the

Cederberg. The other complex, C. grandifolia Eckl. &

Zeyh., is restricted to the southwestern Cape mountains

and Langeberg. In this paper, four of the species complex

varieties are raised to specific status and two new species

belonging to the C. dregeana complex described. Three

other new species belonging to section Multinerviae are

also described.

C. ILICIFOLIA COMPLEX

C. ilicifolia L. has been subdivided into five varieties

based upon the shape of their leaves. Var. cordifolia

(Lam.) Harv. and var. incisa Harv. are doubtfully distinct

from the typical variety. In particular, var. incisa is proba-

bly a shoot from a resprouting plant, the leaves of which

are often more markedly toothed in this species than

mature leaves. Var. cordifolia is possibly worthy of recog-

nition, as it is generally restricted to the southeastern

mountains of the Cape Floristic Region, although there is

intergradation with the typical variety and some indication

that the form may be associated with aridity or altitude.

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

MS. received: 2003-08-26.

Present address: The Royal Horticultural Society Garden, Wisley,

Woking, Surrey GU23 6QB, England, UK.

In contrast, the two varieties described by Weimarck

(1934) are very distinctive, no intermediates have been

found between them and the typical variety, from which

they are allopatric. They are, therefore, here elevated to

the rank of species.

1. C. reniformis ( Weim .) CM. Whitehouse, comb,

et stat. nov. Type: Western Cape, 3321 (Ladismith):

Riversdale Dist., Garcia’s Pass, (-CC), Bolus 11274

(BOL!, lecto., here designated; B, PRE).

C. ilicifolia L. var. reniformis Weim., Monograph of the genus

Cliffortia : 102, fig. 29D (1934).

Erect, tall shrub, up to 3 m high, resprouting after fire

and spreading clonally; dominant main stem with determi-

nate length side branches held horizontally; only forming

long branches; young stems 1.3-2 .6 mm wide, completely

sheathed by leaf bases. Leaves unifoliate, subcircular,

12-20 x 16-26 mm, clasping stem at base, apex sharply

long-acuminate to pungent, 0.2-0 .9 mm long, margins flat,

with 6-11 broad teeth, 2. 8-5.0 mm long; lamina 11-17-

veined from base, tough and rigid, 0.2-0 .5 mm thick, held

at right angles to stem, glaucous, glabrous; sheath 1.4— 6.2

mm long, abaxially glabrous, adaxially glabrous except for

fringe of hairs at apex; stipules similar in texture to lamina,

2.6-5 .9 mm long, fused into a single stipule on reverse side

of stem, margin smooth; petiole absent. Flowers solitary in

axil of undifferentiated leaves; bracteoles hairy on keel,

margins smooth; sepals 3, glabrous. Male flowers: bracte-

oles 3.6-4 .9 mm long; pedicel 0.9-1 .4 mm long; sepals

broadly ovate, 6. 4—9 .2 x 4. 5-6 .6 mm, acute to acuminate at

apex; stamens 51-68; filaments 3.8-6.8 mm long, greenish

white; anthers yellow. F emale flowers: bracteoles 3. 9-5.0

mm long, longer than immature receptacle; sepals broadly

ovate to triangular, 2. 1-2.8 x 1.0-1. 8 mm, erect; carpel 1;

stigma 1.4— 2.4 mm long, greenish white, feathery, hidden

at base of leaves; immature receptacle 1. 7-2.5 x 1. 5-2.3

mm, glabrous, smooth. Achene 8.0-8.7 x 4.8-5 .5 mm,

medium to dark brown, glabrous; ribs rounded. Flowering

time: ± all year, but mature fruits only found between

August and November. Figure 1 .

Habitat: tall, streamside vegetation and wet fynbos of

lower valley slopes, on soils derived from Table Moun-

tain Series in full sun; altitude 300-750 m.

78

Bothalia 34,2 (2004)

FIGURE 1 Lectotype of Cliffortia reniformis.

Distribution', restricted to the Langeberg at Garcia’s

Pass and the immediately surrounding valleys (Figure 2).

Conservation status : highly restricted but very com-

mon and frequently dominant in the area.

Etymology, reniformis means kidney-shaped, refer-

ring to the leaves which are almost circular in outline and

clasp the stem at their base.

This species is very distinctive on account of its

unique growth habit: a single stem often up to 3 m tall

with short branches all the way up, spreading at right

angles to the main stem. The leaves are also diagnostic,

being held at right angles and almost circular in outline

with a leaf-like stipule formed by fusion of the two sep-

arate ones on the reverse side of the stem.

Bredasdorp Dist.*, near Elim, Koude River, (-DA),

Schlechter 9584 (BOL!, lecto., here designated; B. BM,

COI, G-DEL, GRA, HBG, K!, L, P, PRE" S, W, Z).

C. ilicifolia L. var. schlechteri Weim., Monograph of the genus

Cliffortia'. 104 (1934).

Erect, medium, densely branched shrub, up to 1.8 m

high, killed by fire; only forming long branches spread-

ing at right angles; young stems 0.9-2.0 mm wide, com-

pletely sheathed by leaf bases. Leaves unifoliate, subcir-

cular to broadly ovate or oblong, 5.5-10.9 x 5.7-12.0

mm. clasping stem at base, apex sharply long-acuminate

to pungent, 0.3-1 mm long, margins flat or rounded, with

2-4 broad teeth, 0.6-2 .6 mm long; lamina 7-10-veined

from base, tough and rigid, 0.2-0.4 mm thick, held at

right angles to stem, glaucous, glabrous above and

beneath; sheath 1. 4-2.1 mm long, abaxially glabrous,

adaxially glabrous except for fringe of hairs at apex; stip-

ules 1.6-5. 8 mm long, free, margin smooth; petiole

absent. Flowers solitary in axil of undifferentiated

leaves; bracteoles hairy on keel, margins smooth; sepals

3, glabrous. Male flowers: bracteoles 2.7^4 .0 mm long;

pedicel 0.9-1. 5 mm long; sepals broadly ovate, 5-8 x

3.2- 5 .7 mm, acute to acuminate at apex; stamens 28-41;

filaments 2. 8-5.0 mm long, greenish white; anthers yel-

low. Female flowers: bracteoles 2.3-3 .2 mm long, longer

than immature receptacle; sepals ovate to triangular,

1.2- 1. 7 x 0.9-1. 2 mm, erect; carpel 1; stigma 1.1-1. 4

mm long, greenish white, feathery, hidden at base of

leaves; immature receptacle 1.5-1. 8 x 1.0-1. 3 mm,

glabrous. Achene 6.7-8.7 x 3.4-5 .4 mm, medium to dark

brown, glabrous; ribs rounded. Flowering time: ± all

year, but mature fruits only found between May and

November. Figure 3.

Habitat: fynbos on very well-drained, wind-blown

sands and limestone-derived soils in full sun; altitude

0-300 m.

Distribution: widespread on the Agulhas Plain from

Gansbaai to Gouritz River mouth (Figure 2).

Conservation status: widespread and often common,

occurring in several reserves, as well as along road

verges. However, it may be susceptible to local extinc-

tions caused by unnatural fire regimes as it does not

resprout after fire.

2. C. schlechteri (Weim.) C.M.Whitehouse , comb,

et stat. nov. Type: Western Cape, 3419 (Caledon):

Etymology: named after Rudolph Schlechter, who was

the first person to collect this species.

FIGURE 2. — Known distribution of Cliffortia ilicifolia, O; C. reniformis, A; and C. schlechteri, A, in South Africa.

Bothalia 34,2 (2004)

79

: - -i

FIGURE 3.— Lectotype of Cliffortia schlechteri.

Cliffortia schlechteri is unique within the C. ilicifolia

complex as the plants are killed by fire. Whereas it has

the characteristic perpendicular branching and leaf orien-

tation that is similar to C. reniformis , the branching is

much more dense and the plants form rounded bushes.

The leaves are generally the smallest within the complex.

Some forms of C. ilicifolia var. cordifolia do have leaves

as short, but they are much more ovate and usually lack

teeth, whereas C. schlechteri always has leaves that are

generally as broad as long, with at least two teeth.

C. DREGEANA COMPLEX

The C. dregeana CPresl complex still needs much revi-

sion. The species are characterized by their rigid, tough, pun-

gently tipped leaves and large achenes. Weimarck (1934)

divided the species into two varieties, var. dregeana and var.

meyeriana (CPresl) Weim., the former having entire leaves,

whereas the latter has 2-4 stout teeth. However, he admitted

that the boundary between the two varieties was vague.

Furthermore, as populations are found with both entire and

toothed leaves, the varieties cannot be maintained.

However, geographical forms of C. dregeana do exist

and here two such forms are given specific status. C.

ceresana is the most distinct, having much broader

leaves than all other forms within the C. dregeana com-

plex. Recognition of C. ceresana consequently makes C.

acanthophylla , with its thickened margins, geographical-

ly isolated from other populations of C. dregeana and

therefore it too is worthy of specific status.

3. C. acanthophylla C.M. White house, sp. nov., C.

dregeana CPresl affinis, sed marginibus foliorum cras-

sioribus pallidioribus, fructibus majoribus differt.

TYPE.— Western Cape, 3218 (Clanwilliam): Clan-

william Dist., Vogelfontein, (-BA), Levyns 1139 (BOL!,

holo.).

■sjpli

FIGURE 4.— Individual leaves of Cliffortia species, x 2. A, C. acan-

thophylla; B, C. ceresana; C, C. oligodonta ; D, C. prionota.

C. dregeana auct. non C.Presl: Weim.: 109 (1934) pro parte.

C. dregeana C.Presl var. meyeriana auct. non (C.Presl) Weim.:

Weim.: 109 (1934) pro parte.

Erect, medium shrub, up to 1 .8 m high, killed by fire;

only forming long branches; young stems 2.4-3 .0 mm

wide, sometimes tinged reddish, completely sheathed by

leaf bases, glabrous. Leaves unifoliate, narrowly triangu-

lar to lanceolate (Figure 4A), 15-39 x 3. 8-9.0 mm, apex

sharply pungent, E3-1.8 mm long, margins markedly

turned upwards, entire and smooth or with up to 4 long,

straight teeth, 1. 8-4.3 mm long; lamina 6-11-veined

from base, tough and rigid, 0.4-0.6 mm thick, curved

downwards and away from stem, glabrous above and

beneath; sheath 2 .7-4 .9 mm long, abaxially glabrous,

adaxially glabrous except sometimes for fringe of hairs

at apex; stipules 3 .8-5 .5 mm long, free, margin smooth;

petiole absent. Flowers solitary in axil of undifferentiat-

ed leaves; bracteoles hairy on keel, margins smooth;

sepals 3, glabrous. Male flowers : bracteoles 5. 6-6. 5 mm

long; pedicel 1. 0-1.1 mm long, glabrous; sepals broadly

ovate, 5.7-7 .2 x 3.0-3.9 mm, acute to acuminate at apex;

stamens 29 or 30; filaments 4.5-6.5 mm long, greenish

white; anthers yellow. Female flowers: bracteoles

5.8-7 .0 mm long, longer than immature receptacle; sep-

als broadly ovate to triangular, 1. 9^1.2 x 0.8-1. 8 mm,

erect; carpel 1 ; stigma 2.5—4 .2 mm long, greenish white,

feathery, hidden at base of leaves; immature receptacle

2.4-3 .0 x 1.8-24 mm, glabrous, smooth. Athene ovoid

to globose, 7.2-94 x 5.9-74 mm, medium to dark

brown, glabrous; ribs rounded. Figure 5A-C.

Habitat: fynbos on deep, well-drained sands from

Table Mountain Series in full sun; altitude 300-1 600 m.

80

Bothalia 34,2 (2004)

B E

FIGURE 5. — Cliffortia acanthophylla. A, branch with female flowers,

x B, female flower, x 2; C, achene, x 1. C. ceresana. D,

branch with female flowers, x '/2; E, female flower, x 2.

with the rest of the lamina, whereas C. dregeana also has

thickened margins but are not so pronounced. The distri-

butions of the two species do not overlap, and both are

replaced by the more easily discernible C. ceresana in

the intervening mountains. However, some specimens of

C. dregeana from the Hex River Mountains do have

thickened margins similar to C. acanthophylla :

WESTERN CAPE. — 3319 (Worcester): Hex River Mtns, Sentinel

Peak, (-AD), 1 675 m, 15-12-1957, Esterhuysen 27415 (BOL); Hex

River Mtns, Sentinel Camp, (-AD), 1 525 nt, 16-2-1958, Esterhuysen

27567 (BOL); Brandwacht Mtn, (-CB), 4-1929, Stokoe 19852 (BOL).

4. C. ceresana C.M.Whitehouse , sp. nov., habitu

floribus fructibusque C. dregeana C.Presl similis, sed

foliis latioribus plus ovatis vel oblongis, dentibus

foliaribus paucis facile distinguitur.

TYPE.— Western Cape, 3319 (Worcester): Ceres Dist.,

Michell’s Pass, (-AD), Levyns 5827 (BOL!, holo.).

C. ilicifolia auct. non L.: Weim.: 99 (1934) pro parte.

Distribution : northern Olifants River Mountains

around Graafwater and Cederberg between Sneeuberg

and Middelberg (Figure 6) .

Consecution status : limited in its distribution, but well

protected within the Cederberg Wilderness Area, however,

the Graafwater populations are all on private land.

Etymology, acanthophylla means spine-leaf, referring

to the long, hard, straight leaves with a pungent apex.

This species is difficult to separate from C. dregeana.

It is generally more robust with larger leaves and fruits,

but there is overlap between the two. The species appear

most distinct when dried, for C. acanthophylla then

shows well-developed thickened margins that contrast

FIGURE 6.— Known distribution of Cliffortia acanthophylla , A; C.

ceresana , A; and C. dregeana , O, in South Africa.

Erect, medium shrub, up to 1 .8 m high, resprouting

after fire and spreading clonally; only forming long

branches; young stems 1. 5-2.1 mm wide, completely

sheathed by leaf bases. Leaves unifoliate, ovate-elliptic to

oblong (Figure 4B), 10-27 x 5-16 mm, apex sharply long-

acuminate to pungent, 1 .2-3.1 mm long, margins flat with

5-8 broad teeth, 0.8-5.8 mm long; lamina 5-9-veined

from base, tough and rigid, midrib prominent beneath,

0.3-0. 8 mm thick, curved downwards and away from

stem, glabrous; sheath 1.9-3 .5 mm long, abaxially

glabrous, adaxially glabrous except sometimes for fringe

of hairs at apex; stipules 1.0-5 .5 mm long, free, margin

smooth; petiole absent. Flowers solitary in axil of undif-

ferentiated leaves; bracteoles hairy on keel, margins

smooth; sepals 3, glabrous. Male flowers : bracteoles

3. 8-7.0 mm long; pedicel 0.8-1. 3 mm long, glabrous;

sepals broadly ovate, 6.7-10.4 x 3.0-5.9 mm, acute to

acuminate at apex; stamens 21-34; filaments 2.8-5.5 mm

long, greenish white; anthers yellow. Female flowers:

bracteoles 5. 5-6 .5 mm long, longer than immature recep-

tacle; sepals broadly ovate to triangular, 2.8-3 .9 x 0.9-1 .6

mm, erect; carpel 1 ; stigma 1 .8-2.5 mm long, red or some-

times greenish white, feathery, hidden at base of leaves;

immature receptacle 23-3.1 x 1. 8-2.7 mm, glabrous,

smooth. Achene ovoid, 6.4-7 .4 x 4.4-4 .6 mm, medium to

dark brown, glabrous; ribs rounded to sharply acute.

Flowering time: September to January. Figure 5D, E.

Habitat: fynbos on rocky shale slopes and amongst

sandstone rocks at high altitudes, on well-drained soils

derived from Table Mountain Series in full sun; altitude

350-1 850 m.

Distribution: found from the western parts of the Hex

River Mountains, north through the Witzenberg Range to

the Groot Winterhoek Mountains (Figure 6).

Conservation status: common throughout the area in

which it grows, especially at higher altitudes; it is con-

served within the Groot Winterhoek Wilderness Area and

its resprouting habit means that it is not particularly sus-

ceptible to unnatural fire regimes.

Bothalia 34,2 (2004)

Etymology : named after the town of Ceres, where it is

common on the surrounding mountains, and has fre-

quently been collected in Michell’s Pass, which leads to

the town.

Burchell’s specimen of this species was included

within C. ilicifolia by Weimarck (1934) on account of its

broad, toothed leaves. However, in other characteristics

it is closer to C. dregeana , with which it has generally

been placed in herbaria. The broadly oblong, few-

toothed leaves, combined with the limited distribution

range, make this species easily distinguishable.

However, in the south of its range, specimens that are

more intermediate in character with C. dregeana do

occur:

WESTERN CAPE.— 3319 (Worcester): Hex River Mtns, Buf-

felshoek Peak.(-AD), 1 830 m, 26-12-1942, Esterhuysen 8405 (BOL);

Waaihoek Mtn, SE, (-AD), 15-9-1943, Esterhuysen 8970 (BOL); Hex

River Mtns. Sentinel Peak, (-AD), 1 525-1 675 m. 16-2-1958. Ester-

huysen 27571 (BOL).

C. GRAND1FOLIA COMPLEX

The members of the C. grandifolia Eckl. & Zeyh.

complex are all tall, sparsely branched shrubs with large

leaves. Weimarck (1934) divided C. grandifolia into

three varieties based upon the shape of the leaves. The

varieties are easily identifiable, as well as allopatric in

their distributions, especially var. grandifolia, which is

endemic to the Langeberg. Intermediates have not been

If

FIGURE 7.— Lectotype of Cliffortia denticulata.

FIGURE 8.— Known distribution of Cliffortia denticulata , O; C. gran-

difolia, A, and C. recurvata , A, in South Africa.

found between the varieties, although putative hybrids

have been found with other species (e.g. C. grandifolia x

C. lanceolata , C. <& A. Whitehouse 339 and C. denticula-

ta x C. ovalis , C. & A. Whitehouse 340). Therefore the

varieties are here raised to specific rank.

5. C. denticulata ( Weim .) CM. Whitehouse, comb,

et stat. nov. Lectotype: Western Cape, 3319 (Worcester):

Franschhoek, (-CC), in mountains, Schlechter 9240

(BOL!, lecto.; B, BM, COI, G-DEL, GRA, HBG, K!, L,

P. PRE, S,W, Z).

C. grandifolia Eckl. & Zeyh. var. denticulata Weim., Monograph of

the genus Cliffortia : 115, fig. 33C ( 1934).

C. intermedia auct. non Eckl. & Zeyh.: Harv.: 295 (1862) pro parte.

Erect, tall, spindly shrub, up to 3.6 m high; only form-

ing long branches and very sparsely branched, branches

clustered 2-4 together at a node, killed by fire; young stems

1.7-5.25 mm wide, completely sheathed by leaf bases.

Leaves unifoliate, very broadly ovate to almost subcircular,

30-68 x 21— 41 mm, clasping stem at base, apex sharply

long-acuminate, 1 .9-4.0 mm long, margins flat with

52-83, small, straight teeth, 0.3-1. 3 mm long; lamina

11-17-veined from base, relatively soft and chartaceous,

0.2-0 .6 mm thick, curved downwards and away from stem,

glabrous above and beneath; sheath 6.0-10.8 mm long,

abaxially and adaxially glabrous; stipules absent or present

and 1 .6-3.9 mm long, free, margin smooth; petiole absent.

Flowers solitary in axil of undifferentiated leaves; bracte-

oles hairy on keel, margins smooth; sepals 3; glabrous.

Male flowers: bracteoles 14—17 mm long; pedicel 1.1-1 .4

mm long, glabrous; sepals broadly ovate, 23-26 x 6.4—14.5

mm, acute to acuminate at apex; stamens 60-133; fila-

ments 3.5-8.0 mm long, greenish white; anthers yellow.

Female flowers: bracteoles 6. 0-9 .2 mm long, longer than

immature receptacle; sepals broadly ovate to triangular,

3. 2-3 .8 x ± 1.7 mm, erect; carpel 1 ; stigma ±2.5 mm long,

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

ture receptacle ± 3.2 x 2 mm, glabrous, smooth. Athene

medium to dark brown, glabrous; ribs rounded. Figure 7.

Habitat: in damp depressions and beside watercours-

es on well-drained soils from Table Mountain Series in

full sun; altitude 350-1 350 m.

Distribution: Franschhoek to the southern Hottentots

Holland Mountains around Nuweberg (Figure 8).

Conservation status: limited distribution, extent of

occurrence, < 150 km:, but the entire range is conserved

82

Bothalia 34,2 (2004)

( i »

P<.OII«,-5»«/A- '!v'wCc/v>t- Ku)j.cr>.

-

Maws f - t^\d^eixa.tL

-w xlLfa.-Jk_l- it Ul£

tAyf •V.'^'^'^OJVl .VCvU^M Al> ,

n< • ^«A<v . , *«». t<j l 1 . i_„v , T. ?. S£rfir<. .

i&Slb't

long, abaxially glabrous, adaxially with scattered, ad-

pressed hairs F>ut without a fringe of hairs at apex; stip-

ule 0.9-1. 4 mm long, sometimes absent, free, margin

smooth; petiole absent. Flowers solitary in axil of undif-

ferentiated leaves; bracteoles hairy on keel, margins

smooth; sepals 3, glabrous. Male flowers: bracteoles 15-

16 mm long; pedicel 2. 1-2.4 mm long, hairy; sepals

broadly ovate, 19-21 x 7.2-11 .0 mm, acute to acuminate

at apex; stamens 75-120; filaments 6. 8-9. 8 mm long,

greenish white; anthers yellow. Female flowers: bracte-

oles 10.5-11.3 mm long, longer than immature recepta-

cle; sepals ± triangular, 3 .2 — 4 .6 x 0.9-1. 8 mm, erect;

carpel 1; stigma 2-3 mm long, greenish white, feathery,

hidden at base of leaves; immature receptacle 3. 4-3 .9 x

2.7-3.0 mm, hairy towards base, smooth. Achene ellip-

soid, ±8.1 x 4.8 mm, medium to dark brown, glabrous;

ribs rounded. Figure 9.

Habitat: in damp depressions and beside watercours-

es on well-drained soils from Table Mountain Series in

full sun; altitude 300-800 m.

Distribution: confined to the Kogelberg between the

Steenbras and Palmiet River Valleys (Figure 8).

Conservation status: only known from the middle

slopes of the Kogelberg, extent of occurrence, < 75 km ,

and restricted to a very particular habitat where it is

uncommon; being a reseeder it is vulnerable to too fre-

quent fire regimes; however, it is conserved entirely

within the Kogelberg Biosphere Reserve.

FIGURE 9.— Lectotype of Cliffortia recurvata.

Etymology: recurvata means curved backwards, refer-

ring to the leaves that are strikingly curved almost into a

semicircle.

within the Hottentots Holland Nature Reserve; however,

being a reseeder it may be susceptible to local extinction

caused by too frequent fires.

Etymology: denticulata means very small teeth, which

surround the margins of the leaves.

6. C. recurvata {Weim.) C.M.Whitehouse , comb,

et stat. nov. Type: Western Cape, 3418 (Simonstown):

mountains E of Steenbras Valley, (-BB), Feb. 1921 , Stokoe

s.n. (BOL!, lecto., here designated).

C. grandifolia Eckl. & Zeyh. var. recurvata Weim., Monograph of

(he genus Cliffortia : 115, fig. 33F (1934).

Erect, tall, spindly shrub, up to 3.6 m high, only form-

ing long branches and very sparsely branched, branches

clustered 2 -4 together at a node, killed by fire; young

stems 3. 4-4.3 mm wide, completely sheathed by leaf

bases but with upwardly adpressed hairs visible on

young growth. Leaves unifoliate, lanceolate to narrowly

ovate, 64—74 x 20-27 mm, clasping stem at base, apex

pungent, 2.6-4.0 mm long, margins flat, with 90-109

small straight teeth, 1. 4-2.3 mm long, whole leaf often

strongly keeled; lamina 13-18-veined from base,charta-

ceous to tough, 0.25-0.5 mm thick, strongly curved

downwards and away from stem forming almost a semi-

circle, glabrous above and beneath; sheath 5. 2-6 .7 mm

7. C. oligodonta C.M.Whitehouse , sp. nov., similis

C. integerrima Weim. sed foliis longioribus plus

oblongis glaucis facile distinguitur.

TYPE.— Western Cape, 3319 (Worcester): Paarl Dist.,

SE slopes of Wemmershoek Peak, (-CC), C. & A. White-

house 343 (BOL!, holo.).

Erect, medium shrub, up to 1 .2 m high, resprouting

after fire and spreading clonally; only forming long

branches; young stems 2.7-3 .0 mm wide, completely

sheathed by leaf bases. Leaves unifoliate, elliptic-oblong

to ovate (Figure 4C), 22-36 x 9.5-13.5 mm, apex pun-

gent, 1 .4-2.5 mm long, margins Hat, entire or with up to

5 straight teeth, 1. 0-2.7 mm long; lamina 5-7-veined

from base, tough and rigid, 0.5-0.7 mm thick with

midrib occasionally slightly prominent beneath, curved

downwards and away from stem, glaucous, glabrous or

sometimes with a few 0.4-1. 0 mm long hairs along

midrib; sheath 3. 0^1.6 mm long, abaxially glabrous,

adaxially glabrous or with a few adpressed hairs and

fringed at apex; stipules 3.5-5.5 mm long, free, margin

smooth; petiole absent. Flowers solitary in axil of undif-

ferentiated leaves; bracteoles hairy on keel, margins

smooth; sepals 3, glabrous. Male flowers: bracteoles

9.4-11.3 mm long; pedicel 1. 5-2.1 mm long, glabrous;

sepals broadly ovate, 1 1 .4-15.0 x 6.0-7. 8 mm, acute to

acuminate at apex; stamens 30-35; filaments 7.2-11.8

mm long, greenish white; anthers yellow. Female flow-

Bothalia 34,2 (2004)

83

This species was previously determined as C. inte-

gerrima Weim. However, that species was poorly delim-

ited by Weimarck (1934) and encompassed several dif-

ferent entities from the Cape Peninsula, Riviersonderend

Mountains and the Groot Swartberg. The species is dis-

tinctive on account of its relatively broad elliptic-oblong

leaves that are glaucous in colour. Furthermore, the

species has so far not been found beyond the

Wemmershoek Mountains, meaning that confusion with

other species is unlikely. Within sect. Multinerviae, only

C. dregeana , with narrowly lanceolate green leaves,

grows in the same vicinity, but is replaced by C.

oligodonta above about 1 600 m. However, where the

two species grow in close proximity, a putative hybrid

has been found [Paarl Dist., S slopes of peak between

Perdekop and Wemmershoek Peak, C. & A. White house

344 (BOL)] that is intermediate between the two species.

The hybrid has oblong-lanceolate leaves, which are

never toothed, however they are generally broader than

C. dregeana and retain the glaucous coloration of C.

oligodonta.

FIGURE 10.— A, B. Cliffortia prionota : A, branch, x B. female

flower, x 2. C, D, Cliffortia oligodonta: C, branch with female

flowers, x D, female flower, x 2.

ers: bracteoles 5. 6-7 .5 mm long, longer than immature

receptacle; sepals broadly ovate to triangular, 2 .4-3 .2 x

0.8-1. 6 mm, erect; carpel 1; stigma 2.3-3.5 mm long,

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

ture receptacle 2.5-3. 1 x 2.0-2.5 mm, glabrous, smooth.

Achene ovoid. 7. 3-7 .8 x 4. 1-5 .3 mm, medium to dark

brown; glabrous; ribs 16-22, rounded. 0.2-0.4 mm wide.

Flowering time : September to May. Figure 10C, D.

Habitat: south-facing slopes and in the shelter of

rocks at high altitudes, in well-drained rocky soil from

Table Mountain Series; altitude 1 450-1 800 m.

Distribution: very narrow endemic, found only on the

upper slopes of the Wemmershoek Mountains between

Winterberg and Wemmershoek Peak (Figure 11).

Conseiwation status: endemic to a very small area,

extent of occurrence, < 20 km:, but common there at high

altitudes.

Etymology: oligodonta means few teeth, referring to

the leaves which vary from being entire to having up to

five small teeth.

FIGURE 11.— Known distribution of Cliffortia oligodonta. A; C. prio-

nota. O; and C. scandens, A, in South Africa.

8. C. prionota C.M.Whitehouse , sp. nov., a C.

lanceolata Weim. stipulis brevioribus glabris, floribus

masculinis majoribus, staminibus pluribus glabris differt.

TYPE.— Western Cape, 3418 (Simonstown): Caledon

Dist., Kogelberg Nature Reserve, Kogelberg Trail near

where it joins jeep track in upper reaches of Louws River

Valley, (-BD), C. & A. Whitehouse 274 (BOL!, holo.).

Erect, medium to tall shrub, up to 2.5 m high, re-

sprouting after fire and spreading clonally; only forming

long branches; young stems 1. 8-2.8 mm wide, com-

pletely sheathed by leaf bases. Leaves unifoliate, nar-

rowly oblong to lanceolate (Figure 4D), 26^19 x 4.6-8 .5

mm, apex sharply long-acuminate to pungent, 1.8-3 .6

mm long, margins flat or rounded, entire or with 9^f2

straight teeth, 0.8-1. 8 mm long; lamina 5-10-veined

from base, chartaceous to rigid, 0.3-0 .5 mm thick, held

straight or curved upwards and towards stem, glabrous;

sheath 3. 9-5 .5 mm long, abaxially and adaxially

glabrous; stipules 3. 8-6 .2 mm long, free, margin smooth;

petiole absent. Flowers solitary in axil of undifferentiat-

ed leaves; bracteoles hairy or not on keel, margins

smooth; sepals 3; glabrous. Male flowers: bracteoles

6 .8-8 .7 mm long; pedicel 1.3-1. 6 mm long; sepals

broadly ovate, 11.5-13.5 x 4.6-7 .2 mm, acute to acumi-

nate at apex; stamens 40-50; filaments 6.2-9 .7 mm long,

greenish white; anthers yellow. Female flowers: bracte-

oles 6.4-8 .4 mm long, longer than immature receptacle;

sepals broadly ovate to triangular, 2.5-3 .5 x 0.7- 1.5 mm,

erect; carpel 1; stigma 2.4-3 .3 mm long, greenish white,

feathery, hidden at base of leaves; immature receptacle

2.4—3 .2 x 1 .7-2.3 mm, glabrous, smooth . Achene medium

to dark brown, glabrous; ribs rounded. Flowering time:

September to April. Figure 10A, B.

Habitat: in clayish soil on shale bands of Table Moun-

tain Series rocks in full sun, in well-drained soils or

along the edge of damp gullies; altitude 50-350 m.

Distribution: restricted to the lower mountain slopes

of the Palmiet River Valley and behind Kleinmond

(Figure 11).

84

Bothalia 34,2 (2004)

Conservation status: endemic to a small area and

uncommon there forming scattered clonal patches. Popu-

lations behind Kleinmond may be threatened by urban

expansion but the majority of the populations are pro-

tected within the core region of the Kogelberg Biosphere

Reserve and being a resprouter it is not particularly

threatened by unnatural fire regimes.

Etymology: prionota means like a saw, referring to the

leaves, which normally have fine teeth along their edge;

it also alludes to the fact that the species is confined to

the Palmiet River area and palmiet is the common name

for the plant Prionium serratum.

Previously classified with C. intermedia , this species

actually bears closer resemblance to the Langeberg

endemic, C. lanceolata but lacks the hairs on the anthers

that are diagnostic for that species. C. prionota has a very

narrow range, being endemic to the Kogelberg, and for

that area the long, narrowly lanceolate leaves on a clon-

ally, spreading erect shrub, prevent confusion with any

other species. However, the species is comparatively

variable within its range, from having leaves that are

tough, rigid and entire to ones that are relatively soft and

flexible and bearing numerous small teeth. This degree

of variation may be the result of introgression with other

species, such as the sympatric C. heterophylla (e.g.

Levy ns 7042).

9. C. scandens C.M.Whitehouse, sp. nov., a C.

acutifolia Weim. stipulis longioribus glabris, dentibus

foliaribus pluribus, sepalis masculinis angustioribus, sta-

minibus paucioribus glabris differt.

TYPE.— Western Cape, 3419 (Caledon): Caledon Dist.,

Riviersonderend Mtns, kloof near Riviersonderend,

(-BB), Esterhuysen 25075 (BOL!, holo.; K!, NBG!, PRE).

C. dregeana auct. non C.Presl: Weim.: 109 (1934) pro parte.

Scrambling, medium shrub, up to 0.6 m high, densely

branched, forming brachyblasts; young stems 0.6-1. 3

mm wide, glabrous. Leaves unifoliate, oblong to linear or

very narrowly lanceolate, 22-38 x 2.0-3 .2 mm, apex

sharply long-acuminate to pungent, 1.0-1. 8 mm long,

margins turned upwards, with 4-10 straight teeth,

0.5-1. 1 mm long; lamina 1-3-nerved into base, charta-

ceous, 0.2-0. 3 mm thick, curved downwards and away

from stem, with a few 0.7-1. 7 mm long hairs above,

glabrous beneath; sheath 2.5-4.0 mm long, abaxially

glabrous, adaxially glabrous except for fringe of hairs at

apex; stipule 4.2-11.8 mm long, free, margin smooth;

petiole absent. Flowers solitary in axil of undifferentiat-

ed leaves; bracteoles glabrous on keel, margins smooth

to very shortly ciliate; sepals 3, glabrous. Male flowers:

bracteoles 6.4-7 .3 mm long; pedicel 1.0-1. 3 mm long;

sepals broadly ovate, 7.0-9. 1 x 1. 7-2.7 mm, acute to

acuminate at apex; stamens ± 20; filaments 6.5-8. 3 mm

long, red; anthers brownish red. Female flowers: bracte-

oles longer than immature receptacle; sepals oblong to

triangular, 3. 1-3 .4 x 0.4-0. 7 mm, erect; carpel 1; stigma

2. 8-4.7 mm long, red, feathery, receptacle glabrous,

smooth. Achene ovoid to ellipsoid, 4. 1-4 .6 x 1.7-1. 8

mm, medium to dark brown, glabrous; ribs faint, round-

ed. Flowering time: November to February. Figure 12.

FIGURE 12.— Cliffortia scandens. A, branch, x 1; B, leaf, x 2; C, achene,

x 4.

Habitat: fynbos on rocky south-facing slopes; altitude

650-1 250 m.

Distribution: higher south slopes of Riviersonderend

Mountains around Genadendal and Riviersonderend

(Figure 11).

Conservation status: a poorly known species, record-

ed from just two areas of the Riviersonderend Mountains

and apparently very localized there, but the mountain

range as a whole is relatively unexplored; invasive alien

plants and unnatural fire regimes are threats, but the

majority of the mountain range is conserved within a

nature reserve.

Etymology: scandens means scrambling, referring to

the way the plants form a tangled growth.

This species has previously been placed under C.

uncinata or C. acutifolia. However, both those species

are only known from mountains north of Tulbagh. The

species is easily recognized within its distribution range

by its long, narrow, toothed leaves, which form brachy-

blasts, and the plant’s scrambling habit. However, hybrid

swarms between C. dregeana and C. pungens have been

confirmed in the Riviersonderend Mountains (White-

house 2003) and may involve this species as well. Such

hybrids may include the types of C. theodori-friesii

Weim. var. puberula Weim. or C. meyeriana C.Presl.

Much work in this geographical area is needed to identi-

fy the parent and hybrid populations, especially to deter-

Bothalia 34,2 (2004)

85

mine the limits of the species involved and the parents of

the various hybrid populations. Despite this apparent

confusion of species limits, preliminary molecular evi-

dence supports the identification of C. scandens as a dis-

tinct species, as it has both nuclear and chloroplast mark-

ers not found in any other species (Whitehouse 2003).

SPECIMENS EXAMINED

Acocks 15410 (2) K, PRE; 19656 (3) K, PRE; 19765 (3) K, NBG, PRE.

Barker 3325 (6) NBG; 5530 (1) BOL, NBG. Bayliss 964 (2) PRE.

Bohnen 7985 (2) K. NBG, PRE; 8160 (1) NBG, PRE. Bolus 6993 (5)

BOL, NBG, PRE; 11274 (1) BOL, PRE. Bond 1085 , 1344 (3) NBG.

Boucher 1258 (8) K, PRE, (6) NBG, PRE; 5936 (2) NBG. Burchell

8688 (4) K. Burgers 1906 (2) NBG, PRE.

Compton 4165 (5) BOL; 6222 (3) NBG; 6695, 8143, 11989, 12465,

16702, 18802 (4) NBG; 8320, 12984, 18326 (5) NBG; 14119, 19402

(8) NBG; 14491, 16868 (6) NBG; 19051, 23325 (2) NBG; 20643 (4)

BOL, K; 22577 (2) BOL, NBG. Cowling 3174 (2) NBG.

De Vos 1383 (8) NBG. Drege 2927 (3) K.

Esterhuysen s.n. (5) BOL, NBG; 1995, 8244, 9148, 9717, 11594, 27791

(5) BOL; 4169, 5502, 8141, 13142, 27415, 27567 (3) BOL; 7243 (3)

BOL, K; 8176, 9868, 14702, 16143 (4) BOL; 9670 (7) BOL, PRE;

11278 (7) BOL, K, PRE; 11576 (7) BOL, K, NBG, PRE; 12561 (8)

BOL; 15252 (5) BOL. NBG. PRE; 17001 (1) BOL, PRE; 25075 (9)

BOL, K, NBG, PRE; 25337 (9) BOL, PRE; 27620, 27621 (7) BOL;

31466 (9) BOL, K.

Fellingham 81 (2) K, NBG, PRE; 442 (1) NBG.

Galpin 3999 (1) PRE. Gentry & Barclay 19121 (1) PRE. Gillett 4117

(3) K, PRE. Goldblatt 3751 (1) K.

Hanekom 729, 730 (4) K, PRE. Haynes 51 (5) NBG, PRE. Herb.

Forsyth, s.n. (5) K. Hugo 1105 (2) K, NBG, PRE. Hutchinson 1022 (4)

BOL, K.

Johnson 127 (2) BOL, NBG.

Kerfoot 5306 (5) PRE; 6157 (3) NBG. Kruger 1093 (4) NBG; 801 (8)

NBG.

Leighton 11,462 (5) BOL; 759 (6) BOL; 2301 (4) BOL; 2556 (2) BOL,

NBG, PRE; 3301 (2) BOL. Levyns 1139, 2919 (3) BOL; 2293 ( 1 ) BOL;

4946, 5827 (4) BOL; 7042 (8) BOL; 8858 (6) BOL; 9516 (2) BOL.

Linder 4093 (2) K, PRE. Linder in Whitehouse 68 (9) BOL; 216 (6)

BOL.

Maguire 71 (2) NBG. Marloth 3551 (1) PRE; 6158 (4) NBG, PRE.

Morley 27 (2) NBG, PRE. Muir STE10555 (1) NBG; 429 (2) PRE;

4661 (1) NU.

O' Callaghan, Fellingham & Van Wyk 340 (2) NBG, PRE. Oliver 5701

(2) NBG, PRE.

Pillans s.n., 7619 (5) BOL.

Richardson 99 (1) NBG, PRE.

Salter 7561 (3) BOL, K. Schlechter 8514 (3) BOL, K, PRE; 9240 (5)

BOL, K, PRE; 9584 (2) BOL, K, PRE; 9851 (9) PRE. Smith 2750 (1)

K. Stokoe s.n. (6) BOL, K; 7261, 8618 (6) BOL; 407, 413 (8) NBG;

7404 (7) BOL; 16947 (5) K; 19852 (3) BOL.

Taylor 4536a (4) NBG; 6173, 10685 (3) NBG, PRE. H. Thode 2237 (4)

K. PRE. J. Thode 4817 (4) NBG. Thompson 687 (1) NBG. PRE; 849

(1 ) K, NBG, PRE; 1462 (4) K. NBG, PRE; 1730 (2) K, NBG, PRE.

Van der Merwe 1202 (5) NBG, PRE. Van Wyk 684 ( 1 ) NBG, PRE.

C. Whitehouse 12 (7) BOL; 19 (5) BOL; 46 (8) BOL; 92, 225 (2) BOL;

99, 183 (1) BOL; 248 (3) BOL; 269 (4) BOL; 352 (6) BOL. C. & A.

Whitehouse 274 (8) BOL; 308, 313 (4) BOL; 341 (5) BOL; 342, 343

(7) BOL. Willemse 252 (2) NBG, PRE.

REFERENCES

HARVEY, W.H. 1862. Rosaceae. In W.H. Harvey & O.W. Sonder,

Flora capensis 2 : 285-305. Hodges, Smith, Dublin.

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

Ohlsson, Lund.

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

Botaniska Notiser 90: 167-203.

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

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

■

■

Bothalia 34,2: 87-96 (2004)

Taxonomic notes and new species of the southern African genus

Babiana (Iridaceae: Crocoideae)

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

Keywords: Babiana Ker Gawl., B. cuneata sp. nov., B. fragrans comb, nov., B. longiflora sp. nov., B. praemorsa sp. nov., B. regia comb, et stat.

nov.. biogeography, Iridaceae, southern Africa, systematics

ABSTRACT

A member of Iridaceae subfamily Crocoideae. Babiana Ker Gawl. comprises some 80 species from southern Africa.

Field studies have shown the need for several taxonomic and nomenclatural changes, while a number of new species have

been discovered. The type of B.flabellifolia Harv. ex Klatt is a short-tubed plant that matches B. truncata G.J. Lewis, and

that name falls into synonymy. The name B.flabellifolia sensu G.J .Lewis (1959) has been misapplied to long-tubed plants

from the western Karoo which are now renamed B praemorsa sp. nov. In addition. B truncata. as originally circumscribed,

included two species, one short-tubed and a second with a longer tube, which we describe here as B cuneata sp. nov. The

type of B. hypogaea Burch, has also been misinterpreted and matches the species described as B.flavida , which thus falls

into the synonymy of B. hypogaea. A second species, B.falcata G.J .Lewis, closely matches this species and is also reduced

to synonymy. The widespread southern African species long known as B. hypogaea (hypogea sensu G.J .Lewis) matches the

type of B. bainesii Baker and must now be known by that name. Babiana stricta var. erectifolia (G.J .Lewis) G.J .Lewis is

appropriately included in typical B. stricta (Aiton) Ker Gawl. However, var. regia G.J. Lewis is a very different plant and is

treated as a separate species, B. regia comb, et stat. nov., as is long-tubed var. grandiflora G.J .Lewis, which is described as

the new species, B. longiflora sp. nov. Plants included in B. stricta var. sulphurea sensu G.J. Lewis are also included in var.

stricta. We are unable to match the type of Gladiolus sulphureus Jacq.. basionym of var. sulphurea (Jacq.) Baker, with any

known species and the name is thus excluded. Lastly, the name B. disticha Ker Gawl., type of the genus, is superfluous for

Gladiolus fragrans Jacq. and the new combination B fragrans comb. nov. is made, reducing B. disticha to synonymy.

Babiana fragrans Eckl., which was thought to prevent the transfer of G. fragrans, is a nomen nudum and thus invalid and

cannot be taken into account in considerations of nomenclatural priority.

INTRODUCTION

When last revised in 1959 by the South African botanist,

GJ. Lewis, the genus Babiana Ker Gawl. (Iridaceae: sub-

family Crocoideae) was considered to comprise 60 species

in southern Africa and one on the Indian Ocean island of

Socotra. Since the publication of this monograph, two

species, B. lewisiana B.Nord. (Nordenstam 1970) and B.

virginea Goldblatt (Goldblatt 1979) have been described,

and a further two, Antholyza plicata Thunb. and A. ringens

L. have been restored to Babiana as B. thunbergii Ker Gawl.

and B. ringens (L.) Ker Gawl. respectively (Goldblatt 1990).

The latter two species were included in Babiana by most

19th century botanists, notably Ker Gawler (1805) and

Baker (1892; 1896), although not by Lewis (1959), who fol-

lowed Brown ( 1932) in maintaining Antholyza L. for B. rin-

gens, and Anaclanthe N.E.Br. for B. thunbergii and its syn-

onym, A. namaquensis N.E.Br. The Socotran species, B.

socotrana Hook.f., is now known to be allied to Lapeirousia

Pourr. and Savannosiphon Goldblatt & Marais, and has been

referred to the new genus Cyanixia Goldblatt & J.CJVIanning,

as C. socotrana (Hook.f.) Goldblatt & J.C .Manning (Goldblatt

et al. 2004). Currently then, Babiana includes 64 species, all

from southern Africa, mainly the winter rainfall zone of the

south and west of the subcontinent.

Field research, especially since 1995, often conducted

in conjunction with our studies of the pollination biology

of Babiana and other genera of the Iridaceae, has led to

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

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

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

7735 Claremont, Cape Town.

MS. received: 2004-05-24.

the realization that there are several shortcomings in

Lewis’s (1959) account of the genus as well as a number

of undescribed species. In this paper we deal with

changes to the taxonomy of the B. flabellifolia , B.

hypogaea and B. stricta species complexes, describe two

new species, and make two new combinations resulting

from these changes.

Examination of the type specimen of Babiana flabelli-

folia Harv. ex Klatt, most likely collected in northern

Namaqualand, reveals that it does not match the western

Karoo plant that Lewis (1959) associated with this name.

The latter plant, which is distinguished by a long perianth

tube and short filaments, is renamed B. praemorsa sp. nov.

Accordingly, the short-tubed Namaqualand species B.

truncata GJ.Lewis, which closely matches B. flabellifo-

lia, assumes this earlier epithet. Our studies also show that

plants referred by Lewis to B. truncata represent two

species, one largely from Namaqualand with a perianth

tube 18-36 mm long, and a second, described here as B.

cuneata sp. nov., with a perianth tube 40-60 mm long.

Similarly, the type of Babiana hypogaea Burch, (spelled

hypogea by Lewis), collected by William Burchell in

1812, closely matches the species described as B.flavida

by Lewis in general appearance and perianth dimensions,

and differs significantly from the widespread southern

African plant currently called B. hypogea sensu Lewis

(1959). The latter must now assume the name B. bainesii

Baker, and B.flavida GJ.Lewis falls into the synonymy of

B. hypogaea. The revised taxonomy accords with the

treatment by Baker (1896) in Flora capensis.

In addition, taxonomic changes are needed for the

southwestern Cape species, Babiana stricta (Aiton) Ker

Bothalia 34,2 (2004)

Gawl., a complex in which Lewis (1959) recognized five

varieties. Some of these closely resemble the type of B.

stricta but others differ strikingly from that species. No

single exclusive feature links the five varieties, except

for the more or less subequal tepals and, with the excep-

tion of var. grandiflora G.J .Lewis, the relatively narrow,

rigid leaves with a short rough pubescence. Neither of

these features is unique to this complex. Lastly,

Gladiolus fra grans Jacq. is an earlier name for Babiana

di Stic ha Ker Gawl., and we make the nomenclatural cor-

rection in the new combination B.fragrans. The nomen-

clatural and taxonomic consequences of these conclu-

sions are dealt with formally below.

A more extensive account of Babiana is planned for a

future paper in which we will describe additional new

species and present a revised infrageneric classification

of the genus. Including the new species described below

and those in manuscript, Babiana comprises 80 species.

This makes it one of the larger genera of southern

African Iridaceae, after Gladiolus L. (165 species in

southern Africa) and Moraea Mill. (147 species), and

slightly larger than Romulea Maratti (78 species) and

Hesperantha Ker Gawl. (77 species).

1 . The Babiana flabellifolia complex

Defined by its unusual, abruptly truncated leaves and

by its large floral bracts with the inner notched at the tips,

and so provisionally considered a clade by us, the B. fla-

bellifolia complex currently includes three species.

These are the central and northern Namaqualand species,

B. pubescens (Lam.) G.J .Lewis, which has an elongate

perianth tube ± 50 mm long and curved near the apex,

and a dorsal tepal ± 20 mm long; a second long-tubed

species from the western Karoo, identified as B. flabelli-

folia by Lewis (1959); and a widespread, relatively

short-tubed species, named B. truncata by Lewis (1959).

The type of Babiana flabellifolia was collected by the

Rev. Henry Whitehead at an unrecorded locality in

Namaqualand. Whitehead, an Anglican clergyman, was sta-

tioned at Modderfontein near Springbok, and his collections

are assumed to have been made nearby in northern

Namaqualand (Gunn & Codd 1981 ). The type of B. flabelli-

folia, however, exactly matches B. truncata G J.Lewis ( 1 959)

in its short tube, except for conspicuous pubescence on the

leaves and bracts, and is from Namaqualand where B. trun-

cata is common, and where the long-tubed western Karoo

species does not occur. B. truncata is typically glabrescent.

The latter taxon extends from Steinkopf in northern

Namaqualand to the Bokkeveld Plateau near Nieuwoudtville

in the south and its range thus includes that of B. flabellifolia.

We suggest that the glabrescent and pubescent plants with

perianth tubes of similar length represent a single species,

which must carry the earlier name. The conspicuous pubes-

cence of the type specimen is certainly unusual but sparse

pubescence is occasionally encountered in plants matching B.

truncata , especially in the immediate vicinity of Springbok.

We do not regard the hairiness of the type specimen as sig-

nificant enough to treat it as a separate taxon. We rename the

western Karoo plant B. praemorsa here.

In addition to a name change for the short-tubed

Namaqualand species, which must now be called

Babiana flabellifolia , a change to its circumscription

seems necessary. Some plants from the south of the range

of the species have a perianth tube 40-60 mm long and

large subequal tepals. We have concluded that they rep-

resent a separate species, which we here describe as B.

cuneata. This species extends from Lokenberg in the

Bokkeveld Mountains to the Cold Bokkeveld and

Swartruggens near Ceres in the south, and along the

Roggeveld Escarpment to near Laingsburg in the east. B.

cuneata has cupped tepals 26^10 mm long and filaments

15-18 mm long and is thus readily distinguished from

the long-tubed western Karoo species B. praemorsa

(called by Lewis B. flabellifolia-, see Lewis 1959: plate

21), which has a straight perianth tube 40-60 mm long,

horizontally spreading tepals 18-22 mm long, and rela-

tively short filaments, 8-9 mm long. The filaments in the

other long-tubed species in the complex, B. pubescens ,

are 14-15 mm long.

A revised description is provided below for Babiana

flabellifolia (syn. B. truncata ), as well as formal descrip-

tions of the new species B. cuneata and B. praemorsa.

We also lectotypify B. cuneifolia which was treated as a

synonym of B. pubescens by Lewis. Most specimens of

the type, Drege 2627 , actually have a short perianth tube

and accord closely with B. flabellifolia/ truncata .

Babiana pubescens is not known to occur in southern

Namaqualand where the type collection of B. cuneifolia

was made and we suspect that specimens of B. pubescens

from another site were later mislabelled with the Drege

collection number.

Babiana flabellifolia Harv. ex Klatt in Linnaea 35:

380 (1867-68); G.J .Lewis: 98 (1959), misapplied to B.

praemorsa. Type: South Africa, Namaqualand [Northern

Cape, probably near Modderfontein, west of Springbok],

without precise locality or date. Rev. H. Whitehead s.n.

(K, holo.!).

Babiana cuneifolia Baker: 335 (1876), syn. nov. Type: South

Africa, [Northern Cape], Mierenkasteel, karroid slopes, August 1830,

J.F.Drege 2627 (K, holo.!; MO!, P (two sheets), S, iso.).

Babiana truncata G.J.Lewis: 100 (1959), syn. nov. Type: South

Africa, [Northern Cape], 2 miles east of Springbok, 26 July 1950, GJ

Lewis 2203 (SAM, holo.!; SAM, iso.!).

Plants up to 80 mm high. Stem short or entirely under-

ground. Leaves crowded at base, abruptly truncate and

wedge-shaped, pleated, smooth or finely hairy with

shortly velvety margins, or rarely coarsely hairy with

white hairs up to 2.5 mm long; juvenile leaves filiform

with long hairs. Spike compact, 2-several-flowered,

borne at ground level; bracts 15-30 mm long, more or

less membranous or green, smooth or papillate to finely

hairy, inner bracts forked apically, tips tapering into slen-

der awns. Flowers zygomorphic, mauve to violet, tube

cream-coloured or pale yellow and lower lateral tepals

with pale yellow or cream-coloured median markings,

sweetly scented; perianth tube 18-36 mm long, cylindri-

cal, straight; tepals unequal, dorsal suberect or arching

forward, 27-33 mm long, lower tepals slightly shorter,

25-27 mm long. Stamens arched; filaments 15-18 mm

long; anthers 6-8 mm long. Ovary smooth; style dividing

near apex of anthers. Flowering time', mainly June and

July.

Bothalia 34,2 (2004)

89

Distribution : clay and granitic soils, renosterveld and

succulent karroid scrub, in Namaqualand and the western

Karoo, from the Anenous Mountains in the north to

Nuwerus and Koekenaap in the south and on the

Bokkeveld Plateau near Calvinia. Plants cited by Lewis

(1959) ( Compton 11504 and Schlechter 10892 ) from the

Bokkeveld Plateau south of Nieuwoudtville are B.

cune at a.

Babiana cuneata J .C Manning & Goldblatt, sp. nov.

TYPE. — Western Cape, 3219 (Wuppertal): Cold

Bokkeveld, Katbakkies Pass, (-DC), 5 September 2000,

P. Goldblatt & J.C. Manning 11457 (NBG, holo.; K,

MO, PRE, iso.).

Plantae 80-150 mm altae, caule brevi, glabro, foliis

usque ad 6, oblongis (6— )8— 1 5 x 8— 1 8(— 30) mm plicatis

abrupte truncatis glabris vel velutinis, spica compacta 2-

ad 5-(ad 7)-flora, bracteis 25-50 mm longis, interiore ad

apicem furcata, floribus zygomorphis violaceis tepalis

inferioribus albo-notatis, tubo perianthii 40-60 mm

longo, tepalis superioribus 26-3CK-40) x 6-12 mm, fila-

mentis unilateralibus 15-18 mm longis, antheris

6— 8(— 1 1) mm longis, ovario glabro.

Plants 80-150 mm high including leaves. Stem short,

hairless. Leaves crowded at base, up to 6, oblong,

(6— )8— 1 5 x 8—1 8(— 30) mm, leathery, plicate, abruptly

truncate, hairless or finely velvety hairy. Spike compact,

2-5(-7)-flowered, flowers borne at ground level; bracts

25-50 mm long, green with brown attenuate tips, smooth

or sparsely hairy, inner about as long as outer and forked

apically, basal margins sometimes fused around ovary

for ± 2 mm. Flowers zygomorphic, pale bluish to violet,

lower lateral tepals with white median marks outlined

with dark violet to purple below, unscented or sweetly

scented; perianth tube 40-60 mm long, cylindrical,

straight, widening slightly in upper 8 mm; tepals spread-

ing, subequal or three upper slightly larger, 26—30(^40) x

6-12 mm, upper laterals fused to lower for ± 2 mm.

Stamens unilateral; filaments 15-18 mm long, exserted

9-10 mm; anthers 6— 8(— 11) mm long, pale yellow to

cream-coloured. Ovary smooth, shortly stipitate in lower

flowers; style dividing opposite anther bases or apices,

branches ± 6 mm long, arching outward between or over

anthers. Flowering time : August to late September.

Figure 1 .

Distribution : dry rocky sandstone or dolerite slopes

and flats in arid fynbos or renosterveld, from the

Bokkeveld Mountains to the interior Cold Bokkeveld as

far south as Karoopoort, and eastward on the Roggeveld

Escarpment south to the foot of the Witteberg near

Laingsburg (Figure 2).

Diagnosis and relationships ; Babiana cuneata was

previously included by Lewis (1959) in the species that

she called Babiana truncata (now B. flabellifolia) but

comparison of live plants examined in the field shows

that the populations from the south of the range of that

species represent a distinct species, geographically and

edaphically separated from B. truncata. They are recog-

nized by their blue to blue-grey flowers (either fragrant

or apparently unscented) with an elongate perianth tube

40-60 mm long; ± equal, ascending tepals 26-301-40)

mm long; and openly displayed stamens with filaments

15-18 mm long and exserted 9-10 mm from the tube.

The point of division of the style varies. In the type popu-

lation from Katbakkies Pass, the branches separate oppo-

site the base of the anthers but in plants to the south, from

below the Roggeveld Escarpment, the style reaches the

anther apices. Flowering occurs from early August to late

September. Plants here included in B. flabellifolia have

in contrast strongly scented, bilabiate flowers with an

arched dorsal tepal more or less concealing the stamens,

a perianth tube 18-36 mm long, and often dry, submem-

branous, or occasionally green, leafy bracts. In speci-

mens where it is possible to see floral details, the style

divides opposite the anther apices. Flowering in B. fla-

bellifolia occurs from late June through July, rarely in

early August in seasons when the rains are late or condi-

tions are particularly cold.

The flowers of Babiana cuneata closely resemble

those of B. sambucina , which is widespread across dry

parts of the Cape Floral Region, but does not occur in the

Roggeveld and typically has lanceolate, acute or acumi-

nate leaves.

Histoiy: Babiana cuneata was evidently first collect-

ed by Rudolf Schlechter in August 1897 at Papelfontein

(sic.) [Papkuilsfontein], some 16 km south of Nieuwoudt-

ville at the northern end of its range. In 1941 R.H.

Compton collected the species nearby, at Lokenburg in

the Bokkeveld Mountains, a short distance to the south.

The range of B. cuneata falls partly within the arid inte-

rior of the Cape Floral Region but there are also several

recent records from the Roggeveld Escarpment and

Klein Roggeveld to the east. Most collections are from

rocky habitats, with the substrate rock either sandstone

or dolerite. Plants at the southern end of the range, in the

Klein Roggeveld south of Komsberg Pass, have unusual-

ly large flowers with the dorsal tepal ± 40 mm long and

anthers 9-11 mm long, and the style is also unusual in

dividing opposite the anther apices rather than opposite

their bases. These plants are also distinctive in their glau-

cous leaves, which are often 20-30 mm wide, which is

exceptionally broad for the species. Despite these differ-

ences, however, we see no reason at present to recognize

these plants as a separate taxon.

Other material examined

NORTHERN CAPE.— 3119 (Calvinia): Papkuilsfontein (‘Papel-

fontein’), (-AC), 17 August 1897 .Schlechter 10892 (MO); Lokenburg,

(-CA). 29 August 1941 , Compton 11504 (NBG); near Soutpan, (-CD),

24 July 1961, Lewis 5820 (NBG). 3120 (Williston): south of

Middelpos, (-CC), 30 Aug. 2001, Horstmann s.n. (NBG). 3220

(Sutherland): Sutherland, stony ground, (-BC), 15 August 1968,

Hanekom 1094 (NBG); Sutherland-Matjesfontein road at Komsberg

turnoff, (-DC), 19 September 2003, Goldblatt, Maiming & Porter

12310, (K, MO, NBG, PRE); Kleinroggeveld, near Farm Damslaagte,

(-DC), 19 Sept. 2003, Goldblatt & Porter 12312 (MO).

WESTERN CAPE.— 3219 (Wuppertal): turnoff to De Plaat,

(-DC), 14 September 1986, Fellingham 1196 (NBG). 3319

(Worcester): Katbakkies Pass, (-AD), 30 September 1974, Loubser

2250 (NBG); Karoopoort, 3 km north of Sutherland turnoff on Calvinia

road, (-BA), 6 August 2002, Manning 2761 (NBG). 3320 (Montagu):

Karoo Garden, Whitehill, Laingsburg, (-BA), 17 August 1942,

Compton 13389 (NBG); foot of the Witteberg near Laingsburg, (-BD),

1 8 August 2002, Goldblatt & Porter 12070 (MO).

90

Bothalia 34,2 (2004)

FIGURE 1 .—Babiana cuneata. A,

whole plant, B, C, flower: B,

front view; C, 1/s. Scale bar:

10 mm. Artist: John Manning.

Babiana praemorsa Goldblatt & J.C. Manning,

sp. nov. (see Lewis 1959: plate 21; Manning & Goldblatt

1997: 87). Babiana flabellifolia sensu GJ. Lewis: 98

(1959), not of Harv. ex Klatt., misapplied name.

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

Bloukrans Pass, shale Pats, (-DA), 25 August 1976, P.

Goldblatt 3935 (NBG, holo.; K, MO, iso.).

Plantae 50-150 mm altae, caule brevi raro usque ad

20 mm supra terram extenso, foliis oblongo-cuneatis pli-

catis minute villosis, spica 5- vel 6-flora congesta,

bracteis viridis minute villosis 25-50 mm longis, interi-

ore parum breviore, exteriore ad apicem furcata, floribus

zygomorphis atroviolaceis maculis lanceolatis albis vel

cremeis notatis, tubo perianthii 40-60 mm longo cylin-

drico, tepalis subaequalibus usitate 18-22 x 3.5-5 .0 mm

horizontaliter extensis, filamentis unilateralibus suberec-

tis 8-9 mm longis, antheris 5-6 mm longis, ovario

glabro.

Plants 50-150 mm high. Stem short, suberect or

decumbent, rarely reaching 20 mm above ground.

Leaves oblong-cuneate, pleated, minutely hairy, in a

spreading fan; juvenile leaves linear, ± 1 mm wide, silky

hairy. Spike congested, 5 or 6-flowered; bracts green,

minutely hairy, 25-50 mm long, inner slightly shorter

than outer and forked apically with acuminate tips.

Flowers weakly zygomorphic, dark violet with white to

cream-coloured spear-shaped marks often edged with red

or dark blue on lower lateral tepals, unscented or rarely

slightly sweet-scented; perianth tube 40-60 mm long,

cylindrical, straight; tepals subequal, spreading horizon-

tally, mostly 18-22 x 3. 5-5.0 mm. Stamens suberect; fila-

ments 8-9 mm long, exserted 4-5 mm; anthers 5-6 mm

long, violet or yellow. Ovary smooth; style dividing

Bothalia 34,2 (2004)

91

FIGURE 2.— Distribution of Babiana cuneata.

opposite upper half of anthers or shortly beyond them,

branches 2-3 mm long. Flowering time : June-July.

Distribution : dolerite outcrops in the Calvinia District

of the western Karoo, from the Hantamsberg to Bloukrans

Pass, often growing in rock crevices in dolerite pavement

where the corms are secure from predation.

Diagnosis and relationships : in her account of

Babiana , Lewis (1959) treated B. praemorsa as B.fla-

bellifolia , a species described by F.W. Klatt in 1867-68,

based on a specimen, from ‘Namaqualand Minor’. This

is certainly not the same as the plants from the Calvinia

District with which she associated the name. The type

specimen (a single plant) at the Kew Herbarium has

flowers with a short, obliquely funnel-shaped perianth

tube, ± 18 mm long with the narrow part ± 10 mm long,

and broadly cuneate leaves that are densely covered with

long hairs on the major veins and margins. Except for the

prominently hairy leaves and slightly shorter tube, this

specimen corresponds exactly to the Namaqualand plant

that Lewis called B. truncata, now reduced to synonymy

in B . flabellifolia .

One of four species of the Babiana flabellifolia com-

plex, B. praemorsa is recognized by its flowers with a

straight, cylindrical perianth tube 40-60 mm long, and

firm, narrow, spreading tepals, 18-22 mm long. The peri-

anth is deep violet rather than blue as in the related long-

tubed species of the alliance and they are perhaps most

like those of the central Namaqualand B. pubescens, with

which it is easily confused. Babiana praemorsa can be

distinguished from this and another long-tubed species,

B. cuneata from the dry interior of northern Western

Cape, by its short filaments, 8-9 mm long, exserted 4-5

mm from the perianth tube, and the tepals spreading at

right angles to a nearly straight tube of uniform diameter.

As well as occurring in the Calvinia District, Babiana

praemorsa was cited by Lewis as growing at ‘Upington,

Namaqualand’ ( Orpen s.n ., NBG), and at Whitehill

( Compton 13389 , NBG) south of Laingsburg. Whitehill

was the site of the first Karoo Botanical Garden, no

longer extant. The latter plant is B. cuneata from the

southern limit of its range. The Upington record is, how-

ever, almost certainly an error in labelling the prove-

nance of the plant, as Babiana praemorsa is evidently a

narrow endemic of dolerite outcrops in the Calvinia

District. Since it was treated by Lewis, B. praemorsa has

been collected at several localities not known to her,

extending the known range of the species.

Other material examined

NORTHERN CAPE. — 3119 (Calvinia): Hantamsberg summit

plateau, in cracks in dolerite outcrops, (-BD), 3 September 1994,

Goldblatt & Manning 9961 (MO); Kareeboomfontein, (-DA), 30

August 1974, Hanekom 2368 (K, M, MO, P, PRE).

2. The Babiana hypogaea complex

Recognized by the partly underground spike, the

flowers thus with a subterranean ovary, the slender, lin-

ear to falcate leaves, and, according to Lewis (1959), a

stipitate ovary, the Babiana hypogaea complex included

three species in Lewis’s (1959) account of Babiana.

These were B.falcata, B.flavida and B. hypogaea (for no

apparent reason spelled hypogea by Lewis). Examination

of type material of B. hypogaea , collected by William

Burchell in August 1812, and described by him in 1824,

shows the slender leaves and pale flower with tepals

darkly pigmented along the midline that closely corre-

spond to what Lewis called B. flavida. That plant must

therefore assume the earlier name B. hypogaea , whereas

plants with longer leaves and blue to violet flowers in

which the lower tepals are marked with white, and which

were associated with the name B. hypogea by Lewis

(1959) and later authors (e.g. Solch 1969; Goldblatt

1993), must be called B. bainesii. Confusingly, in the

protologue of B. hypogaea, Burchell (1824) described

the flower colour as blue ( caeruleis ), possibly an error or

the result of basing the description on his pressed speci-

mens in which the tepals have a dull mauve cast, as do

all dry specimens of the species. This is a result of the

purple pigmentation on the reverse of the tepals, masking

the dull yellow colouring of the inside of the tepals. Our

revised taxonomy corresponds to the treatment of the

complex by Baker (1896) in Flora capensis.

The so-called stipitate ovary described by Lewis

(1959) in Babiana hypogaea is misleading. Species of

Iridaceae with an inflorescence (typically a spike) borne

at or below ground level often have shortly stipitate

lower flowers and we suggest that this is simply a devel-

opmental aberration of the inflorescence due to its posi-

tion. This feature is associated with the underground

inflorescence in other Iridaceae, including Crocus L.,

Duthiastrum MP.de Vos and Romulea, and even some

other Babiana species with spikes borne at ground level

(e.g. B. cuneata and B. sambucina (Jacq.) Ker Gawl.)

and does not in itself mark the complex as sharing a

synapomorphy other than the underground spike.

Babiana bainesii and B. hypogaea are morphological-

ly similar in their underground spikes with long-tubed

flowers arising below the ground, and in their narrow,

linear leaves. The significant features of Babiana

92

Bothalia 34,2 (2004)

hypogaea are the pale yellow to buff flowers with a tube

30^-0 mm long, and the shorter, often sparsely hairy to

almost smooth leaves, sometimes with inclined to pros-

trate blades. The blades are often visibly constricted" and

apparently flexible at the base, which suggests an

inclined to prostrate orientation even when they appear

upright in pressed specimens. In contrast, B. bainesii has

larger flowers with a tube (40-)50-60(-70) mm long,

blue-mauve to violet flowers with prominent white

markings edged in darker blue on the lower lateral tepals,

and densely hairy leaves, mostly 15-25 mm long.

Misunderstanding the identity of the type collection,

Lewis (1959: 112), united B. bainesii with B. hypogaea

and described B. flavida for pale yellow- to buff-flow-

ered plants largely of Bushmanland, that mostly flower

in the winter and early spring. This species matches

closely Burchell’s type collection in its smaller flower

size, colour and other details, as well as in its habitat and

winter to spring flowering.

Babiana hypogaea occurs west and south of the range

of B. bainesii , in the northwestern Great Karoo,

Bushmanland, and adjacent southeastern Namibia. It

mostly flowers in winter and early spring, June to

September (rarely as late as December, according to the

type of B. flavida ), in contrast to February to April or

May for B. bainesii. The flowering time and ranges of

both species overlap to some extent in the Kimberley-

Vryburg area of Northern Cape and North-West but there

is no indication that they converge in floral morphology

along the area of overlap. Flowering in Babiana hypo-

gaea appears to be related to rainfall timing, normally

coming into bloom in winter and early spring in response

to autumn showers but blooming in summer when occa-

sional good early summer rains fall across Bushmanland

and the Upper Karoo.

While Babiana hypogaea and the plant described by

Lewis (1959) as B. flavida are essentially identical,

another species of Bushmanland with similar flowers, B.

falcata G.J. Lewis was distinguished by its short leaves,

25-70 mm long and 1. 5-3.0 mm wide, that are falcate,

firm, strongly plicate, and somewhat pungent. Lewis

contrasted the species with B. flavida (i.e. B. hypogaea),

which she described in her key to the species as having

leaves erect or decumbent but not firm or falcate. The

range of specimens now available make the distinction

untenable. Leaf shape ranges without any clear division

from the short, falcate, and firm type in the west to the

suberect, more prominently hairy type in the east. We

therefore, include B. falcata in B. hypogaea. Some col-

lections even show a range of leaf shapes from linear and

erect, to sword-shaped, to falcate.

Babiana hypogaea Burch., Travels 2: 415 (1824).

Antholyza hypogaea (Burch.) Klatt in Abhandlungen der

Naturforschenden Gesellschaft zu Halle 15: 345 (1882).

Type: South Africa, [Northern Cape|, Pellat Plains near

Takun (or Litakun), 24 August 1812, W..1 . Burchell 2241

(K, lecto.!, here designated; B, P, iso.).

Babiana flavida G.J. Lewis: 1 17 (1959), syn. nov. Type: South Africa,

[North-West |, Warrenton, December 1922, C.G. Adams 819 (BOL.

holo.!).

Babiana falcata G.J .Lewis: 115 (1959), syn. nov. Type: South

Africa, [Northern Cape], 20 miles east of Springbok, red sand flats, 8

September 1950, W.F. Barker 6671 (NBG, holo.!).

Plants acaulescent, 50-80 mm high, often growing in

tufts with leaves up to 150 mm long. Stem subterranean,

simple or with short or vestigial lateral branches. Leaves

linear to falcate or sword-shaped, often bent near base,

sometimes inclined toward ground or evidently prostrate,

25-150 x 1.5-3.0 mm, slightly pleated, sparsely hairy to

finely long-haired, somewhat pungent. Spike arising

below ground level, with 1 or 2 flowers per branch,

seemingly in a 2-6-flowered spike; bracts smooth, more

or less membranous below, green in upper third where

they reach above ground, 25-50 mm long, inner as long

as or slightly exceeding outer and forked apically with

attenuate tips. Flowers zygomorphic, greenish yellow to

buff, flushed with pale brown or mauve outside, tepals

darker in midline especially when dry, lower tepals with

pale nectar guides edged with reddish arrow-shaped

marks near base and with dark reddish streaks in throat,

strongly scented; perianth tube emerging from below

ground, cylindric, widening at throat, 30^10 mm long;

tepals subequal, dorsal 35-42 mm long, lower tepals

joined to upper laterals for up to 6 mm, forming a promi-

nent lip, lower tepals 30-35 mm long. Stamens unilater-

al; filaments 15-18 mm long; anthers 8-11 mm long.

Ovary smooth, shortly stipitate in lower flowers; style

dividing close to anther apices. Flowering time', mainly

June to September, occasionally December to May.

Distribution: flats on red sand plains in the Upper

Karoo, Bushmanland, and southeastern Namibia.

Additional collections made since the publication of

Lewis’s monograph represent range extensions into south-

ern Namibia where B. hypogaea in its current sense was

previously known only from a collection near Graspoort.

The new record from near Kimberley ( Leistner 2635),

identified by Lewis as B. falcata subsequent to the publi-

cation of her monograph of Babiana, is a particularly close

match to the type collection of B. hypogaea.

Other material examined

NAMIBIA.— 2716 (Witpiitz): Farm Witpiitz Sud, (-DA), 1983,

Lavranos 21225 (NBG); Spitskop, (-DC), 15 July 1986, Van Berkel

556 (NBG). 2816 (Aus): Farm Kubub, (-CB), 9 Sept. 1973, Giess

12858 (K, M, NBG). 2718 (Griinau): Farm Carolina, red sandy flats,

(-AD). 17 May 1972, Giess & Miiller 12041 (K, M). 2818 (Warmbad):

Farm Sperlingspiitz, (-CA), sandy plains, 16 May 1963, Giess. Volk &

Bleissner 7009 ( K.M.MO).

NORTHERN CAPE. -2724 (Taung) Andalusia, sandboden,

(-DD), 9 July 1942, Giess 298 (M). 2824 (Kimberley): 7 miles north-

east of Kimberley, (-DB), 20 August 1961 .Leistner 2635 (K, M, PRE);

4 miles west of Kimberley, red sand, (-DB), 21 July 1963, Leistner

3148 (K, M, PRE).

Babiana bainesii Baker in Journal of Botany

1876: 335 (1876). Type: South Africa, [Gauteng], ‘Gold

Fields’, Witwatersrand, near Johannesburg, 1870, J.T.

Baines s.n ., (K, lecto.!, here designated).

Babiana schlechteri Baker: 865 (1901), syn. nov. Type: South Africa,

[Mpumalanga], Witbank, 22 December 1893, R. Schlechter 4055 (Z,

holo.).

Bothalia 34,2 (2004)

93

Babiana hypogaea [hypogea] var. longituba G.J .Lewis: 115 (1959),

syn. nov. Type: South Africa, [Limpopo], Mount Myorul, 29 March

1894, R. Schlechter 4729 (BOL, holo.!; K, iso.!).

Babiana hypogaea [ hypogea ] var. ensifolia G.J .Lewis: 1 14 (1959),

syn. nov. Type: South Africa, Eastern Cape, near Murraysburg, August

1879, W. Tyson 312 (SAM. holo.!; K. SAM, iso.!).

See Lewis (1959) and Goldblatt (1993) for complete synonymy

(under the name B. hypogea).

Plants mostly 150-250 mm high (leaves only), some-

times growing in tufts. Stem underground, often

branched. Leaves linear to sword-shaped, pleated, much

exceeding flowers, 15-25 x 3-10 mm, densely hairy,

scabrid, or virtually smooth. Semi-spike compact, 2-8-

flowered, borne below ground level, lateral flowers usu-

ally with short stalks; bracts membranous to papery with

dry, rusty tips or dry and rusty throughout, sparsely hairy,

mostly 35-60 mm long, inner slightly shorter than outer

and forked at tips. Flowers zygomorphic, shades of blue

to violet or mauve with tepals paler (rarely white) toward

edges, lower lateral tepals with white markings often

edged in dark blue, usually sweetly scented; perianth

tube mostly 40-60(-70) mm long, cylindric with

expanded throat; dorsal tepal 40-50 mm long, upper lat-

erals joined to lower for ± 6 mm to form a lip, lower

tepals 25 — 40 mm long. Stamens unilateral; filaments

10-15 mm long; anthers 8— 10(— 1 2) mm long. Ovary

smooth, shortly stipitate in lower flowers; style usually

dividing more or less opposite anther apices, branches ±

6 mm long. Flowering time : mainly in summer, February

to April, occasionally in December or January; southern

Karoo populations flower in August or September.

Distribution: stony or sandy slopes and flats in dry

grassland and bush, across summer-rainfall southern

Africa from Murraysburg and Carnarvon in the Upper

Karoo to Zimbabwe and southern Zambia in the north-

east, and to northern Namibia in the northwest.

The most widespread species of the genus, Babiana

bainesii extends from the central Upper Karoo to south-

ern Zambia (Lewis 1959; Goldblatt 1993), thus having a

range greater than the rest of the genus. It grows in grass-

land on soils ranging from deep Kalahari sands to stony

slopes and flats, and typically flowers from late summer

until late autumn, the exact timing in dry areas probably

depending on rainfall in areas of low annual rainfall.

With their narrow, erect leaves, plants are difficult to see

among the grass where they grow, but when in bloom can

often be located by the strong, sweet scent of the bluish

to violet flowers.

Lewis (1959) recognized two additional varieties of

B. hypogea : var. ensifolia G.J.Lewis from the Upper

Karoo, and var. longituba G.J. Lewis from the mountains

of Limpopo. The latter was distinguished by its densely

hairy leaves and bracts, and elongate perianth tube up to

70 mm long. The long tube in these plants is only the

extreme of a range that includes many plants with a tube

55-65 mm long (the type of B. bainesii from the

Witwatersrand has a tube nearly 60 mm long). Near

Haenertsburg in Limpopo, close to the type locality of

var. longituba , plants have a perianth tube 35-59 mm

long ( n = 15) (P.J.D. Winter pers. comm.). Information

now available indicates that var. longituba does not merit

taxonomic recognition and we reduce it to synonymy in

B. bainesii.

Babiana hypogea var. ensifolia has comparatively

short, broad leaves, 50-140 x 10-14 mm, unlike the nar-

row, sparsely to densely hairy, narrowly sword-shaped to

linear leaves, mostly 150-250 x 3-10 mm, of typical B.

bainesii. It approaches in general appearance the largely

southern Cape species, B. sambucina but has the entirely

subterranean inflorescence, short lateral branchlets, and

rusty bracts of B. bainesii. These plants flower in early

spring, August and September. Var. ensifolia is poorly

documented and requires additional study before its sta-

tus can be established. It may be recognized or not at the

discretion of the reader but we hesitate to make the new

combination in B. bainesii until more is learned about

these plants from the central parts of the Upper Karoo.

Babiana schlechteri. described by J.G. Baker in 1901 ,

and based on a collection from Witbank, east of

Johannesburg, does not differ in any significant way

from the type of B. bainesii from near Johannesburg.

3. The Babiana stricta complex

Lewis (1959) recognized five varieties in the south-

western Cape species, Babiana stricta , all with the inner

bracts divided to the base and a densely hairy ovary, fea-

tures present in many other species. Listed in the se-

quence presented by Lewis, the varieties and their sig-

nificant taxonomic features are as follows;

1 . var. stricta— leaves stiff, narrow, with short rough

pubescence; stem suberect; flowers zygomporphic, blue;

tepals subequal and spreading, lower three with white

markings; anthers unilateral [incorrectly said by Lewis to

be symmetrically arranged], blue, arrow-shaped with

connective wider toward base; style mostly dividing

opposite the anther bases.

2. var. erectifolia — similar to var. stricta but flowers

white to cream-coloured, also with spreading tepals,

lower with pale yellow markings; anthers unilateral, dark

blue-black, with connective broader below.

3. var. sulphurea — similar to var. stricta but flowers

yellow with unilateral dark blue anthers with connective

broader toward the base;

4. var. regia — strikingly different from the above

three varieties— -leaves stiff, narrow, with short pubes-

cence; stem spreading horizontally; flowers radially

symmetric, blue with dark red centre; tepals cupped;

anthers linear, dark brown, without visible connective;

style short, dividing below bases of anthers.

5. var. grandiflora— leaves broad with soft pubescence;

stem suberect; flowers subactinomorphic, blue-mauve;

tepals spreading, perianth tube 22-32 mm long; anthers

unilateral, dark blue, linear with narrow connective visible;

style dividing opposite middle and apex of anthers.

We see no merit in this taxonomy, in which Babiana

stricta has a wider circumscription than any other species

of the genus and seems to comprise disparate elements.

Var. grandiflora, which we have re-collected at its type

locality north of Piketberg, evidently represents a distinct

94

Bothalia 34,2 (2004)

species. Its broad, soft-textured leaves, and faintly scent-

ed flower with relatively long perianth tube, hollow to the

base, suggests that it is more closely related to B. disticha,

the type of the genus. Likewise, the features of var. regia

are so distinctive that it too must represent a separate

species, not obviously immediately related to B. stricta.

In contrast to these two examples, we see no reason to

separate Babiana stricta var. erectifolia from var. stricta , and

thus unite them. Lastly, plants assigned to var. sulphurea

seem to us merely cream-coloured to pale yellow-flowered

variants of B. stricta but we cannot identify the type of

Gladiolus sulphureus, the basionym of var. sulphurea.

Babiana stricta (Aiton) Ker Gawl. in Curtis’s

Botanical Magazine 17: t. 621 (1803a). B. stricta var. stricta

Lewis: 40 ( 1959). Gladiolus strictus Aiton, Hortus kewensis

1: 63 (1789). Type: South Africa, without precise locality,

illustration in Curtis's Botanical Magazine 17: t. 621 (1803a)

(neo., here designated).

Babiana stricta var. erectifolia (G.J .Lewis) G.J .Lewis: 43 (1959). B

erectifolia G.J .Lewis: 3 (1938), syn. nov. Type: South Africa, [Western

Cape], Brand Vlei, near Worcester, Sept. 1932, GJ. Lewis s.n. (holo.!,

NBG2686I32 in BOL).

The complete synonymy of Babiana stricta is present-

ed by Lewis (1959: 40) and is not repeated here. Note that

we also include B. stricta var. sulphurea sensu G.J .Lewis

(based on Gladiolus sulphureus Jacq.) in B. stricta but do

not regard the type, an illustration in Jacquin’s leones

plantarum rariorum (1793), as matching B. stricta and

are in fact unable to recognize the plant illustrated.

As here circumscribed, Babiana stricta comprises

plants with narrow, fairly stiff, roughly hairy leaves and a

zygomorphic flower with unilateral stamens and anthers

slightly sagittate with the connective expanded in the

lower half. Lewis described the flower as actinomorphic

or almost so, but we have seen no plants in the field that

have such flowers, nor does the type illustration show an

actinomorphic flower. The tepals are subequal in length

but the dorsal is held somewhat apart from the others and

the lower three bear contrasting markings, either white to

cream-coloured, edged in a darker colour when the peri-

anth is predominantly blue or pink, or yellow when the

perianth is otherwise cream-coloured. The perianth tube

is 12-18 mm long and almost filiform, the walls tightly

enveloping the enclosed style so that nectar is forced into

the slightly wider distal portion of the tube. The stamens

are erect, but clearly unilateral, with anthers facing the

lower tepals and the style is held against the back of the

anthers. Plants with symmetrically disposed stamens as

illustrated by Lewis evidently exist but are not usual.

Babiana regia (G.J. Lewis) Goldhlatt & J C. Man-

ning, comb, et stat. nov.

Babiana stricta var. regia G.J. Lewis in Journal of South African

Botany , Suppl. 3: 45 (1959). Type: South Africa, [Western Cape],

Bottelary road near Stellenbosch, September 1950, G.J . Lewis & D.K.

Davis 22/6 (SAM, holo.!, iso.!).

As outlined above, Babiana regia has strikingly marked

tepals, deep blue with a red base. Although known since at

least the early 19th century, when it was collected by J.F.

Drege (Lewis 1959), it was long confused with B. rubro-

cyanea (Jacq.) Ker Gawl., which has a similarly coloured

flower but unilateral stamens and unusual broad style

branches. Lewis realized that these plants were different

species and recognized B. stricta var. regia for plants with

symmetrically disposed stamens and short, slender style

branches. However, it sits uncomfortably in B. stricta

because the flower is radially symmetric, the tepals nearly

equal and cupped even when fully open, and the brown

anthers are linear, lacking the expanded connective of B.

stricta. Vegetatively the two also differ, for B. regia has an

arching stem nearly horizontal above, whereas the stem of

B. stricta is suberect.

Babiana regia is restricted to the Western Cape fore-

lands between Malmesbury and Stellenbosch and favours

sandy soils. As far as we are aware, it persists today at

just one site, adjacent to the Farm Joostenbergkloof, on

the northwest foothills of the Joostenberg, between

Durbanville and Paarl. It co-occurs with other rare species,

notably Geissorhiza purpurascens and Hesperantha spi-

cata subsp./Am/oxa in a small area heavily infested with

Australian Acacia species. We suspect that all three

species will soon be extinct unless some action can be

taken to conserve them.

Babiana longiflora Goldblatt & J .C .Manning, sp.

nov.

TYPE.— Western Cape, 3218 (Clanwilliam): ± 28 km

north of Piketberg, (-CC), 31 August 2002, Goldblatt &

Porter 12129 (NBG, holo.; MO, iso.).

Babiana stricta var. grandiflora G.J .Lewis: 46 (1959). Type: South

Africa, [Western Cape], 18 miles north of Piketberg, 28 July 1950, G.J.

Lewis 2197 (SAM, holo.!, iso.!).

Plantae usitate 150-200 mm altae, caule erecto saepe

ramoso, foliis oblongis vel lanceolatis plicatis sparse vil-

losis, spica 6- ad 10-flora inclinata, bracteis viridis ad

apicem brunneis pubescentibus 16-20 mm longis, interi-

ore parum breviore, exteriore usque ad basem diviso,

floribus zygomorphis lilacinis usque atromalvinis tepalis

inferioribus albo-notatis, tubo perianthii 25-30 mm

longo cylindrico ad apicem expanso, tepalis subaequal-

ibus 20-24 mm longis adscendentibus, filamentis unilat-

eralibus leviter arcuatis ± 1 5 mm longis, antheris ± 6 mm

longis purpureis, ovario dense piloso.

Plants mostly 150-200 mm high; stem erect, often

branched, velvety hairy. Leaves lanceolate to oblong,

mostly 9-12 mm wide, slightly pleated, fairly soft-tex-

tured, sparsely hairy. Spike slightly inclined, 6-10-flow-

ered; outer bracts mostly 16-20 mm long, ± obtuse,

green with dry, rusty tips, densely hairy; inner bracts

slightly shorter than outer, divided to base. Flowers

zygomorphic, lilac to deep mauve, lower tepals each

with spear-shaped dark basal mark, white in centre, or

rarely with narrow median white stripe, with faint rose

scent; perianth tube slender, straight, widening in upper

third, 25-30 mm long; tepals subequal, ascending, 20-24

mm long, dorsal about 1-3 mm longer than lower, upper

laterals joined to lower for ± 2 mm. Stamens unilateral,

slightly arched, mauve, ± 15 mm long; anthers dark pur-

Bothalia 34,2 (2004)

95

pie, ± 6 mm long, with narrow connective visible. Ovary

densely hairy; style dividing between middle and apex of

anthers, branches 3^1 mm long, expanded and densely

ciliate at tips, exceeding anther apices. Flowering time :

mainly mid August to mid September.

Distribution : sandstone outcrops in transitional fyn-

bos-renosterveld. B. longifilora is known only from

rocky flats at the foot of the Piketberg and Porterville

Mountains of Western Cape.

Diagnosis and relationships: Babiana iongifiora has

been rarely collected, mostly at the same locality at the

foot of the Piketberg or close by, and we consider its

affinities to be have been misunderstood. It seems to us

not to be closely allied to B. stricta nor to any of its four

varieties that were recognized by Lewis. In particular, it

has fairly broad leaves that are only slightly plicate and

with soft pubescence, unlike the fairly rigid, narrow and

deeply pleated, ± erect and shortly hairy leaves of B.

stricta. Plants have long-tubed, pale purple flowers with

subequal, ascending tepals, and long, rather blunt outer

bracts. We suspect that it may be most closely related to

B. fragrans, which has comparable, soft-textured, fairly

broad leaves with long hairs, and large floral bracts rusty

only at the tips. B. Iongifiora can be recognized among

the species of section Babiana by the short floral bracts,

the inner divided to the base, a perianth tube 25-30 mm

long (relatively long for the section) and dark violet sta-

mens reaching to about the middle of the dorsal tepal.

Other long-tubed species with inner bracts divided to the

base, .6. ecklonii and B. latifolia , are unlike B. Iongifiora

in their dark violet perianth, the tube 30^-7 mm long,

curved at the apex, and the lower tepals joined to one

another for at least 4 mm (5-7 mm in B. ecklonii ), thus

forming a strongly bilabiate perianth with horizontally

extended lower tepals and the dorsal erect. Similar in

vegetative features, B. fra grans has strongly scented

flowers with a perianth tube 18-20 mm long, broader,

usually pale blue (rarely pale yellow) tepals, and the

style dividing opposite the lower third of the anthers.

We include plants from the foot of the Porterville

Mountains at Twenty Four Rivers, some 40 km east of the

type locality here. These plants have dark blue flowers

but otherwise closely resemble other specimens of

Babiana Iongifiora, notably in their straight perianth tube

± 30 mm long, hollow to the base and containing nectar.

Other material examined

WESTERN CAPE.— 3218 (Clan william): 18 miles north of Piket-

berg, (-CC), 28 July 1950, Barker 6371 (NBG). 3318 (Cape Town):

Twenty Four Rivers, sandy alluvium, (-BB), 1 September 1992 .Goldblatt

& Manning 9363 (MO, NBG).

The new combination, Babiana fragrans for the type

of the genus, B. disticha

In Lewis’s (1959) account of what she called Babiana

plicata, the type species of Babiana, she cited as syno-

nyms both B. disticha Ker Gawl. and Gladiolus fragrans

Jacq. Bullock (1961) in his review of Lewis’s mono-

graph of the genus pointed out that B. plicata is a super-

fluous name for Gladiolus fragrans and suggested that

the correct name for the plant was B. disticha. The earli-

er name G. fragrans was not considered by either Lewis

or Bullock as available for transfer to Babiana because

of the homonym, B. fragrans Eckl. (1827), a synonym of

B. nana (Andrews) Spreng. Babiana fragrans Eckl.,

however, is a nomen nudum and thus is invalid and can-

not be taken into consideration in questions of priority of

species epithets. The name for the type species of

Babiana thus becomes B. fragrans. We have examined

the type of G. fragrans, an illustration in the Hortus

schoenbrunnensis , and concur with Lewis that this

species and B. disticha are synonyms.

Babiana fragrans (Jacq.) Goldblatt & J .C Man-

ning, comb. nov.

Gladiolus fragrans Jacq., Plantarum rariorum Horti Caesarei

Schoenbrunnensis descriptiones et icones 1: t. 14 (1797). Type: South

Africa, [Western Cape], without precise locality or collector, cultivated

in Vienna, illustration in Jacq., Hortus Schoenbrunnensis t. 14 (1797).

Babiana plicata Ker Gawl.: t. 576 (1802), nom. illeg. superfl. pro

Gladiolus fragrans Jacq. ( 1797). Type: illustration in Curtis’s Botanical

Magazine 16: t. 576 (1802), South Africa, [Western Cape], without

locality.

Babiana disticha Ker Gawl.: t. 626 (1803b). Type: South Africa,

[Western Cape], without locality, illustration in Curtis’s Botanical

Magazine 17: t. 626 (1803b).

See Lewis (1959: 53) for additional synonyms of the species.

A relatively unspecialized species, Babiana fragrans

is recognized by the subequal, spreading tepals with the

dorsal only slightly longer than the lower, and small pale

markings edged in darker blue or purple on the lower

tepals. The perianth tube is about as long as or slightly

longer than the dorsal tepal and the erect, usually

branched stem bears spikes of up to 10 flowers. The

suberect stamens are unilateral and the anthers parallel

and contiguous, and usually dark blue. The soft-textured,

hairy leaves are weakly pleated and often oblong rather

than the usual sword shape of most species of the genus.

Both Ker Gawler (1802), when describing the synonym

B. plicata, and Lewis (1959) remarked on the strong,

pleasing fragrance, likened by Lewis to that of a carna-

tion. Somewhat variable across its range, B. fragrans

from the interior southwestern Cape has a more strongly

bilabiate flower with the dorsal tepal up to 5 mm longer

than the lower and the upper lateral tepals are united for

a short distance with the lower, thus forming a more pro-

nounced lip than is evident in plants from the Cape

Peninsula and nearby.

The confused pre-1800 taxonomic history of Babiana

plicata was outlined in detail by Lewis (1959) and is not

repeated here.

ACKNOWLEDGEMENTS

Support for this study by grants 6704-00 and 7103-01

from the National Geographic Society is gratefully

acknowledged. Collecting permits were provided by the

Nature Conservation authorities of Northern Cape and

Western Cape, South Africa. We thank Ingrid Nanni and

Lendon Porter for their assistance and companionship in the

field and Roy Gereau for revising our Latin descriptions.

96

Bothalia 34,2 (2004)

REFERENCES

AITON. W. 1789. Hortus kewensis, vol. 1 . Nicol, London.

BAKER, J.G. 1876. New Gladioleae. Journal of Botany, London 14:

333-339.

BAKER, J.G. 1892. Handbook of the Irideae. Bell, London.

BAKER. J.G. 1896. Irideae. In W.T. Thiselton-Dyer, Flora capensis 6:

7-171. Reeve, London.

BAKER, J.G. 1901. Beitrage zur kenntnis der Afrikanischen-Flora.

Bulletin de I’ Herbier Boissier, ser. 2, 1: 865.

BROWN, N.E. 1932. Contributions to a knowledge of the Transvaal

Iridaceae. Part II. Transactions of the Royal Society of South

Africa 20: 261-280.

BULLOCK, A.A. 1961. Review: The genus Babiana. Kew Bulletin 15: 168.

BURCHELL, W.J. 1824. Travels in the interior of southern Africa, vol.

2. Longman, Hurst, Rees. Orme, Brown & Green, London.

ECKLON, C.F. 1827. Topographisches Verzeichniss der Pflanzen-

sammlung von C.F. Ecklon.

GOLDBLATT, P. 1979. New species of Cape Iridaceae. Journal of

South African Botany 45: 81-89.

GOLDBLATT, P. 1990. Status of the southern African Anapalina and

Antholyza (Iridaceae), genera based solely on characters for

bird pollination, and a new species of Tritoniopsis. South

African Journal of Botany 56: 577-582.

GOLDBLATT, P. 1993. Iridaceae. In G.V. Pope, Flora zambesiaca

12,4: 1-106.

GOLDBLATT, P„ MANNING, J.C., DAVIES, T.J., SAVOLAINEN, V.

& RESHAI, S. 2004. Cyani.xia, a new genus for the Socotran

endemic. Babiana socotrana (Iridaceae subfamily Crocoideae).

Edinburgh Journal of Botany 60: 517-532.

GUNN, M. & CODD, L.E. 1981. Botanical exploration of southern

Africa. Balkema, Cape Town.

JACQUIN, N.J. VON. 1793. leones plantarum rariorum 2: t. 239.

Vienna.

JACQUIN, N.J. VON. 1797. Plantarum rariorum Horti Caesarei

Schoenbrunnensis descriptiones et icones 1: t. 14. Vienna.

KER GAWLER. J. 1802. Babiana plicata. Curtis's Botanical Maga-

zine 16: t. 576.

KER GAWLER, J. 1803a. Babiana stricta. Curtis’s Botanical Maga-

zine 17: t. 621.

KER GAWLER, J. 1803b. Babiana disticha. Curtis’s Botanical Maga-

zine 17: t. 626.

KER GAWLER, J. 1805. Ensatorum ordo. In J. Konig & J. Sims, An-

nals of Botany 1: 219-247.

KLATT, F.W. 1867-68. Diagnoses Iridearum novarum. Linnaea 35:

380.

KLATT, F.W. 1882. Erganzungen und Berichtigungen zu Baker’s

Sy sterna Iridacearum. Abhandlungen der Naturforschenden

Gesellschaft zu Hcdle 15: 44-404.

LEWIS, G.J. 1938. Eight new Iridaceae from the Cape Province.

Journal of South African Botany 4: 1-1 1 .

LEWIS. G.J. 1959. The genus Babiana. Journal of South African

Botany , Suppl. 3.

MANNING, J.C. & GOLDBLATT, P. 1997. Nieuwoudtville , Bokkeveld

Plateau and Hantam. South African Wild Flower Guide 9.

Botanical Society of South Africa, Cape Town.

NORDENSTAM, B. 1970. Notes on South African Iridaceae:

Lapeirousia and Babiana. Botaniska Notiser 123: 431^143.

SOLCH, A. 1969. Iridaceae. In H. Merxmiiller, Prodromus einer Flora

von Siidwestafrika. 155: 6-10. Cramer, Lehre.

Bothalia 34,2: 97-102 (2004)

Studies in the genus Riccia (Marchantiales) from southern Africa. 27.

Riccia lanceolata and R. radicosa now also locally reported

S.M. PEROLD*

Keywords: Riccia lanceolata Steph., Riccia radicosa Pearson, sporangia, spores, thallus

ABSTRACT

Riccia lanceolata and R. radicosa are newly reported from southern Africa. Riccia lanceolata is quite widespread in

sub-Saharan Africa, whereas R. radicosa has only been collected once before, namely in the Democratic Republic of the

Congo (Zaire). Both species are assigned to the group ‘Squamatae’, section Riccia , subgenus Riccia, but are easily distin-

guished from one another.

Riccia lanceolata Steph. in Hedwigia 27: 110(1 888).

INTRODUCTION

Riccia lanceolata was originally described by Stephani

(1888) from a specimen collected by Newton (G) in

Canne, Dahomey (Benin). It was subsequently shown to

be widespread in sub-Saharan Africa, Jones (1957)

reporting it from Nigeria and Uganda, while Jones &

Harrington (1983) mention its occurrence in Ghana as

well. Perold (1993) described and illustrated a specimen

collected by Frahm in Rwanda, whereas Fischer (1995),

besides illustrating it, also provided notes on its commu-

nities and distribution patterns in Rwanda. Perold (1995)

extended the list to include Ivory Coast, Malawi,

Mozambique, Niger, Tanzania and Zambia. The present

report is, however, the first record for southern Africa,

where it was collected in South Africa, near the Gauteng-

Mpumalanga border and in Fimpopo (Northern Province),

at Penge (Figure 1).

DIAGNOSTIC CHARACTERS

Although the thalli of R. lanceolata are more robust

than those of R. atropurpurea Sim, there is a close

resemblance between the two species: both are dorsally

glaucous green and have wavy, hyaline thallus margins.

In R. lanceolata however, the finely reticulate dorsal

surface lacks any deposit of salts, as is usually found in

R. atropurpurea and its purple flanks often appear to be

vertically ‘striped’, due to the ‘leading’ hyaline margins

of the overlapping scales. The spores are quite large,

sometimes up to 120/<m in diameter, triangular-globular,

polar and reddish brown, with the wing distinct, ± 5 pm

wide and faintly granular with the marginal angles

unperforated. The surface is often incompletely reticu-

late on both spore faces: the distal face is convex, the are-

olae, (5)6-8 across, variously incomplete and not

extending to the margin, (10-) 12-22 pm wide, the cen-

tral walls up to 5 pm high and raised at the nodes, the

outer walls low and often absent. The proximal face has

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

MS. received: 2004-05-31.

FIGURE 1 .—Localities of Riccia lanceolata, •; and R. radi-

cosa, ■ , in southern Africa.

a well-defined triradiate mark and the facets are mostly

with poorly developed areolar walls or only with discon-

nected low walls, occasionally almost smooth.

It should be stressed, however, that the ornamentation

of the spores of even the type specimen (Jones 1957: 219)

of this species, Newton 6 (G !), shows some variation

(Figure 2A-C), as do others as well, e.g. LA. Lye B320

(E, ex Herb. Jones!) (Figure 2D-F). Jones & Harrington

(1983), in an extreme example, described ‘the outer face

as areolate over the whole of its surface’ in Woodell’s col-

lection from Ghana. Micrographs of one of the South

African specimens are included (Figure 3A-C). Previous

publications including spore micrographs of R. lanceola-

ta are Perold (1993, 1996) and Fischer (1995).

South African specimens examined

MPUMALANGA.— 2529 (Witbank): near Gauteng/Mpumalanga

border, 20 km N of Witbank Railway Station, on dirt road to Zaaihoek,

turnoff to Farm Kwarsspruit and ± 5 km along rough track to Olifants

River, (-CA), on soil in vlei area, between grass, 20-02-1994, Perold

3189,3194 (PRE).

LIMPOPO (Northern Province).— 2430 (Pilgrim’s Rest): Penge,

western slope, on soil overlying quartzite, 1994, Swartz CHI 3682

(PRE).

98

Bothalia 34,2 (2004)

FIGURE 2. — SEM micrographs of Riccia lanceolata spores, A-C, Newton 6 (G!): A, distal face; B, C, proximal face. D-F, C.A.

Lye B320 (E, ex Herb. Jones): D, distal face; E, F, proximal face. A, x 410; B, x 400; C, x 450; D, F, x 460; E, x 455.

Riccia radicosa Pearson in Natuurwetenschappe-

lijk tijdschrift 4; 142 (1922).

INTRODUCTION

The description of this new species was originally

presented by W.H. Pearson at the 20th Vlaamsche

Natuur-, Wis- en Geneeskundig Congres August 1921,

in Mechlin (Belgium) and the proceedings were pub-

lished in the following year (1922) in Natuurweten-

schappelijk tijdschrift 4: 142. It had been collected by H.

Vanderyst in September 1910 in the former Belgian

Congo (later Zaire, now the Democratic Republic of the

Congo), at Mpie, Territoire de Mushie (± 3°S, 16°5'E).

Jones ( 1957) studied an isotype specimen held at BR,

adding some details to the original description and illus-

trating it with, inter alia, good drawings of the spores. In

1995 Perold briefly referred to this species (BR speci-

men) and later, (Perold 1996) published two SEM micro-

graphs of the spores.

Until recently, the Vanderyst s.n. specimens (MANCH,

holo.!; BR, iso.!) were the only known examples of this

species. Fortunately, in May 1995 Mrs Colleen Mann-

heimer of the Windhoek Herbarium, collected bryo-

phytes in the Caprivi Strip, Namibia, at a branch of the

Zambezi River near Schuckmannsburg (1724 DB).

Mannheimer 134 proved to be a mixed gathering of R.

FIGURE 3.— SEM micrographs of Riccia lanceolata spores, P. Swartz CH13682 (PRE). A, B, distal face; C, proximal face. A,

x 450; B, x 445; C, x 425.

Bothalia 34,2 (2004)

99

crinita Taylor and, on closer inspection, of R. radicosa.

The collection site was in a floodplain that is seasonally

waterlogged and the substrate was wet, loamy clay,

where grasses and sedges, as well as algae also grew. The

vegetation type at this moist, tropical, northeast corner of

the ‘Strip’ is forest savanna and woodland, the altitude ±

900 m above sea level and the annual rainfall 700-750

mm (Barnard 1998). This locality is quite a considerable

distance from where the type specimen was found,

although this does not rule out its possible occurrence in

the intervening countries of Angola and Zambia, where

bryophytes are undercollected.

DESCRIPTION

Thalli small, annual, scattered, dorsally light grey,

dull, finely reticulate, once or twice to three times sym-

metrically or asymmetrically furcate; main branches up

to 6 mm long, narrow at base, widening to ± 2 mm before

bifurcating, diverging by ± 60°, secondary, and if present,

tertiary segments by ± 40°. Branches (Figure 4A) 4.5

mm long, including terminal segments, 2. 0-2 .5 x ± 1.5

mm, linear-oblong, gradually tapering toward blunt or

wedge-shaped apices when remaining unbranched, oth-

erwise apically widening if bifurcating again; distally

with one or two sharp grooves, the latter soon converg-

ing and proximally single, eventually becoming only

shallowly indented medianly; margins thin, hyaline, scal-

loped; flanks (Figure 4N) covered with overlapping

scales, partly hyaline, partly deep purple-red and gener-

ally shiny; ventral face bearing dense mat of rhizoids;

when dry, thalli dorsally white, occasionally with faint

green blotches; margins also white, incurved, revealing

scales along sides, sometimes their ‘leading’ hyaline

edge forming a white tuft. In cross section toward apex

of single branch (Figure 4G), mid-dorsally narrowly V-

shaped, only 0.39 mm thick from bottom of groove to

ventral face, at slightly sloping sides 0.7 mm thick, width

across 1.2 mm, i.e. 1.7 times wider than thick, ventrally

rounded; in section of bifurcating branch at ± 1 .25 mm

from apices (Figure 4H), deep dorsal groove on either

side separating off rounded, central hump of tissue, 0.43

mm high, laterally wings arched, flanks rising obliquely

to steeply, up to 0.75 mm thick, width across 1 .14 mm,

ventrally rounded; in progressively proximal sections

(Figure 41, J) central hump becoming smaller and even-

tually wedge-shaped, wings not arched, sloping oblique-

ly; in next section (Figure 4K) groove single, dorsally V-

shaped, 0.41 mm thick from bottom of groove to ventral

face, flanks steep, up to 0.9 mm thick, width across ±

1.14 mm, nearly 1.3 times wider than thick; in section

toward base (Figure 4L) medianly only shallowly indent-

ed, wings laterally shortly expanded, flanks obliquely

rising to slightly recurving, ± 0.54 mm thick, width

across 1 .2 mm.

Dorsal epithelium with cells hyaline, in one layer,

rounded above (Figure 4C, E), and often encrusted with

some deposit, 25-A5 x 27 .5 — 45 .0 pm, some collapsing

and sunken in the middle, forming ‘ringed’ cells (Figure

4D), or shallow cups (Figure 4F), air pores between

epithelial cells small, mostly triangular; margins (Figure

4M) acute, hyaline, quite delicate, in cross section only

one cell projecting above epithelium; assimilation tissue

± >/2 thickness of thallus, to slightly more than '/2, up to 10

rectangular cells attached end to end in each row, 45-65

x 37.5-50.0 pm, separated by narrow air canals; storage

tissue forming the remaining thickness of thallus, cells

crowded together, ovoid, ± 52.5 x 65 pm\ outer cells of

flanks and ventral face greenish brown, not clearly thick-

ened; rhizoids arising from ventral epidermal cells and

from lower ‘front’ or ‘leading’ edge of scales, 10.0-22.5

pm wide, mostly smooth, but some tuberculate. Scales

bicoloured (Figure 40, P), quite fragile; except for loose,

‘leading’ edge, the rest closely adherent to thallus flanks

and difficult to detach intact, overlapping and consisting

of the ‘leading’, hyaline part, 450-550 pm high and

400-590 pm wide, at the ‘front’ margin sloping oblique-

ly outward and down, with outer row of cells mostly rec-

tangular, 30.CM-2.5 x 37.5-50.0 pm, adjacent and near-

by cells 4-6-sided, 62.5-100.0 x 25^-0 pm, these hya-

line cells surrounding inner group of ± 35 or more, pur-

ple-red cells of similar shape and size; at rear of coloured

cell group, hyaline cells mostly rectangular, 62.5-87.5 x

37.5- 50.0 pm, and joined end to end in rows; then fused

to flanks of thallus, where cells also joined together at

sides for 12 to 14 rows (up to ± 400 pm wide), some-

times tom assimilation filaments remaining stuck to

inner surface; at the top a row of laterally fused margin-

al cells, short-rectangular, 42.5-57.5 x 30-40 pm,

exceeding thallus margin, sometimes slightly dusted

with fine, powdery deposit.

Monoicous. Antheridia inconspicuous, with short

necks turned white, emerging from slightly sunken, faint,

brownish flecks in a row along middle of branch.

Arche gonia with thin, dark red necks along midline of

same or a different branch, persistent and toward base

projecting from bulging sporangia in close proximity, ±

500 pm wide. Spores (Figure 5A-F) 90-120 pm diam.,

triangular-globular, polar, brown to reddish brown,

ridges more deeply coloured, wing undulating, 7.5-10.0

pm wide, marginal angles not perforated or notched,

margin crenulate; distal face convex, with 7 or 8 angular

or rounded areolae across diameter, 7.5-15.0 x 7.5-15.0

pm, bordered by ridges, usually raised and thickened,

rarely faint, almost always with tubercles at nodes where

joining, up to 5 //m high and stout, occasionally wider

and truncate, then plate-like; proximal face with distinct

triradiate mark, ends of rays extending across wing, each

of 3 facets with 11 or 12 areolae, round or angular,

7.5- 10.0 x 7.5-12.5 pm, ridges quite low, but slightly

raised at nodes, sometimes vestiges of areolar ridges

reaching partly across wing.

DISCUSSION

It appears that Riccia radicosa is a very rare species,

but presumably, it could also be confined to undercol-

lected areas. It can be recognized by its rather small size,

by the deeply grooved apical branches of the thalli, by

the closely adherent, overlapping, hyaline and purple-red

scales that scarcely exceed the scalloped thallus margins,

by the dense mass of ventral rhizoids (for which it had

been aptly named), as well as by the distinct ornamenta-

tion of the winged spores.

The Mannheimer collection of R. radicosa is a small

one, consisting of only about 14 thalli at various stages of

100

Bothalia 34,2 (2004)

FIGURE 4 . — Riccia radicosa , C. Mannheimer 134. A, repeatedly bifurcating thallus branches; B, copy of Pearson’s ‘radiate’ thalli in his plate

XIII, fig. I ; C. intact dorsal epithelial cells of thallus and air pores from above; D, collapsed, 'ringed' dorsal epithelial cells and air pores

from above; E, c/s intact dorsal epithelial cells and upper cells of assimilation tissue (at this magnification, air canals not visible); F, c/s

collapsed dorsal epithelial cells and some assimilation tissue. G-L, c/s thallus branch; G, apical part with deep, narrow, single groove; H-J,

doubly grooved part; K, wider, shallowly grooved part; L, basal part, M, c/s thallus at margin, with overlapping scales on outside (left) and

dorsal cells and cell columns on inside (right); N, flank of thallus with scales seen from side, broken line indicating cut edge; O, P,

bicoloured scales, proximally fused to cells covering flanks. Scale bars: A, 2 mm; B, 10 mm; C-F, 50 /an; G-L, 500 /vm; M, 100 ;mi; N,

250 /an; O, P, 250 /an.

Bothalia 34,2 (2004)

101

FIGURE 5. — SEM micrographs of Riccia radicosa spores, C. Mannheimer 134. A-D, distal face; C, side view; D, part of distal

face with areolae and wing. E, F, proximal face: F, side view. A, x 475; B, x 540; C, x 550; D, x 830; E, x 460; F, x 560.

maturity, some with sporangia containing ripe spores, oth- are said to be 15 mm long. It has been reproduced here in

ers juvenile. There are no rosettes like Pearson’s (1922) Figure 4B, at the same ‘nat.’ (= natural) size. Since the

‘radiate’ drawing in his plate XIII. fig. 1 , where the ‘fronds' rosette measures 15 mm in diameter, the radius is 7.5 mm

FIGURE 6. — SEM micrographs of Riccia radicosa spores, Vanderyst s.n. (BR). A-C, distal face: B, C, side view. D-F, proximal

face: E, F, side view. A, x 535; B, F, x 540; C, x 495; D, x 470; E, x 515.

102

Bothalia 34,2 (2004)

and represents a ‘main' branch. This does not exceed by

much the longest branches in Mannheimer 134, at 6 mm.

The absence of rosettes in the latter specimen could per-

haps be ascribed to a shortened growth period on an unsta-

ble alluvial substrate. In addition, some of the smaller thal-

li were partly overgrown by algae. Which may have had a

detrimental effect. At the time of study, nine years had

elapsed since this specimen was collected. On wetting a

small portion, the thalli did not fully recover their form, the

flanks remaining somewhat inflexed.

Jones (1957) refers to the MANCH specimen (which

he did not see) as the type, since Pearson was resident in

Manchester. Pearson clearly kept the portion of

Vanderyst’s collection, which was sent to him by Mr

Naveau, whom he thanks for the opportunity of examin-

ing the collection (the rest of the specimen is held at BR).

In the holotype specimen, there is very little soil mixed

with the abundant rhizoids underneath the thalli, which

appear to have been pressed, as the flanks have been flat-

tened. The scales are nearly black, with the hyaline ‘lead-

ing’ parts barely visible. In Pearson’s illustration (plate

XIII, fig. 2), the scales, when seen from above, are drawn

as cross-hatched squares separated by white rectangles,

and he describes them as ‘subquadrate, distant, dark pur-

ple'. He also notes that they ‘are not imbricate as in R.

nigrella DC., but are remote’. Jones (1957) on the other

hand, depicts the scales along one of the flanks, as seen

from the side, in his fig. 5b; they are clearly overlapping,

with a black patch at the lower, rear comer, the rest

remaining ± white. He also remarks that the ‘ventral

scales are imbricate or locally distant, deep violet with

broad hyaline margin’. The scales are undoubtedly over-

lapping, but because they are so difficult to detach with-

out tearing, neither of the previous two authors attempt-

ed to illustrate them individually. It would appear that in

the type specimens, the walls of the flanks underneath

the scales are also purple.

Pearson’s ‘dimensions’ for the spores are incorrect,

but this is probably due to a typographical error, as the

decimal point is in the wrong position, i.e. not ‘.9 mm. .1 .

mm. x .85 mm.’, but are meant to be 90-100 x 85 p m,

with ‘6 to 7 areolae across surface’. Jones’s measure-

ments are ‘80-95 //m diam. with 8 or 9 areolae across the

face’, but he concedes that ‘the spores are so opaque that

it is by no means easy to ascertain the exact number’. My

measurements of the BR spores are 90-115 pm and are

illustrated in Figure 6A-F.

The above two species both belong to subgenus

Riccia , section Riccia and to group ‘Squamatae’. How-

ever, R. lanceolata is more robust than R. radicosa, its

scales are also difficult to detach from the flanks, and are

deep violet to mauve, with a hyaline margin, but with

some gradual shading, not clearly delimited to a purple

blotch as in R. radicosa. Its winged spores are variably

incompletely reticulate on both faces, as opposed to

completely reticulate in R. radicosa.

ACKNOWLEDGEMENTS

I sincerely thank the referees for their kind advice,

also the curators of BR, G, Herb. Jones (E) and MANCH

for kindly lending specimens to PRE. A special word of

thanks to Mrs Colleen Mannheimer, curator of WIND,

for collecting bryophytes and presenting them as gifts to

PRE. I also extend my gratitude to Mrs M. Steyn for the

drawings, Ms D. Maree for typing the manuscript, Mrs S.

Turck for the layout of the spore micrograph plates, and

Mrs S.S. Brink for improving the line drawing.

REFERENCES

BARNARD, P. (ed.). 1998. Biological diversity in Namibia: a

country study. Namibian National Biodiversity Task-

force, Windhoek.

FISCHER, E. 1995. The genera Ricciocarpos and Riccia

(Hepaticae, Ricciaceae) in Rwanda. Fragmenta Floris-

tica et Geobotanica 40: 93-1 1 1 .

JONES, E.W. 1957. African Hepatics XIII. The Ricciaceae in

tropical Africa. Transactions of the British Bryological

Society 3: 208-227.

JONES, E.W. & HARRINGTON, A.J. 1983. The hepatics of

Sierra Leone and Ghana. Bulletin of the British Museum

( Natural Histoiy), Botany, ser. 1 1 : 215-289.

PEARSON, W.H. 1922. Notes on a collection of Hepaticae

from Belgian Congo (ex herb. R. Naveau, Antwerp). In

Proceedings of the 20th Vlaamsche Natuur-, Wis- en Ge-

neeskundig Congres, August 1921, Mechlin, Belgium.

Natuurwetenschappelijk tijdschnft 4: 142, 143.

PEROLD, S.M. 1993. Taxonomic results of the BRYOTROP

Expedition to Zaire and Rwanda. 7. Ricciaceae. Tropical

Bryology 8: 55-68.

PEROLD, S.M. 1995. A survey of the Ricciaceae of tropical

Africa. Fragmenta Floristica et Geobotanica 40: 53-91 .

PEROLD, S.M. 1996. Studies in the Ricciaceae of sub-Saharan

Africa: a provisional key to the currently known species.

Bothalia 26: 95-123.

STEPHANI, F. 1888. Hepaticae africanae. d) Aus verschiede-

nen Theilen des westlichen Afrikas. Hedwigia 27: 110,

111.

Bothalia 34,2: 103-113 (2004)

Notes on African plants

VARIOUS AUTHORS

IRIDACEAE

A NEW SPECIES OF THEREIANTHUS (CROCOIDEAE) FROM WESTERN CAPE, SOUTH AFRICA,

NOMENCLATURAL NOTES AND A KEY TO THE GENUS

The genus Thereianthus (Iridaceae: Crocoideae) was

established by Lewis (1941) to accommodate the small-

er, blue or mauve-flowered species included by Baker

(1896) in Watsonia P.Mill. subgenus Beilia Baker. Dif-

ferences between the two genera that were highlighted

included the solitary basal leaf in Thereianthus versus a

basal fan of several leaves in Watsonia and the straight,

rather than curved perianth tube. Lewis (1941) also noted

that the stamens were inserted in the mouth of the tube

instead of some distance below it. This difference is most

evident when species of Thereianthus are compared with

omithophilous species of Watsonia. At the time she com-

mented on the similarity between Thereianthus and the

genus Micranthus, an observation that was later support-

ed by cytological studies, when it was shown that the two

share a unique karyotype with x=10 (all 2n = 20)

(Goldblatt 1971a). Recent molecular studies of chloro-

plast DNA regions have confirmed the sister relationship

between Thereianthus and Micranthus , which together

are sister to the monospecific Cape genus Pillansia

(Reeves et al. 2001). In a later molecular study using

additional chloroplast DNA regions, Thereianthus and

Micranthus are sister to Pillansia plus Watsonia (Gold-

blatt et al., in press).

Thereianthus is endemic to the southwestern Cape,

where it is found mainly on soils derived from sandstone

in montane habitats. It is unusual among Cape Iridaceae

in flowering during the hot summer months, mostly

November to January (Manning et al. 2002). Seven

species of Thereianthus were recognized by Lewis (1941)

and the taxonomy of the genus has remained unchanged

since then, apart from minor nomenclatural matters

(Goldblatt 1971b; 1989). This is an uncommon situation

among the Iridaceae of the Cape Lloral Region, where

new species are still regularly discovered. It was thus not

surprising when an unusual Thereianthus was collected

from high up in the poorly explored Riviersonderend

Mountains of the Caledon District in 1999. Comparison

with the known species of the genus indicated that it rep-

resented an undescribed species, which we included in

Cape plants , an account of the Cape flora, and elsewhere

as Thereianthus sp. 1 (Goldblatt & Manning 2000;

Manning et al. 2002). With additional material now

available, we are able to formally describe the species.

Thereianthus montanus J .C .Manning & Gold-

blatt, sp. nov.

TYPE.— Western Cape, 3419 (Caledon): Riviersonder-

end, Pilaarkop, ridge WNW of peak, (-BB), 31 January

2004, E.G.H. Oliver 12197 (NBG, holo.; MO, iso.).

Plantae 200-350 mm altae, cormo globoso 7-10 mm

diam., tunicis fibrosis, caule erecto eramoso, cataphyllis

papyraceis rubro-brunneis, foliis 3, inferiore basale

laminis linearibus 2-3 nervosis, spica erecta dense 7- ad

10-florum, bracteis breve imbricatis supra siccis brun-

neis externa 6-8 mm longa interna ± 1.5 mm breviore,

floribus violaceis albis notatis, perianthio subactinomor-

pho, tubo 22-27 x 1.2-1 .5 mm cylindrico, tepalis subae-

qualibus, anguste ellipticis ad lanceolatis patentibus

9-15 x 3. 5-5.0 mm, staminibus unilateralibus, filamen-

tis 6-8 mm longis erectis, antheris 4-5 mm longis pur-

pureis, styli ramis furcatis + 2.5 mm longis.

Plants 200-350 mm high. Corm globose, 7-10 mm

diam.; tunics of fine-textured, netted fibres accumulating

with age and forming a neck around base of stem. Stem

erect, flexed outward above sheath of second leaf and

inclined ± 30°, unbranched, 1.0-1. 5 mm diam. below

spike. Cataphylls 2, dry and papery, reddish brown.

Leaves 3, lower one basal, blade reaching or exceeding

spike, drying from tip at flowering, linear, 150-300 x

2. 5-2 .8 mm, thick-textured, without distinct midrib and

with 2 or 3 equally prominent veins, margins not thick-

ened when fresh, upper two leaves cauline, inserted

respectively on lower and upper thirds of stem, the second

sheathing for half to two thirds its length with short blade,

15-25 mm long, upper entirely sheathing or with blade up

to 7 mm long. Spike erect, compact, densely 7-10-flow-

ered; bracts shortly imbricate, green and leathery below,

dry and brown in upper half, outer 6-8 mm long, obtuse to

truncate, inner ± 1.5 mm shorter, notched apically.

Flowers violet; lower three or all tepals each with spear-

shaped, purple median streak near base, throat and lower

part of tube white, unscented; perianth sub-actinomorphic;

tube straight or slightly arching in upper ± 5 mm, 22-27 x

1.2-1. 5 mm, cylindrical and widening slightly in upper ±

5 mm; tepals narrowly elliptical to lanceolate, subequal,

spreading and slightly cupped, 9-15 x 3. 5-5.0 mm.

Stamens unilateral; filaments erect, 6-8 mm long, exsert-

ed 3-4 mm from top of tube; anthers purple, 4—5 mm long;

pollen violet. Ovary ovoid, 2.0-2.5 mm long; style arch-

ing over stamens, dividing between base and middle of

anthers, branches recurved, divided for ± half their length,

±2.5 mm long. Capsules and seeds unknown. Flowering

time: late January to Lebruary. Ligure 1 .

Distribution and biology: known from a single popu-

lation on steep, south-facing slopes of Pilaarkop in the

Riviersonderend Mountains (Ligure 2). Plants of

Thereianthus montanus are scattered in moist, loamy soil

in short, grassy fynbos at an altitude of ± 1 500 m.

Lrequent summer cloud, driven by strong southeasterly

winds, is a feature of many of the high, coastal moun-

104

Bothalia 34,2 (2004)

FIGURE I — Thereianthus montanus. A, whole

plant; B, flower, front view; C, I/s flower; D,

outer (left) and inner (right) bracts. Scale

bar: 10 mm. Artist: John Manning.

FIGURE 2.— Distribution of Thereianthus montanus, •; 71 longicollis,

▲ ; and 7. spicatus, O.

tains in the southwestern Cape, particularly Pilaarkop. At

this altitude the soil around the plants is still moist in

midsummer when they flower. Several other plant

species are endemic to the moist, south-facing slopes of

Pilaarkop and adjacent peaks, including Gladiolus

stokoei and Nivenia dispar (Iridaceae), Lonchostoma

esterhuyseniae (Bruniaceae), and a number of Erica spe-

cies, among them E. alfredii, E. columnaris and E.

orthiocola.

The pollination biology of species of Thereianthus is

poorly known, although the variation in floral morpholo-

gy among the species indicates some diversity in pollina-

tion strategies (Manning etal. 2002). Short-tubed T. race-

mosus (Klatt) GJ. Lewis is believed to be pollinated by

hopliine scarab beetles (Goldblatt et al. 1998), and papil-

ionid butterflies ( Papilio demodocus) have been seen vis-

iting flowers of T. spicatus (L.) GJ. Lewis (original obser-

vation). The long-tubed flowers of T. longicollis (Schltr.)

GJ. Lewis are evidently adapted for pollination by long-

proboscid flies (Goldblatt & Manning 2000) and a simi-

lar inference can be drawn for T. montanus. Its long-

tubed, violet flowers accumulate nectar only in the lower

few millimetres, which is consistent with pollination by

long-proboscid flies with mouthparts 20-25 mm long.

History: a single flowering stem of the species was

first collected by botanists Ted and Inge Oliver in

February 1999. The type material was collected by Dr

Ted Oliver five years later, in January 2004.

Diagnosis and relationships : the relationships of

Thereianthus montanus lie with T. spicatus and T. longi-

collis, with which it shares thick, linear leaves without a

distinct midrib, and flowers with a cylindrical perianth

tube at least as long as the tepals and erect, unilateral sta-

mens. It is distinguished from T. spicatus by its fine-tex-

tured corm tunics, 7-10-flowered spikes, short, obtuse or

truncate bracts, 6-8 mm long, violet flowers with peri-

anth tube 22-27 mm long and short anthers, 4-5 mm

long. The corm tunics in T. spicatus are coarse-textured.

Bothalia 34,2 (2004)

105

the spikes are mostly more than 10-flowered and up to

25-flowered, the bracts are longer, (9-) 12- 18 mm long,

and the pale blue to mauve flowers have a perianth tube

1 2— 1 6(— 20) mm long with anthers 5-6 mm long. T.

longicollis resembles T. montanus in its long perianth

tube, 25^15 mm long but in other respects is similar to T.

spicatus. The three species occupy complimentary

ranges, replacing one another along the coastal moun-

tains of the southwestern Cape, with the two longer-

tubed species at the edges of the range of T. spicatus. T.

longicollis extends from the Grootwinterhoek Mountains

as far south as Tulbagh Waterfall; T. spicatus is distribu-

ted from just southeast of Tulbagh to the Palmietrivier

Mountains above Kleinmond, with a single record from

the western Riviersonderend Mountains above Gena-

dendal; and T. montanus is restricted to the eastern end of

the Riviersonderend Mountains (Figure 2). A fourth

species in this alliance of long-tubed species, T. ixioides

G.J .Lewis, is distinguished by its compact, subcapitate

spike of white flowers with a filiform perianth tube

clasping the style throughout its length and lacking any

nectar, and a short style that divides opposite the base of

the anthers.

Conservation status: known from a single small popu-

lation high on the Riviersonderend Mountains, the spe-

cies is not currently under any threat and must be regard-

ed as Vulnerable (B1 + 2c, D).

Other material examined

WESTERN CAPE.— 3419 (Caledon): Riviersondered, Pilaarkop, ridge

WNW of peak. (-BB). 26 Feb. 1999, E.G.H. & l.M. Oliver 11228 (NBG).

tus L.: 37 (1753). Type: South Africa, Western Cape,

without precise locality, collector unknown (holo.,

LINN!).

Thereianthus spicatus var. linearifolius G.J .Lewis: 40 (1941), syn.

nov. Type: South Africa, Western Cape, mountains near Franschoek,

T.M. Salter 2973 (holo., BOL!).

Collections of T. minutus (Klatt) G.J .Lewis (= T.

lapeyrousioides (Baker) G.J .Lewis) that have accumulat-

ed since the publication of Lewis’s (1941) account of the

genus render the recognition of the variety T. lapey-

rousioides var. elatior G.J. Lewis unnecessary. Var. ela-

tior was recognized for populations of the species from

the mountains around Ceres in the north of the range and

is separated from the typical variety by its taller, more

slender habit and shorter bracts, ± 5 mm vs. 6-8 mm long

in var. lapeyrousioides. Subsequent collections of the

species from Ceres and near Clanwilliam in the north

accord with var. elatior in these features but more exten-

sive collections from Bain’s Kloof in the centre of the

species’ range, comprise a full range of intermediates

between the two varieties. Indeed, collections actually

seen by Lewis herself, such as Loubser 866 (NBG) and

Middlemost 1638 (NBG) from Bain’s Kloof, comprise

plants that exhibit the whole range of variation found in

the species, from slender individuals with leaves 1 .5 mm

wide and bracts 5 mm long to others with leaves 5 mm

wide and bracts 11 mm long. A taxonomic separation

between the northern and southern populations does not

accurately reflect the biological situation as we under-

stand it and var. elatior is reduced to synonymy in T.

minutus.

Reduction of Thereianthus spicatus var. linearifolius and

T. lapeyrousioides var. elatior

Lewis (1941) distinguished two varieties within

Thereianthus spicatus: var. linearifolius GJ.Lewis,

with narrowly linear or subeterete leaves as long as or

longer than the stem, and smaller flowers in which the

perianth tube is as long as or slightly longer than the

tepals; and var. spicatus with linear leaves up to 6 mm

wide and mostly shorter than the stem, and larger flow-

ers with the perianth tube 12 mm long and slightly

shorter than the tepals. There are now numerous collec-

tions available, e.g. Boucher 377 , Hansford 242 (NBG),

in which plants with leaves shorter than the stem have

the perianth tube longer than the tepals, whereas many

plants with leaves longer than the stem have leaves as

wide as in var. spicatus, e.g. De Vos 2307 , Jordaan 1141

(NBG). Moreover, specimens cited by Lewis as var. lin-

earifolius encompass the entire geographic range of the

species. Examination of specimens collected since

Lewis published her account of the genus confirms the

widespread occurrence of narrow-leaved plants but

fails to show any correlation between leaf width and

flower size or tube length. Narrow-leaved plants are

often just younger plants within a population or merely

local forms and do not warrant recognition at any taxo-

nomic rank. Finding no merit in var. linearifolius, we

reduce it to synonymy.

Thereianthus spicatus (L.) GJ.Lewis in Journal

of South African Botany 7: 39 (1941). Gladiolus spica-

The seeds of T. minutus are highly unusual and appear

to be unique in the genus. Seeds of T. bracteolatus and T.

spicatus are similarly angular with the funicle more or

less shortly prolonged and somewhat flap-like, and the

testa lightly rugulate or reticulate with the cell surface

colliculate. They measure 1-2 mm long and are black or

blackish brown (Figure 3 A, B). In sharp contrast, seeds

of T. minutus are fusiform with a thread-like funicle

about as long as the body of the seed and hooked at the

end, and the testa is longitudinally rugose with the cell

surface smooth. The body of the seeds measures 1. 5-2.0

mm long and is pale reddish brown (Figure 3C). Seeds of

the remaining species are not known.

Thereianthus minutus (Klatt) GJ.Lewis in

Journal of South African Botany 7: 43 (1941). Type:

FIGURE 3. -Seeds of The-

reianthus. A. T. brac-

teolatus ( Goldblatt

& Manning 9567,

NBG): B. T. spica-

tus ( SU360 , NBG),

C, T. minutus (Bolus

4010 , NBG). Scale

bar: 1 mm. Artist:

John Manning.

106

Bothalia 34,2 (2004)

South Africa, Western Cape, Tulbagh Waterfall, Ecklon

& Zeyher Irid. 189 (lecto., B!, designated by Goldblatt

1989: 143; isolecto., MO!, S!).

Thereianthus lapeyrousioides var. elatior G.J .Lewis: 38 (1941),

syn. nov. Type: South Africa, Western Cape, Ceres hills, L.Guthrie

2208 (holo., BOL!).

Key to Thereianthus species

la Perianth tube short, 1-2 mm long, less than half as long as tepals and included in bracts; bracts 3-5 mm long; pollen exine reticulate

T. racemosus

lb Perianth tube well developed, at least 5 mm long, as long as or longer than bracts; bracts 3-18 mm long; pollen exine perforate:

2a Leaves sword-shaped to falcate, with definite midrib; flowers reddish purple; seeds fusiform with thread-like funicle T. minutus

2b Leaves linear and subterete, or terete, without definite midrib; flowers white, mauve or bluish purple; seeds (as far as known) angular

without thread-like funicle:

3a Spike lax with bracts short, 3^1 mm long, distant, less than half as long as internodes; leaves terete, ± filiform, less than 1 mm

diam.; perianth tube 12-20 mm long T. juncifolius

3b Spike dense with bracts 6-18 mm long, at least two intemodes long; leaves terete or flattened but not filiform, at least 1 .5 mm diam.;

perianth tube 10-45 mm long:

4a Perianth tube more than twice as long as bracts, 22^15 mm long:

5a Perianth tube 25 — 45 mm long; corm tunics of coarse fibres; anthers ± 6 mm long; bracts 12-18 mm long T. longicollis

5b Perianth tube 22-27 mm long; corm tunics of fine fibres; anthers 4—5 mm long; bracts 6-8 mm long T. montanus

4b Perianth tube slightly longer than, to about twice as long as bracts, 10— 16(— 20) mm long:

6a Perianth tube filiform throughout, not widening above, clasping style; inflorescence compact, subcapitate; flowers white,

rarely pale mauve; style dividing opposite base of anthers T. ixioides

6b Perianth tube wider, cylindrical and dilating above, not clasping style; inflorescence elongate; flowers usually blue to violet or

lilac, rarely whitish; style reaching to at least middle of anthers:

7a Tepals with 3 inconspicuous veins with spreading side branches; leaves with distinct and fairly prominent veins; flowers facing

upwards with tepals spreading more or less horizontally; stamens erect with anthers facing lower tepals; filaments ±

4 mm long T. spicatus

7b Tepals with 3-5 prominent veins without side branches; leaves with veins not or scarcely visible; flowers facing sideways with

tepals spreading almost vertically; stamens held horizontally with anthers facing dorsal tepal; filaments ± 10 mm long . .

T. bracteolatus

ACKNOWLEDGEMENT

We gratefully recognize support for field work from

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

Geographic Society.

REFERENCES

BAKER, J.G. 1896. Irideae. In W.T. Thistelton-Dyer, Flora capensis 6:

7-171. Reeve, Kent.

GOLDBLATT, P. 1971a. Cytological and morphological studies in the

southern African Iridacea e. Journal of South African Botany 37:

317^460.

GOLDBLATT, P. 1971b. A new species of Gladiolus and some nomen-

clature! changes in the Iridaceae. Journal of South African

Botany 37: 229-236.

GOLDBLATT, P. 1989. The genus Watsonia. A systematic monograph.

Annals of Kirstenbosch Botanical Garden 17: 1-148.

GOLDBLATT, P. & MANNING, J.C. 2000. The long-proboscid fly

pollination system in southern Africa. Annals of the Missouri

Botanical Garden 87: 146—170.

GOLDBLATT, P„ BERNHARDT, P. & MANNING, J.C. 1998. Polli-

nation of petaloid geophytes by monkey beetles (Scarabaeidae:

Rutelinae: Hopliini) in southern Africa. Annals of the Missouri

Botanical Garden 85: 215-230.

GOLDBLATT, P„ DAVIES, J„ SAVOLAINEN, V., MANNING, J.C.

& VAN DER BANK, M. In press. Phylogeny of Iridaceae sub-

family Crocoideae based on plastid DNAs. Proceedings of the

Monocots III Conference.

LEWIS, G.J. 1941. Iridaceae. New genera and species and miscella-

neous notes. III. The new genus Thereianthus. Journal of South

African Botany 7: 33 — 43 .

LINNAEUS, C. 1753. Species plantarum , edn 1 . Salvius, Stockholm.

MANNING, J„ GOLDBLATT, P. & SNIJMAN, D. 2002. The color

encyclopedia of Cape bulbs. Timber Press, Portland, Oregon.

REEVES, G., CHASE, M.W., GOLDBLATT, P„ RUDALL, P.J., FAY,

M.F., COX, A.V., LEJEUNE, B. & SOUZA-CHIES, T. 2001.

Molecular systematics of Iridaceae: evidence from four plastid

DNA regions. American Journal of Botany 88: 2074-2087.

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

* 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: 2004-03-08.

ASTERACEAE

DICOMA PICT A , THE CORRECT NAME FOR CYPSELODONTIA ECKLON IAN A

In 1838 De Candolle described the genus Cypse-

lodontia with one species, C. eckloniana DC. (Figure 4)

from the Uitenhage District in Eastern Cape (Figure 5).

Since then, no specimen of this taxon was collected or

identified in any South African herbarium. Several

authors referred to the genus (Harvey 1865; Bentham

1873; Hoffmann 1889; Marloth 1932; Phillips 1951;

Dyer 1975; Herman et al. 2000; Herman 2003) but

Bremer (1994), in his massive work on the genera of the

Asteraceae, did not mention the genus. When a previous

manuscript on the subject came back from referees, it

was pointed out that Cypselodontia had already been

sunk under Dicoma\ It could not be established who was

the first person to place the genus in synonomy, but in an

appendix containing an alphabetical list to genera in The

biology and chemistry of the Compositae, vol. 2, edited

by Hey wood et al. ( 1977), Cypselodontia was placed in

synonomy with Dicoma as Dicoma picta (Thunb.) Druce

(Figure 6). The introductory sentence to this appendix

stated that ‘the genera as recognized by the authors of the

Bothalia 34,2 (2004)

107

FIGURE 4 .—Cypselodontia eckloniana in the Geneve Herbarium (G-

DC), Ecklon 270.

various tribal reviews, as given in this volume,’. A.L.

Cabrera compiled the chapter on the Mutisieae (pp.

1040-1066), but nowhere did he state the synonomy of

Cypselodontia.

FIGURE 6.— Portion of a Dicoma picta specimen in the National

Herbarium, Pretoria (PRE), collected by Erasmus ex Marloth

13720.

In view of the obscure publication of the synonomy

and because it has been overlooked for so many years, it

was decided to publish this short note for general infor-

mation:

Dicoma picta ( Thunb .) Druce in Report of the

Botanical Exchange Club of the British Isles 1916: 619

(1917); Wilson: 385 (1923); N.Netnou & P.PJ .Herman:

210 (2003). Type: Cape, Thunb erg. no further informa-

tion.

Levssera picta Thunb., Prodromus plantarum capensium 2: 160

(1800).

Cypselodontia eckloniana DC.: 286 (1838); Harv.: 123 (1865),

Benth.: 332 (1873), O.Hoffm.: 204 (1889), Marloth: 236 (1932),

E. Phillips: 806, 807 (1951); R.A.Dyer: 690 (1975); P.P.J.Herman et al.:

108

Bothalia 34,2 (2004)

132 (2000); P.P.J .Herman: 209 (2003). Type: Eastern Cape, Uitenhage

District, Ecklon 270 (G-DC, holo.!; PRE, photo!).

Dicoma radiaia Less.: 278 ( 1830). Type: Cape, Mundt & Made, no

further information.

ACKNOWLEDGEMENTS

Mrs S. Brink is thanked for the scanning of the photo-

graphs and herbarium material. Ms H. Steyn is thanked

for the distribution map, and the referees for valuable

comments on the first draft of this manuscript.

REFERENCES

BENTHAM, G. 1873. Compositae. In G. Bentham & J.D. Hooker,

Genera plant arum 2. Reeve, London.

BREMER, K. 1994. Asteraceae. cladistics and classification. Timber

Press, Portland, Oregon.

DE CANDOLLE, A.P. 1838. Mantissa Compositarum. Prodromus 7,1 :

286. Treuttel & Wiirtz, Paris.

DRUCE, G.C. 1917. Nomenclatorial notes: chiefly African and

Australian. Report for 1916. Report of the Botanical Exchange

Club of the British Isles 1916: 601-653.

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

vol. 1: 690. Botanical Research Institute, Pretoria.

HARVEY, W.H. 1865. Compositae. Flora capensis 3: 123, 124.

HERMAN, P.P.J. 2003. Cypselodontia DC. In G. Germishuizen & N.L.

Meyer, Plants of southern Africa: an annotated checklist.

Strelitzia 14: 209. National Botanical Institute, Pretoria.

HERMAN, P.P.J., RETIEF, E„ KOEKEMOER, M. & WELMAN,

W.G. 2000. Asteraceae. In O.A. Leistner, Seed plants of south-

ern Africa: families and genera. Strelitzia 10: 132. National

Botanical Institute, Pretoria.

HEYWOOD, V.H., HARBORNE, J.B. & TURNER, B.L. (eds). 1977.

The biology and chemistry of the Compositae, vol. 2. Academic

Press, London.

HOFFMANN, K.A.O. 1889. Compositae. Die natiirlichen Pflanzen-

familien 4,5: 87-387.

LESSING, C.F. 1830. De synanthereis herbarii regii berolinensis.

Linnaea 5: 278.

MARLOTH, H.W.R. 1932. The flora of South Africa, vol. 3. Weldon &

Wesley, Cape Town.

NETNOU, N. & HERMAN, P.P.J. 2003. Dicoma Cass. In G. Ger-

mishuizen & N.L. Meyer, Plants of southern Africa: an anno-

tated checklist. Strelitzia 14: 209, 210. National Botanical

Institute, Pretoria.

PHILLIPS. E.P. 1951. The genera of South African flowering plants,

edn 2. Memoirs of the Botanical Survey of South Africa No. 25:

806,807.

THUNBERG, C.P. 1800. Prodromus plantarum capensium Pars 2.

Edman, Uppsala.

WILSON, F.C. 1923. Revision of the genus Dicoma. Kew Bulletin

1923: 377-388.

P.P.J. HERMAN* and M. KOEKEMOER*

* National Botanical Institute, Private Bag X10L0001 Pretoria.

MS. received: 2004-02-20.

PTERIDOPHYTA

A NEW COMBINATION IN LASTREOPSIS (TECTARIACEAE) FROM MADAGASCAR

Dryopteris perrieriana C.Chr., a species restricted to

Madagascar, was first described and published by

Christensen in 1925. Christensen placed the species in

subgenus Eudryopteris , but noted that it differs from all

other species in this subgenus in general habit, and in the

pubescence of the rachis.

In spite of these remarks by Christensen (1925), the

placing of the species was never challenged, but accept-

ed by Tardieu-Blot (1958) and Fraser-Jenkins (1986).

Fraser-Jenkins (1986) included D. perrieriana in a small

group of species he could not place in his classification.

In 1981, however, Fraser-Jenkins made a note on the

holotype suggesting it belongs to Ctenitis C.Chr.

Holttum (1983) reviewed the Mascarene Island species

of Ctenitis, but this species was not included. A combi-

nation in this genus appears never to have been made.

Whilst working on a taxonomic review of African and

Madagascan Dryopteris , I had the opportunity to study

the holotype of D. perrieriana housed in the British

Museum Herbarium (BM), and an isotype in the Paris

herbarium (P). The holotype consists of what appears to

be a basal pinna fragment only, whereas the isotype con-

tains more complete material. These collections exhibit

sufficient features for the species not to be included in

Dryopteris as currently defined. The ctenitoid hairs

occurring along the lamina axes and veins, the continu-

ous lamina axes sulci which are densely set with short

ctenitoid hairs, the prominent ridges of the axes sulci

which are continuous with the thickened lamina margin,

and the centrally ridged sulci of at least the main lamina

axes, place D. perrieriana in the genus Lastreopsis Ching,

as defined by Ching (1938) and Tindale (1965).

Tindale (1965), in her monograph of Lastreopsis , makes

no mention of this species. Based on the evidence provid-

ed above and the fact that no name exists for the species in

Lastreopsis , I propose the following combination;

Lastreopsis perrieriana (C.Chr.) J.P.Roux, comb.

nov.

Dryopteris perrieriana C.Chr. in Notes pterido-

logiques 16: 176, t. VIII (1925). Type: Madagascar:

Foret d’Analamazoatra 800 m, Dec., M.H. Perrier de la

Bdthie 6093 (BM!, holo., P000349499L iso.).

ACKNOWLEDGEMENTS

My appreciation is extended to the Keeper of the

British Museum Herbarium, and the Curator of the Her-

bier National de Paris, for making their Dryopteris mate-

rial available to me for study.

REFERENCES

CHING, R.C. 1938. A revision of the Chinese and Sikkim-Himalayan

Dryopteris with reference to some species from neighbouring

regions. Bulletin of the Fan Memorial Institute of Biology 8:

157-268.

CHRISTENSEN, C. 1925. Fougeres de Madagascar, recoltes de M.H.

Perrier de la Bathie. Supplement. Notes pteridologiques 16:

157-198, t. VIII.

Bothalia 34,2 (2004)

109

FRASER- JENKINS, C.R. 1986. A classification of the genus Dry-

opteris (Pteridophyta: Dryopteridaceae). Bulletin of the British

Museum (Natural History). Botany 14: 183-218.

HOLTTUM, R.E. 1983. The fern-genera Tectaria. Heterogonium and

Ctenitis in the Mascarene Islands. Kew Bulletin 38: 107-130.

TARDIEU-BLOT, M.-L. 1958. Polypodiacees ( senso lato). Dennstaed-

tiacees-Aspidiacees. In H. Humbert, Flore de Madagascar et

des Comores (Plantes Vasculares) Famille 5,1: 1-391, figs 1-51.

Paris.

TINDALE, M.D. 1965. A monograph of the genus Lastreopsis Ching.

Contributions from the New South Wales National Herbarium

3: 249-339.

J.P. ROUX*

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

7735 Claremont, Cape Town.

MS. received: 2004-03-23.

FABACEAE

A NEW SPECIES OF ACACIA (MIMOSOIDEAE) FROM THE PROVINCE OF LIMPOPO, SOUTH AFRICA

INTRODUCTION

In March 1972, W.F. Stuurman collected material of a

peculiar, large-leaflet Acacia Mill, near Burgersfort, in

the province of Limpopo (then eastern Transvaal), South

Africa. Burgersfort and its surrounds form part of the

Sekhukhuneland Centre of Endemism (Van Wyk &

Smith 2001), a region traditionally inhabited mainly by

people of the Pedi cultural group. No other specimens

matching the nine sheets of Stuurman WS4 were found in

subsequent years (Ross 1975, 1978, 1979). The identity

of this Acacia remained uncertain and the collection in

some respects vanished into obscurity. In the course of

taxonomic work on the genus Acacia in southern Africa,

the first author embarked on a dedicated search for plants

matching the Stuurman collection. An area within a 10

km arc north of Burgersfort was targeted, especially an

isolated quartzite mountain surrounded by rock strata of

the prevailing Bushveld (Igneous) Complex. In March

2003, plants matching the original collection were dis-

covered in the designated area. Field observations and a

comparative morphological study confirmed that the

plants represent a distinct new species of Acacia sub-

genus Acacia that is here formally described. Ross

(1975, 1978, 1979) provides comments on and a provi-

sional description of Stuurman W34 as an ‘insufficiently

known species’. The present contribution has benefited

greatly from Ross’s pioneering work.

Acacia sekhukhuniensis PJ.H.Hurter, sp. nov.,

ab A. robusta floribus luteis (non cremeis), anthesi medio

aestate (non primo vemifera), leguminibus chartaceis

(non lignosis) semina pauciora minora continentibus, et

foliolis maioribus cum nervis prominentibus in superfi-

cie abaxiali differt. Etiam ab A. karroo foliis glandu-

laribus, foliolis valde maioribus, pedunculo longiori,

corolla breviori et leguminibus inter seminibus non con-

strictis differt.

TYPE.— Limpopo, 2430 (Pilgrim’s Rest): Sekhukhune-

land, Farm Schlickmannskloof 258KT, 1 324 m, (-CB),

26-11-2003, T. Mukoma & P.J.H. Pturter 17 (PRE, holo.;

NBG, PRU, iso.).

Illustration: Ross: 145, fig. 130 (1979).

Small tree up to 3 .5 m tall, trunk slender. Branches fasti-

giate, ascending, striate, blackish brown; new growth con-

spicuously striate, ferruginous. Stipules in pairs, spines-

cent, 30-50 mm long, white, glabrous, attenuate, pungent.

antrorse, basally thickened. Bark coarsely flaking or split-

ting to reveal a reddish brown inner layer on young shoots.

Leaves dull green, glabrous, bipinnately compound, pulvi-

nus yellow at maturity; petiole sulcate, 0.5-20.0 mm long;

primary leaves with large, raised, crateriform, petiolar nec-

tary gland, often absent in secondary leaves; rachis sulcate,

0-20 mm long, terminating in a short, rigid, persistent,

deflexed hook or claw, at least one nectary gland at junc-

tion of distal pinna pair; rachillae 1 pair on secondary

leaves, 2 to 3 pairs on primary leaves, distichous, 30-65

mm long, midrib raised, dotted with small raised glandular

structures between each leaflet pair, terminating in a short,

rigid, persistent, deflexed hook or claw; leaflets' distichous,

5-9 pairs per rachillae, oblong to obovate-oblong, 8-18 x

3-9 mm, entire, eglandular, apex mucronulate, terminal

pairs largest, asymmetrical, base oblique, midrib and usu-

ally several other basal and lateral veins + raised and con-

spicuous on abaxial surface only. Inflorescences capitate,

globose, bright yellow, 6-16 mm diam., borne on new

growth, fascicled on axillary peduncles; peduncle 24-45

mm long, glabrous below involucel, or with few random,

small yellow glands above involucel, glutinous when

young, often with shards of villose hairs; involucel

bracteate, 1.0-2 .2 mm long, halfway to two thirds up the

peduncle, dotted with small red glands, androgenous flo-

rets always present between bracts. Flowers bright yellow,

dichlamydeous. Bracteole spatulate, glandular, apices with

sparse villose hairs. Calyx campanulate, pentamerous,

glabrous, 2. 0-2 .5 mm long, ascending, apices weakly

pubescent, clasping corolla. Corolla campanulate, pen-

tamerous, membranaceous; lobes ± united, free above

calyx, ligulate, 2.5-3 .2 mm long, ascending, edges and

apex papillate. Stamens numerous; filaments 6.0-7 .5 mm

long. Ovary slightly stipitate, ventricose, septate, 2-3 mm

long, surface with a few pusticulate waxy globules; style

2-3 mm long. Pods dehiscent, falcate, complanate, valves

venous, 40-115 x 6-11 mm, 4— 10-seeded, sparsely cov-

ered with a few non-pustular glands, margins not constrict-

ed between seeds at maturity. Seeds elliptic, 6-11 x 4—6

mm, areole elliptic, 4—6 x 2-4 mm. Figure 7.

Diagnostic features and affinities: Acacia sekhukhu-

niensis is unlikely to be confused with any other species

of Acacia in Africa; in the field it superficially resembles

Albizia anthelmintica on account of its exceptionally

large leaflets. Ross (1975, 1979) provisionally compared

A. sekhukhuniensis with both A. karroo Hayne and the

sympatric A. robusta Burch, subsp. robusta: he also allud-

ed to the possibility that it might be of hybrid origin.

However, A. sekhukhuniensis differs markedly from both

these taxa. The inflorescences and flowers of A. robusta

110

Bothalia 34,2 (2004)

FIGURE 1.— Acacia sekhukhuniensis P.J.H. Hurter. A, primary leaf, x 0,9; B, secondary leaf, x 0.9; C, rachillae, x 4.4; D, c/s rachillae, x 17.5;

E, leaflet, x 2.6; F, petiole, x 4.4; G,, G2, stipules, x 0.9; H, inflorescence, x 1.7; I, bracteole, x 8.7; Ji J2 floret, x 8.7; K, calyx, x 8.7; L,

corolla, x 8.7; M, gynoecium, x 17.5; N, pod, x 0.9; O, seed, x 1 .7. Drawn by S. Burrows.

Bothalia 34,2 (2004)

subsp. robust a are white or pale cream-coloured, com-

pared with yellow in A. sekhukhuniensis . Moreover, flower-

ing in A. robusta subsp. robust a takes place in early

spring (September-October), whereas A. sekhukhuniensis

flowers in midsummer (December-January). The papery

pods of A. sekhukhuniensis are quite different from the

much larger and robust woody pods of A. robusta subsp.

robusta and contain much smaller and less robust seeds.

Leaflets of A. sekhukhuniensis are at least twice the size

of the largest ones recorded for A. robusta subsp. robusta

and in addition have somewhat raised veins on the abax-

ial surface, a feature not found in the latter.

Acacia sekhukhuniensis also differs conspicuously

from A. karroo in the morphology of the leaves, inflores-

cences, flowers and pods. Leaflets of A. sekhukhuniensis

are up to ten times larger than those of A. karroo and in

addition have somewhat raised veins on the abaxial sur-

face, a feature not found in the latter. Secondary leaves

of A. sekhukhuniensis always consist of only two glan-

dular rachillae, whereas those of A. karroo are eglandu-

lar and always consist of three or more pairs. In A. kar-

roo the peduncle is conspicuously shorter and less robust

than that of A. sekhukhuniensis . Flowers of A. karroo are

very different from those of A. sekhukhuniensis in that

the corolla lobes are fused above the calyx to form a

short tube with lobes reflexed, whereas in A. sekhukhu-

niensis the corolla lobes are much shorter, ascending and

not fused into a short tube above calyx. Pods of A.

sekhukhuniensis are much wider than those of A. karroo

and unlike the latter, are not constricted between the

seeds.

In the field. Acacia sekhukhuniensis could potentially

also be confused with the sympatric A. exuvialis , mem-

ber of a distinct group of southern African acacias with

conspicuous glandular-glutinous pods, the glands being

sessile, pustular, and usually dark-coloured (Ross 1971).

Also in this group are A. borleae , A. nebrownii , A. per-

mixta, A. swazica, A. tenuispina, and A. torrei. Acacia

sekhukhuniensis , however, differs markedly from all

these taxa in having much larger leaflets — larger than in

any other known African member of Acacia subgenus

Acacia — rachillae with a raised midrib, prominent

glands between each leaflet pair and pods that are much

more robust and neither covered by sessile, pustular

glands, nor glutinous.

Ross’s (1975, 1979) allusion to the possibility that A.

sekhukhuniensis might be of hybrid origin seems

unfounded as no likely putative parents are to be found

in the area and the population does not exhibit Mendelian

patterns of variation expected in a hybrid population.

Rather, A. sekhukhuniensis appears to be a palaeoendemic

species of a relictual nature.

Distribution and habitat, at present, A. sekhukhunien-

sis is known only from the type locality (Figure 8), an iso-

lated, flat-topped quartzite mountain near the northeast-

ern boundary of Sekhukhuneland. Biogeographically this

locality falls within the Sekhukhuneland Centre, a region

rich in endemic plants (Van Wyk & Smith 2001). The

recently described A. robbertsei P.P.Swartz (Coates

Palgrave 2002) as well as several other undescribed

species of Acacia (unpublished data) are known to be

endemic to this local centre of endemism. Plants of the

new species occur gregariously in open woodland and

wooded grassland on quartzite ridges locally overlain by

deep deposits of what appears to be a relictual Kalahari-

type sand. Much of the vegetation in the area has been

impacted on negatively by human activities; an estimated

60% has been transformed by urban sprawl and subsis-

tence farming lots, with overgrazing by domestic live-

stock prevalent throughout. A. sekhukhuniensis is consid-

ered to be threatened and could in future be assessed as

Critically Endangered [World Conservation Union

(IUCN) 2001), mainly due to its extremely small extent

of occurrence and the fact that much of Sekhukhuneland

is under considerable threat from urban sprawl, heavy

grazing pressure and extensive, destructive mining activ-

ities, mainly for various heavy metals and dimension

stone.

Etymology, the specific epithet refers to the geo-

graphical area where this species occurs, namely

Sekhukhuneland, a region named for King Sekhukhune I

(1814-1882) of the Bapedi tribe.

Other specimen examined

LIMPOPO.— 2430 (Pilgrim’s Rest): Sekhukhuneland, 10 km N of

Burgersfort, (-CB), W.F. Stuurman W34 (PRE, PRU).

ACKNOWLEDGEMENTS

We are indebted to the curators of PRE and PRU for

access to their herbarium collections. Miss R. Luyt, Mr

W.C. Froneman and Mr T. Mukoma for technical assis-

tance, Dr H.F. Glen for the Latin diagnosis, Sandie

Burrows for the line drawing, SABONET, the National

Botanical Institute and University of Pretoria for finan-

cial support.

REFERENCES

COATES PALGAVE, M. 2002. Keith Coates Palgrave Trees of south-

ern Africa, edn 3. Struik, Cape Town.

ROSS, J.H. 1971. The Acacia species with glandular glutinous pods in

southern Africa. Bothalia 10: 351-354.

ROSS. J.H. 1975. Fabaceae, subfamily 1: Mimosoideae. Flora of south-

ern Africa 16,1. Department of Agricultural Technical Services,

Pretoria.

112

Bothalia 34,2 (2004)

ROSS, J.H. 1978. A curious Acacia from the Eastern Transvaal. Trees

in South Africa 100: 30-32.

ROSS, J.H. 1979. A conspectus of the African Acacia species. Memoirs

of the Botanical Survey of South Africa No. 44. Botanical

Research Institute, Pretoria.

VAN WYK, A.E. & SMITH, G.F. 2001 . Regions offloristic endemism

in southern Africa: a review with emphasis on succulents.

Umdaus Press, Pretoria.

WORLD CONSERVATION UNION (IUCN). 2001. IUCN Red List

categories and criteria: version 3.1. IUCN Species Survival

Commission. IUCN, Gland, Switzerland and Cambridge, UK.

P.J.H. HURTER*f and A.E. VAN WYK**

*Lowveld Garden, National Botanical Institute, P.O.Box 1024, 1200Nelspruit.

t 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: 2004-03-24.

HYACINTHACEAE

ORN1THOGALUM LA1KIP1ENSE. A SYNONYM OF DRIMIA MACROCARPA

The species Oniithogalum laikipiense L.E. Newton

was recently described from Kenya (Newton 2003).

Examination of the protologue and the ample illustra-

tions that accompany it leave no doubt that the species

has been incorrectly assigned to the genus Oniithogalum

L. and is in fact a species of Drimia Jacq. In its broad

sense, Drimia is distinguished from all other genera of

Hyacinthaceae by its spurred, often deciduous bracts and

short-lived flowers with caducous perianth, each lasting

less than a day (Manning et al. 2004). All of these criti-

cal features are evident in Oniithogalum laikipiense ,

along with the loose, scale-like bulb tunics and hysteran-

thus flowering that is characteristic of many species of

Drimia. Within the genus Drimia, O. laikipiense is allied

to the small group of species previously segregated in the

genus Thuranthos C.H.Wright, defined by the nodding

flowers borne on elongate pedicels, reflexed tepals, and

distinctive stamens in which the lower part of the fila-

ments converge over the ovary to form a cage-like struc-

ture. The dark maculae on the leaf sheaths of O. laikipi-

ense are also highly characteristic, and have otherwise

been recorded in Drimia only among this group of

species. Two species in this group are known from

Kenya, D. indica (Roxb.) Jessop and D. macrocarpa

Stedje, separated essentially on the basis of size. D.

macrocarpa is a more robust plant (up to 0.9 m tall vs 0.5

m), with larger flowers (tepals 15-24 mm long vs 6-12

mm) and capsules (20-24 mm long vs 8-18 mm), borne

on longer pedicels (30-42 mm long vs 12-30 mm)

(Stedje 1987). Comparison with the dimensions given

for O. laikipiense leaves no doubt that this species is

conspecific with D. macrocarpa and it is accordingly

reduced to synonomy. The status of D. macrocarpa in

relation to the southern African D. angustifolia Baker is

uncertain and the two may prove to be conspecific on

further study.

Drimia macrocarpa Stedje in Nordic Journal of

Botany 7: 664 (1987). Type: Tanzania, Mpanda Dist.,

Uruwira, Richards & Arasululu 26126 (holo., K).

Oniithogalum laikipiense L.E. Newton: 18 (2003), syn. nov. Type:

Kenya, Laikipia Plateau, Roberts sub Newton 5567 (holo., K; iso., EA).

REFERENCES

MANNING, J„ GOLDBLATT, P. & FAY, M. 2004. A revised generic

synopsis of Hyacinthaceae in sub-Saharan Africa, based on

molecular evidence, including new combinations and the new

tribe Pseudoprospereae. Edinburgh Journal of Botany 60:

533-568.

NEWTON, L.E. 2003. A new Ornithogalum from Kenya. Plantsman,

n.s., 2,1: 18-20.

STEDJE, B. 1987. A revision of the genus Drimia (Hyacinthaceae) in

East Africa. Nordic Journal of Botany 7: 655-666.

J.C. MANNING*

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

7735 Claremont, Cape Town.

MS. received: 2004-06-15.

ZAMIACEAE

TYPIFICATION OF ENCEPHALARTOS

In 1926 Pilger lectotypified Encephalartos with E.

cajfer (‘Typische Art: E. cajfer (Thunb.) Lehm.’). This

was accepted in the paper edition of Farr, Leussink &

Stafleu (1979) as well as its updated electronic version

(2002). The syntype material of E. cajfer is extant as

two sheets of leaves and parts of male and female cone

material, in the Thunberg Herbarium in UPS (Dyer

1966: 33).

In 1992 Stevenson again lectotypified Encephalartos,

but with E. friderici-guilielmi, claiming that Pilger’s lec-

totypification was invalid because it was published

‘before and without a type concept’.

I dispute Stevenson’s interpretation, and submit that

Pilger’s lectotypification is perfectly in order and should

be upheld.

Stevenson’s ( 1992) claim that no type concept existed

in 1926, is unfounded. At the very first International

Botanical Congress (De Candolle 1867) it was defined in

article 54 of the Laws of Botanical Nomenclature: ‘If a

genus contains a section or some other division which,

judging by its name or by its species, is the type or ori-

gin of the group, the name is reserved for that part of it’.

It would appear that it was not generally applied in so

many words, but it would certainly seem that numerous

Bothalia 34,2 (2004)

113

practising taxonomists accepted the principle. Thus at a

meeting of the American Association for the Advance-

ment of Science held in Rochester, New York, in 1892, a

set of nomenclatural rules which included application of

the type method, was presented and discussed. These

were further refined at a meeting at Philadelphia in 1904

(Anon. 1904). It appears that the failure of the Inter-

national Botanical Congresses in Vienna (1905), Brus-

sels (1910), London (1924), and Ithaca (1926) to adopt

the type concept stemmed not so much from aversion to

the method as from anti-American sentiment. The type

method was widely applied, but its application was not com-

pulsory according to the International Rules of Botanical

Nomenclature. Between 1919 and 1926 Sprague in the UK

and Hitchcock in the USA, published a series of articles

on typification (Hitchcock 1919, 1925, 1926, 1929;

Sprague 1920, 1921, 1923, 1924, 1926), and this led to

the acceptance of the type principle at the International

Botanical Congress in Cambridge in 1930; and its inclu-

sion in the third edition of the International Rules of

Botanical Nomenclature (Briquet & Rendle 1935). The

history of the adoption of the type method into the Inter-

national Code of Botanical Nomenclature is admirably

related by Lawrence (1951).

Furthermore, the type concept is retroactive, and is

today applied to species described at the very starting

point of botanical nomenclature.

Its mention in the Laws accepted at the very first

International Botanical Congress in 1867 (De Candolle

1867) proves that European taxonomists were familiar

with the type method, and at the time of Pilger’s publi-

cation, the publications cited above prove that taxono-

mists were beginning to accept the concept of type-based

taxa. Nevertheless Pilger’s acceptance or otherwise of

the type concept is irrelevant, as the current edition of the

International Code of Botanical Nomenclature (Greuter

et al. 2000) clearly states (Art. 7.11) 'designation of a

type is achieved ... if the type is definitely accepted as

such by the typifying author [and] if the type element is

clearly indicated by direct citation including the term

‘Type’ ... or an equivalent’. Pilger’s citation of a 'typis-

che Art’ can only be read as complying with these re-

quirements. Art. 9.17 then states 'The author who first

designates a lectotype or neotype must be followed', and

no limiting starting date is stipulated. Stevenson’s argu-

ment about the lack of a type concept at the time is irrel-

evant in terms of the Code. Pilger’s action is in line with

the requirements of the Code, and must be followed. The

only permissible reason for rejecting Pilger’s action

would be if it were in ‘serious conflict’ with the proto-

logue, which Stevenson hasn’t proved.

ACKNOWLEDGEMENTS

Drs R.K. Brummitt of the Royal Botanic Gardens, Kew;

H.F. Glen of the KwaZulu-Natal Herbarium, Durban; and

W. Greuter of the Botanischer Garten und Botanisches

Museum Berlin-Dahlem, are thanked for their opinions.

REFERENCES

ANON. 1904. Code of botanical nomenclature. Bulletin of the Torrey

Botanical Club 31: 249-290.

BRIQUET, J. & RENDLE, A.B. (eds). 1935. International Rules of

Botanical Nomenclature, edn 3. Jena.

DE CANDOLLE, A. 1867. Lois de la nomenclature botanique. Geneve.

English translation: Weddell, H.A. 1868. Laws of botanical

nomenclature , together with an historical introduction and a

commentary. London.

DYER, R.A. 1966. Cycadaceae, Zamiaceae. In L.E. Codd. B. de Winter

& H.B. Rycroft, Flora of southern Africa 1: 3-34 [33]. Botani-

cal Research Institute, Pretoria.

FARR, E.R., LEUSSINK, J.A. & STAFLEU, F.A. 1979. Index nominum

genericorum (plantarum) 1: 611 .Regnitm vegetabile 100. Bohn,

Scheltema, & Holkema, Utrecht.

FARR, E.R., LEUSSINK, J.A. & STAFLEU, F.A. 2002. Index

nominum genericorum (plantarum ).

httm://www.ravenel .si .edu/botany/ing

GREUTER, W„ MCNEIL, J„ BARRIE, F.R., BURDET, H.M., DE-

MOULIN, V., FILGUEIRAS, T.S., NICOLSON, D.H.. SILVA.

P.C., SKOG, J.E., TREHANE, P.. TURLAND, N.J. &

HAWKSWORTH, D.L. 2000. International Code of Botanical

Nomenclature (Saint Louis Code). Regnum vegetabile 138.

Koeltz, Konigstein.

HITCHCOCK, A.S. 1919. Type-basis code of botanical nomenclature.

Science 49: 333-336.

HITCHCOCK, A.S. 1925. Methods of descriptive botany. New York.

HITCHCOCK, A.S. 1926. A basis for agreement on nomenclature at

the Ithaca Congress. American Journal of Botany 13: 291-300.

HITCHCOCK, A.S. 1929. The relation of nomenclature to taxonomy.

Proceedings of the International Congress of Plant Science 2:

1434-1439.

LAWRENCE, G.H.M. 1951. Taxonomy of vascular plants . Macmillan,

New York.

PILGER, R . 1 926. Cycadaceae. In A. Engler, Die natiirlichen Pflanzen-

familien edn 2, 13: 44—82 [79]. Engelmann, Leipzig.

SPRAGUE. T.A. 1920. Plant nomenclature: some suggestions. Journal

of Botany (London) 59: 153-160.

SPRAGUE, T.A. 1921 . The nomenclature of plant families. Journal of

Botany (London) 60: 69-73.

SPRAGUE, T.A. 1923. Suggestions for a World-Code of plant nomen-

clature. Science 57: 207.

SPRAGUE, T.A. 1924. Proposed changes in the International Rules.

Journal of Botany (London) 62: 196-198.

SPRAGUE. T.A. 1926. Standard-species. Kew Bulletin 1926: 96-100.

STEVENSON, D.W. 1992. A formal classification of the extant cycads.

Brittonia 44: 220-223.

P. VORSTER*

* Department of Botany and Zoology, University of Stellenbosch,

Private Bag XI , 7602 Matieland, Stellenbosch, South Africa.

MS. received: 2004-05-25.

.

Bothalia 34,2: 115-126(2004)

Extrafloral nectaries in Combretaceae: morphology, anatomy and

taxonomic significance

P.M. TILNEY* and A.E. VAN WYK**

Keywords: anatomy, Combretaceae, extrafloral nectary, Laguncularia C.F.Gaertn., Lumnitzera Willd., Meiostemon Exell & Stace, morphology,

Pteleopsis Engl., Quisqualis L., Terminalia L.

ABSTRACT

Extrafloral nectaries (EFNs) in members of the Combretaceae are nectaries not involved with pollination and occurring

on vegetative structures; they are believed to attract ants to protect plants against herbivory by other insects. In the

Combretaceae EFNs are reported in species of Terminalia L. and Pteleopsis Engl., putative EFNs in Meiostemon Exell &

Stace and Quisqualis L., and an absence of EFNs in Combretum Loefl. and Lumnitzera Willd. EFNs in the family are gen-

erally spherical in shape and may be raised, level with the surface or somewhat concave. They are similar in the Terminalia

and Pteleopsis species where they display varying degrees of internal zonation and are composed of small cells; those

species observed in the field were all found to have functional EFNs. In Meiostemon tetrandrum (Exell) Exell & Stace,

Quisqualis indica L., Q. littorea (Engl.) Exell and Q. parviflora Gerrard ex Sond., apparent EFNs lack internal zonation and

are composed of enlarged cells; confirmation is required as to whether these are functional. The formation of EFNs appears

to be highly flexible. They are usually essentially associated with new growth but their occurrence is sporadic and they do

not appear on every leaf or every branch of a plant. The distribution of EFNs on leaves, when present, is of taxonomic sig-

nificance to separate species of Pteleopsis and Terminalia ; otherwise the presence or absence and distribution of EFNs are

too variable and sporadic in occurrence to be of taxonomic significance at the species level. Indiscriminate use of the terms

gland and domatium instead of EFN, and possible confusion with damage caused by other organisms, has probably con-

tributed to many of these structures not previously being recorded as EFNs. Floral and extrafloral nectar samples of T

phanerophlebia Engl. & Diels differed in sugar composition.

INTRODUCTION

The Combretaceae is a pantropical family of 20 genera

and 500 species (Mabberley 2000). The largest genera are

Combretum Loefl. (250 tropical species) and Terminalia

L. (150 tropical species).

The great structural diversity of EFNs and their occur-

rence in various plant taxa have been documented in sev-

eral reviews (e.g. Delpino 1886-1889; Zimmermann

1932; Schnell etal. 1963; Bentley 1977; Elias 1983; Fahn

1988). They have only been reported in two families of

the Myrtales, viz. the Melastomataceae where they

appear to be poorly known and the Combretaceae (Elias

1983). There are relatively few references to EFNs in the

Combretaceae and, as far as we have ascertained, there

are no reports on the efficacy of EFNs in the family in

attracting ants to protect the plants against herbivory. In

an extensive study of extrafloral glands in several groups

of tropical plants, Schnell et al. (1963) make no mention

of the family. The Combretaceae is one of 55 families list-

ed by Metcalfe & Chalk (1979) as having EFNs.

Only a limited number of anatomical studies of EFNs

have been carried out on members of the Combretaceae.

These have been largely confined to various species of

Terminalia and to Laguncularia racemosa (L.) C.F.Gaertn.

(Von Hohnel 1882; Zimmermann 1932; Biebl & Kinzel

1965; Belin-Depoux 1978, 1989, 1993; Ramakrishna &

Rajashekara 1981 ). Mention has been made of the possible

* Department of Botany, Rand Afrikaans University, P.O. Box 524,

2006 Auckland Park, Johannesburg.

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

University of Pretoria, 0002 Pretoria.

MS. received: 2003-09-23.

occurrence of EFNs in species of Meiostemon Exell &

Stace, Pteleopsis Engl, and Quisqualis L. (Tilney 2002).

Foliar nectaries were suspected by Elias (1983) to

occur in many additional species of Combretaceae which

our observations in the field and on herbarium material

have confirmed. Furthermore, it seems that the terms

EFNs (or simply glands) and domatia (small structures

including pits, hair tufts, etc. associated with veins, often

found on leaves of woody dicotyledons and known to har-

bour mites) have sometimes been used indiscriminately

in the family, especially in the early literature, and that

possibly some confusion also exists with galls or other

markings caused by animal damage. For these reasons,

and to investigate their structure more fully, a morpho-

logical and anatomical study of mainly EFNs, but includ-

ing some domatia and galls, was undertaken. The purpose

of this paper is therefore to document the occurrence of

EFNs in the Combretaceae, to give an account of their

morphology and anatomy, and to report on the taxonom-

ic significance of the distribution of EFNs on leaves. In

addition, a comparison was made between the sugar com-

position of secretions of floral and extrafloral nectaries of

a single species to ascertain whether or not the same type

of insect is likely to visit both kinds of nectaries.

MATERIALS AND METHODS

All specimens of every species of the southern

African genera of Combretaceae as well as all those of

more northerly species available at PRE and PRU were

studied for the presence and distribution of EFNs,

including approximately 130 Combretum spp. and

Lumnitzera . All the southern African species with EFNs

were used for the anatomical study as well as selected

116

Bothalia 34,2 (2004)

TABLE 1 .—Specimens used in the anatomical study of Combretaceae extrafloral nectaries, domatia and galls, with an indication of EFN mor-

phology and predominant shape of the cells of the secretory tissue

a-elongated, anticlinally elongated; p-elongated. periclinally elongated; seer., secretory; CP, ML Coates Palgrave.

examples of other species. Single specimens of Termi-

nalia catappa L. and T. mollis Lawson with domatia and

galls respectively were also included for comparison.

The sources of this material and authorities for species

names are given in Table 1 . Fresh material was preserved

in FAA; dried material was first rehydrated in distilled

water and then placed in FAA. Leaf samples with EFNs

of several Meiostemon, P tele op sis , Quisqualis and

Terminalia species were examined with Cambridge

Stereoscan 240 and Jeol JSM 5600 scanning electron

microscopes after being coated with platinum and gold

respectively. Transverse sections of leaf portions bearing

EFNs were prepared using a freezing or Reichert sliding

microtome and staining with alcian blue and safranin, or

by embedding in GMA, sectioning with an ultramicro-

tome and staining according to the periodic acid-

Schiff/toluidine blue method (Feder & O’Brien 1968).

Photographs were taken with a Leitz diaplan microscope

using Pan F film ASA 50 and Agfa APX 25 film. Slides

are housed at JRAU. Field observations were made over

a number of years on the EFNs of members of the

Combretaceae whenever possible.

Nectar samples were taken from the EFNs of a tree

of T. phanerophlebia growing in the grounds of the

University of Pretoria on 13 November 2001 . Samples

were also taken from the flowers of the same tree on 9

November 2002. They were collected as spots on

Whatman no. 1 filter paper and air-dried. The nectar

was then eluted with distilled water (3x15 //l), using a

centrifuge. HPLC was performed isocratically at a flow

rate of 2.5 ml min 1 on a ‘Waters Sugarpack’ column,

with acetonitrile-water (87:13) as eluent. The use of a

refractive index detector allowed the accurate calcula-

tion of the sugar composition, using peak height and 8

mg ml 1 of fructose, glucose and sucrose as external

standards.

RESULTS

To date, EFNs, as well as ‘glands of unknown func-

tion’ (see below), have been found in Laguncularia

C.F.Gaertn., Meiostemon , Pteleopsis, Quisqualis and

Terminalia but not in Combretum and Lumnitzera. With

the Meiostemon and Quisqualis species, the glands pre-

sent are assumed in this paper to be EFNs but field stud-

ies are needed to confirm this and also to ascertain

whether or not they are functional. In Lumnitzera race-

mosa Willd., at least two types of secretory structures of

unknown function are present on the leaves (Tilney

2002). Since this plant is a mangrove, other types of

secretory structures, e.g. salt glands, may well be in-

volved.

Ants and other insects were observed at one or more

times during the period November to early January on

the extrafloral glands of the young newly-formed leaves

of Terminalia brachystemma Welw. ex Hiern, T. mollis,

T. phanerophlebia (Figure 1 A, B), T. prunioides M.A.Law-

son and T. sericea Burch, ex DC. On many of the leaves

of T. phanerophlebia a white deposit, presumably sugar

crystals left after evaporation of water, was visible on the

glands on a very hot day (Figure 1C). Crystals were also

seen on T. sericea in association with the EFNs in the

SEM study (Figure ID)-. Ants were observed on Pte-

leopsis myrtifolia (M. A. Lawson) Engl. & Diels in No-

vember and January.

Distribution

In all the taxa with EFNs studied, these glands do not

appear on every leaf of every branch, thus limiting their

taxonomic significance. They are usually most conspicu-

ous, and probably only functional or at least optimally

functional, on new growth. When they become non-

Bothalia 34,2 (2004)

117

FIGURE 1. — EFNs of Terminalia species. A-C, T. phanerophlebia : A, leaves with ants, indicating presence of functional EFNs; B, ant on active

EFN: C, EFN with white sugary deposit visible on hot days following evaporation; mite (arrowed) can be seen in domatium. D, SEM of

portion of T. sericea EFN with crystals. Scale bars: A, 20 mm; B, C, 1 mm; D, 50 pm.

secretory in older leaves, there appears to be some

shrinkage frequently making them more difficult to

detect, especially in herbarium specimens.

The typical positions of the EFNs on the leaves in

species of Laguncularia, Meiostemon , Pteleopsis,

Quisqualis and Terminalia are shown in Figure 2. In

Laguncularia racemosa glands, whose structure and

function needs to be ascertained, were also included

because of their apparent similarity in herbarium materi-

al to certain known EFNs. In T. prunioicles a pair of

EFNs may occasionally be present on the petiole itself

rather than at the base of the leaf blade but are then never

conspicuous.

FIGURE 2. — Abaxial surfaces of leaves in taxa of Combretaceae showing variation in position of EFNs and related structures. A. Laguncularia

racemosa ; B. Meiostemon tetrandrum ; C, Pteleopsis anisoptera ; D, P. myrtifolia ; E, Quisqualis indica\ F, Q. parviflora ; G, Terminalia cat-

appa\ H, T. phanerophlebia', I. T. prunioides. EFNs are shown as black dots, domatia as circles and glands of unknown function as circles

with dots. Scale bar: 50 mm.

Bothalia 34,2 (2004)

118

FIGURE 3.— SEM of morphology of EFNs in members of Combretaceae. A, B, Pteleopsis myrtifolia, surface or somewhat concave EFNs. C-E,

raised EFNs: C, Quisqualis parviflora\ D, Terminalia pharterophlebia ; E, P. anisoptera. F, surface/raised EFN of T. gazensis. Scale bars:

A, 1 mm; B, E, 100 //m; C, 50 /<m; D, F, 200 pm.

Morphology

EFNs of members of the Combretaceae are generally

spherical in shape and may be raised, level with the surface

or somewhat concave (Figure 3). Species studied of

Meiostemon, Quisqualis and some of Terminalia (especial-

ly members of sections Platycarpae and Psidioides — see

Table 1 ) and Pteleopsis anisoptera (Fawson) Engl. & Diels

usually fit into the ‘Hochnektarien’ category of Zim-

mermann (1932)— referred to as ‘elevated’ by Elias (1983).

The other species studied of Terminalia (generally belong-

ing to section Abbreviatae — see Table 1) and Pteleopsis

myrtifolia , would be more suitably placed into the

‘Flachnektarien’ category of Zimmermann (1932)— ‘sur-

face’ gland of Elias (1983). Some of the glands may be slit-

shaped in Terminalia species (e.g. T. sericea). In T.

prunioides the EFNs are frequently a reddish colour but are

nevertheless inconspicuous.

Anatomy of EFNs in t/s

In species of Terminalia the nectaries are more or less

spherical structures (Figure 4A-F). Internal zonation

varies in degree but usually becomes more marked with

Bothalia 34,2 (2004)

FIGURE 4. — Terminalia species: t/s of EFNs showing varying degrees of internal zonation. A, T. mollis ; B, T. prunioides ; C, T. catappa ; D, T.

brachystemma\ E, T. trichopoda ; F, T. randii. G, T. mollis with outer portion of EFN showing epidermal layers and adjacent cells rich in

tanniniferous substances; H, peripheral portion of EFN of T. stenostachya with abundant crystals (some arrowed). Scale bars: A, D-H, 200

pm\ B, 100 pm\ C, 400 pm.

age. The nectariferous tissue may be a rounded, oval, variously shaped, but predominantly ± isodiametric cells

heart-shaped or irregular mass of small, tightly-packed, with thin walls and dense cytoplasm, which may become

120

Bothalia 34,2 (2004)

FIGURE 5.— Combretaceae taxa: t/s of EFNs or putative EFNs. A, Pteleopsis myrtifolia ; B, EFNs of Meiostemon tetrandrum composed of broad

cells; C, Quisqualis indica with cells distinctly aligned; D, Q. littorea', it is not clear whether structures such as this represent scar tissue,

ns, nectary sheath; vt, vascular tissue. Scale bars: A, C, 200 /( m; B, D, 100 //m.

tanniniferous with age (particularly in Terminalia section

Psiclioides ; see Table 1). This is sometimes surrounded

by a region of slightly larger parenchymatous cells with

sparser cytoplasm but numerous calcium oxalate crystals

(Figure 4H). Vascular tissue is associated with the nec-

tariferous tissue and the phloem usually appears well

developed. The epidermal cells overlying the nectary

may or may not resemble those found elsewhere in size,

cell wall thickness and contents. They may well be secre-

tory when young but frequently become thick-walled

and tanniniferous with age, as do a few layers of adjacent

cells, which in species such as T. catappa and T. mollis

(Figure 4G) form a distinctive area. Stomata and a nec-

tary sheath are absent.

Pteleopsis anisoptera and P. myrtifolia EFNs are simi-

lar to those of Terminalia species and consist of a

roundish to discoid mass of small, thin-walled, variously

shaped cells lacking visible intercellular spaces (Figure

5A). The contents of the cells comprising the nectarifer-

ous tissue are densely granular towards the base becom-

ing non-granular towards the outside, i.e. the abaxial sur-

face. The surface of the gland is frequently level with the

rest of the epidermis but occasionally concave. Stomata

are absent in this region. The epidermis overlying the

gland is initially much less tanniniferous than the rest of

the epidermis. The epidermal cells are very similar to the

underlying cells and may also be secretory. With age, the

epidermal cell walls, together with those of a few layers

of adjacent glandular cells, become lignified and the cell

interiors become filled with darkly-staining tannins.

Vascular tissue is associated with the base of the nectary

but does not appear to differ from tissue elsewhere in the

leaf. No nectary sheath is present but druses, some very

large, are found in the parenchymatous tissue in the vicin-

ity of the gland.

Quisqualis indica EFNs are quite different (Figure

5C), with groups of about six to ten superimposed layers

of broad, elongated cells, separated by groups of narrow,

columnar cells in close proximity to, but not obviously

connected to, the vascular tissue. The nectariferous tissue

is separated from adjacent tissue by a nectary sheath com-

posed of one or two layers of more rounded cells. The

epidermis does not appear to be secretory. Stomata may

be present. Druses (calcium oxalate) of varying sizes are

present in the nectariferous tissue and sheath. The vascu-

lar tissue close to the nectariferous tissue sometimes

appears to be better developed than elsewhere. The glands

in Q. littorea are much smaller and are composed of

broad cells only, most of which are not elongated anticli-

nally (Figure 5D). The cell walls of the epidermis and

underlying layers stain more intensely (suberised?). A

nectary sheath is absent. In Q. parviflora the mesophyll,

including the palisade, in localized areas tends to under-

go divisions resulting in fairly large cells. Intercellular

spaces are absent and there is no clear cell alignment.

In Meiostemon tetrandrum , the EFNs are composed

of about two layers of relatively large, anticlinally elon-

Bothalia 34,2 (2004)

121

FIGURE 6.— Combretaceae taxa: domatia and galls. A, domatium of Terminalia phanerophlebia with mite (arrowed); B, SEM of hairy domatium

of Quisqualis littorea: C, leaf of T. mollis , showing galls (some arrowed) in positions frequently occupied by domatia and EFNs: D. SEM

of pit domatium of T. catappa, with stomata (some arrowed) clearly visible; E, galls with stalks (arrowed) and rich in tanniniferous sub-

stances in t/s of leaf of T. mollis', F, pit domatium in t/s of leaf of T. catappa. Scale bars: A, E, 500 pm: B, F, 250 pm: C, 20 mm; D, 100

pm.

gated cells (Figure 5B). Druses are uncommon. A nectary

sheath is absent. Stomata were observed. The epidermis,

which appears to be non-secretory, becomes highly tan-

niniferous with age as do the cells in contact with it.

Domatia and galls

The typical distribution of domatia in representative

taxa under study is shown in Figure 2. The appearance

(SEM) of domatia in Terminalia phanerophlebia, Quis-

qualis littorea and T. catappa is illustrated in Figure 6A,

B and D respectively. The latter is also shown in t/s in

Figure 6F. The distribution of galls in T. mollis and their

structure in t/s is depicted in Figure 6C and E respective-

ly-

Nectar

The composition of the nectar produced by the EFNs

of Terminalia phanerophlebia was 35% fructose, 34%

glucose and 31% sucrose with no additional peaks in-

dicative of other sugars being present. This differed

markedly from that of the floral nectar which was 65%

fructose, 21% glucose and 14% sucrose (Figure 7).

122

Bothalia 34,2 (2004)

Floral Extra-floral

FIGURE 7.— Comparison of sugar composition between floral and

extra-floral nectar of Terminalia phanerophlebia. f, fructose; g,

glucose; s, sucrose.

DISCUSSION

Our field observations on five Terminalia species (T.

brachystemma, T. mollis, T. phanerophlebia , T. pruni-

oides and T. sericea) confirmed ant and other insect

activity on the extrafloral glands in the growing season.

Later observations on some of these species revealed no

such activity indicating that EFNs may be solely (or opti-

mally) functional on young growth. This is in line with

the statement of Elias et al. (1975) that foliar nectaries

are generally active when the leaves are young. Bentley

(1977) cites the studies of a number of researchers,

working on families other than the Combretaceae, who

also found that EFNs are active on the younger portions

of plants, usually being mature and active long before the

associated organ is fully developed. Galetto & Ber-

nardello (1992), referring to the work of Holldobler &

Wilson (1990), state: ‘There is evidence that plants time

their secretions in a way that enhances the protective role

of the nectaries’. Jolivet (1996) points out that sugars are

generally more concentrated in EFNs than in the phloem

partly due to evaporation. In his earlier work on

Clerodendrum fragrans Willd. (Jolivet 1985), he noted

that evaporation could transform the liquid into crystal-

lized sugars on the nectary surface as was found on

Terminalia phanerophlebia in the present study (Figure

1C). It is noteworthy that when EFNs are observed in

nature, they usually appear dry which may create the

false impression that they are non-secretory. A tactile

stimulation is not necessary to trigger the release of the

secretion as EFNs of T. phanerophlebia leaves without

ants, placed by us in closed containers with a high

humidity, produced droplets within minutes. This liquid

gradually dissolved any white deposit (sugar crystals)

that was present. The EFNs of T. amazonica are also

active as Schupp & Feener (1991) noted the presence of

ants. Ants were also observed by us on Pteleopsis myrti-

folia during the growing season. No field observations

were made by us at this time of year on Meiostemon

tetrandrum, P. anisoptera, Quisqualis littorea or on Q.

parviflora. Meiostemon, Pteleopsis and Terminalia

species are deciduous (frequently tardily so); Quisqualis

species are evergreen. It would thus be informative to see

in the case of Q. parviflora at what times it is frequented

by ants, if the EFNs are functional. No ant or other insect

activity was observed at any time of the year on the

glands of Q. indica.

Distribution

No EFNs were observed in any of the Combretum

species examined and there do not appear to be any

reports of such structures in members of the genus in the

literature except on the petiole in C. argenteum Bertol.

(Zimmermann 1932). The listing of this species is thus

considered doubtful and may well be based on confusion

with domatia as the latter do occur in members of the

genus Combretum. The mangrove, Lumnitzera race-

mosa, also appears to lack EFNs. This species, which

resembles the other strict combretaceous mangrove,

Laguncularia racemosa, very closely in leaf anatomy

(Biebl & Kinzel 1965; Tilney 2002), thus lacks the peti-

olar EFNs typical of the latter species. Von Hohnel

(1882) refers to two pairs of glands at the apex of the

petiole of Laguncularia racemosa-, the lower pair on

small ‘warts’ but the upper pair not prominent. Belin-

Depoux (1993) does not make a size distinction and

refers to one to three small prominences on the adaxial

side of the petiole of this species. The presence of EFNs

in a mangrove seems surprising and observations on their

efficacy would be very informative. Glands of unknown

function, and only functional in young leaves, have been

reported by Biebl & Kinzel (1965) and Stace (1965a) in

Laguncularia racemosa. The distribution of these glands

forms a characteristic pattern (Figure 2A) which is par-

ticularly conspicuous in older leaves, and appears to be

of taxonomic value. Tomlinson (1986) suggests that

these may function temporarily as hydathodes in young

leaves, secreting water or mucilage. However, the

anatomical structure is similar to what we found in some

of the EFNs of the southern African taxa (notably

Terminalia spp. e.g. T. trichopoda Diels) and we recom-

mend that their function be investigated further.

The present study confirmed the presence of EFNs in

Terminalia but their distribution differs from most previ-

ous reports in other species of the genus. In the

Combretaceae paired petiolar nectaries, sometimes sim-

ply referred to as ‘glands’, have been reported in un-

named species of Terminalia, Conocarpus L., Anogeis-

sus Wall, and Laguncularia by Bentham & Hooker as

cited by Solereder (1908); T. brasiliensis (authority not

indicated), C. procumbens L. and L. racemosa (Zim-

mermann 1932); T. mucronata Craib & Hutchinson

(Belin-Depoux 1978); an unnamed Terminalia species

(Exell 1978); T. arjuna Wight & Arn. (Ramakrishna &

Rajashekara 1981), and in T. argentea Mart. & Zucc. and

also T. brasiliensis Camb. (Oliveira & Leitao-Filho

1987). According to Ramakrishna & Rajashekara (1981),

the EFNs of T. arjuna have a prominent stalk which was

not observed in any of the species in the present study. In

T. paniculata Roth, Von Hohnel (1882) describes the

glands as large ‘warts’ (‘Warzen’), over 1 mm in height

and breadth, occurring on the underside of the lamina on

both sides of the base of the midrib. Liben (1965) refers

to EFNs being present on the lamina of T. griffithsiana

Liben, and Van Wyk et al. (2000) and Coates Palgrave

(2002) on the lamina of certain southern African

Bothalia 34,2 (2004)

123

Terminalia species. In the present study, in some species

of Terminalia , such as T. catappa , glands were found to

be next to the midrib near the petiole insertion but were

not confined to this position. In such cases, those at the

base of the midrib tended to be the largest and most

prominent. In other species their distribution appeared

completely sporadic without any fixed pattern.

Repeated observations on living material of the in-

digenous species, Terminalia phanerophlebia, revealed

occasional glistening areas on some of the veins. These

areas were not associated with any visible proliferation

of leaf tissue as is found in conventional EFNs. However,

it appeared as if a nectar-like substance was being secret-

ed. This raises the question as to whether nectar secretion

could be independent of the presence of distinct glands.

In the Combretaceae this requires further investigation

since non-structural nectaries, which cannot be recog-

nized macroscopically and which exude nectar through

stomata, have been reported in many plants of other

families (Fahn 1979).

So-called ‘inconspicuous bullae’, two or three in num-

ber and positioned ‘on or near laterals’ are mentioned in

connection with Pteleopsis anisoptera but not with P.

myrtifolia by Carr (1988). These ‘bullae’ we have identi-

fied as EFNs (Figure 3E). When present, we observed

2-6 of these glands per leaf, usually in pairs (Figure 2C).

In P. myrtifolia , EFNs may also occur but there is almost

invariably only a single pair on the lamina (Van Wyk &

Van Wyk 1997; Coates Palgrave 2002) in the characteris-

tic position shown in Figures 2D and 3 A. When present,

these EFNs are a useful taxonomic character.

In the indigenous species Quisqualis parviflora , the

EFNs are very small and they often appear to be close to

one another. The alien, Q. indica , has EFNs which are

similarly very small but are visible as purple spots which

could easily be dismissed as being insect damage. The

EFNs appeared to be non-secretory and no ants were

observed on the plant. This raises the possibility that

feeding activity of ants may be necessary to induce the

secretion of nectar in this species. This would provide an

effective mechanism to economize on the secretion of

nectar, an obviously expensive commodity for the plant

to produce. If this is the case, there is the possibility of a

substance being secreted to attract ants but these aspects

also need further study in situ.

It is thus noteworthy that in the case of taxa such as

Pteleopsis anisoptera , P. myrtifolia and Laguncularia

racemosa the position of the EFNs is genetically fixed

but in other taxa (e.g. Terminalia phanerophlebia ) there

is a more flexible system with their formation appearing

to be opportunistic. In all cases, however, plants appar-

ently have some control over the formation of these

structures (their presence or absence) as well as their

functioning.

Morphology

Zimmermann (1932) reported two types of EFNs on

the underside of the leaf in two different species of

Terminalia , viz. ‘Hochnektarien’ (raised nectaries) in T.

paniculata , and ‘Flachnektarien’ (nectaries flush with the

surface) in T. catappa. Pascal et al. (2000) prefer the

term ‘nectary with an apical depression’ rather than ‘ele-

vated’ EFN to describe the stalkless pit or cup-shaped

nectaries of some legumes which resemble externally

those of some Pteleopsis and Terminalia species (notably

members of the section Platycarpae). In particular the

scanning electron micrograph of the secretory structure

of the legume Inga feuillei DC. resembles that of the

Terminalia species studied and of Pteleopsis anisoptera.

In the present study, although the glands of the Meio-

stemon and Quisqualis species (Figure 3C) are similarly

raised to those of the Terminalia species (Figure 3D) and

Pteleopsis anisoptera (Figure 3E), they differ markedly

in size and appearance.

Anatomy

Combretaceous EFNs possess a specially differentiat-

ed nectariferous tissue; thus they belong to the structural

type (Fahn 1979). However, as was mentioned earlier in

connection with Terminalia phanerophlebia , there is the

possibility of the non-structural type also occurring. The

EFNs of the species of Terminalia and Pteleopsis studied

are very similar particularly in having nectariferous tis-

sue composed of very small cells but differ from those of

Quisqualis and Meiostemon which are similar to one

another and have secretory tissue in the form of anticli-

nally elongated palisade-like cells. The latter two gen-

era—as well as the others — do not, however, have elon-

gated epidermal cells which are typical of epidermal

glands (Schnell et al. 1963; Haberlandt 1965; Wilkinson

1979). The EFNs of the Combretaceae usually have large

numbers of calcium oxalate crystals associated with

them (Figure 4H) as is reported frequently in various

taxonomic groups of plants (Wilkinson 1979). Pteleopsis

myrtifolia EFNs have the same subepidermal ring of lig-

nified and tanniniferous cells as in T. catappa and T. mol-

lis (Figure 4G). The EFNs of T. catappa as described by

Zimmermann (1932) and shown in our sections, show a

similarity with those of T. mollis (Figure 4A, C). Those

of T. mucronata (Belin-Depoux 1989) also resemble the

EFNs particularly of members of the section Platy-

carpae. The ‘nectary sheath’ of Zimmermann (1932)

appears to correspond to the ‘median zone’ of Belin-

Depoux (1989). In Laguncularia racemosa , EFNs also

consist of differentiated regions (Von Hohnel 1882;

Biebl & Kinzel 1965), and Belin-Depoux (1993) similar-

ly identifies three histological zones in longitudinal sec-

tions. Von Hohnel (1882) reports a similar structure in T.

paniculata to that in L. racemosa apart from an absence

of suberisation in the outer sheath of the former.

EFNs vs. domatia vs. galls

Confusion and difficulties in distinguishing between

domatia and glands (EFNs) in many taxa, at least in

herbarium material, have been recognized for many

years and pointed out by authors such as Jacobs (1966),

Wilkinson (1979) and Brouwer (1985). Stace (1965a)

mentions the possibility of the mites which inhabit

domatia enlarging the domatia, thus making identifica-

tion of the structures even more difficult. Mites were

observed in the domatia of Terminalia phanerophlebia

(Figure 6A). All the major types of domatia are recorded

by Stace (1965b) in the Combretaceae where they are

124

Bothalia 34,2 (2004)

important diagnostic features in several genera. He

makes no mention of EFNs in the family as such but

records gland-like structures, found not very frequently,

in the domatia of Buchenavia parvifolia Ducke and T.

archboldiana Exell (Stace 1965a). We suspect that many

structures thought to be domatia are, in fact, glandular

and would more accurately be described as EFNs. If

some of these structures are actually EFNs the sugar, in

addition to the shelter, could possibly contribute to the

high degree of insect infestation and damage in many

leaves. Bentley (1977) cites examples where an increase

in the rate of secretion caused by sucking insects is posi-

tively correlated with increased infestation levels and it

would be informative to ascertain whether this is applic-

able in members of the Combretaceae.

The fact that EFNs and domatia may occupy similar

positions on the leaf blade not only results in misinter-

pretation but it may also indicate that the structures are

very closely related in their ontogeny, even to the extent

of being interchangeable. The relative abundance of the

one or other may well be due to the developmental stage

of the leaf. However. Wilkinson (1979) states: 'Since

EFNs sometimes occur in the same type of situation as

domatia, i.e. in the vein axils on the abaxial surfaces of

leaves, the difference of presence or absence of glands is

probably of doubtful significance’.

Although EFNs and domatia may be located in simi-

lar positions, in our experience it is usually possible to

distinguish between them with low magnification.

Anatomically the distinction is very clear as can be seen

in Terminalia catappa (Figures 4C; 6F). Brouwer (1985)

gives a comparison between the structure of 'undoubted’

domatia and EFNs. In our study of the domatia of T. cat-

appa no tanniniferous substances were visible in the

vicinity of the domatia and no narrow columnar epider-

mal cells or hypodermis were evident (Figure 6F).

Brouwer (1985) also states that lysigenous cavities in

Lumnitzera racemosa have been misinterpreted as doma-

tia but no suggestion is made as to their nature.

Jacobs (1966) cites the work of authors who report the

presence of galls in the typical position of domatia. This

appears to be the result of gall-forming mites preferen-

tially attacking the positions normally occupied by the

domatia. Despite similarities (position, shape, the pres-

ence often of hairs and of mites), however, galls are gen-

erally irregular in extension and cell structure, and the

cells ‘disturbed’, whereas domatia are regular in exten-

sion and cell structure, and the cells intact (Jacobs 1966).

Furthermore he quotes Fundstroem’s (1887) statement

that in domatia a palisade layer is present and tannin

absent, the opposite of which occurs in galls. Brouwer

(1985) and Belin-Depoux (1989) both record the pres-

ence of tannins (as well as calcium oxalate crystals) in

EFNs. If domatia may ‘increase the liability of nerve

axils to action on the part of gall-mites’ (Jacobs 1966), it

seems likely that EFNs would be even more effective in

this regard. In the present study galls were frequently

observed in the position of EFNs (Figure 6C). Transverse

sections of the galls of Terminalia mollis (Figure 6E)

show that they differ markedly in structure from the

EFNs (Figure 4A). The galls are stalked (and therefore

more raised) unlike the EFNs. They are also unique in

having one or two large central cavities. Tannins are

abundant in the surrounding cells but calcium oxalate

crystals are essentially absent. In South African Com-

bretaceae some of the most complex galls are caused by

members of the Fepidoptera rather than by mites

(Scholtz 1978). Three types of Fepidopteran galls are

recorded on the stems and shoots of Terminalia sericea

and one type in the fruits of Combretum molle R.Br. ex

G.Don in South Africa. It is evident from Figure 6E that

the galls in this specimen are not mite-induced but prob-

ably Fepidopteran galls.

Nectar

Galetto & Bernardello (1992), reporting on an analy-

sis of the chemical composition of the extrafloral nectar

of about 60 plant species, found that three or more sug-

ars were usually present, as is generally the case in floral

nectar where sucrose, glucose and fructose are the most

common (e.g. Fahn 1988). In Turnera ulmifolia F.

(Turneraceae), the floral nectar is sucrose-dominant,

whereas the foliar nectar is a balanced solution of

sucrose, glucose and fructose (Elias et al. 1975). Pate et

al. (1985) noted that in the cowpea [Vigna unguiculata

(F.) Walp.], the sucrose: glucose: fructose weight ratios

of nectar were similar in the two structurally dissimilar

types of EFNs but differed markedly in balance between

sugars and other organic solutes. Fiala & Maschwitz

(1991), in studies of Macaranga Thouars (Euphorbi-

aceae), found that the secretion of the marginal glands of

myrmecophytic species lacked sugar unlike that of the

non-myrmecophytic species. Freitas et al. (2001) studied

the composition of nectar produced by floral and EFNs

at different stages of development in Croton sacro-

petalus Mtill.Arg. (Euphorbiaceae). The sugars in the

nectar of all nectary types were mainly monosaccharides

with a predominance of glucose over fructose. The only

exception was the post-floral nectar (nectar produced

after anthesis during fruit development) that mostly had

sucrose and a greater proportion of fructose than glucose.

Hardly anything is known about the movement of

nectar during the secretory process. For example, since

intercellular spaces are essentially lacking, what role

does symplastic transport play? It is also not clear how

the nectar moves from the nectariferous tissue to the leaf

surface. The permeability of the cuticle needs to be

investigated and also, in those species where they are

present, the possible role of stomata. This requires fur-

ther study of live material .

In the present study the sugar composition was inves-

tigated only in the floral and extra-floral nectar of

Terminalia phanerophlebia . The EFNs are very active in

this species (Figure 1 A). In both cases the nectar is com-

posed of only three sugars, fructose, glucose and sucrose

(Figure 7). The markedly different ratios of these sugars

obtained in floral and extrafloral nectar indicate that dif-

ferent insects would probably be attracted. Our own

casual observations suggest that the balanced-sugar nec-

tar is presumably preferred by ants and the fructose-domi-

nant nectar by the pollinators (especially flies) but this

aspect obviously requires further study. The very high

fructose level of the floral nectar is unusual; fructose-

dominant nectar has so far only being found in Aster-

Bothalia 34,2 (2004)

125

aceae, the genus Crassula (Crassulaceae) and in Wel-

witschia (B-E. van Wyk pers. comm.). Bemardello et al.

(1994) found that the floral nectar sugar composition was

remarkably consistent in the different stages throughout

the lifetime of the flowers of Combretum fruticosum

(Loefl.) Stuntz. The nectar contained a very small amount

of sucrose (less than 0.6%) and a predominance of hex-

oses. Fructose and glucose were in a relative ratio of ±

1 : 3.5 respectively. This thus differs markedly from the

sugar composition of both the floral and extrafloral nec-

tar in Terminalia phanerophlebia . Further studies are

obviously needed to determine whether the unusual sugar

composition found in the floral nectar of T. phanero-

phlebia is characteristic of other related taxa and the

extent to which the sugar composition of foliar nectar

may be of taxonomic significance.

CONCLUSIONS

Where EFNs are known to be present, they are only

functional, or optimally functional, on young leaves.

They can be very small (as in most southern African

Combretaceae) and easily overlooked unless specifical-

ly searched for. In many taxa their occurrence on a plant

is sporadic, thus limiting their taxonomic significance.

When present, the EFNs in Pteleopsis myrtifolia occur

as single pairs in a characteristic position on the lamina

which readily enables their identification. In P. ani-

soptera , although the EFNs also tend to be arranged in

pairs, many of the gland-bearing leaves have two or

three pairs. Glands (EFNs?) on the leaves of Lagun-

cularia racemosa form a characteristic pattern. In

Terminalia catappa there are usually two prominent

EFNs, one on either side of the midrib at the base of the

leaf, whereas in other species (e.g. T. phanerophlebia)

they are not necessarily prominent in this position. All

southern African species of Terminalia except occasion-

ally T. prunioides appear to lack paired petiolar glands

(not to be confused with paired glands at the junction

between petiole and lamina). In this latter species where

they were observed on the petiole, they were never con-

spicuous. Apparent structural EFNs occur in Meioste-

mon and Quiscptalis species, but it is not known whether

they are functional. Ants were, however, seen on young

leaves of five Terminalia species and were also

observed on a plant of Pteleopsis myrtifolia. EFNs are

absent in Combretum and probably also in Lumnitzera

racemosa.

The macroscopic distinction between EFNs, domatia

and galls is often not clear. Structures, previously regard-

ed as domatia, especially on the lamina of Terminalia

species, may well be EFNs. Since EFNs (and galls) are

frequently in the position of domatia, it seems that it is

not essential to differentiate between them and that they,

just like domatia, can be used for taxonomic purposes.

Microscopically they can be readily distinguished. The

known microscopic structure of the EFNs of the

Terminalia and Pteleopsis species shows a basic similar-

ity but with somewhat varying degrees of zonation. The

structure of the putative EFNs of Meiostemon tetran-

drum and the Quisqualis species studied, on the other

hand, differs in cell size, arrangement and zonation from

those of the Terminalia and Pteleopsis species.

In Terminalia phanerophlebia, the difference in sugar

composition between the nectar of foliar and floral nec-

taries may have the effect of attracting different insects.

ACKNOWLEDGEMENTS

We thank the curators and staff of the Herbarium,

Royal Botanic Gardens, Kew (K) and the National

Herbarium, Pretoria (PRE) for allowing us to examine

herbarium material during study visits and for the gall-

bearing material of Terminalia mollis. Part of this study

was done by the first author at the Jodrell Laboratory,

Royal Botanic Gardens, Kew, and the staff is thanked for

the use of facilities and their assistance. Meg Coates

Palgrave and Tony Abbott are thanked for supplying

some of the material used in the study and the Natal

Herbarium (NH) for the fresh specimens of T. catappa.

The sugar analyses were kindly made possible by Prof.

B-E. van Wyk of the Rand Afrikaans University.

REFERENCES

BELIN-DEPOUX, M. 1978. Contribution a l’etude des glandes petio-

laires du Terminalia mucronata Craib. et Hutch. (Combre-

taceae). Revue de Cytologie et de Biologie Vegetales, le

Botaniste 1: 295-307.

BELIN-DEPOUX, M. 1989. Des hydathodes aux nectaires foliaires

chez les plantes tropicales. Bulletin de la Societe botanique de

France 136, Actualites botaniques 3, 4, 151-168.

BELIN-DEPOUX, M. 1993. Importance des nectaires extra-floraux

dans les interactions plantes-fourmis. Acta Botanica Gallica

140: 183-205.

BENTLEY, B.L. 1977. Extrafloral nectaries and protection by pugna-

cious bodyguards. Annual Review of Ecology and Systematics

8: 407^127.

BERNARDELLO, L„ GALETTO, L. & RODRIGUEZ. I.G. 1994.

Reproductive biology, variability of nectar features and pollina-

tion of Combretum fruticosum (Combretaceae) in Argentina.

Botanical Journal of the Linnean Society 114: 293-308.

BIEBL. R. & KINZEL, H. 1965. Blattbau und Salzhaushalt von

Laguncularia racemosa (L.) Gaertn.f. und anderer Man-

grovebaume auf Puerto Rico. Osterreichische Botqnische

Zeitschrift 112: 56-93.

BROUWER, Y.M. 1985 .Domatia — morphology, anatomy and biologi-

cal significance. Ph.D. thesis. University of Queensland.

CARR, J.D. 1988. Combretaceae in southern Africa. The Tree Society

of Southern Africa, Johannesburg.

COATES PALGRAVE. M. 2002. Keith Coates Palgrave Trees of south-

ern Africa, edn 3. Struik, Cape Town.

DELPINO, F. 1886-1889. Funzione mirmecofila del regno vegetale.

Prodromo d’una monografia delle piante formicarie. Memorie

della R. Accademia delle Scienze dell' Istituto di Bologna 7:

215-323; 8: 601-650; 10: 115-147.

ELIAS, T.S. 1983. Extrafloral nectaries: Their structure and distribu-

tion. In B. Bentley & T.S. Elias, The biology of nectaries:

174-203. Columbia University Press, New York.

ELIAS, T.S. , ROZICH. W.R. & NEWCOMBE. L. 1975. The foliar and

floral nectaries of Turnera ulmifolia L. American Journal of

Botany 62: 570-576.

EXELL, A.W. 1978. Combretaceae. In E. Launert, Flora zambesiaca 4:

100-183. Flora Zambesiaca Managing Committee. London.

FAHN, A. 1979. Secretory tissues in plants. Academic Press, London.

FAHN, A. 1988. Tansley Review No. 14. Secretory tissues in vascular

plants. New Phytologist 108: 229-257 .

FEDER. N. & O'BRIEN, T.P. 1968. Plant microtechnique: some prin-

ciples and new methods. American Journal of Botany 55:

123-142.

FIALA, B. & MASCHWITZ, U. 1991. Extrafloral nectaries in the

genus Macaranga (Euphorbiaceae) in Malaysia: comparative

studies of their possible significance as predispositions for

myrmecophytism. Biological Journal of the Linnean Society

44: 287-305.

FREITAS, L„ BERNARDELLO, G„ GALETTO, L. & PAOLI, A.A.S.

2001. Nectaries and reproductive biology of Croton sar-

126

Bothalia 34,2 (2004)

copetalus (Euphorbiaceae). Botanical Journal of the Linnean

Society 136: 267-277 .

GALETTO. L. & BERNARDELLO, L.M. 1992. Extrafloral nectaries

that attract ants in Bromeliaceae: structure and nectar composi-

tion. Canadian Journal of Botany 70: 1101-1106.

HABERLANDT. G. 1965. Physiological plant anatomy , reprint edn.

Today and Tomorrow’s Book Agency, New Delhi.

HOLLDOBLER. B. & WILSON, E.O. 1990. The ants. Springer-

Verlag, Berlin.

JACOBS, M. 1966. On domatia— the viewpoints and some facts, I, II

& III. Koninklijke N ederlandse Akademie van Wetenschappen.

Proceedings Series C, 49: 275-316.

JOLIVET, P. 1985. Les decouvertes recentes en myrmecophilie des

plantes. Tendances actuelles et interpretations. Actes du

Congres J.H. Fabre Paris : 41-52.

JOLIVET, P. 1996. Ants and plants: an example of coevolution, en-

larged edn. Backhuys, Leiden.

LIBEN, L. 1965. Note sur quelques Combretaceae ( Combretum LoefL,

Terminalia L. et Strephonema Hook.f.) du Congo, du Rwanda

et du Burundi. Bulletin van de Rijksplantentuin Brussel 35:

167-184.

LUNDSTROEM, A.N. 1887. Pflanzenbiologische Studien II. Die

Anpassungen der Pflanzen an Thiere. Nova Acta Regiae

Societatis Scientiarum Upsaliensis iii 2,6: 1-74.

MABBERLEY, D.J. 1997. The plant-book. A portable dictionary of the

vascular plants, edn 2. Cambridge University Press, Cam-

bridge.

METCALFE, C.R. & CHALK, L. 1979. Anatomy of the dicotyledons,

vol. 1 , edn 2. Clarendon Press, Oxford.

OLIVEIRA, PS & LEITAO-FILHO, H.F. 1987. Extrafloral nectaries:

their taxonomic distribution and abundance in the woody flora

of cerrado vegetation in southeast Brazil. Biotropica 19:

140-148.

PASCAL, L.M., MOTTE-FLORAC, E.F. & MCKEY, D.B. 2000.

Secretory structures on the leaf rachis of Caesalpinieae and

Mimosoideae (Leguminosae): implications for the evolution of

nectary glands. American Journal of Botany 87: 327-338.

PATE, J.S.. PEOPLES, M.B., STORER, P.J. & ATKINS, C.A. 1985.

The extrafloral nectaries of cowpea (Vigna unguiculata (L.)

Walp.) II. Nectar composition, origin of nectar solutes, and nec-

tary functioning. Planta 166: 28-38.

RAMAKRISHNA, T.M. & RAJASHEKARA, G. 1981. Foliar nec-

taries in Terminalia arjuna (Combretaceae). Current Science

50: 1072-1074.

SCHNELL, R„ CUSSET, G. & QUENUM, M. 1963. Contribution a

I'etude des glandes extra-florales chez quelques groupes de

plantes tropicales. Revue Generale de Botanique 70: 269-341.

SCHOLTZ, C.H. 1978. Notes on plant galls formed by Lepidoptera in

the Transvaal. Journal of the Entomological Society of Southern

Africa 41 : 45^19.

SCHUPP, E.W. & FEENER, D.H. 1991 . Phylogeny.lifeform, and habi-

tat dependence of ant-defended plants in a Panamanian forest.

In C.R. Huxley & D.F. Cutler, Ant-plant interactions: 175-197.

Oxford LIniversity Press, Oxford.

SOLEREDER, H. 1908. Systematic anatomy of the dicotyledons, vol.

1 : 347. Clarendon Press, Oxford.

STACE, C.A. 1965a. Cuticular studies as an aid to plant taxonomy.

Bulletin of the British Museum (Natural Histoiy), Botany 4:

3-78.

STACE, C.A. 1965b. The significance of the leaf epidermis in the tax-

onomy of the Combretaceae I: a general review of tribal, gener-

ic and specific characters. Botanical Journal of the Linnean

Society 59: 229-252.

TILNEY, P.M. 2002. A contribution to the leaf and young stem anato-

my of the Combretaceae. Botanical Journal of the Linnean

Society 138: 163-196.

TOMLINSON, P.B. 1986. The botany of mangroves. Cambridge

University Press, Cambridge.

VAN WYK, B. [A.E.] & VAN WYK, P. 1997. Field guide to trees of

southern Africa. Struik, Cape Town.

VAN WYK, B. [A.E.], VAN WYK, P. & VAN WYK, B-E. 2000.

Photographic guide to trees of southern Africa. Briza Publi-

cations, Pretoria.

VON HOHNEL. F. 1882. Beitrage zur Pflanzenanatomie und

Physiologie V. Zur Anatomie der Combretaceen. Botanische

Zeitschrift 40: 177-182.

WILKINSON, H P. 1979. Extrafloral nectaries. In C.R. Metcalfe & L.

Chalk, Anatomy of the dicotyledons, vol. 1, edn 2: 124—130.

Clarendon Press, Oxford.

ZIMMERMANN, J.G. 1932. Uber die extrafloralen Nektarien der

Angiospermen. Beihefte zum Botanischen Centralblatt 49:

99-106.

Bothalia 34,2: 127-140 (2004)

Cape heaths in European gardens: the early history of South African

Erica species in cultivation, their deliberate hybridization and the

orthographic bedlam

E.C. NELSON* and E.G.H. OLIVER**

Keywords: botanical history, Cape heaths, garden history, hybrids, nomenclature

ABSTRACT

This paper discusses the horticultural history of southern African Erica spp. in Europe, and especially in Britain, dur-

ing the late eighteenth and the early decades of the nineteenth century. We note evidence for the deliberate hybridization of

the so-called Cape heaths by European horticulturists, in particular by the English nurseryman William Rollisson and by the

Very Rev. William Herbert. We discuss some of the nomenclatural consequences of the naming by miscellaneous botanists

and nurserymen of the hundreds of new Erica species and hybrids, emphasizing the proliferation of eponyms. An appendix

tabulates eponyms and their numerous orthographic variants published before 1835 within Erica , and provides the correct

orthography for these epithets.

INTRODUCTION

The early history of the discovery of the flora of south-

ern Africa is well summarized by Gunn & Codd (1981),

and a brief historical sketch of the description and depic-

tion of Cape species of Erica prefaces Baker & Oliver

( 1968). This paper concentrates on the horticultural histo-

ry of southern African Erica spp. in Europe, notes evi-

dence for deliberate hybridization of the so-called Cape

heaths by European horticulturists and discusses some of

the nomenclatural consequences of the naming of these

plants by miscellaneous botanists and nurserymen.

The exploration of southern Africa by Europeans com-

menced, at least as far as extant historical documents

record, with the voyages of Portuguese navigators in the

late fifteenth century. With the discovery of this new route

to the East Indies, Dutch, French and English vessels soon

joined the Portuguese and began to sail around the south

of Africa into the Indian Ocean. The formation of the

Honourable East India Company in England and

Vereenigde Oost-indische Compagnie in the Netherlands

added further impetus and soon resulted in the first pub-

lished accounts of the natural history of the region. The

directors of these companies frequently instructed the cap-

tains of their vessels to bring back any objects that they

thought would be of interest, not just commercially, and so

by the early seventeenth century, at least, specimens of

plants and animals from southern Africa were being

brought to Europe. Undoubtedly, numerous botanical

curiosities, including seeds, from the Cape of Good Hope

reached Europe in the sixteenth century but they were not

recorded in books or manuscripts and so have left no trace.

The earliest botanical specimen from the southernmost

part of Africa illustrated in a European publication was an

* Tippitiwitchet Cottage, Hall Road, Outwell, Wisbech PE14 8PE,

Cambridgeshire, United Kingdom.

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

7735 Claremont, Cape Town.

MS. received: 2004-03-09.

inflorescence of Protea neriifolia R.Br. depicted by

L’Ecluse (olim Clusius) in Exoticorum libri decern (Clusius

1605; Kerkham 1988; Rourke 1980). L'Ecluse also first

reported an instance of a Cape species in cultivation in a

European garden, that being 'Omithogalum aethiopicum’,

the bulbs of which had been collected by sailors at the Cape

of Good Hope (Clusius 1611; Gunn & Codd 1981: 13). The

inflorescence of Protea neriifolia is a tough object which

will not disintegrate easily, and so its survival as a curiosi-

ty is not unexpected. Bulbs, corms and rhizomes, while

perishable, stood a reasonable chance of remaining viable

and reaching European collectors despite the numerous

hazards of sea voyages in that era. Thereafter, southern

African plants, especially those native to the vicinity of the

Cape of Good Hope, became increasingly familiar, even

commonplace, in European gardens, and numerous botani-

cal works contained illustrations of Cape species, predomi-

nantly bulbs, succulents and plants with tough rhizomes.

While Erica spp. are abundant in the coastal peripheries

likely to have been visited by sailors (Oliver & Oliver

2000), they are not so easy to preserve. Heaths resent dis-

turbance and are unlikely to have survived transport alive

at that period. This was not yet the age of scientific expe-

ditions staffed by individuals who knew how to preserve,

by pressing and drying specimens of plants such as Erica ,

or how to care for living plants on ocean-going vessels.

Furthermore, the collection of seeds was probably a ran-

dom happening, a seaman or curious traveller taking a

chance when he saw some ripe fruits— gathering and keep-

ing the minute seeds of Erica spp. would have required

deliberate actions at the right time of the year. Bulbs and

succulents were much easier to gather— they could be

obtained in quantity and the chance taken that a few would

survive the rigours of an Atlantic voyage in a sailing ship

which, given the vagaries of winds and currents, usually

took around three months.

THE FIRST CAPE HEATHS IN CULTIVATION IN EUROPE

There is evidence from the late seventeenth or early

eighteenth century that at least one Cape species of Erica

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Bothalia 34,2 (2004)

129

had been successfully cultivated in the Netherlands.

Wijnands (1983) recorded that a painting by Maria

Moninckx, dated between 1686 and 1706, contained in

volume 8 of the unpublished ‘Afteeken ingen van ver-

scheyden vreemde gewassen; in de Medicyn-Hoff der

Stadt Amsteldam' , depicts E. curvirostris Salisb., a species

that grew in the vicinity of the Dutch settlement at the Cape

of Good Hope. The painting cannot be precisely dated and

was not published by Casper Commelin (Wijnands 1983:

94. 95), but its existence proves that before 1707 at least

one Cape Erica had bloomed in Dutch gardens.

The earliest published illustrations of Cape Erica

spp. date from the same period. Plukenet (1700) includ-

ed three in Almagesti botanici mantissa : E. plukenetii

L., E. curviflora L. and E. bruniades L. (Baker & Oliver

1968). Almagesti botanici mantissa was one of four vol-

umes which Henrey (1975: 1: 140-142) described as a

‘great catalogue of plants’. There is no evidence the

Erica spp. were illustrated from living plants — on the

contrary, the engravings seem to portray specimens that

were plucked from the wild and pressed. This accords

with what is known about the plants described and

depicted in Plukenet’s books. Many of the illustrations

were done from specimens in his own herbarium that

had been obtained from a worldwide network of corre-

spondents (Henrey 1975: 1: 142). The same applies to

Seba’s Locupletissimi rerum naturalium (1734-1765)

— Cape Erica spp. were illustrated from Seba’s Kunst-

kamer , from pressed specimens, not from garden plants.

Indisputable evidence that Cape Erica spp. were in

cultivation, at least in specialized collections, by the

1730s, is contained in the herbarium of George Clifford’s

garden at Hartecamp, the Netherlands, and in the associ-

ated book by Linnaeus (1738: 147, 148). According to

Linnaeus, Clifford grew five different African heaths,

including E. cuniflora (Linnaeus 1738: 148), which

Plukenet (1700) had illustrated.

Cape Erica spp. are not easy for northern European

gardeners to grow, even today with modem computer-

controlled glasshouses. The vast majority cannot be cul-

tivated outdoors all year round because they are frost-

sensitive. They require lime-free soil, and when cultivat-

ed in containers, the watering regime has to be carefully

regulated. The fact that they had to be kept indoors at least

during the late autumn, winter and early spring meant that

only gardeners possessing heated conservatories could

attempt their cultivation. In the early eighteenth century,

greenhouses were heated by either free-standing stoves

or smoked-filled flues built into the walls. Furthermore,

especially in winter, these buildings were poorly venti-

lated and had low light levels, two more factors that mil-

itated against success in the cultivation of these plants.

However, as the design of greenhouses and conservato-

ries improved, especially once roof slates were replaced

with glass panes, and as gardeners became more familiar

with the plants’ cultural requirements, more species were

successfully grown. Indeed, during the first half of the

nineteenth century, the skills needed for the successful

cultivation of Cape heaths had been so well mastered that

these plants became extraordinarily fashionable.

The horticultural progress went in tandem with botan-

ical exploration of the Cape of Good Hope.

BOTANICAL COLLECTORS

Linnaeus (1753) published 12 names for species of

Erica from southern Africa— but Species plantarum was

not a catalogue of plants in gardens. The number of

named species increased steadily in the three following

decades (Figure 1). By the end of the 1760s, about 63

Cape Erica spp. (including those originally placed in

Blaeria) had been given Latin binomials. The 1770s saw

27 more names published, and during the 1780s another

54 Erica names appeared in print, almost 40 of these

being coined by Thunberg (1785).

During the late eighteenth century. Cape Erica spp.

were introduced into European gardens, principally by

seed gathered in the wild. In 1789 William Aiton pub-

lished Hortus kewensis, for which Daniel Solander sup-

plied the descriptions of seven new species of Erica from

the Cape; one proved to be an already described species,

but six were previously unknown. These had all been

raised from seed collected by Francis Masson (1741-

1805), who was the first collector to make an impact in

the horticultural, as distinct from the purely botanical,

aspects of Cape Erica spp. The new species originated

from his first visit to the Cape, which had lasted two and

a half years, from October 1772 to March or April 1775.

According to Masson’s (1776) own accounts of his 'three

journeys from Cape Town into the southern parts of

Africa’ , most of his Erica seeds were obtained during his

first extended expedition which started in December

1772 and ended in late January or early February 1773

(Masson 1776; Bradlow 1994). Of that first expedition,

from Cape Town via Paarl, Franschhoek, Stellenbosch

and Swartberg to Swellendam and back, Masson (1776;

Bradlow 1994: 109) stated: ‘It was on this journey that I

collected the seed of the many beautiful species of Erica

which, I find, have succeeded so well in the Royal

Garden at Kew’. Solander (Aiton 1789) attributed to

Masson the collection or introduction of 18 Cape species

during 1774; two of his collections are dated 1773 and

three are dated 1775. In all. more than half of the 41

Erica spp. listed at Kew in 1789 had been raised from

seed obtained by Masson at the Cape.

Masson returned to England in 1775, and 11 years

later, in 1786. went back to the Cape where he remained

for more than nine further years. While it appears that he

did not travel as extensively during this second visit, the

extended sojourn clearly gave Masson the opportunity to

plan seed-collecting trips; unlike the ‘passing’ collector,

he could go to a particular place at different seasons,

gathering specimens in flower for making into herbarium

specimens and subsequently returning to collect fresh,

ripe seeds for introducing the particular plant into gar-

dens. By 1792, six years after his return to Cape Town,

the specially built greenhouse in the Royal Garden at

Kew contained more than 80 different Erica spp.

(Hepper 1989; Bradlow 1994). Aiton (1811) attributed a

further 63 Erica spp. to Masson, collected and intro-

duced between 1787 and 1795, some through James

Lee’s Hammersmith nursery (Willson 1961), making

Masson’s total around 86.

In a contemporary commentary on the influx of Cape

heaths due to Masson’s explorations, Thomas Marty n, pro-

fessor of botany at the University of Cambridge, in his edi-

130

Bothalia 34,2 (2004)

tion of Philip Miller’s The gardener’s dictionary (Martyn

‘1807’) observed that Miller ‘so late as the year 1768 ...

[had] not [listed] one of those beautiful natives of the Cape

of Good Hope, which now form so great an ornament to

our green-houses or dry stoves’. In contrast, Martyn

(‘1807’) listed 83 species, and noted that James Donn,

curator of the Cambridge Botanic Garden, ‘has collected

upwards of sixty of these Cape heaths, which he cultivates

this year 1795; though he was appointed to his office only

at Michaelmas 1794'. Donn’s own Hortus cantabrigiensis

(1796) accounts for 59 named Cape heaths.

Other collectors were active for private or commercial

patrons. William Paterson, sent to the Cape in 1777 by the

Countess of Strathmore to collect for her garden, does not

feature in Hortus kewensis. He remained at the Cape until

1780, making four plant-collecting trips, and sent seeds

back to various contacts in England including James Lee

(Willson 1961). While the introduction of no Erica spp. is

attributed to him, Paterson was commemorated in E. pater-

sonii Andrews. (Contrary to Coats’s footnote (1969: 257),

William Paterson was indeed the same person who was

afterwards governor of New South Wales.)

Emperor Joseph II of Austria sent two gardeners,

Georg Scholl and Franz Boos, to the southern hemi-

sphere to collect for the royal gardens in Vienna. They

met Francis Masson at the Cape and went on a brief col-

lecting trip with him. Boos left in February 1787 for

Mauritius, leaving Scholl at the Cape to continue col-

lecting. Boos returned to the Cape briefly a year later and

then travelled back to Austria with a large collection of

specimens, apparently including living plants. However,

once again Scholl remained behind: indeed, he was

‘obliged to remain ... contrary to his orders’ (see Bradlow

1994) in Cape Town until 1799 because he was unable to

get a passage on a ship that would also carry his living

plants. For a while Scholl lived under the protection of

Colonel Robert Jacob Gordon (1743-1795), in whose

garden he cared for the plants he had collected. From

these he harvested seeds which were sent to Vienna

(Gunn & Codd 1981: 170, 171). When he did return to

Vienna in 1799, Scholl evidently also had a substantial

consignment of live plants including Erica spp.

(Hitchcock 2003; Oliver 2003), a most remarkable

achievement because, as far as is known, all other eigh-

teenth and early nineteenth century introductions were

effected by seed. Scholl’s plants do not appear to have

had as much horticultural impact as Masson’s or Niven’s

(see below), and no new species were described from his

or Boos’ herbarium specimens (Dulfer 1965). Scholl

died in Vienna soon after his return from the Cape and

was commemorated by the English nurseryman Conrad

Loddiges (1804, as E. schollii) in E. schoiliana:

‘This name has been given in memory of our friend the late

George Scholl, sent by the Emperor Joseph II to collect plants

in the vicinity of the Cape of Good Hope. He sent home great

quantities, and in 1799, after remaining twelve years in South

Africa, returned with a large cargo to Vienna, where a few years

afterwards he died [Loddiges 1821].’

(Regel (1842) regarded E. schoiliana (as E. schollii)

as a putative hybrid between E. empetrina L. (as E.

empetrifolia L.) and E. plumosa Thunb., whereas Dulfer

(1965: 101) placed E. schoiliana in synonymy under E.

amoena Wendl.)

As far as introduction of Cape heaths into cultivation

in England is concerned, the next, and most prolific col-

lector represented at Kew before 1811 was James Niven

(± 1774-1827) who collected in the Cape region between

1798 and 1812 (Nelson & Rourke 1993), returning

briefly to England in 1803. His first patron was George

Hibbert, but after 1803 he collected for a consortium in-

cluding the Empress Josephine of France and the London

nursery of Messrs Lee and Kennedy. Aiton (1811) attrib-

uted 31 Erica spp. to Niven, dating the introductions

between 1799 and 1804. His introductions were un-

doubtedly more numerous but accurate data are not

available because most subsequent publications do not

attribute individual species to particular collectors (see

Appendix 1 for a list of Erica spp. introduced by Messrs

Lee & Kennedy). Niven collected thousands of herbari-

um specimens — his Erica series bear numbers up to 297

but there are many instances of individual numbers being

repeated (for instance, seven sheets numbered 21 are

known, and these represent three distinct species from

three separate localities). Of course he would also have

collected species that had already been introduced by

Masson. An indication of the extent of his seed gather-

ings can be obtained by analysing his so-called Erica

manuscript (now in the Royal Botanic Gardens, Kew)

which is, in fact, an hortus siccus that he compiled, pre-

sumably to enable him to link herbarium specimens gath-

ered from plants in bloom with seeds gathered weeks or

months later. The extant manuscript, which relates to his

earlier visit to the Cape, is incomplete yet contains about

250 descriptions of heaths; 98 of the entries include a

note about seed, usually indicating the month or months

when this was ripe. Occasionally Niven noted that seed

was not obtained: under ‘ Erica smithiana' he recorded

that seed was ‘near ripe or gone April’ [f. 59r]; of anoth-

er unnamed species he recorded, ‘Good seed is scarcely

to be had in the winter months’ [f. 41r]; and while seed of

‘ Erica thunbergii ’ was ripe in December, Niven wrote

‘not got’ in the manuscript [f. 59'].

Few subsequent collectors matched Masson’s and

Niven’s combination of gathering both herbarium spec-

imens and fresh seed. Aiton (1811) credited John

Roxburgh (± 1777- ± 1817) with the introduction of two

Erica spp. There is no doubt that he knew Niven and

worked alongside him when both men were at the Cape

between 1798 and 1804. Young Roxburgh had arrived

early in 1798, accompanied by his father, William

(1751-1815) who remained at the Cape only until

October 1799 (Gunn & Codd 1981: 302, 303). At least

one of the Roxburghs provided names for new Erica

spp.; Salisbury (1802) credited John with 20 manuscript

names, while Niven’s herbarium specimens bear at least

30 other names attributed simply to ‘Roxburgh’.

Niven’s Erica manuscript also contains references to

names coined by Roxburgh, but it is not now possible to

determine unequivocally whether this was father or

son — William is the more likely person to have coined

names.

Salisbury ( 1 802) also credited Jacob (? James) Mulder,

about whom nothing more is known, with collecting

Cape heaths, but Mulder seems only to have gathered

herbarium specimens.

Bothalia 34,2 (2004)

131

By the end of the 1810s, the flow of Erica seeds into

Europe seems to have virtually ceased. Following

Niven’s return to Britain in 1812, there was apparently

no-one actively seeking new Cape heaths for introduc-

tion, with one enigmatic exception. Andrews (post

1823), when describing E. baueri, recorded that “We

found this new species ... in bloom at the Hammersmith

nursery in the autumn of 1823, under the specific title of

Bauera [sic] ... Throughout this extensive Genus of

plants, we do not know any one it resembles at present:

but in three or four years time the case may be different,

Mr Lee having a collector now in the interior of the Cape

in search of novelties; and should his attempt in explor-

ing that botanic mine prove successful, we may expect

an importation of many distinct species and beautiful

varieties’.

The name of the collector has not been determined

beyond doubt, nor what he may have introduced, if any-

thing. It is possible that the individual was James Bowie

whose interest in Cape heaths is well demonstrated by an

article, written by him (Bowie 1826), relating his obser-

vations on the plants in their natural habitats to their cul-

tivation in English gardens. Bowie returned to the Cape

“on his own account' (Coats 1969: 264: Gunn & Codd

1981: 101) in 1827 — there is no explicit statement about

him collecting for Lee. but it is not at all unlikely that

Lee was among the clients Bowie may have secured.

Undoubtedly one reason for the falling off in intro-

ductions was the cost of sending, equipping and main-

taining a collector at the Cape of Good Hope for not less

than a year. There was, however, a much less expensive

way of ensuring a supply of beautiful new heaths — arti-

ficial hybridization.

‘SILENTLY AND SUCCESSFULLY’: THE DELIBERATE

HYBRIDIZATION OF CAPE ERICA

Hitherto there has been no written account of the history

of deliberate hybridization of Erica spp. in European

gardens, although William Rollisson’s work (see below)

on Cape heaths was the first ‘extensive program of

breeding new ornamental plant varieties’ (Elliott 2001;

see also Elliott 2004: 237). Gorer (1978: 64), unaware of

Rollisson’s successful pioneering attempts at cross-polli-

nation, was, like others, misled and wrote:

‘It is possible that William Rollisson, who was later to

found a nursery that specialised in Cape Heaths, travelled there

[Cape of Good Hope] at the start of his career. Hortus Kewensis

[Aiton 1811] credits him with the introduction of fifteen

species between 1796 and 1800 and if he did not collect them

himself it is not clear how he could have obtained them.’

In fact, the list of Erica spp. credited to Rollisson by

Aiton (1811) included 16 names, and all are now gener-

ally acknowledged as applying to hybrids, and it is clear

that Rollisson never ventured to the Cape, nor did he

maintain a collector there. Gorer (1978: 138) did note

Herbert’s work on raising hybrids of Gladiolus and of

Erica but did not expand on the latter. While both

Bentham (1839) and Regel (1842) make it clear that they

considered certain ‘species’ were hybrids, the origins of

many have remained vague or entirely unknown.

William Rollisson of Tooting

The most prominent name in nineteenth-century

accounts of European-raised Erica hybrids is that of

William Rollisson (± 1765-1842), founder of the Spring-

field Nursery, Upper Tooting, Surrey, which was famous for

its orchids. (Rollisson’s surname was, and still is, very fre-

quently misspelled Rollinson.) Undoubtedly, he was the

first to raise artificially created hybrids of the Cape heaths

but, for commercial reasons, he never announced his

achievement. His priority is made clear by two sources.

Firstly, by a short note written by his eldest son George (±

1799-1879) and published in The Gardeners' Chronicle on

8 July 1 843 in which were listed the names of more than 90

hybrids raised by William Rollisson (Rollisson 1843). The

second source, which contains the most remarkable

account of early attempts to cross-breed Cape Erica spp.,

was written by W.H. Story (1848) who was himself an

important hybridizer, although he is an obscure person who

is not given an entry in Desmond & Ellwood (1994). Story

wrote:

‘It is now about fourteen years [i.e. ± 1834] since I became

imbued with a taste for this interesting pursuit; imparted, no

doubt, by the frequent instructive conversations I used to have

with my friend the late Mr Rollisson, of Tooting, than whom a

more persevering hybridiser of the Erica never lived. For forty

years and upwards he silently and successfully carried on his

favourite pursuit, introducing, during that long period, most (I

was going to say all) of the choicest and most favoured vari-

eties now in cultivation.’

In 1826, two lists of Messrs Rollisson’s heaths were

published in The Gardener's Magazine (Rollisson 1826a,

b); the latter was a month-by-month account of those in

bloom. These lists included 288 names. No hybrids are

mentioned, nor is any explicitly indicated, unless the

plant called ‘Rollisson's Blanda’ is counted. Rollisson’s

second list was followed by James Bowie’s (1826) arti-

cle about the cultivation of Cape heaths.

The Hon. and Very Rev. William Herbert

Whereas Rollisson remained ‘silent’ , The Hon. and Very

Rev. William Herbert (1778-1847), one-time dean of

Manchester, an authority on Amaryllidaceae and Iridaceae,

was one of the first to write about ‘hybridization amongst

vegetables’ (Herbert 1847; Steam 1952). In a paper (Her-

bert 1847: 87) written shortly before his death there is a lit-

tle-known paragraph about Cape Erica hybrids:

‘We learn that most of the fine heaths of South Africa are very

local. About thirty years ago I announced that I had crossed E.

vestita coccinea with jasminiflora (though Mr Salisbury fancied

they were of two separate genera, on account of the shape of the

seed-pods); and it is now ascertained that Mr. Rollisson, of

Tooting, raised E. jasminiflora by mule impregnation between E.

Aytoni [sic] and ampullacea, and several others, of which no wild

specimens have been found, and kept his secret until his death for

the sake of profit. The genus Erica not yielding its pollen till the

anthers are forcibly touched, and having the stigma therefore

extremely likely to be hybridized in a wild state, there seems rea-

son to believe that the species have been multiplied on the African

wastes not merely by the variation of soil and position, but still fur-

ther by the intermixture of the various forms which had so arisen.'

Herbert first announced his work on Erica when he

spoke about hybrids at a meeting of the Horticultural

132

Bothalia 34,2 (2004)

Society of London on 7 July 1818. The published version

of this paper (Herbert 1818: 196) concluded with this

scanty comment: ‘and the new heaths I have already

obtained, are most distinct and remarkable, the individu-

als of each new species [sic] being perfectly uniform’.

However, in his monograph on Amaryllidaceae, Herbert

(1837) was more explicit: ‘Several most beautiful mule

Gladioli and Ericae, which had been raised at Mitcham

between the years 1808 and 1814, and removed from

thence to Spofforth, have also flowered there, but had not

been made known to the public till the year 1819 [sic]’

(Herbert 1837: 356). (Herbert was appointed Rector of

Spofforth, Yorkshire, in 1814.)

Remarkably, while the names given to many of Rol-

lisson’s hybrids are known — there are more than 170

entries "in The Heather Society’s database— not a single

record of any of Herbert’s hybrids has been traced and

there is no Erica bearing his name. The explanation lies

in the fact (see below) that Herbert lost some (if not all)

of his Cape heath hybrids when he moved from Mitcham

to Spofforth.

Herbert (1837: 374) made some remarkable com-

ments about pollination biology in Cape species of Erica

and drew some equally remarkable conclusions:

‘In the tubular African heaths the pollen remains confined,

unless the anthers are touched by something inserted, as the

point of a pin or the proboscis of an insect, when they spring

asunder and discharge it. This genus, therefore, affords greater

facility of intermixing, and it is probable that some of the native

species, which are said to be quite local, have been produced by

accidental intermixing ... Amongst other crosses of Ericae, I

obtained at Mitcham many plants from two very dissimilar,

namely, from Jasminiflora by vestita coccinea, which had the

foliage slender and near an inch long. The late Mr. Salisbury

had conceived that those two species, being distinguished by a

shorter and a large and more pointed pod, were referable to two

distinct genera to which he had accordingly assigned names,

and he told me that I should fail in my attempts to cross them;

which was answered by shewing him the seedlings then sever-

al inches high. They were all lost on, or soon after, removing to

Spofforth before they had flowered, though one of them was

above a foot high.’

Not everyone approved of Herbert’s work: ‘Soon after

the publication of [my] communication to the

[Horticultural] Society, I was accosted by more than one

botanist with the words, ‘I do not thank you for your

mules,’ and other expressions of like import, under an

impression that the intermixture of species which had

been commenced, and was earnestly recommended to

cultivators, would confuse the labours of botanists, and

force them to work their way through a wilderness of

uncertainty’ (Herbert 1837: 336).

The facility with which hybrids could be created by

artificial cross-pollination of Cape Erica spp. was evi-

dently widely appreciated in the first half of the nine-

teenth century. Regarding the plethora of hybrids, a

unidentified reviewer using the initials E.M., writing in

The Gardeners’ Chronicle on 14 January 1843 about a

recently published monograph (Regel 1842) on heaths,

commented:

‘The number [of species] enumerated is 335, besides vari-

eties, but Mr. Regel includes as species many of the garden

hybrids, which are now multiplied almost indefinitely, so as to

render almost fruitless any attempt at describing them on paper.

Of two hybrid seedlings raised from the same parents, one will

often have more of the character of one of the parents than the

other; and thus a complete description of all the hybrids in cul-

tivation would have to include a separate account of almost

every individual raised. It appears to us that it would have been

better, even for garden purposes, to have enumerated as species

only such permanent ones as are really so in a botanical sense;

and after each species to have referred to the hybrids which

either are known to have that species as one of their parents, or

to have such an affinity to it to make that probable.’

Thus we have to view the population of Cape heaths

in cultivation in Europe during the last decade of the

eighteenth and first half of the nineteenth centuries as

comprising a vast assemblage of wild-collected, seed-

raised species, multifarious seedlings that were primary

hybrids, and, undoubtedly, innumerable backcrosses.

The concomitant variation presented new challenges,

especially in identification and naming.

Other hybridizers

Other British nineteenth-century nurserymen and garden-

ers who raised, or claimed to have created, hybrid Cape

heaths were William McNab (1780-1848), author of the first

English-language manual on the cultivation of Cape heaths

(McNab 1832), and superintendent of Edinburgh Botanic

Garden (E. xmacnabiana, E. xneillii)\ Andrew Turnbull

(1804-1886), gardener at Bothwell Castle, Lanarkshire,

Scotland, whose hybrids included E. xtortuliflora and sever-

al others which were given ‘fancy’ (i.e. cultivar) names;

W.H. Story, the originator of E. xspenceriana, who was

quoted above and about whom so little is recorded; and

Messrs Lee and Kennedy, Vineyard Nurseries, Hammer-

smith, whose introductions include at least two putative

hybrids E. xbandonii and E. xcoventryi. Among the others,

we single out for mention John Williams, gardener to John

Willmore of Oldford, near Birmingham, who raised E.

xpseiidovestita , E. xpyriformis and E. xwillmorei (a much

confused and misapplied name; see Nelson & Oliver 2003).

Maund & Henslow (1839), noting that E. xpseiidovestita

was ‘one amongst many hundred varieties raised by the

same indefatigable cultivator’, ventured the opinion that

‘generating, as it may be termed, numerous attractive plants,

like that which we now figure, may be counted amongst the

benefits accruing to society by the united zeal of a liberal

proprietor and intelligent gardener.’ When an illustration of

E. xmurrayana, one of Turnbull’s crosses, was published,

Joseph Paxton (1844) remarked that ‘The practice of hybri-

dising Heaths has now been carried to a considerable extent

by some cultivators, and our pages [Magazine of botany]

have contained figures of several excellent seedlings. But no

one will assert that there is not an ample field for further

improvement in this way, since the genus is so very large,

and the species so varied’.

There can be little doubt, for comments by various

authors including Regel (1842), that hybrids were also

produced in gardens in France and Germany but detailed

histories are wanting.

While the principal purpose in creating the hybrids

was to ensure a continuing supply of novelties for cus-

tomers, towards the end of the nineteenth century, nurs-

erymen were considering the possibility of deliberately

Bothalia 34,2 (2004)

133

breeding better plants. One of the aims of the breeders of

Cape heaths was to produce flowers that were not cov-

ered by gum because the sticky flowers trapped dust and

became unsightly; for example, among heaths listed in

the horticultural press during 1878 was a ‘gumless’

seedling named E. xdennisoniana ( Journal of horticul-

ture and cottage gardening 60: 91).

By the late 1870s, the number of distinct Cape heaths

in collections had diminished very substantially: in 1874

Joseph Dalton Hooker wrote, ‘Many years ago the Cape

Heaths formed a conspicuous feature in the greenhouses

of our grandfathers, and in the illustrated horticultural

works of the day .... These have given place to the cul-

ture of soft-wooded plants— Geraniums, Calceolarias,

Fuchsias, &c.; and the best collections of the present day

are mere ghosts of the once glorious Ericeta of Woburn,

Edinburgh, Glasgow, and Kew. A vast number of the

species have indeed fallen out of cultivation ... .No less

than 186 species of Erica were cultivated [at Kew] in

1811, now we have not above 50, together with many

hybrids and varieties [Hooker 1874]'.

Instead, nurserymen, at least those around London,

were concentrating on mass production of a few selected

Cape heaths for the winter pot-flower market. Hooker

(1874) remarked that ‘a few easily propagated hybrids

for decorative purposes are all that are to be seen of this

lovely tribe in most of the best establishments of

England', while a report in The garden early in 1879

noted that growers were known to sell more than two

hundred thousand Cape heaths in the pre-Christmas peri-

od—‘About Christmas time the markets are crammed

with little bushy plants’ (Anon. 1879). The writer com-

mented that in some English nurseries Cape heaths were

growing as ‘freely as scarlet pelargonium’. A little more

than a decade later. Watson (1892) made similar remarks.

Hybrids between European (‘hardy’ ) and Cape heaths

Having so easily achieved hybrids between the exotic

Cape Erica spp., it was not unnatural for European hor-

ticulturists to speculate about the possibilities of crossing

these with the less showy but frost-hardy European

species. The idea was certainly current in the mid- 1800s

as shown by this comment about ‘crossing splendid

species of South Africa with the hardy natives’ in The

Gardeners' Chronicle of 17 March 1849: ‘I need not say

that a successful result would be most interesting’ (‘A

Devonian’ 1849; see also ‘A Lancastrian’ 1849). George

Gordon (1863) wrote on the same topic in 1863 without

adding anything new.

In fact, there is no evidence that crosses between the

Cape and European species were tried, let alone effected,

during the nineteenth century. Only in the late twentieth

century was this achieved and primary hybrids were

announced between E. arborea L. and E. baccans L. (E.

xafroeuropaea D.C.McClint.) (Jones 1988; McClintock

1998, 1999). Other reported examples are (the African

species being named first) E. bergiana L. x spiculifolia

Sm. [McClintock 1998; at least two clones of this are

presently cultivated in Europe and western North

America (Kay 2003: fig. 2)], E. curvirostris x arborea ,

E. baueri Andrews [as bauera \ x australis L., and E. sub-

divaricata PJ.Bergius x carnea L. (Jones 1988). In the

1970s John Crewe-Brown, in Johannesburg, attempted

various crosses between European and African species

including E. caffra L. x australis (in Joyner 1979).

PROLIFERATION OF EPONYMS & ORTHOGRAPHIC BEDLAM

In the decades immediately following publication of

Species plantarum (Linnaeus 1753), the choice of names

for the new Erica spp. was uncomplicated. A principal

character, or the superficial resemblance to plants of

another familiar genus, or a toponym provided the

majority of the epithets. Only one species was named

after an individual and that was Linnaeus’s E. plukenetii ,

commemorating Dr Leonard Plukenet who, as noted, had

illustrated the species in 1700.

By the end of the 1770s, six more individuals had

been commemorated by Linnaeus, by his son (Linnaeus

‘1781’) and by L.J. Montin (1774): PJ. Bergius in E.

bergiana ; James Petiver in E. petiverr, Anders Sparrman

in E. sparrmanif Carl Thunberg in E. thunbergii.

Linnaeus the younger also encapsulated Francis Mas-

son’s name in E. massonii, and named E. monsoniana

after Lady Ann Monson; both these epithets were pub-

lished in 1782 (Linnaeus ‘1781’). Still, the majority of

specific epithets were related to a distinctive characteris-

tic of the new species.

The sudden and continuing influx after 1772 of sub-

stantial numbers of new Erica spp. presented a major

problem for European botanists. Studying, describing

and naming the countless new plants that Masson and

Niven, in particular, had collected, was evidently beyond

the capacities of the botanists then active at least in

Britain — and the Cape of Good Hope was not the only

place yielding undreamt of botanical riches. Com-

pounding the difficulties for the botanists were the very

numerous seedlings of unknown species of Erica that

were sprouting in gardens throughout Britain, France,

Germany and other countries. Nurserymen who had

obtained seeds or, subsequently, seedlings required

names for these so that they could sell the plants. The

botanists were either overwhelmed or indifferent, and so

the naming of the Cape heaths became a haphazard

process with horticulturists, ill-versed in, or entirely

ignorant of, the principles of botanical nomenclature,

taking over the primary role of coining and applying

names. Not only did the nurserymen give names to their

plants, they also printed those names in catalogues —

their businesses demanded this — with accompanying

descriptions, no matter how brief, and not infrequently

the given names were misspelled or misprinted. The

results approached bedlam with as many as nine differ-

ent variants of a single epithet (as prime example is E.

savileae Andrews, see Nelson & Scarbrough 2003).

It is clear from manuscripts and published catalogues

dating from the first decade of the 1800s that many Cape

heaths had been given names in gardens and nurseries

without the normal accompaniment of detailed, compar-

ative study. It is also evident that collectors, including

Francis Masson (1776; see Bradlow 1994), James Niven

and the Roxburghs, coined names in the field and used

them to label their pressed specimens and seed packets.

134

Bothalia 34,2 (2004)

Many of these names persisted and were published, usu-

ally without accompanying descriptions, in nurserymen’s

catalogues and garden lists. For example, the name

'Erica hibbertii' appears on James Niven’s herbarium

specimens (no. 171); on one of these, in the herbarium of

Trinity College Dublin (TCD), the epithet is attributed to

Niven. The particular epithet was employed at the

Dublin Society’s Botanic Gardens, Glasnevin, in 1802,

undoubtedly attached to a plant purchased from Messrs

Lee and Kennedy (Nelson & McCracken 1987), and it

appeared in print at least a year before Andrews formal-

ly published the binomial in Coloured engravings of

heaths (1805). This is not an isolated example. The name

E. humea was first printed in 1809, yet a description was

not formally published in The Botanical Cabinet

(Loddiges 1817-1833) until a decade later when the epi-

thet was rendered E. humeana. Likewise, E. scholliana

was described in 1821 , but that name, as E. schollii, was

in print no less than 17 years earlier.

There were other problems. Horticulturists in differ-

ent countries produced lists and' catalogues without

knowing whether a particular seedling Erica had already

been named elsewhere— they had neither the expertise,

nor resources, nor time to carry out the necessary en-

quiries and research. Furthermore, voucher herbarium

specimens were hardly ever preserved so that when

botanists eventually came to sort out the nomenclatural

tangle, they had no material for comparison. Add to that

an opinionated, acerbic English botanist: Richard

Anthony Salisbury (1761-1829).

Richard Salisbury

To his credit, Salisbury did attempt to produce a prop-

er monograph of the genus (Salisbury 1802), but he had

some contrary opinions about names, rejecting any that

commemorated individuals, and when in his opinion a

descriptive epithet was not appropriate he simply substi-

tuted one that he deemed better.

In Rees’ The Cyclopaedia, the president of the

Linnean Society, James Edward Smith (1809), in what

amounts to a precis of Salisbury’s monograph, applaud-

ed Salisbury’s attempt to arrange the species in natural

subdivisions. In fact, Salisbury had a remarkable, pre-

scient view of Erica, as Smith acknowledged. Salisbury

(1802) had rejected Linnaeus’ genus Blaeria, and clus-

tered together 15 species ‘having but four stamens’,

some of which had formerly been in Blaeria :

‘Nor can we much object to the union of these two genera,

except that in so vast a tribe, we may be glad of even so slight a cir-

cumstance as number to make a genus, when there is moreover

such a difference in habit as this very arrangement of Mr.

Salisbury’s implies. If indeed a few species of Erica, here and there

in the different natural subdivisions, were tetrandrous, no-body

would think of separating them for that reason [Smith 1809].’

However, Smith was not impressed by some of

Salisbury’s nomenclatural decisions. Regarding E. nigri-

ta ‘of Linn|aeus] and Thunberg, so called from its con-

spicuous dark anthers, elegantly contrasted with the

white corolla and calyx’, Smith noted that Salisbury ‘is

pleased to denominate | this ] volutceflora' . The name of

‘our elegant English’ E. tetralix L. was ‘changed by Mr.

Salisbury to botuliformis , sausage-shaped. ... Next comes

by itself our common British cinerea [L.] ... more happi-

ly, but without any necessity, altered to mutabilis'.

However, E. milleflora Bergius was, in Smith’s view,

‘well retained by Mr. Salisbury, in preference to the inac-

curate on e, paniculata, given by Linnaeus’.

Another to object to Salisbury’s alteration of estab-

lished names was Carl Willdenow (1809: 415), in a cata-

logue of the Royal Botanic Gardens, Berlin. Noting that

he was deliberately omitting ‘several species [of Erica]

grown from seeds, the flowers of which I have not yet

seen’, he reproached Salisbury thus:

‘The most illustrious Salisbury ... illustrated the genus of

Erica with many observations and, using this occasion,

reproached me with having accepted inappropriate and false gar-

deners’ names. Names of plants [that are] now widespread, just

as [those] applied long since, are to be scrupulously preserved. In

the same way, whatever the naming of a plant is, it suffices if

only its character can have fixed [distinguishing] marks. The

changes which the illustrious Salisbury has adopted in his

Prodromus [Salisbury 1796], in this dissertation [Salisbury

1 802] and in other places always lead to the detriment of science.

Only under pressing necessity have I altered plants' names to

others’. [Translated from the original Latin by P. H. Oswald.]

Eponyms

Returning to the matter of specific epithets derived

from persona] names, at this period there were no rules

governing the formation of such names. Smith (1809)

made this most pointed comment at the end of his entry

on Erica for Rees’ The Cyclopaedia :

‘We trust, that those who may have occasion to describe new

heaths in future, will, as far as they are competent, keep the

above arrangement in view, as they will find it very instructive,

however they may differ from Mr. Salisbury in principles of

nomenclature. To this subject we have scarcely found it neces-

sary to advert, except occasionally, and we rather pass over in

silence what we cannot approve. This botanist rejects all names

of persons as applied to the species of any genus. Mr. Dryander

once began a more useful reformation of such names of Erica;,

making those which commemorate a writer on the subject end

in ana, as Sebana\ those which apply to a collector only, in the

genitive case, as Massoni. The greatest and most correct infor-

mation is still to be expected from this able botanist, in the

intended new edition of Mr. Aiton’s Hortus Kewensis , where the

genus of Erica must always make a principal figure.’

The topic of the correct termination for epithets de-

rived from personal names has been vexatious ever

since. Lindley (1832) commented on it and, as Steam

(1973) pointed out, several attempts have been made to

differentiate the use of substantive and adjectival (with

the termination -anus, -a, -urn) epithets along the lines

enunciated by Dryander and Lindley, but they have never

been successful. Commenting on Lindley ’s remarks,

Steam (1973: 294) noted that ‘most of those who then

and thereafter named new species paid no attention what-

ever to it; probably they never knew such a distinction

had been proposed’.

In the second edition of Hortus kewensis the princi-

ples described by Smith ( 1809) were followed, with, for

example, the Erica epithets archeria amended to

archeriana, hibbertia to hibbertiana, lambertia to lam-

bent iana, and even monsoniana to monsoniae. Yet this

Bothalia 34,2 (2004)

135

did not, and does not, resolve the difficulties— under the

current rules of nomenclature [International Code of

Botanical Nomenclature, (St Louis Code): Greuter et al.

2000], it is not permissible to change the termination of

an epithet except to correct certain errors (for example, if

the gender is inappropriate); misinterpretation of the role

of the person commemorated is not deemed to be an

error. Changing archeria to archeriana is forbidden but

amending the termination to correct the gender is

required— archerae is the correct orthography.

Examining the earliest use of personal names for epi-

thets within Erica (Appendix 2), it is clear that their

authors (Linnaeus father and son, and Montin) were

abiding by this, then unwritten, distinction: petiveri,

sparrmanii and thunbergii after Petiver (author),

Sparrman (collector) and Thunberg (collector), whereas

bergiana and monsoniana were in compliment to

Bergius and Lady Monson. It was the advent of essen-

tially horticultural publications that precipitated a large

increase in personal epithets, and the ensuing bedlam.

The abundance of unnamed Erica spp. following espe-

cially Masson’s and Niven's plant introductions from the

Cape provided many opportunities to coin names. While

the majority of new species were given names that encap-

sulated a distinctive morphological character, the need for

horticulturists especially to recognize patronage also led

to a proliferation of personal epithets.

Henry Charles Andrews issued the first fascicle of his

monumental four-volume Coloured engravings of heaths

on 15 October 1794 (Cleevely & Oliver 2002: 264); he

was later to issue a six-volume edition, a 'green-house

companion', titled The heathery , starting in 1805

(Cleevely et al. 2003). In the first 17 fascicles of Coloured

engravings of heaths, all published before 1800, he used

the following names (their original orthography being

retained here) to dedicate species to William Paterson (E.

pattersonia) in 1795, William Aiton ( E . aitonia) in 1796,

Sir Joseph Banks (E. banksia ) and James Lee (E. leea) in

1797, and James Walker (E. walkeria ) in 1799. By 1830

when he had completed publication, Andrews had added

yet further personal epithets (see Appendix 2). In all,

Andrews (1794—1830) commemorated in the names of

African Erica spp. more than two dozen individuals few

of whom had any connection with the Cape of Good

Hope. Towards the conclusion of Coloured engravings of

heaths, Andrews (± 1823) himself commented, ‘In the

genus Erica it is sometimes very difficult to find an appro-

priate or un-occupied specific title’.

Other horticulturists joined suit. Conrad Loddiges, to

give just one example, published the following personal

epithets within Erica in The Botanical Cabinet (Loddi-

ges 1817-1833): E. bonplandia (1819); E. bowieana

(1824); E. celsiana (1830); E. cliff ordiana (1817); E.

humeana (1819); E. parmentierii (1818); and E. schol-

liana (1804).

One frequent feature of the names deriving from horti-

cultural works published at the end of the eighteenth cen-

tury and during the first half of the nineteenth century is

the use of eponyms with the termination -ia (or -a). In the

past this has been interpreted as indicating that the author

was using a pre-existing generic name in apposition to the

name Erica (Oliver & Oliver 2002; 43). For example, it

might be thought that in coining E. banksia, Andrews was

employing the younger Linnaeus’s generic name Banksia

(Proteaceae), or in the case of E. hibbertia that he was

using the generic name Hibbertia (Dilleniaceae). Whereas

the generic names Aitonia, Banksia, Bauera, Hibbertia

and others (see Table 1) already existed when Andrews

published E. aitonia, E. banksia, E. bauera and E. hibber-

tia, there have never been genera named, for example,

Bandonia, Coventrya, Savileia, Shannonea, Templea or

Uhria. Examination of the chronology of other names

makes clear that they also are not generic names used in

apposition but are substantival epithets with incorrect ter-

minations. Archeria and Nivenia were coined after the

publication of these epithets for species of Erica—

Ventenat published Nivenia (Iridaceae) in 1808 (Ventenat

1808); Hooker published Archeria (Epacridaceae) in 1860

(Hooker 1860).

Under the present International Code of Botanical

Nomenclature (St Louis Code) (Greuter et al. 2000),

such names have to be corrected— they have incorrect

terminations. The -ia has to be replaced with the appro-

priate genitive ending. This means that it is essential to

establish the gender of the person who was commemo-

rated. The variants’ spellings can be treated as ortho-

graphical variants because only one nomenclatural type

is involved , and under Article 6 1 (Greuter et al. 2000 ) the

one variant ‘that conforms to the rules’ needs to be speci-

fied and retained. Oliver & Oliver (2002) have corrected

the terminations for E. banksii, E. baueri and E. hibber-

tii, and the remaining names are corrected in Appendix 2.

CONCLUSION

Erica is one of the largest genera among flowering

plants containing ± 860 species; about 760 of these are

endemic in southern Africa (Oliver & Oliver 2003), ± 50

are recorded in Madagascar and adjacent islands, while

another 30 Erica species inhabit Africa north of the

Zambezi (excluding the extreme northwest) (see Oliver

2000: fig. 15). Around 20 species, plus three naturally

occurring hybrids, are recognized in Europe, Macaro-

nesia and adjacent parts of northwest Africa, and western

Asia.

Given these facts alone, it is not surprising that during

the past 250 years, since the publication of Species plan-

tarum (Linnaeus 1753), many more than 860 names have

been bestowed on Erica species. The database for the

‘African’ (‘non-hardy’) section of the International reg-

ister of heather names (Nelson & Small 2004) dramati-

cally demonstrates the nomenclatural bedlam to which

we have referred. While the total of all names reported

on the International Plant Names Index website

(http://www.ipni.org) for Erica (as redefined in Oliver

2000) and its component genera is around 3 350, the

database compiled by The Heather Society now includes

more than 6 100 entries for names. More than two-thirds

of these (± 4 200 = 69%) are records comprising only a

specific epithet, whereas ± 1 650 record names of taxa at

the rank of subspecies or below (27%). Only ±310 cul-

tivar names are in this part of the database (5%).

136

Bothalia 34,2 (2004)

TABLE 1 .—Eponymous specific epithets in Erica', epithets with the termination -ia or -a (see Appendix 2 for

details) which correspond with published generic names giving the respective dates of publication

In stark contract, the ‘hardy’ section of the database

contains just 175 records (7%) comprising only a specif-

ic epithet in Erica , whereas more than 1 800 cultivar

names (77%) are included (Nelson & Small 2000).

We cannot estimate the exact number of artificial

hybrids that were named during the nineteenth century

but it is incontrovertible that from the 1790s until at

least the 1840s, some European horticulturists were

adept at artificial pollination of Cape heaths and raising

seedlings of hybrid origin. It is also clear that hybrids

were often surreptitiously introduced into collections

without their origins being recorded. Almost 1 600

names at the rank of variety are on record for African

Erica species, and the vast majority of these were names

given to plants raised in gardens — in this context the

term variety is equivalent, in the majority of instances,

to the modem term cultivar.

ACKNOWLEDGEMENTS

This paper arose from work carried out by the first

author for the second volume of the International regis-

ter of heather names which contains data on African

species and their cultivars (Nelson & Small 2004). The

research work for the register was funded jointly by The

Heather Society and the Stanley Smith Horticultural

Trust. Our thanks are due to Philip Oswald for translat-

ing crucial Latin texts.

REFERENCES

AITON, W. 1789. Hortus kewensis. Nicol, London.

AITON, W.T. 1811. Hortus kewensis , edn 2. Longman, Hurst. Reeves,

London.

ANDREWS, H.C. 1794- ± 1828. Coloured engravings of heaths. 4

volumes. The author, London.

ANDREWS, H.C. [post 1823]. Erica bauera [.s/'c ] . Coloured engrav-

ings of heaths, vol. 4: [t. 221].

ANDREWS, H.C. [± 1823]. Erica peltata. Coloured engravings of

heaths, vol. 4: [t. 254].

ANON. 1879. Plant culture for market. The garden 15: 99.

BAKER, H.A. & OLIVER, E.G.H. 1968. Ericas in southern Africa.

Purnell, Cape Town & Johannesburg.

BENTHAM, G. 1839. Erica. In A.-P. de Candolle, Prodromus system-

atis naturalis regni vegetabilis, vol. 7, 2: 614-693. Treuttel &

Wiirtz, Paris.

BOWIE, J. 1826. Hints for the better cultivation of the Cape Heaths,

derived from observations of their nature, soils, and situations.

The Gardener's Magazine 1: 363-366.

BRADLOW, F.R. 1994. Francis Masson's account of three journeys at

the Cape of Good Hope 1772-1775. Tablecloth Press, Cape Town.

CLEEVELY, R.J., NELSON, E.C. & OLIVER, E.G.H. 2003. More

accurate publication dates for H.C. Andrews’ The Heathery,

particularly volumes 5 and 6. Bothalia 33: 195-198.

CLEEVELY, R.J. & OLIVER, E.G.H. 2002. A preliminary note of the

publication dates of H.C. Andrews’ Coloured engravings of

heaths (1794-1830). Archives of natural history 29: 245-264.

CLUSIUS, C. 1605. Exoticorum libri decent. Plantin, Antwerp.

CLUSIUS, C. 1611. Curae porteriores, sen plurimarum non ante cog-

nitarum, aut descriptarum stirpium. Plantin, Antwerp.

COATS, A.M. 1969. The quest for plants. A history of horticultural

explorers. Studio Vista, London.

DESMOND, R.G.C. & ELLWOOD, C. 1994. Dictionary of British and

Irish botanists and horticulturists. Revised edn. Taylor &

Bothalia 34,2 (2004)

137

Francis, & The Natural History Museum. London.

'DEVONIAN’, (A). 1849. Hybrid heaths. The Gardeners’ Chronicle ,

17 March 1849: 166.

DONN, J. 1796. Hortus cantabrigiensis, or a catalogue of plants,

indigenous and foreign cultivated in the Walkerian Botanic

Garden, Cambridge. Donn, Cambridge.

DULFER, H. 1964. Revision der stidafrikanischen Arten der Gattung

Erica L. 1 , Teil. Annalen des Naturhistorischen Museums, Wien

68: 79-147.

DULFER, H. 1965. Revision der stidafrikanischen Arten der Gattung

Erica L. 2. Teil Forsetzung von Ann. Bd. 67. Annalen des

Naturhistorischen Museums, Wien 68: 25-177.

ELLIOTT, W.B. 2001 . Flora, an illustrated history of the garden flower.

Scriptum-Cartago, London.

ELLIOTT, W.B. 2004. The Royal Horticultural Society, a history

1804-2004 . Phillimore, Chichester.

GORDON, G. 1863. Hints for raising a hardy race of cross-bred heaths.

Journal of horticulture and cottage gardening 4 ( new ser.): 43,44.

GORER, R. 1978. The growth of gardens. Faber & Faber, London.

GREUTER. W.. McNEILL. J.. BARRIE, F.R., BURDET, H. M.. DE-

MOULIN, V., FILGUEIRA, T.S., NICOLSON, D.H., SILVA,

P.C., 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.

GUNN. M. & CODD. L.E. 1981. Botanical exploration of southern

Africa. Balkema, Cape Town.

GUTHRIE, F. & BOLUS, H. 1905. Erica. In W.T. Thistelton-Dyer,

Flora capensis 4,1: 4-315. Reeve, London.

HENREY. B. 1975. British botanical and horticultural literature be-

fore 1800. 3 volumes. Oxford University Press. London.

HEPPER, F.N. (ed.). 1989. Royal Botanic Gardens Kew: gardens for

science and pleasure. Her Majesty's Stationery Office, London.

HERBERT, W. 1818. Instructions on the treatment of the Amaryllis

longifolia ... with some observations on the production of hybrid

plants. Transactions of the Horticultural Society of London 3:

187-196.

HERBERT, W. 1837. On crosses and hybrid intermixture in vegetables.

In Amaiyllidacece: 335-380. Ridgway, London.

HERBERT, W. 1847. On hybridization amongst vegetables. Journal of

the Horticultural Society of London 2: 1-28, 81-107.

HITCHCOCK, A. 2003. Erica verticillata is brought back from the

brink of extinction. Yearbook of The Heather Society: 45-50.

HOOKER. J.D. 1860. Archeria. The botany of the Antarctic voyage of

H.M. Discovery ships Erebus and Terror in the years 1839- 1843,

vol. 3 .Flora Tasmaniae part i: 262, t. 80, 81 . Reeve, London.

HOOKER, J.D. 1874. Erica chamissonis. Curtis's Botanical Magazine

100: t. 6108.

JONES, A.W. 1988. A trip to Holland and Germany. Bulletin of The

Heather Society 4, 4: 6-8.

JOYNER. P. 1979. [South West Group news]. Bulletin of The Heather

Society 2, 17: 5.

KAY, S. 2003. Trying to create a garden in Connemara. Yearbook of

The Heather Society 2003: 11-20.

KERKHAM, A.S. 1988. Southern African botanical literature

1600-1988. South African Library, Cape Town.

'LANCASTRIAN'. (A). 1849. Hardy heaths. The Gardeners' Chronicle

31 March 1849: 197. 198.

LINDLEY, J. 1832. An introduction to botany. Longman, Rees, Orme,

Brown, Green & Longman, London.

LINNAEUS. C. 1738. Hortus cliff ortianus . [sine nom.], Amsterdam.

LINNAEUS, C. 1753. Species plantarum. Salvius, Stockholm.

LINNAEUS, C. ‘178U [1782], Supplementum plantarum. Impensis

orphanotrophei , Braunschweig.

LODDIGES, C. 1804. Catalogue of plants. Loddiges, Hackney.

LODDIGES, C. 1817-1833. The Botanical Cabinet. 20 volumes.

Loddiges, Hackney.

LODDIGES, C. 1821 . Erica scholliana . The Botanical Cabinet 6: t. 538.

E.M. (Name unknown), 1843. Review: Die Kultur und Aufzahlung der

... Eriken, by E. Regel. The Gardeners’ Chronicle 14 January

1843:24.

McCLINTOCK, D.C. 1998. Erica xkrameri: the hybrid between E.

carnea and E. spiculifolia. Yearbook of The Heather Society

1998: 26.

McCLINTOCK, D.C. 1999. Cape heaths crossed with European

species. The New Plantsman 6, 4: 207.

McNAB. W. 1832. A treatise on the propagation, cultivation, and

general treatment of Cape heaths, in a climate where they

require protection during the winter months. Thomas Clark,

Edinburgh.

MARTYN, T. ‘1807’. Erica. In The gardener's and botanist’s dictio-

nary ... by the late Philip Miller ... . Volume I, part II (Cla-I).

Rivington, London. [Although the title-page is dated 1807,

there is evidence (Nelson 2004) that portions of this part were

issued at least a decade earlier.]

MASSON, F. 1776. An account of three journeys from Cape Town into

the southern parts of Africa. Philosophical transactions of the

Royal Society 66: 268-316. [Reprinted in Bradlow (1994):

103-135.]

MAUND, B. & HENSLOW, J. S. 1839. Erica pseudovestita. The Botanist

3: t. 104.

MONTIN, L.J. 1774. Erica thunbergii. Nova acta Regiae Societatis

Scientiarum Upsalensis 2: 290-292.

NELSON. E.C. 2004. Publication dates of (early) parts of Thomas

Martyn’s edition of Philip Miller's Gardener's and botanist’s

dictionary. Society for the History of Natural History newsletter

No. 79:16, 17.

NELSON, E.C. & McCRACKEN, E.M. 1987. The brightest jewel. A

history of the National Botanic Gardens, Glasnevin, Dublin.

Boethius Press, Kilkenny.

NELSON, E.C. & OLIVER, E.G.H. 2003. Understanding Erica xw ill-

mo rei, a nineteenth century English garden hybrid. Bothalia 33:

149-154.

NELSON, E.C. & ROURKE, J.P. 1993. James Niven (1776-1827), a

Scottish botanical collector at the Cape of Good Hope: his hor-

tus siccus at the National Botanic Gardens, Glasnevin, Dublin

(DBN), and the Royal Botanic Gardens, Kew (K). Kew Bulletin

48: 663-682.

NELSON, E.C. & SCARBROUGH, Earl of. 2003. Erica savileae, the

Countess of Scarbrough's heath: a mystery resolved? Yearbook

of The Heather Society 2003: 31-36.

NELSON. E.C. & SMALL, D.J. (eds). 2000. International register of

heather names. Volume 1 [in 4 parts] Hardy cultivars and

European species. The Heather Society, Creeting St Mary.

NELSON. E.C. & SMALL, D.J. (eds). 2004. International register of

heather names. Volume 2 [in 4 parts] African species, hybrids

and cultivars. The Heather Society, Creeting St Mary.

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

species with indehiscent and partially dehiscent fruits.

Contributions from the Bolus Herbarium No. 19.

OLIVER, E.G.H. 2003. Footnote [to Hitchcock (2003)]. Yearbook of

The Heather Society 2003: 50.

OLIVER, E.G.H. & OLIVER, I.M. 2002. The genus Erica (Ericaceae)

in southern Africa. Bothalia 32: 37-61 .

OLIVER, E.G.H. & OLIVER, I.M. 2003. Ericaceae. Strelitzia 14:

424-451.

OLIVER, I. & OLIVER, E.G.H. 2000. Ericas of the Cape Peninsula.

Protea Atlas Project, National Botanical Institute, Cape Town.

PAXTON, J. 1844. Erica murrayana . Magazine of Botany 1 1 : 77, 78.

PLUKENET, L. 1700. Almagesti botanici mantissa. The author, London.

REGEL, E. 1842. Die Kultur und Aufzahlung der ... Eriken. Ver-

handlungen des Vereins zur Beforderung des Gartenbaues in

den Koniglich Preufiischen Staaten 16: 221-331.

ROLLISSON, G. 1843. Hybrid plants. The Gardeners' Chronicle 8 July:

461.

ROLLISSON, Messrs. 1826a. Ericas in flower from October 1st to

November 26th, in the Tooting Nursery. The Gardener's

Magazine 1: 88.

ROLLISSON, Messrs. 1826b. List of Cape heaths which have been in

flower in the Tooting Nursery in each month of the year. The

Gardener's Magazine 1: 366-375.

ROURKE, J. 1980. The proteas of southern Africa. Purnell, Cape Town.

SALISBURY, R.A. 1796. Prodromus stirpium in horto ad Chapel

Allerton vigentium. The author, London.

SALISBURY, R.A. 1802. Species of Erica. Transactions of the Linnean

Society 6: 316-388.

SCHUMANN, D„ KIRSTEN. G. & OLIVER, E.G.H. 1992. Ericas of

South Africa. Femwood Press, Vlaeberg.

SEBA, A. 1 734(— 1 765). Locupletissimi rerum naturalium. Jannson-

Waesberg, Amsterdam.

138

Bothalia 34,2 (2004)

SMITH. J.E. 1809. Erica. In A. Rees, The Cyclopaedia. Longman,

Hurst, Rees, Orme & Brown, London.

STEARN, W.T. 1952. William Herbert’s ‘Appendix’ and ‘Amarylli-

daceae’. Journal of the Society for the Bibliography of Natural

History 2: 375-377.

STEARN, W.T. 1973. Botanical Latin. History, grammar, syntax, ter-

minology and vocabulary , edn 2. David & Charles, Newton

Abbot.

STORY, W.H. 1848. Seedling heaths. The Florist 1848: 314-316.

THUNBERG, C.R 1785. D. D. dissertatio botanica de Erica. Edman,

Uppsala.

VENTENAT. E.R 1808. Nivenia. Decas generum novorum. Dufarb,

Paris.

WATSON, W. 1 892. Plant notes. Erica hyemalis. Garden and Forest 5:

136-138.

WIJNANDS, D.O. 1983. The botany of the Commelins. A taxonomical,

nomenclatural and historical account of the plants depicted in

the Moninckx atlas and in the four books by Jan and Caspar

Commelin on the plants in the Hortus Medicus Amstelodamen-

sis 1682-1710. Balkema, Rotterdam.

WILLDENOW, C.L. 1809. Enumeratio plantarum horti regii botanici

berolinensi. Tabema Libraria Scholae Realis, Berlin.

WILLSON, E.J. 1961. James Lee and the Vineyard Nursery Ham-

mersmith. Hammersmith Local History Group, London.

WRIGHT, S.J. & NELSON, E.C. 2004. Diana of the heathers.

Heathers ( Yearbook of The Heather Society) 1 : 42-51 .

APPENDIX 1 .—‘List of Erica’s not in Your List of Hort. Kew.’

The series of ‘Inwards Books’ recording plants received at the Royal (Botanic) Gardens, Kew, provides some fas-

cinating information about the development of the Kew collections. A letter, dated 24 October 1808 (‘Inwards Book’,

f. 259), from Messrs Lee & Kennedy of Hammersmith, London, evidently was a response to a request, probably from

William T. Aiton, for lists of plants that the nursery had introduced. While not explicit in the letter it is possible that

this list was sought by Aiton as he then would have been preparing the second edition of Hortus kewensis (1811). Lee

& Kennedy explained that ‘Our busy Season is come upon us and we are not able to accomplish the List you desire

of what we have introduced into this Country ...’. However, they did furnish this ‘... List of Erica’s not in Your List of

Hort. Kew.’ (‘Inwards Book’, ff 262, 263). [* All the names following consimilis are bracketed by these dates.]

Kalmiflora

lanuginosa

Bothalia 34,2 (2004)

139

APPENDIX 2. — Epithets in Erica published before 1835 commemorating individuals

Article 61 of the International Code of Botanical Nomenclature (St Louis) (Greuter et al. 2000) applies to ortho-

graphic variants that are based on one nomenclatural type. Article 61.2 states that ‘For the purpose of this Code,

orthographical variants are the various spelling, compounding, and inflectional forms of a name or its epithet (includ-

ing typographical errors), only one nomenclatural type being involved.’ Article 61.3 states that ‘If orthographical

variants of a name appear in the original publication, the one that conforms to the rules and best suits the recom-

mendations of Art. 60 is to be retained; otherwise the first author who, in an effectively published text ..., explicitly

adopts one of the variants and rejects the other(s) must be followed'.

The identities of the persons commemorated in the eponyms listed below may differ from those published in other

sources [e.g. Schumann et al. (1992), and miscellaneous lists published in issues of the Yearbook of The Heather

Society], The names (and aristocratic titles) given below have been determined by ECN, by careful comparison of

protologues with biographical details of the individuals concerned.

140

Bothalia 34,2 (2004)

Bothalia 34,2: 141-153(2004)

Vegetation of high-altitude fens and restio marshlands of the

Hottentots Holland Mountains, Western Cape, South Africa

EJJ. SIEBEN*t,C. BOUCHER* & L. MUCINA*

Keywords: canonical correspondence analysis. Cape Wetlands Fynbos. phytosociology, plant communities, syntaxonomy

ABSTRACT

Seepages occurring at high altitudes in the Hottentots Holland Mountains (HHM) (Western Cape Province, South Africa)

were subject to a phytosociological survey. Releve sampling method and classification procedures of the floristic-sociological

(Braun-Blanquet) approach as well as numerical data analyses (numerical classification and ordination) were used to reveal syn-

taxonomic patterns and characterize the position of the syntaxa along major environmental gradients. Nine plant communities

were recognized, three of which were classified as associations, following formal syntaxonomic and nomenclatural rules of the

floristic-sociological approach. Most of the studied mire communities were dominated by low-growing clonal restios

(Restionaceae), whereas some consisted of other types of graminoids. The most important species determining the structure (and

function) of the mire communities on sandstones of the HHM include restios Anthochortus crinalis, Chondropetalum deustum,

C. mucronatum, Elegia intermedia, E. thyrsifera, Restio subtilis, R purpurascens, cyperoids Epischoenus villosus, Ficinia argy-

ropa, grasses Ehrharta setacea subsp. setacea, Pentameris hirtiglumis as well as shrubs Berzelia sqnarrosa, CUffortia tricuspi-

data. Erica intervallaris and Grubbia rosmarinifolia. Protea lacticolor and Restio perplexus dominate a rare shale band seep-

age community. There are two major groups of communities —the fens (dominated by carpets of Anthochortus crinalis and other

low-growing species) and the restio marshlands (mosaics of low tussocks of Restio subtilis and tall Chondropetalum mucrona-

tum). The degree of soil (and water) minerotrophy was found to be the most important differentiating feature between the mire

(fen and restio marshland) communities studied. The soils in the centre of mires were found to have high contents of peat and

showed very little influence from the underlying sandstone. The

al soil derived from the sandstone or shale bedrock.

INTRODUCTION

The Cape mountain ranges are important catchments

for high-quality drinking water. A vast quantity of this

water is stored in the basement rocks before it is released

into river systems via seepages (Hewlett 1982). Seepage

is a very general term for an area where water percolates

through the upper soil layers (mostly lying over a layer

of impenetrable rock) and usually forms the source of

rivers. It includes the majority of wet slopes as well as

many mires of the Cape mountains. The quality of water

is largely determined by the soil conditions that percolat-

ing water encounters prior to its emergence at the surface

and on its way to a river, hence seepages are of great

importance to river ecosystems (Bosch et al. 1986).

However, the total surface area of seepages that influ-

ence the water quality of rivers is variable and many

seepages can be out of touch with the river system for a

long time. This was called the Variable Source Area

Concept by Hewlett (1961). After heavy rains the Source

Area of a river expands and much water, that was stored

in basement rocks or in fens for a long time, moves into

the river. During its storage in the seepage areas, water is

stained by tannins leached from decaying plant litter.

This explains the brown colour that is characteristic of

many of Western Cape rivers, particularly after heavy

rains (King & Day 1979; Dallas & Day 1993).

Some seepages located in high-altitude areas supporting

fynbos vegetation have soils rich in peat— accumulated organ-

ic material (Gore 1983)— they can be classified as mires.

* Department of Botany & Zoology, University of Stellenbosch,

Private Bag XI , 7602 Matieland, Stellenbosch, South Africa,

t author for correspondence; E-mail: e.j.j.sieben@buwa.nl

MS. received: 2001-10-15.

soils along the mire margins had a greater admixture of miner-

These are defined as wet, swampy habitats characterized by

peaty soils, regardless of the chemico-physical properties of

the peat and water captured in the peaty soils. Concepts such

as bogs and fens refer to specific types of mires (Gore 1983).

The term bog refers to the strictly ombrotrophic (rain-fed)

and usually oligotrophic mires found in places with high pre-

cipitation, whereas fens are the mires (minerotrophic or tran-

sitional) that are fed to a larger extent by water that has per-

colated through the mineral substrate. The Cape high-altitude

mires generally qualify as fens; true bogs are rare in the south-

ern hemisphere, although they do occur on some steep south-

facing slopes in Western Cape mountains. In the mountains,

mires are found at the sources of the rivers and in watersheds.

Riparian mires can be formed in places where the floodplain

is very wide and the area remains inundated long after a flood

has receded.

There are four types of seepages linked to the drainage

network of riverine systems recorded in the Cape moun-

tain ranges, namely:

1, well-drained slope seepages supporting soils of the

Femwood form (Fry 1987); this type shows a high level

of variability and can be characterized by the increased

presence of Bruniaceae; Campbell (1986) in his structur-

al classification of the Fynbos Biome, classified the veg-

etation of these seepages as Wet Ericaceous Fynbos;

2, low-altitude valley seepages characterized by high soil

water levels and peaty Champagne soils (Fry 1987);

Boucher (1978) described the Erica-Osmitopsis Seepage

Fynbos in this habitat in the Kogelberg Biosphere Reserve.

A subtype of the low-altitude valley seepages occurs on

temporarily wet sandy soils. It is dominated by Elegia

filacea (Taylor 1978);

142

Bothalia 34,2 (2004)

3, the high-altitude fens, situated at the sources of rivers:

Campbell (1986) has classified the vegetation of this

habitat as Sneeukop Azonal Restioid Fynbos and

Otterford Wet Proteoid Fynbos;

4, in order to distinguish between the different degrees of

minerotrophy in the mires described in this study, we

want to introduce the term restio marshlands for the bet-

ter-drained sites at the edges of mires of the Fynbos

Biome, in contrast to the ‘fens’ being situated in the cen-

tre of the mire. The restio marshland supports a vegeta-

tion structural type called Sneeukop Azonal Restioid

Fynbos (Campbell 1986).

Most of the African swamps (including mires and

other types of marshlands) are dominated by grasses and

sedges (Van Zinderen Bakker & Werger 1974; Weisser &

Howard-Williams 1982; Thompson & Hamilton 1983;

Rogers 1995, 1997). The fens and restio marshlands of

the Fynbos Biome are conspicuously different due to the

dominance of (often endemic) Restionaceae (Campbell

1986). Despite their peculiarity, the wetland ecosystems

of the Cape mountain ranges have received little atten-

tion (Boucher 1988; Rogers 1997) and hence deserve a

closer look.

This study describes vegetation types found in the

rare and poorly studied high-altitude fens and restio

marshlands located in the region of richest precipitation

in Western Cape— the Hottentots Holland Mountains

(HHM). The floristic composition of these vegetation

types and their relationship to major ecological factors

are the main foci of this paper.

MATERIAL AND METHODS

Study area: location, climate, geology and soils

The majority of vegetation samples used for this

paper were recorded from the HHM (between 33° 56' S

and 34° 03' S latitude and 18° 57' E and 19° 09' E longi-

tude). This area is situated between the towns of

Stellenbosch, Franschhoek and Grabouw in Western

Cape, South Africa— the region with the highest rainfall

in Western Cape and possibly also in the entire South

Africa. There are many peaks reaching above 1 000 m

altitude, where numerous and extensive mires have

developed. We have added some additional vegetation

samples from the Du Toitskloof Mountains, which are

located to the north of the HHM and some from the

Groenland Mountains, southeast of the HHM.

Most of the fens in the HHM are found in the Palmiet

River catchment, although the Riviersonderend and

Eerste Rivers are fed by water from fairly extensive mire

systems. Two other rivers originating in the area, the

Berg River and the Lourens River, barely receive water

from mires as they originate on very steep mountain

headwalls.

The climate of the Fynbos Biome in the southwestern

Cape is classified as mediterranean, with hot, dry sum-

mers and mild, wet winters. In the Koppen (1931 ) system

the climate of the area is classified as Csb— having

mesothermal (C) climate with a warm, dry summer and

average temperatures above 22°C and relatively wet win-

ters (sb). Most mountains of the Fynbos Biome have a

rainfall between 1 000 and 2 000 mm mean annual pre-

cipitation (MAP), but in the wettest areas (such as the

HHM) it might exceed 3 000 mm (Schulze 1965). Most

low-lying localities receive much less — up to 750 mm

near the coast, and mostly less than 400 mm MAP in the

intermontane valleys (Fuggle & Ashton 1979). The

mountains in the Fynbos Biome play a major role in

influencing precipitation and evaporation. Extremely

high regional variation in precipitation (Figure 1) occurs

due to the windward-leeward geomorphological dichoto-

my in the mountains and the fact that winds can sweep

unhindered over the coastal plains (Deacon et al. 1992).

Schulze (1965) described a strong gradient in rainfall

with increasing altitude — 50 mm of precipitation

increase per 300 m increase in altitude. The regional

rainfall pattern in the mountains is variable, depending

on aspect of slope as well as frequency and strength of

northwesterly and southwesterly winds. Mountains can

Mean Annual Precipitation (MAP)

1 8°48'

33°48' -T1

34*00'-

34*12'

Rainfall (in mm)

0-800

801 -1200

1201 -1600

1601 -2000

2001 - 2400

■H 2401 - 2800

— 2801 - 3600

No Data

FIGURE 1 .—Grid with Mean Annual

Precipitation in Hottentots

Holland Mountains (Source:

Computing Centre for Water

Research; grid data comput-

ed from weather data from

nine official weather stations

in vicinity, extrapolated using

methodology of Dent et al.

1987).

Bothalia 34,2 (2004)

143

also receive much precipitation (not registered in the rain

gauges) from mist (Kerfoot 1968; Fuggle & Ashton

1979) associated with the summer trade winds, locally

known as ‘southeasters’.

Sixty per cent of the precipitation falls in the wettest

four months spanning June to September. This precipita-

tion is mostly in the form of rain, but snow regularly

occurs in winter on the mountain peaks (Fuggle &

Ashton 1979). Wind speeds are the highest in winter on

the mountain tops (Fuggle 1981). This is in contrast with

the wind patterns in the surrounding lowlands, showing

the highest speed in summer.

The geology of the HHM is dominated by sandstones

and shales of the Table Mountain Group, especially at the

higher altitudes. Some of the valleys are underlain by

Achaean Cape Granite and Malmesbury Group shales,

but no mires were recorded on these latter substrates. The

Table Mountain Group sandstones (TMS) are mainly

from the Nardouw Formation (De Villiers et al. 1964).

High-altitude shale bands of the Cederberg Formation

accompanied by tillites of the Pakhuis Formation occur

throughout the area. The shale bands are situated at high-

er altitudes than most of the seepage areas and virtually

all the vegetation types (with one exception) described in

this study, are reported from sandstone.

Soils of the mires are classified as belonging to the

Champagne Form (Soil Classification Working Group

1991). The amount of peat is variable; towards the outer

edges of seepages, the soil contains a greater portion of

minerotrophic material (mainly sand) originated from

sandstone. Here the prevailing soil form is the Femwood

Form. In this study, we have used soils as a major crite-

rion for the location and delimitation of the fens.

Methods of data collection

Vegetation was studied by means of plot sampling.

The plots were laid out in selected fen and marshland

habitats in a representative way (Westhoff & Van der

Maarel 1978). The size of most of the plots was 10x10

m (sometimes less in the case of small vegetation

stands), as this was considered to be sufficient to record

the species richness. In each plot, each plant species

was recorded and the cover-abundance of each was esti-

mated using the modified Braun-Blanquet sampling

scale (Barkman et al. 1964). Thirty-three plots were

recorded in the HHM. We also added a further two sam-

ples from the Du Toitskloof Mountains merely to point

out that similar communities also occur in the neigh-

bouring mountain ranges (Appendix 2). Soil samples

were only collected from the topsoil (A-horizon). The

particle size distribution of the soils is determined for

the fractions finer than 2 mm 0. After drying and grind-

ing the soil to break up any coagulation, the soils were

shaken through a set of sieves. The sieves of the fol-

lowing raster size were used: 500 /Am (to separate

coarse sand), 106 /<m (to separate medium sand) and 63

H m (to separate fine sand). The fraction that passes

through all the sieves is composed of silt and clay.

Acidity and resistance were measured in water-saturat-

ed soils using a pH meter (Orion 420A) and a conduc-

tivity meter (YSI 3200). The fraction of organic matter

was measured by titration according to the Walkley-

Black method (Walkley 1935; Non-Affiliated Soil

Analysis Work Committee 1990).

A list of environmental variables and other relevant

information is presented in Table 1 .

TABLE 1 .—Explanatory variables used in the canonical ordination. Data type: I: interval scale; R: ratio-scale variables, N: nominal-scale variable

144

Bothalia 34,2 (2004)

Methods of data handling and presentation

The vegetation samples were stored in a database in the

format of the National Vegetation Database (Mucina et al.

2000) and using the database-management software

Turboveg (Hennekens 1996b; Hennekens & Schaminee

2001) . The original cover- abundance data was trans-

formed into percentage format using Turboveg. The data

were classified using Two-way Indicator Species Analysis

(TWINSPAN) (Hill 1979) followed by manual table-sort-

ing using MEGATAB 2.0 (Hennekens 1996a), aimed at

improvement of coincidence between the groups of

releves and groups of species. Differential species for

communities and their groups were identified on the basis

of fidelity. The differential species, constant companions

and the dominant species are identified in each case. A dif-

ferential species is a species that can be used to differenti-

ate between one community or a group of communities

and the rest at the same syntaxonomic level (Westhoff &

Van der Maarel 1978; Mucina 1993). The difference in the

frequency of more than two presence classes (40%) was

taken as sufficient for a species to be considered to be dif-

ferential for one of the communities under comparison.

However, a species can also be ranked as differential on

the basis of distribution of cover values (dominant versus

non-dominant) among releves of the communities under

comparison. A dominant species is a species that is con-

stant and has an average cover of more than 25%

(Westhoff & Van der Maarel 1 973). A constant companion

is a species that occurs in more than 60% of all the sam-

ples in a community and is not considered differential at

the same time. Less frequent species recorded in plots are

listed in Appendix 1 . The communities are described here

at the level of association or rankless vegetation type com-

parable to the level of association. We refrained from

defining units of the higher syntaxonomic ranks due to

limited extent of the data and local character of the study.

We have used only two informal groups of the communi-

ties based on the habitat characteristics (fens vs. restio

marshlands).

The relationship between the environmental data and

the vegetation data was determined using multivariate

techniques. We follow 0kland (1996), who made a case

for validity of use of both ordination and constrained

ordination as complementary approaches, both direct and

indirect gradient analyses were performed The four vari-

ables, coarse sand, medium sand, fine sand and silt,

together comprise 100% in every soil sample so they are

not independent from each other. According to the rec-

ommendations of Ter Braak & Smilauer (1998) this sort

of (compositional) data, has to be log-transformed prior

to analysis. Correspondence Analysis (CA) was adopted

as the indirect gradient analysis technique, while

Canonical Correspondence Analysis (CCA) was used to

perform the direct gradient analysis (see Ter Braak 1986;

Jongman et al. 1987 for details on the techniques). The

Canoco 4 program suite (Ter Braak & Smilauer 1998)

was used to perform CA and CCA.

Nomenclature of taxa and plant communities

The nomenclature of plant species follows Germis-

huizen & Meyer (2003). Three of the well-sampled plant

communities were named according to the rules for syn-

taxomic nomenclature (Weber et al. 2000). Vernacular

names, using a combination of important taxa and veg-

etation structure (Edwards 1983), were coined for all

plant communities as well.

RESULTS

The fens (and most of the restio marshlands) of the

region have a high cover of Restionaceae. Only a few seep-

age types are dominated by grasses or sedges, such as

Carpha glomerata, Epischoenus spp., Isolepis prolifer and

Pennisetum macrourum. Some Restionaceae typically

occurring in the seepages are Anthochortus crinalis,

Chondropetalum mucronatum, Elegia thyrsifera and Restio

subtilis. Two graminoids such as Ehrharta setacea subsp.

setacea and Epischoenus villosus are also common.

The following Community Groups and Communities

have been revealed in our data:

Community Group A: Fens

Fens form the wettest parts of the seepages— they are

poorly drained and contain much peat. The low-grown restio

Anthochortus crinalis is usually dominant and forms dense

mats in between the tussocks of cyperoid Epischoenus villo-

sus. Five fen communities were distinguished: the

Communities Al and A2 occur on steep slopes and experi-

ence somewhat better drainage than the flat-habitat fen com-

munities (Communities A3, A4 and A5).

Community Al: Protea lacticolor-Hippia pilosa Tall

Shrubland

(Table 2, releves 1 , 2)

The Community A 1 is peculiar due to its link to shale

bands. Protea lactic olor is the dominant species and forms

a dense shrubbery 2-4 m tall. This is the only form of pro-

teoid fynbos recorded on the slope seepages in HHM. The

herb layer covers over 80% and is dominated by tussocks

of Epischoenus villosus and mats of Restio perplexus.

Other important species include Senecio umbellatus,

Seriphium plumosum ( = Stoebe plumosa), Hippia pilosa

and Oxalis truncatula. The stands of this community were

recorded on the eastern slopes of Somerset Sneeukop at a

very high altitude (about 1 400 m). The habitat receives a

very high annual rainfall (more than 3 300 mm), some of it

in the form of snow, which might persist longer on the

southern than on the northern slopes of the mountain. A

dense mist blanket covers the mountain especially in sum-

mer. It is purported to contribute a considerable additional

amount of ambient precipitation (Marloth 1903). Campbell

(1986) refers to this vegetation (in structural terms) as

Otterford Wet Proteoid Fynbos.

Community A2: Elegia thyrsifera-Centella eriantha

Short Closed Herbland

(Table 2, releves 3, 4)

This community is found near the sources of the

Lourens River on the western slopes of Somerset

Sneeukop (1 100 m) at high altitudes and receives a high

Bothalia 34,2 (2004)

145

precipitation. The upper herb layer is formed by the dom-

inating Elegia thyrsifera , whereas the lower herb layer is

formed by a multitude of species, such as Senecio umbel-

latus , Hippia pilosa and Erica curviflora. This communi-

ty, like the Protea lacticolor-Hippia pilosa Tall Shrub-

land, is quite atypical for the seepages of the studied area.

The differential species, Carpacoce spermacocea, Centella

eriantha, Othonna quinquedentata and Ursinia eckloniana,

are all more common in typical ericaceous fynbos (Sieben

2003). The Elegia thyrsifera-Centella eriantha Commu-

nity occurs on steep slopes on sandstone.

Community A3: Anthochortus crinalis-Elegia inter-

media Tall Closed Restioland

(Table 2, releves 5-8)

Scientific name: Anthochorto crinalis-Elegietum inter-

mediate ass. nova hoc loco

Holotypus: Table 2, releve 7

This is an extremely species-poor seepage communi-

ty, which is limited to the Dwarsberg Mountains in the

Berg River catchment. The dominant vegetation stratum

is a dense layer of Elegia intermedia, which grows 1 .2 to

1.5 m tall. Linder (1987) has not recorded this species

outside the Cape Peninsula, but Kruger (1978) found it

on the Dwarsberg. In this study, it was recorded in several

other locaties in the HHM. The lower herb layer of the

community is dominated by Anthochortus crinalis and is

less dense. Dwarsberg receives more than 3 000 mm

rainfall per annum and the community is found in the

wettest, extremely peaty habitats, surrounded by stands

of the Eicinio argyropae-Epischoenetum villosi and Tetra-

rio capillaceae-Restietum subtilis. Epischoenus villosus,

Senecio crispus and the moss Campylopus stenopelma as

well as the species mentioned above are the only species

present in the vegetation.

Community A4: Ficinia argyropa-Epischoenus villo-

sus Short Closed Restioland

(Table 2, releves 9-14)

Scientific name: Ficinio argyropae-Epischoenetum

villosi ass. nova hoc loco

Holotypus: Table 2, releve 9

This is one of several seepage communities dominat-

ed by the restio Anthochortus crinalis. This clonal spe-

cies forms dense mats and is often found intertwined

with Ehrharta setacea subsp. setacea, Cliffortia tricuspi-

data and Senecio crispus. The vegetation is much short-

er than in the previous communities. The tallest restio

present is Elegia grandis, which grows taller than 0.5 m

together with the tussocks of Epischoenus villosus. In the

limited open spaces, low-grown Ficinia argyropa and

Anthoxantum tongo can be found. This community as

well as the Community A5, resemble similar vegetation

described from the Table Mountain (Glyphis et at. 1978:

Erica mollis Fynbos Community; Laidler et al. 1978:

Restio-Hypolaena Subcommunity). Both communities

are associated with peaty soils that are waterlogged for

most of the time. Ficinia argyropa-Epischoenus villosus

community also represents a typical form of what is

described by Campbell (1986) as Sneeukop Azonal

Restioid Fynbos. In one releve, Carpha glomerata was

recorded as the dominant species — a situation usually

encountered in wet habitats at lower altitudes.

Community A5: Restio bifurcus-Anthochortus crinalis

Short Closed Restioland

(Table 2, releves 15-18)

A further seepage type is also dominated by

Anthochortus crinalis. Floristically and structurally this

community resembles the previous one closely and it

might also be considered as a subassociation of the

Ficinio argyropae-Epischoenetum villosi. The main dif-

ference is in the absence of Senecio grandiflorus and

Ficinia argyropa. but this community also has some dif-

ferential species of its own, such as Gladiolus carneus,

Restio bifurcus , Restio corneolus, Tetraria capillacea

and Chondropetalum mucronatum (the last-named

reaches far above the dominant herb layer). Notable is

the occurrence of Pr ionium serration — a typical shrub in

Cape mountain streams (Sieben 2003).

Community Group B: Restio Marshlands

The habitats supporting all four communities of Com-

munity Group B are better drained than those of Com-

munity Group A. The soils are largely of minerotrophic

origin and the peat content is low. They are often found

on the edges of the mires and the water drains into the

fens. The restio marshlands are richer in species than the

fens. The Communities B1 and B2 show transitional fea-

tures between restio marshlands and fens, through occur-

rence of species such as Senecio crispus.

Community Bl: Platycaulos depauperatus Short Closed

Restioland

(Table 2, releves 19-21)

This is the only seepage type dominated by restio

Platycaulos depauperatus, which forms dense green

mats and is the most conspicuous differential species of

this community. The tussock-forming Restio subtilis is

the co-dominating element. Together they form the lower

herb stratum. One emergent 1.5 m tall restio, Chondro-

petalum mucronatum, forms its own stratum. Epischoe-

nus villosus, Elegia neesii and Tetraria capillacea are

significantly shorter. Apart from the eponymous, the

only other differential species of this community are the

grass Pentameris hirtiglumis and the geophyte Kniphofia

tabularis, flowering after a fire. We believe that this

community, although only recorded in our study in three

samples, due to being visible after a recent fire, is quite

common in the Palmiet River catchment. It seems to

occupy an intermediate ecological position between the

Ficinio argyropae-Epischoenetum villosi Association

(from peaty soils) and the Tetrario capillaceae-Restie-

tum subtilis Association (from more minerotrophic

soils). The localities of this community receive less rain-

fall than other seepage types, with a MAP of just over

2 000 mm.

146

Bothalia 34,2 (2004)

Bothalia 34,2 (2004)

147

148

Bothalia 34,2 (2004)

Bothalia 34,2 (2004)

149

Community B2: Erica autumnalis-Restio purpuras-

cens Tall Closed Restioland

(Table 2, releves 22, 23)

This is one of the two communities described from

riparian mires. From the point of view of hydrology and

species composition these communities are closely relat-

ed to the Restio Marshlands, hence they are classified as

such in this study. These communities occur along the

highest reaches of the rivers (in this study all were sam-

pled in the Palmiet River catchment) and have a mixture

of seepage and riparian elements making up very

species-rich communities, both in comparison with other

seepage types as well as with riparian communities. The

Erica autumnalis-Restio purpurascens community has a

more prominent tall herb layer than most of the seepage

communities. The dominant species is Restio purpuras-

cens, but Elegia racemosa and E. thyrsifera are also

abundant. In the lower stratum, Anthochortus crinalis is

conspicuous. Some species such as Hippia pilosa and

Senecio crispus are shared with Community Group A.

This type can best be described as a Restioland. because

small and big restio species are its most important struc-

tural constituents. It is difficult to determine the differen-

tial species in a situation where releves are few. but

Aristea bakeri, Cliffortia oralis, Elegia racemosa. Hippia

pilosa and an unidentified species of Asteraceae, might

serve as possible candidates. Eponymous Erica autum-

nalis is endemic to the HHM. The community occurs in

the highest reaches of the Wesselsgat River, where river-

banks are steep and rocky.

Community B3: Grubbia rosmarinifolia-Restio aff.

versatilis Medium Closed Shrubland

(Table 2, releves 24—26)

This is the other type of riparian mire found along

high altitude streams. The main difference from the pre-

vious type is the shape of the banks, which are flatter and

less rocky in this vegetation. The dominant small restio

here is Restio aff. versatilis, compared with Antho-

chortus crinalis in community B2.

This community has a tall herb stratum (reaching

1 .0-1 .5 m), dominated by the shrubs Berzelia squarrosa,

Brunia alopecuroides and Grubbia rosmarinifolia and

restioids Restio purpurascens and Chondropetalum

mucronatum. The most closely related riparian commu-

nity is the Erico-Tetrarietum crassae (Sieben 2003),

which shares many Erica species with the Grubbia ros-

marinifolia-Restio aff. versatilis Closed Shrubland. The

most closely related seepage community is the Tetrario

capillaceae-Restietum subtilis. A species that is shared

with this community is Restio aff. versatilis, which is the

dominating element of the ground layer. The Grubbia

rosmarinifolia-Restio aff. versatilis Community is typi-

cal of situations where river banks are not steep and there

is a lot of lateral seepage. It can be described as a shrub-

land because of the high cover of shrubs of Ericaceae and

Bruniaceae. There are many differential species, most of

which are shared with ericaceous fynbos and the

Erico-Tetrarietum crassae in particular. These are.

amongst others, Brunia alopecuroides, Berzelia squar-

rosa, Erica fastigiata, Grubbia rosmarinifolia and Restio

bifidus.

Community B4: Tetraria capillacea-Restio subtilis

Short to Tall Closed Restioland

(Table 2, releves 27-35)

Scientific name: Tetrario capillaceae-Restietum sub-

tilis ass. nova hoc loco

Holotypus: Table 2, releve 27

This is the most common type of seepage community

in the area, which can be characterized by the absence of

Senecio crispus. The dominant restio is Restio subtilis,

with Anthochortus crinalis and Restio aff. versatilis as

co-dominants. A typical characteristic is the mosaic

formed by patches of low vegetation of small restios and

sedges (Restio subtilis, R. aff. versatilis, Tetraria capil-

lacea and Epischoenus villosus ) and patches of tall veg-

etation consisting only of Chondropetalum mucronatum.

This species does not resprout after fires, like many other

seepage species, but regenerates from seed. It tends to

dominate the community, because the old plants form a

thick litter layer on the soil beneath it, which seems to

prohibit other (aggressively spreading) clonal species

from growing there.

Differential species of this community are few,

because most species are shared with the Grubbia ros-

marinifolia-Restio aff. versatilis Closed Shrubland.

Diagnostic features are mostly the dominance of Restio

subtilis and the occurrence of Chrysithrix species. As in

the case of the riparian seepage types, this community

contains numerous shrub species, such as the differential

species Grubbia rosmarinifolia and Berzelia squarrosa,

but they do not grow very tall.

This community occurs on more minerotrophic soils

than the former communities. Nevertheless, the soils are

very acidic and highly organic. In one case it was found

in a riparian zone and it is closely related to the riparian

seepage types described above. The community

described by Boucher (1978) as C hondropetalum-Restio

Tussock Marsh seems to be quite similar, but the domi-

nant small restio in the Kogelberg is not Restio subtilis

but R. ambiguus and many other species are absent in the

Kogelberg community.

Gradient analyses

The most important environmental factors that come

out of the CCA (Figure 2) are slope and altitude. This is

mainly due to the outlier communities of A1 and A2,

which are located at a higher altitude and on steeper

slopes than any other of the mire communities. They also

have, together with the riparian mire communities B2

and B3, the highest values for rockiness. It is interesting

to see that there is a sharp contrast in the fraction of soil

particle sizes: the fraction of coarse sand is an important

environmental variable and the communities with a high

fraction of coarse sand are the riparian mires (B2 and B3)

150

Bothalia 34,2 (2004)

FIGURE 2.— Biplot of constrained

ordination (CCA) of mire

vegetation of Hottentots Hol-

land Mountains, featuring

Axes 1 and 2 with position of

community samples and se-

lected environmental vari-

ables. Comm. A 1 and Comm.

A2, O; Comm. A3, +; Comm.

A4, □; Comm. A5, T;

Comm. B2 and Comm. B3,

X; Comm. B3, A; Comm. B4,

o.

and restio marshlands (B4) which are the most

minerotrophic mires in the mire system. On the other

hand, there are the communities which have a high frac-

tion of fine sand and silt, which represent the fens in the

middle of the mire system where peat formation occurs

(especially A3, but also A4 and A5). The axis that is

formed by the variable of coarse sand, fine sand and silt

reflects a gradient in minerotrophy or in waterlogging.

The fens (A3, A4 and A5) have the highest values for

organic matter contents and soil depth. They are also

mainly found at the higher altitudes because this is where

the highest rainfall occurs.

DISCUSSION

One of the most important questions that is raised

from the results of this study is where the high-altitude

mires of the Fynbos Biome fit into the world-wide typol-

ogy of fens and mires. Although the mires regularly form

the sources of the rivers, they are clearly very different

from the European spring ecosystems (Zechmeister &

Mucina 1994). Swamps and bogs with a high graminoid

cover are found extensively in the boreal zone of the

northern hemisphere (Sjors 1983) and the vegetation

cover of swamps and bogs in Africa is also mostly domi-

nated by graminoids (Thompson & Hamilton 1983).

Gore (1983) distinguishes between ombrotrophic and

minerotrophic mires, based on the origin of the water. In

ombrotrophic seepage, a thick layer of peat has devel-

oped and there is no more contact with the mineral sub-

strate. The water originates exclusively from rain, which

results in very oligotrophic conditions. The water from

minerotrophic mires seeps through the mineral substrate

into the mire, so it is richer in nutrients than the om-

brotrophic mire. Ombrotrophic mires can only exist in

very humid climates such as the blanket bogs of the

British Isles and the elevated bogs of northern Europe;

they are quite rare in the southern hemisphere. Actually,

the distinction between ombrotrophic and minerotrophic

mires is more like a gradient, an idea expressed by Sjors

(1983). Ombrotrophic mires are on the one extreme of

this gradient and all mires that do not feed exclusively on

rainwater make up the rest of this gradient.

Sjors (1983) also gives a more detailed subdivision of

European mires: topogenous mires (influenced by stag-

nant water), soligenous mires (influenced by seepage),

limnogenous mires (influenced by floodwaters) and

ombrogenous mires (influenced by rainwater). Solige-

nous and limnogenous mires are both associated with

rivers. Soligenous mires form around springs and lim-

nogenous mires occur in the floodplains along the lower

reaches of rivers. The mires described in this study are all

of the soligenous type. The different communities

described in this study are situated along a gradient from

dry to moist. In the Ficinio argyropae-Epischoenetum

villosi Association, occurring in the centre of the mire,

water stagnates more because the drainage is slow. On

the edges, the Tetrario capillaceae-Restietum subtilis

Association, which has a faster drainage, will prevail.

Further towards the margins, communities dominated by

Chondropetalum deustum can occur, but these were not

recorded during this study. Two communities can occur

towards the centre of very wet mires, namely the

Anthochorto crinalis-Elegietum intermediae Association

or the Isolepis prolifer-Bulbinella nutans Tall Closed

Sedgeland described from the Du Toitskloof Mountains.

Both communities are extremely poor in species,

because of the specific stresses that occur under water-

logged conditions. It is clear that this gradient, from

Bothalia 34,2 (2004)

151

well-drained to poorly drained or from the edge to the

centre of the mire, is also very prominent in the ordina-

tion diagrams. This gradient was also found by Bragazza

& Gerdol (1999) in some mires in the southeastern Alps.

The other distinguishing feature that shows clearly in the

ordination diagrams is the importance of the substrate, as

can be seen from the Protea mundii-Hippia pilosa Tall

Shrubland from the shale band.

A conspicuous thing about the vegetation of mires of

the Fynbos Biome is that they are dominated by the clon-

al restios A. crinalis, P. depauperatus and R. subtilis

(Linder 1985). Because these species tend to cover

everything, the vegetation is relatively poor in species.

Clonal reproduction is often coupled to environmental

plasticity, so the species can tolerate slight differences in

the environment. The diversity of microsites is much

bigger than the species richness suggests (Price &

Marshall 1999). The tall restio C. mucronatum only

regenerates from seed after fires and usually occurs in

large, dense monotypic stands when mature. It does not

support much vegetation underneath it. The dead mater-

ial from previous generations can form dense accumula-

tions of debris and this creates a very unfavourable sub-

strate for other species.

In marshes elsewhere in the world, clonal sedges and

grasses take the place of the clonal Restionaceae record-

ed in this study. It is generally accepted that the clonal

growth form is an adaptation to the stress of waterlog-

ging. This is confirmed by the investigations by Souku-

pova (1994) of three clonal graminoids. After waterlog-

ging there is an increase in clonal modules. Specht

(1981) reviews many of the problems that sclerophyllous

plants have to overcome in seasonally waterlogged areas .

It has become clear from this study that the mires in

South Africa are very different from those in the northern

hemisphere. Although they are vulnerable to predicted cli-

mate change (Rutherford et al. 1999). there is very little

knowledge about the fens and mires of the southern hemi-

sphere. In order to be able to make general statements

about mire ecosystems, more attention should be paid to

the mire ecosystems in countries like South Africa.

REFERENCES

BARKMAN. J.J.. DOING. H. & SEGAL, S. 1964. Kritische Bemer-

kungen und Vorschlage zur quantitativen Vegetationsanalyse.

Acta Botanica Neerlandica. 13: 394-419.

BOSCH. J.M.. ALLETSON. D.J.. JACOT GUILLARMOD. A.F.M.G.,

KING, J.M. & MOORE, C.A. 1986. River response to catch-

ment conditions. In J.H. O’Keeffe, The consen’ation of South

African rivers. South African National Scientific Programmes

Report No. 131.

BOUCHER. C. 1978. Cape Hangklip area. II. The vegetation. Bothalia

12: 455-497.

BOUCHER, C. 1988. Wetland vegetation of the Fynbos Biome. In

J.M.King, Hydrology and hydrobiology in the Fynbos Biome :

34-40. Ecosystems Programmes Occasional Report No. 26,

CSIR. Pretoria.

BRAGAZZA. L. & GERDOL, R. 1999. Hydrology, groundwater

chemistry and peat chemistry in relation to habitat conditions in

a mire on the southeastern Alps of Italy. Plant Ecology 144:

243-256.

CAMPBELL. B.M. 1986. A classification of the mountain vegetation

of the Fynbos Biome. Memoirs of the Botanical Survey of South

Africa No. 50: 1-121.

DALLAS, H.F. & DAY, J.A. 1993. The effect of water quality variables

on riverine ecosystems: a review. Water Research Commission

Report TT 61/93, Pretoria.

DEACON, H.J., JURY, M.R. & ELLIS, F. 1992. Selective regime and

time. In R.M. Cowling, The ecology of fynbos: nutrients, fire

and diversity. 6-22. Oxford University Press, Cape Town.

DENT, M.C., LYNCH, S.D. & SCHULZE, R.E. 1987. Mapping mean

annual precipitation and other rainfall statistics over southern

Africa. Agricultural Catchments Research Unit Report No. 27.

Water Research Commission Report No. 109/1/89.

DE VILLIERS, J„ JANSEN, H. & MULDER, M.P. 1964. Die geolo-

gie van die gebied tussen Worcester en Hermanus. Geological

Survey, Pretoria.

EDWARDS, D. 1983. A broad-scale structural classification of vegeta-

tion for practical purposes. Bothalia 14: 705-712.

FRY, M. ST. L. 1987. A detailed characterization of soils under differ-

ent fynbos-climate-geology combinations in the southwestern

Cape. M.Sc. thesis. University of Stellenbosch.

FUGGLE. R.F. 1981 . Macro-climatic patterns within the Fynbos Biome.

Final Report. National Programme for Environmental Sciences,

Fynbos Biome Project, Cape Town.

FUGGLE, R.F. & ASHTON, E.R. 1979. Climate. In J. Day. W.R.

Siegfried, G.N. Louw & M.L. Jarman, Fynbos ecology: a pre-

liminary synthesis. South African National Scientific Program-

mes Report No. 40: 7-15.

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

Africa: an annotated checklist. Strelitzia 14. National Botanical

Institute, Pretoria.

GLYPHIS, J„ MOLL. E.J. & CAMPBELL, B.M. 1978. Phyto-

sociological studies on Table Mountain, South Africa. 1. The

Back Table . Journal of South African Botany 44: 281-289.

GORE, A.J.P. (ed.). 1983. Ecosystems of the World 4. Mires: swamp,

bog, fen and moor. A. General studies. Elsevier, Amsterdam.

HENNEKENS. S.M. 1996a. MEGATAB — a visual editor for phytosoci-

ological tables. Version 1 .0. User’s Guide. Giesen & Geurts, Ulft.

HENNEKENS, S.M. 1996b. TURBOVEG: software package for input,

processing, and presentation of phytosociological data. Version

July 1996. User’s guide. IBN-DLO & Lancaster University,

Wageningen & Lancaster.

HENNEKENS, S.M. & SCHAMINEE, J.H.J. 2001. TURBOVEG, a

comprehensive data base management system for vegetation

data. Journal of Vegetation Science 12: 589-591.

HEWLETT, J.D. 1961. Soil moisture as a source of base flow from

steep mountain watersheds. Southeastern Experimental Station,

USDA Forest Service Paper 132.

HEWLETT, J.D. 1982. Principles of forest hydrology. University of

Georgia Press, Athens, Georgia.

HILL, M.O. 1974. Correspondence analysis: a neglected multivariate

method. Applied Statistics 23: 340-354.

HILL. M.O. 1979. TWINSPAN—a FORTRAN program for arranging

multivariate data in an ordered two way table by classification

of individuals and attributes. Cornell University, Ithaca, New

York.

JONGMAN, R.H.G., TER BRAAK, C.J.F. & VAN TONGEREN,

O.F.R. 1987. Data analysis in community and landscape ecolo-

gy. Pudoc, Wageningen.

KERFOOT, O. 1968. Mist precipitation on vegetation. Forestry

Abstracts 29: 8-20.

KING, J.M. & DAY, J.A. 1979. Hydrology and hydrobiology. In J.

Day, W.R. Siegfried, G.N. Louw & M.L. Jarman, Fynbos ecol-

ogy: a preliminary synthesis. South African National Scientific

Programmes Report No. 40: 27—42.

KOPPEN, W. 1931. Grundrifi der Klimakunde. De Gruyter, Berlin.

KRUGER. F.J. 1978. South African heathlands. In R.L. Specht,

Ecosystems of the World 9. Heathlands and related shrublands.

B. Analytical studies: 19-80. Elsevier, Amsterdam.

LAIDLER, D.. MOLL, E.J., CAMPBELL, B.M. & GLYPHIS, J. 1978.

Phytosociological studies on Table Mountain, South Africa. 2.

The Front Table . Journal of South African Botany 44: 29 1-295 .

LINDER, H.P. 1985. Conspectus of the African species of Restion-

aceae. Bothalia 15: 387-503.

LINDER, H.P. 1987. The evolutionary history of the Poales/Restio-

nales— a hypothesis. Kew Bulletin : 42: 297-318.

MARLOTH, R. 1903. Results of experiments on Table Mountain for

ascertaining the amount of moisture deposited from the south-

east clouds. Transactions of the Royal Philosophical Society of

South Africa 14: 403^108.

MUCINA, L. 1993. Nomenklatorische und syntaxonomishe

Definitionen, Konzepte und Methoden. In L. Mucina, G.

152

Bothalia 34,2 (2004)

Grabherr & T. Ellmauer, Die Pflanzengesellschaften Osterre-

ichs. Teil I Anthropogene Vegetation: 19-28. Gustav Fischer,

Jena.

MUCINA, L„ BREDENKAMP, G.J., HOARE, D.B. & MCDONALD,

D.J. 2000. A national vegetation database for South Africa.

South African Journal of Science 96: 497, 498.

NON- AFFILIATED SOIL ANALYSIS WORK COMMITTEE. 1990.

Handbook of standard soil testing methods for advisory pur-

poses. Soil Science Society of South Africa, Pretoria.

0KLAND, R.H. 1996. Are ordination and constrained ordination alter-

native or complementary strategies in general studies? Journal

of Vegetation Science 7: 289-292.

PRICE, E.A.C. & MARSHALL, C. 1999. Clonal plants and environ-

mental heterogeneity. Plant Ecology 141: 3-7.

ROGERS, K.H. 1995. Riparian wetlands. In G.I. Cowan, Wetlands of

South Africa: 41-52. Department of Environmental Affairs and

Tourism, Pretoria.

ROGERS, K.H. 1997. Freshwater wetlands. In R.M. Cowling, D.M.

Richardson & S.M. Pierce, Vegetation of southern Africa :

322-347. Cambridge University Press, Cambridge.

RUTHERFORD, M.C., POWRIE, L.W. & SCHULZE, R.E. 1999.

Climate change in conservation areas of South Africa and its

potential impact on floristic composition: a first assessment.

Diversity and Distribution 5: 253-262.

SCHULZE, B.R. 1965. Climate of South Africa . Part 8. General sur-

vey. Government Printer, Pretoria. ■

SIEBEN, E.J.J. 2003. The riparian vegetation of the Hottentots

Holland Mountains, Western Cape, South Africa. Ph.D. thesis,

University of Stellenbosch.

SJORS, H. 1983. Mires of Sweden. In A.J.P. Gore, Ecosystems of the

World 4. Mires: swamp, bog, fen and moor. B. Regional studies :

69-94. Elsevier, Amsterdam.

SOIL CLASSIFICATION WORKING GROUP 1991. Grondklassi-

fikasie. Een taksonomiese sisteem vir Suid-Afrika. Department

of Agriculture Affairs, Pretoria.

SOUKUPOVA, L. 1994. Allocation plasticity and modular structure in

clonal graminoids in response to waterlogging. Folia

Geobotanica & Phytotaxonomica 29: 227-236.

SPECHT, R.L. (ed.). 1981 . Ecosystems of the World 9. Heathlands and

related shrublands. B. Analytical studies. Elsevier, Amsterdam.

TAYLOR, H.C. 1978. Capensis. In M.J.A. Werger, Biogeography and

ecology of southern Africa: 171-229. Junk, The Hague.

TER BRAAK, C.J.F. 1986. Canonical Correspondence Analysis: a new

eigenvector technique for multivariate direct gradient analysis.

Ecology 67: 1167-1179.

TER BRAAK, C.J.F. & SMILAUER, P. 1998. CANOCO Reference

Manual and User's Guide to Canoco for Windows. Software for

Canonical Community Ordination (version 4). Microcomputer

Power , Ithaca, New York.

THOMPSON, K. & HAMILTON, A. C. 1983. Peatlands and swamps of

the African continent. In A.J.P. Gore, Ecosystems of the World

4. Mires: swamp, bog, fen and moor. B. Regional studies:

331-373. Elsevier, Amsterdam.

VAN ZINDEREN BARKER, E.M. & WERGER, M.J.A. 1974. En-

vironment, vegetation and phytogeography of the high-altitude

bogs of Lesotho. Vegetatio 29: 37—49.

WALKLEY. A. 1935. An examination of methods for determining

organic carbon and nitrogen in soils. Journal of Agricultural

Science 25: 598-609.

WEBER. H.E., THEURILLAT, J.-P. & MORAVEC, J. 2000. Inter-

national Code of Phytosociological Nomenclature, edn 3. Journal

of Vegetation Science 11: 739-768.

WEISSER, P. J. & HO WARD- WILLI AMS, C. 1982. The vegetation of

the Wilderness Lakes system and the macrophyte encroachment

problem. Bontebok 2: 19 — 40.

WESTHOFF, V. & VAN DER MAAREL, E. 1978. The Braun-

Blanquet approach. In R.H. Whittaker, Classification of plant

communities: 287-399. Junk, The Hague.

ZECHMEISTER, H. & MUCINA, L. 1994. Vegetation of European

springs: high-rank syntaxa of the Montio-Cardaminetea.

Journal of Vegetation Science 5: 385-399.

APPENDIX 1 . — List of less frequent species having one or two occurrences in the releve table. Sequence: name of

species, the field code of the releve(s) and cover-abundance (in brackets)

Agathosma pentachotoma 262 (2a)

Askidiosperma chartaceum 262 (2a)

Askidiosperma esterhuyseniae 164 (2b), 202 (+)

Berzelia lanuginosa 262 (+)

Blechnum tabulare 131 (+)

Bobartia gladiata 259 (+)

Chironia decumbens 262 (r)

Cliffortia graminea 165 (1)

Cliffortia ruscifolia 233 (2a)

Corymbium conge stum 131 (1)

Corymbium cymosum 131 (+)

Dicranoloma billardieri 123 (2a)

Disa tripetaloides 123 (2m), 127 (2a)

Edmondia pinifolia 131 (+)

Ehrharta ramosa 259 ( 1 )

Ehrharta setae ea subsp. uniflora 111 (2m)

Epischoenus complanatus 131 ( 1 ) , 20 1 (+)

Epischoenus gracilis 262 (2a)

Erica longifolia 262 (r)

Euchaetis glabra 201 ( 1 )

Euryops abrotanifolius 232 (2m), 233 (1)

Felicia cymbalariae 111 ( 1 )

Ficinia cf. involuta 259 (3)

Ficinia sp. 233 ( 1 )

Fissidens plumosus 1 1 1 (2m)

Geissorhiza umbrosa 131 (+)

Helichrysum cymosum 233 (2a)

Hymenophyllum peltatum 131 (2m)

Hypochaeris radicata 259 (+)

Ischyrolepis triflora 262 ( 1 )

Isolepis digitata 1 35 (2m)

Kogelbergia verticillata 127 (+)

Lobelia jasionoides 134 (+)

Lycopodiella caroliniana 123 (+)

Osmitopsis afra 241 (+)

Oxalis nidulans 202 (+)

Pentameris thuarii 233 (2b)

Pentaschistis pallida 135 (+)

Protea cynaroides 128 (+), 233 (r)

Psoralea aculeata 241 (2a)

Raspalia virgata 136 (2a)

Restio bifarius 131 (+)

Restio intermedius 128 (3)

Restio obscurus 262 (1)

Restio pedicellatus 259 (+), 262 (3)

Schizaea tenella 127 (1)

Senecio coleophyllus 134 (2a)

Senecio pubigerus 233 (1)

Senecio rigidus 131 (+)

Sonchus oleraceus 259 (+)

Staberoha cernua 234 (1)

Staberoha vaginata 124 (+), 189 (+)

Seriphium plumosum (juvenile) 188 (r)

Tetraria pillansii 201 (1), 259 (2a)

Tetraria thermalis 1 24 (r)

Todea barbara 11) ( 1 ), 259 (2a)

Ursinia dentata 233 (1), 241 (+)

Utricularia hisquamata 163 (+)

Villarsia capensis 163 (1), 164 (+)

Wahlenbergia procumbens 201 ( 1 ), 24 1 (2b)

Watson ia borbonica 241 (r).

APPENDIX 2.— Selected data on sampled vegetation properties and geographical location of two releves in Du Toitskloof Mtns and releves in Hottentots Holland Mtns. No., releve number; Com., community; FC, field code; As., aspect; SI.,

slope in (.leg tees; Area, sampled area (m~); CE2, cover of shrub layer (%); CE 1 , cover of herb layer (%); CEO, cover of moss layer (%); HE2, height of shrub layer (m); HE I , average height of herb layer (m); SR, species richness

Bothalia 34,2 (2004)

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Bothaiia 34,2: 155-173 (2004)

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Sass, Ms D. Senior Provisioning Admin. Clerk II. Leave

& IOD admin.

Williams, Mrs L.R. Dip.(Sec.). Senior Personnel Officer

II. Recruitment

FINANCE DIRECTORATE (AFIN)

CAPE TOWN

Hughes, W.S.G. B.Com., C.A.(SA). Director. Chief Financial Officer

Rawoot, N.A. B.Acc.Sc. Deputy Director: Financial management

Maholwana, S. B.Com.(Hons). Assistant Director. Financial management

Neuwirth, Ms E.V. Assistant Director. Staff benefits

Cassiem, Ms S. Chief Accounting Clerk. Salaries

Cooper, S.K. Accounting Clerk II. Creditors

Goodman, Mrs I.W. State Accountant. Supervisor: Creditors

Jacobs, F.H. Senior Accounting Clerk II. Staff benefits

Jacobs, Ms S. State Accountant. Financial admin.: Gene-

ral ledger

Kolbe, Ms D.V. Senior Accounting Clerk I (contract

worker)

Mcontsi, Ms N. N.Dip.(Acc.). Senior Accounting Clerk

I. Creditors

Mirkin, Ms Y.A. Senior Secretary III

Paulse, Mrs D.W.S. Dip.(Bookkeep.), Dip.(Sec.). Chief

Accounting Clerk III. Creditors

Potgieter, Ms G. Clerk. Salaries (contract worker)

ENVIRONMENTAL EDUCATION DIRECTORATE (EENT/GP)

PRETORIA

Qwathekana, Ms N.M. B.A.(Hons), B.A.(Eng., Geog. & Film Studies III),

Dip. (Education). Director

Symonds, Ms A.M. N.Dip.(Nature Cons.), H.E.D. Deputy Director

Adams, Ms E.M. Senior Provisioning Admin. Officer. Centre

Manager

Eyssell, Ms A. B.Sc. (Hons) (Hort.). Outreach greening

Horticulturist

Maphuta, Mrs M.S. Specialist Cleaner, Assistant to centre

manager

Mathaba,T.C. Environmental Education Officer. Garden-

based programme

Novellie, Mrs E. H.E.D. B.Sc.(Hons) (Zool. & Mammo-

logy). Principal Environmental Education Officer

GOLD FIELDS CENTRE -CAPE TOWN (EECT)

Fullard, D. B.Sc. Ed., B.Ed. (Hons). Assistant Director. Environmental Education

Hey, Ms S.J. B.A. (Geog.), HDE. Senior Environmental Education Officer.

Garden-based programme

Mgodeli, W.M. Driver II

September, Ms M. Senior Provisioning Admin. Clerk II. Admin, support

WALTER SISULU NBG (EENT)

Molefe, Ms K.E. Dip. (Nature Cons.). Senior Environmental Education Officer. Outreach education

Moore, Mrs J.M. N.H.Dip.(Sec.). Senior Provisioning Admin. Clerk II. Admin, support (part time)

Bothalia 34,2 (2004)

157

BIODIVERSITY POLICY AND PLANNING DIRECTORATE (BDIR)

Maze, Ms K.E. M.Sc. Director. Biodiversity and Action Planning

Baloyi, Ms I. Senior Secretary IV

THREATENED SPECIES PROGRAMME (YDBR/TS)

PRETORIA

Foden, Ms W. Programme Manager (contract worker)

Keith, M. Research Scientist (contract worker)

Pillay, Ms D. Red List Officer (contract worker)

Victor, Ms J.E. M.Sc., H.Dip.(Joum.). Principal Agricultural Scientist.

Taxonomy of Rutaceae, Asclepiadaceae

C.A.P.E. PROJECT

CAPE TOWN

Barnett, Dr M. Programme Developer (contract worker)

Cadman, Dr M.J. Bioregional Programme Co-ordinator. Eastern Cape (contract worker)

Magasela, Ms B.B. Communication Manager (contract worker)

Sandwith, T. Programme Co-ordinator (contract worker)

GARDENS DIRECTORATE— ADMIN STAFF (GDIR)

PRETORIA

Willis, C.K. M.Sc. (Cons. Biol.)— Chief Director: Gardens and Horticultural Services

Heilgendorff, J.P. N.H.Dip.(Hort.). Gardens IT Manager

Els, Ms L. N.Dip.(Sec.). Senior Secretary III

URBAN CONSERVATION -CAPE TOWN (GDIR/UC)

Davis, G.W. Ph.D. Assistant Director: Communication. Project management and fund raising

Goldman, Ms T. Project manager (contract worker)

Peter, L.M. Dip.(Hort.). Principal Communications Officer. Project management

INTERPRETATION

Roff, J. Cert.(Envir. Interpr. & Ed.). Communication Officer. Interpretation (Pietermaritzburg)

PLANNING, MAINTENANCE & DEVELOPMENT-CAPE TOWN (BPMD)

Linde, D.C. N.T.C.III(Civil & Structural: Building) N.T.C.III (Inspector of Works: Building).

M.S.A.I.D. Cert. Estate Agency. Control Inspector of Works

Arendse, D.S. Senior Handyman. Building maintenance

Manasse, S.P. Dip.(Masonry). Artisan Foreman. Building maintenance

Peck, W.I. Senior Handyman. Building maintenance

CURATORS

Behr, Ms C.M. Curator: Pretoria NBG

Britz, R.M. Curator: Lowveld NBG (Nelspruit)

Gavhi, M.P. Curator: Free State NBG (Bloemfontein)

Le Roux, P.H. Deputy Director. Curator: Kirstenbosch NBG (Cape Town)

Oliver, I.B. Curator: Karoo Desert NBG (Worcester)

Tarr, B.B. Curator: Natal NBG (Pietermaritzburg)

Turner, Ms S.L. Curator: Witwatersrand NBG (Roodepoort)

Xaba, Ms A.C. Curator: Harold Porter NBG (Betty’s Bay)

158

HAROLD PORTER NBG— BETTY’S BAY (GHPG)

Xaba, Ms A.C. N.DipJHort.). Curator

Bothalia 34,2 (2004)

Abrahamse, SJ. Principal Foreman. Garden

Arendse, Ms M. Auxiliary Services Officer II . Access control

Bebe, Ms N. Cleaner I

Bezuidenhout, Mrs H.M. Provisioning Admin. Officer

Carolus, Ms BJ. N.H.Dip.(Hort.) Agricultural Develop-

ment Technician. Horticulture

Forrester, Ms J.A. N.T.C.III(Hort.). Chief Agricultural

Development Technician. Horticulture

October, Ms R.P. Dip. (Education). Senior Auxiliary Ser-

vices Officer. Plant records and asset register

Samuels, Ms D.C. Cleaner II

Simpson, H.A.S. Auxiliary Services Officer II. Access

control

Van Wyk, A.B. Artisan. Building maintenance

KAROO DESERT NBG -WORCESTER (GKAR)

Oliver, I.B. N.Dip.(Hort.), N.Dip.(Public Relations & Admin). Control Agricultural Technician. Curator

Ashworth, Mrs E.H. Senior Provisioning Admin. Clerk III.

Admin, support

Mtetwa, A.M. Artisan. General maintenance

Makubalo, F.N. Senior Foreman. Nursery

Mpeke, Ms E.N. Specialist Cleaner

Simani, D.K. Principal Foreman. Garden

Van Tonder, M. (volunteer)

Viljoen, D.M. N.Dip.(Hort.). Chief Agricultural Develop-

ment Technician. Collections

Voigt, W.E. N.Dip.(Hort.). Senior Agricultural Develop-

ment Technician. Garden development

Williams, C. (student)

KIRSTENBOSCH NBG— CAPE TOWN (GKBC)

Le Roux, P.H. Dip. (Forestry), N.Dip.(Hort.), N.Dip.(Parks & Recr.), Cert.(Turf Management). Deputy Director:

Garden Management. Curator

Goldschmidt, S.M. B.A.(Soc.)(Hons.). Assistant Director. Personnel Manager

Adams, T.D. B.Tech.(Hort.). Senior Agricultural Development Technician. Supervisor: Greenhouse

Hitchcock, A.N. N.H.Dip.(Hort.). Control Agricultural Technician. Nursery Manager

Morkel, A.T. N. Dip. (Nature Cons.). Control Agricultural Technician. Estate Manager

Notten, Ms A.L. B.Sc., N.Dip.(Hort.). Chief Agricultural Development Technician. Supervisor: Seed room

Trautman, C.E. Artisan. Supervisor: Workshop

Adonis, A. Principal Foreman. Dell & ericas

Adonis, SJ. Senior Foreman. Aliens vegetation control

Arends, Ms S.J. Principal Auxiliary Services Officer. Plant

records

Badenhorst, Ms C.L. (student)

Botha, P.A. N.H.Dip.(Hort.). Chief Agricultural Develop-

ment Technician. Seed research

Boyana (Magija), Ms N.F. N.DipJHort.). Senior Agricul-

tural Development Technician. Outreach programme

Brown, B.M. N.DipJHort.). Agricultural Development

Technician. Nursery

Burring, J.H. Kirstenbosch Scholar

Crowie, R.W. Senior Foreman. General garden

Crowie, Mrs U.M. Accounting Clerk II. Seed room

Duncan, G.D. N.Dip.(Hort-). Control Agricultural Develop-

ment Technician. Bulbs

Engelbrecht, F. Senior Provisioning Admin. Clerk II. Stores

Engelbrecht, Mrs L.D. Control Auxiliary Services Officer.

Plant records

Grace, T. Senior Provisioning Admin. Clerk III. Stores &

admin, support

Harrower, A.D. Ball Agreement. Greenhouse Supervisor

(Contract worker)

Hope, C.F. Senior Handyman. Construction

Jacobs, H.C. Senior Foreman. Plant production

Jaftha, Ms D.L. Accounting Clerk II. Seed room

Jamieson, Mrs H.G. N.Dip.(Parks & Rec.). Chief Agricul-

tural Development Technician. Restiol Asparagus

(part time)

Jodamus, Ms N.L. N.Dip.tHort.). Senior Agricultural

Development Technician. Annuals

Kamalie, Ms S. Senior Typist. Receptionist

Kayster, G J. Senior Foreman. Construction

Kuscus, G.W. Senior Foreman. General maintenance

Manuel, I.P. Senior Foreman. Seed room

Masango, J. (student)

Mathys, Mrs S.S.B. Senior Accounting Clerk III. Revenue

and visitor statistics

Matthews, I.N. Senior Foreman. Estate & trails

Mbambezeli, N.G. N.Dip.(Hort). Agricultural Develop-

ment Technician. Trees & shrubs

Mitchells, G. Senior Foreman. Mowers

Morris, J.N.M. Senior Foreman. Proteas

Newman, W. Artisan. Mechanical workshop

Nonkenge, Ms S. (student)

Picane, Ms S. Auxiliary Services Officer II. Tissue culture

Prins, F.B. Security Officer III

Rudolph, A. Security Officer III

Smith, Mrs A. Senior Provisioning Admin. Clerk II. Admin,

support

Solomons, T.C. Senior Security Officer II

Tamboer, J.S. Senior Foreman. Nursery services

Tuckledoe, R. (student)

Van Gusling, EJ. Senior Foreman. Mowers

Van der Walt, Mrs L.E. N.DipJHort.). Chief Agricultural

Development Technician. Herbaceous collections

(part time)

Van Jaarsveld, E J. M.Sc., N.Dip.(Hort.). Control Agricul-

tural Technician. Succulents

Van Rooy, K. Senior Foreman. Annuals

Bothalia 34,2 (2004)

159

Van Wyk, F. Principal Auxiliary Services Officer II. Lable

Maker

Viljoen, Ms C.C. N.Dip.(Hort.). Chief Agricultural Develop-

ment Technician. Plant production

Xaba, PA. N.Dip.(Hort.). Overberg Useful Plants Project

(contract worker)

Yohane, T.E. (student)

VISITORS CENTRE -CAPE TOWN (GKBC/VC)

Struys, Ms S. Dip.(Market. Manag.), B. A .(Directing). Principal Communications Officer. Centre Manager

August, T. Senior Auxiliary Services Officer. Information

services

Fredericks, Ms N.C.E. Senior Auxiliary Services Officer.

Visitors’ Centre. Information services

Jacobs, A.P. Chief Auxiliary Services Officer. Visitors’

Centre. Information services

Jacobs, Ms B.C. Dip.(Travel & Tourism), Dip.(Compu-

ter Business). Senior Admin. Clerk II . Facilities Co-

ordinator

Malan, Ms C.E. B.Sc.(Hons). Principal Communication

Officer: Tour co-ordinator

Petersen, J.E. Senior Provisioning Admin. Clerk. Events

co-ordinator

Williams, G.C. Senior Auxiliary Services Officer. Infor-

mation

LOWVELD NB G — NELS PRUIT (GLOW)

Britz, R.M. N. Dip. (Forestry). Control Agricultural Technican. Curator

Froneman, W.C.F. N.T.C.III(Hort.), N.Dip.(Nature Cons.

& Man.), N.Dip.(Parks & Rec. Admin.), N.T.C.III

(Hort.). Control Agricultural Technician. Nursery

management & garden development

Hurter, P.J.H. B. Sc .(Hons). Control Agricultural Techni-

cian. Cycad conservation

Khoza, Ms P.K. Cleaner I

Le Roux, Ms L.N. N.H.Dip.(Nature Cons.). Principal

Auxiliary Services Officer II. Interpretation

Maqungo, Ms V.L.B. Auxiliary Services Officer. Kiosk

Musweli, K.J. Senior Foreman. New development

Ngwenya, PS. Auxiliary Services Officer II. Kiosk

Sibanyoni, Ms S.M. Cleaner II

Van der Walt, Mrs G.A.M. Senior Provisioning Admin.

Clerk III

Xozumti, M.M. Principal Foreman. Supervisor

NATAL NBG- PIETERMARITZBURG (GNAT)

Tarr. B.B. N.Dip.(Parks & Rec. Admin.), Advanced Dip.(Adult Educ.). Control Agricultural Technician. Curator

Dlungwane, T.R. Senior Foreman. Garden maintenance

Johnson, Ms I. M.Sc., H.E.D. Chief Agricultural Develop-

ment Technician

Nonjinge, S.H.B. N.T.C.HI(Hort.). Chief Agricultural De-

velopment Technician

Sibiya, Ms C.P.T. Cleaner II

Van der Merwe, Mrs M.E.H. Senior Provisioning Admin.

Clerk III

Zimu, M.J. Principal Foreman. Garden

FREE STATE NBG -BLOEMFONTEIN (GFSG)

Gavhi, M.P. N.Dip.(Hort.). Curator

Barnard, Ms A.D. Admin. Clerk (part time) (contract

worker)

Harris, Ms S. N.Dip.(Hort.). Senior Agricultural Develop-

ment Technician

Mbolekwa, L.M. Foreman. Garden

Nakanyane, R.B. Principal Foreman. Garden

Raditlhare, Mrs E.M. Cleaner II

Sebolai, R.P.A.N. Senior Handyman. General maintenance

Thaele, Mrs M.E. Cleaner II

Tshabalala, Ms B.N.W. (student)

PRETORIA NBG (GPTA)

Behr, Ms C.M. B.Sc.(Hons). Control Agricultural Development Technician. Curator

Baloyi, K J. Senior Auxiliary Services Officer II. Informa-

tion Officer

Baloyi , M .S . Dip .( IBM ) , Dip .(PTM ) , Dip .( Payroll Admin .) .

Senior Provisioning Admin. Clerk I. Leave records

and H.R. support

Baloyi, S.J. Senior Handyman

Bell, Ms C.

Creighton, Ms D.D. Senior Provisioning Admin. Clerk III.

Admin, support

Keyter, B.A. Senior Security Officer II

Klapwijk, N.A. N.Dip.(Hort.), N.Dip.(Plant Prod.), N.Dip.

(Diesel Fitting).Control Agricultural Development

Technician

Kutama, B.T. Senior Foreman. Garden development

Lithudza, E.F. Chief Agricultural Development Technician

Maeta, C.J. Senior Agricultural Development Technician

Makgobola, Ms M.R. Auxiliary Services Officer II. Recep-

tion & admin, support

160

Bothalia 34,2 (2004)

Mariri, Ms M.A. Cleaner II

Matthews, A.V. Chief Agricultural Development Technician

Modisha, M.D. Cleaner II

Mphaka, Ms N.F. Student Horticulturalist

Noku, Y.A. Principal Foreman. Irrigation and machine ope-

rators

Solomons, Ms C.V. Senior Auxiliary Services Officer.

Plant records clerk

Venter, W.A. N.T.C.II. Senior Artisan. Workshop and general

maintenance

WALTER SISULU NBG— ROODEPOORT (GWIT)

Turner, Ms S.L. B.Sc.(Hons), N.Dip.(Hort.). Control Agricultural Technician. Curator

Aubrey, Mrs AE. B.Tech.(Hort.). Chief Agricultural Develop-

ment Technician. Plant records, interpretation, informa-

tion (part time)

Baloyi, S J. Handyman. Stores

Dlamini, M.D. N.Dip.(Hort.). Agricultural Development

Technician

Hankey, A J. N.Dip.(Hort.). Control Agricultural Technician.

Garden, estate, collections, nursery

Head, Mrs S.E. Dip.(Shorthand & Typing). Provisioning

Admin. Officer

Mamosebo, M.A. Factotum

Manyikana, T.M. Factotum

Mmola, Mrs B.E. Cleaner II

Ndou, A.P. Senior Auxiliary Services Officer II. Garden

information

Ndzondo, Ms N.L. Senior Provisioning Admin. Clerk I

Ndzondo, Mrs P.G. Cleaner II

Nedambale, M.P. Senior Foreman. Garden

Nemalili, M.E. Senior Foreman. Machines and vehicles

Nenungwi, M.S. Senior Foreman. Nursery

RESEARCH DIRECTORATE (RDIR)

PRETORIA

Smith, Prof. G.F. Ph.D., F.L.S. Chief Director: Research & Scientific Services

Arnold, T.H. Head: Data Management (Pretoria)

Crouch, N.R. Head: Ethnobotany Unit (Durban)

Donaldson, J.S. Ph.D. (Zoology) Director: Kirstenbosch Research Centre (Cape Town)

Koekemoer, Ms M. Curator: National Herbarium (Pretoria)

Leistner, O.A. D.Sc. F.L.S. Scientist (contract worker)

Marais, Mrs A.C. Senior Provisioning Admin. Officer

Meyer, Mrs N.L. B.Sc.(Hons). Agricultural Development Technician (contract worker)

Roux, J.P. Curator: Compton Herbarium (Cape Town)

Singh, Ms Y. Curator: Natal Herbarium (Durban)

Steenkamp, Ms Y. Regional Project Co-ordinator: SABONET (Pretoria)

Steyn, Mrs E.M.A. D.Sc. Specialist Scientist. Embryology, anatomy, taxonomy

Wolfson, Mrs M.M. Ph.D. Director: Research Support Services

NATAL HERBARIUM— DURBAN (RHED)

Singh, Ms Y. M.Sc., H.E.D. Senior Agricultural Scientist. Taxonomy of Araceae, Hypoxidaceae. Curator

Apollos, Ms C.E. Senior Provisioning Admin. Clerk I

Dimon, Ms Z.Y. B.Sc. Data Capturer (SABONET contract

worker)

Govender, Mrs N. B.Sc. (Hons) Data Capturer. Chironia

systematics (SABONET contract worker)

Hlongwane, Mrs N.C. Cleaner II & messenger

Mbonambi, B.M. Groundsman II. Garden maintenance

Nathoo, Ms M. B.Sc. Data Capturer (SABONET contract

worker)

Ngwenya, A.M. Senior Agricultural Development Tech-

nician. Herbarium Officer. Plant identification and

information, Zulu Botanical Knowledge Project

Mazibuko, J.V.G. Senior Auxiliary Services Officer. Herba-

rium Assistant

Noble, Mrs H-E. Senior Provisioning Admin. Clerk III

Williams, Ms R. B.Sc.(Hons), H.E.D. Principal Agricul-

tural Scientist

ETHNOBOTANY UNIT-DURBAN (RETH)

Crouch, N.R. Ph.D. Assistant Director. Ethnobotany of southern African flora

Bothalia 34,2 (2004)

161

NATIONAL HERBARIUM -PRETORIA (RHEN)

Koekemoer, Ms M. Ph.D. Deputy Director. Herbarium management.

Taxonomy of Asteraceae: Gnaphalieae

Bredenkamp. Mrs C.L. Ph.D. Principal Agricultural Scientist. Assistant Curator:

Public relations. Taxonomy of Vitex, Passerina, Malvaceae, Sterculiaceae,

and other related families

Herman. P.P.J. M.Sc. Principal Agricultural Scientist. Assistant Curator: Personnel.

Taxonomy of Asteraceae

Mothogoane, M.S. Principal Auxiliary Services Officer. Assistant Curator:

Herbarium assistants. Wing C

Sebothoma, P.N. Cert.(Sec.). Principal Auxiliary Services Officer. Assistant Curator:

Service room. Plant identifications co-ordinator

Van Rooy, J. Ph.D. Chief Agricultural Scientist. Assistant Curator: Technical staff.

Taxonomy and biogeography of mosses

Anderson, J.M. Ph.D. Specialist Scientist. Molteno Palaeo-

flora, Gondwana Alive

Arendse. Ms S. African Plant Initiative Programme (con-

tract worker)

Archer Mrs C. M.Sc. Principal Agricultural Scientist. Tax-

onomy of Cyperaceae, monocotyledons (general)

Archer, R.H. Ph.D. Principal Agricultural Scientist. Tax-

onomy of mainly Celastraceae, Euphorbiaceae

Bester, S.P. M.Sc. Senior Agricultural Scientist. Tax-

onomy of Apocynaceae, Ericaceae, Rutaceae

Burgoyne, Ms P.M. M.Sc. Principal Agricultural Scien-

tist. Mesembryanthemaceae

Dimpe, Ms K.M.C. Graphic designer (contract worker)

Fish, Mrs L. B.Sc. Principal Agricultural Scientist. Tax-

onomy of Poaceae. Collections manager

Glen, H.F. Ph.D. Specialist Scientist. Taxonomy of trees,

herbarium for cultivated plants, and botanical col-

lectors

Glen. Mrs R.P. M.Sc. Control Agricultural Development

Technician. Taxonomy of ferns, water plants

Gotzel, Ms A. Senior Provisioning Admin. Clerk III

Govender, Ms M. Agricultural Development Technician

Grunyuza, Ms T. Senior Herbarium Assistant. African

Plant Initiative Programme (contract worker)

Jordaan, Mrs M. M.Sc. Principal Agricultural Scientist.

Taxonomy of Celastraceae: Celastroideae

Kgaditsi, T.W. Senior Auxiliary Services Officer. Speci-

men mounter, general assistant

Klopper, Ms R.R. M.Sc. Senior Agricultural Scientist. Pte-

ridophyta and selected monocotyledonous families

Makgakga, M.C. B.Sc. Agricultural Development Tech-

nician. Wing B

Makgakga, K.S. Senior Auxiliary Services Officer. Her-

barium Assistant. Encoding plant specimens, data

capturing, labels typist

Manamela, Ms M.T. B.Sc.(Hons). Agricultural Develop-

ment Technician. Information Officer

Maserumule, M.K. Auxiliary Services Officer II. Wing B

Masombuka. Ms A.S. Senior Auxiliary Services Officer.

Herbarium Assistant. Wing A

Meyer. J.J. H.E.D. Chief Agricultural Development Tech-

nician. Bioprospecting Project

Mothapo, M.A. Senior Auxiliary Services Officer

Nkoane, Ms G.K. Senior Auxiliary Services Officer. Loans,

exchanges, gifts, parcelling, stores

Nkonki, Mrs T. B.Sc.(Hons). Senior Agricultural Scien-

tist. Fabaceae taxonomy. Wing B

Perold, Mrs S.M. Ph.D. Scientist. Taxonomy of Hepati-

cae (contract worker)

Phahla, T.J. Senior Auxiliary Services Officer. Specimen

mounter of cryptogams, packer

Ready, Mrs J.A. N.Dip.(Hort.). Principal Auxiliary Ser-

vices Officer. Plant identifications, Helichrysum.

Wing D

Rampho, Ms E.T. B.Sc. Senior Agricultural Development

Technician

Retief, Miss E. M.Sc. Principal Agricultural Scientist.

Taxonomy of Boraginaceae, Verbenaceae, Lami-

aceae, Asteraceae, Rubiaceae, Geraniaceae, Oxalid-

aceae, Vitaceae

Smithies, Mrs S.J. M.Sc., Dip.Ed.(Moray House). Chief

Agricultural Development Technician. Taxonomy

of Scrophulariaceae, Pedaliaceae, Bignoniaceae,

Lentibulariaceae, Gesneriaceae, Orobanchaceae

Steyn, Ms C.C. Principal Auxiliary Services Officer. Label

typist, service room support

Swelankomo, Ms N. B.Sc.(Hons). Senior Agricultural De-

velopment Technician. Taxonomy of gymnospenns,

Convolvulaceae

Welman, Ms W.G. M.Sc. Principal Agricultural Scientist.

Taxonomy of Convolvulaceae, Solanaceae, Cucurbit-

aceae, Asteraceae: Senecioneae, Acanthaceae

Winter, P.J.D. Principal Agricultural Scientist

DATA MANAGEMENT -PRETORIA (RPDC)

Arnold, T.H. M.Sc. Principal Data Technologist. Assistant Director.

Computer database application especially in taxonomy

Botha, Mrs A.G. Chief Auxiliary Services Officer. Ad-

ministrative Assistant

De Wet, Mrs B.C. B.Sc .(Computer Science), B. A., H.D.L.S.

Principal Agricultural Datametrician

Neveling. Mrs V.H. Principal Typist I

Smit, G.C. NT Workstation 4, NT Server 4. Chief Net-

work Controller

Snyman, Mrs E.E. B.Sc. N.Dip.(Comp. Data Proc.). Senior

Agricultural Development Technician

Steyn, Ms H.M. Botanical Information Officer (contract

worker)

162

SABONET— PRETORIA (YSGE/IS)

Steenkamp, Ms Y. Assistant Director. Regional Co-ordinator

Bothalia 34,2 (2004)

Malan, Ms E. Admin. Officer (contract worker)

Smith, Dr T. Project Assistant

Williams, V.J. (contract worker)

RESEARCH SUPPORT SERVICES— PRETORIA (EDIR)

Wolfson, Mrs M.M. Ph.D. Director. Physiology/Ecophysiology of Poaceae, carbon uptake metabolism,

allocation in response to environmental and management stress

Liebenberg, Mrs E.J.L. Head: Research Support Services, Publications

Mapiyeye, P. Project Co-ordinator, SAB SP (contract worker)

Naicker, K. Head: Admin., Human Resources and OHASA

Potgieter, Mrs E. Principal Librarian

Van Wyk, E. M.Sc. Project manager, Millenium Seed Bank Project

ADMINISTRATION, HUMAN RESOURCES AND OHASA -PRETORIA (RPTA)

Naicker, K. Dip.(Bookkeeping), Cert.(Sales & Market. Manag.),

H.Cert.(Prac. Accounting). Assistant Director

Bosheilo, M.S. Cleaner II

Khumalo, N.P. Senior Registry Clerk II

Ledwaba, Mrs D.M. Senior Registry Clerk I

Malefo, R.P. Cleaner II

Maphuta, Mrs M.S. Cleaner II

Marakalala, M.C. Senior Telekom Operator I

Marule, P.M. Aetisan

Nkosi, Mrs M.P. Specialist Cleaner

Phaahla, M.C. Cleaner II

Pretorius, Ms M.A. Senior Provisioning Admin. Clerk II

Ramsey, Y.K. Handyman. Building maintenance

Randima, Ms G.D. Cleaner II

Sithole, A.M. Cleaner II

Thibela, A. Senior Foreman. Supervisor: Cleaning ser-

vices

Tloubatla, J.M. Driver II. Courier services

PUBLICATIONS -PRETORIA (RPUB)

Liebenberg, Mrs EJ.L. M.Sc. Control Agricultural Technician. Cytotaxonomy. Manager

Brink, Mrs S.S. Dip. (Typing). Chief Typesetter. Typeset-

ting, layout, word processing

Condy, Ms G.S. M.A. Chief Industrial Technician. Botani-

cal artist

Du Plessis, Mrs E. B.Sc.(Hons), S.E.D. Chief Language

Practitioner. Technical editor. Editing, translating,

layout

Germishuizen, G. M.Sc. Assistant Director. Editor

Mapheza, T.P. Senior Provisioning Admin. Clerk III.

Bookshop Manager

Momberg, Mrs B.A. B. Sc. (Entomology & Zoology).

Principal Language Practitioner. Technical editor.

Editing, layout (part time)

Maree, Ms DJ. H.E.D. Senior Computer Operator.

Nkosi, P.B. Senior Provisioning Admin. Clerk I. Book-

store

Turck, Mrs S. B.A .(Information Design). Control Indus-

trial Technician. Graphic design

MARY GUNN LIBRARY -PRETORIA (RLBP)

Potgieter, Mrs E. B.Libr. Principal Librarian

Fourie, Mrs A.L. B.A., H.Dip.Libr.Sci. Principal Librarian (part time)

Shipalana, Ms K.M. N.Dip. Library Assistant II

MILLENIUM SEED BANK PROJECT -PRETORIA

Van Wyk, E. M.Sc. Principal Agricultural Scientist. Project manager

Klein, R.G. Plant Collector (contract worker)

Nkuna, L.A. B.Envir.Sc. Senior Agricultural Development Technician (contract worker)

Bothalia 34,2 (2004)

163

KIRSTENBOSCH RESEARCH CENTRE (RREL)

CAPE TOWN

Donaldson, J.S. Ph.D. (Zoology). Director

Morkel, Ms L. N.Dip.(Office Admin.). Senior Secretary IV. Personal Assistant to Director

COMPTON HERBARIUM-CAPE TOWN (RHEC)

Roux, J.P. N.T.C.III(Hort.), F.L.S., Ph.D. Deputy Director. Collections Manager.

Systematics of Pteridophyta

Manning, J.C. Ph.D. Senior Specialist Scientist. Research Leader, Systematics.

Systematics of Iridaceae and Hyacinthaceae; anatomy

Chesselet, Ms P.C.M. M.Sc. Principal Agricultural Scientist

Cupido, C.N. M.Sc. Principal Agricultural Scientist

Cupido, Mrs C.S. Senior Auxiliary Services Officer II.

Technical Assistant

Davids, Ms N. Data Capturer (contract worker)

Engelbrecht. Ms M. Data Capturer (contract worker)

Foster, Mrs S.E. Senior Secretary IV

Kurzweil, H. Ph.D. Specialist Scientist. Systematics of

southern African terrestrial orchids

Leith, Mrs J. Senior Provisioning Admin. Clerk III

Marinus, Ms E.D.A. Chief Auxiliary Services Officer.

Herbarium Assistant

Paterson-Jones, Mrs D.A. (nee Snijman) Ph.D. U.E.D.

Specialist Scientist. Systematics of Hypoxidaceae;

cladistics

Smith, Mrs M. Data Capturer (contract worker)

Williams, Mrs V.J. Data Capturer (contract worker)

CLIMATE CHANGE

Midgley, G.F. Ph.D. Principal Specialist Scientist. Plant ecophysiology, modelling

Hughes, G.O. GIS Researcher (contract worker)

Kgope, B.S. M.Sc. Principal Agricultural Scientist. Plant ecophysiology

Motete, Ms N. B.Sc.(Bot. & Ed.), M.Sc.(Envir. Biol.), Dip.(Science Ed.).

Senior Agricultural Scientist

CONSERVATION BIOLOGY

Donaldson, J.S. Ph.D.(Zoology). Cycad biology

Bosenberg, J. de Wet. B.Sc.(Hons). Chief Agricultural De-

velopment Technician. Cycad biology. Conserva-

tion farming

Ebrahim, I. N.Dip.(Hort.). Custodians for Rare and En-

dangered Flowers Programme (CREW) (contract

worker)

Marinus, E.M. N.Cert.(Building & Structures). Principal

Auxilliary Services Officer. Conservation farming

Nanni, Ms I. B.Sc., H.E.D. Control Agricultural De-

velopment Technician. Project Co-ordinator

Petersen, Ms A. B.Sc. (Hons). Senior Agricultural Develop-

ment Technician. Land use and vegetation map-

ping

Raimondo, Ms D. Manager: Custodians for Rare and En-

dangered Flowers Programme (CREW) (contract

worker)

Rouget, M.J.F. Ph.D. GIS Manager. Spatial modelling

(contract worker)

INVASIVE SPECIES PROGRAMME (YISP/CT)

Musil, C.F. Ph.D. Senior Specialist Scientist. Ecophysiology, modelling

Arnolds, Ms J.L. Chief Auxiliary Services Officer

Brown, N.A.C. Ph.D. Specialist Scientist. Seed research

Parker-Allie, Ms F. (nee Parker) M.Sc. Agricultural Scientist

Snyders, S.G. Senior Auxiliary Services Officer II. Greenhouse, maintenance

LANDSCAPE ECOLOGY

Rutherford, M.C. Ph.D., Dip.(Datamet.). Deputy Director. Modelling, global change

Parenzee, Ms H.A. Dip.(Ed.). Senior Provisioning Admin. Clerk III

Powrie, L.W. M.Sc. Chief Information Technology Advisor. Spatial modelling, databases

Rebelo, A.G. Ph.D .(Zoology). Principal Agricultural Scientist. Protea Atlas Project

164

Bothalia 34,2 (2004)

HARRY MOLTENO LIBRARY (RRLC)

Rutherford, M.C. Ph.D., Dip.(Datamet.). Deputy Director

Jagger, B.W. B.A.(Soc. Sc.), PGDip.Lis. Librarian

Ovens, Ms C.S.H. Ph.D.(Inf.Sc.), Dip.Datametrics. (contract librarian)

Reynolds, Ms P.Y. M. A .(Inf .Sc.), B.Proc., Dip.Datametrics. Principal Librarian.

NBI Web Site Manager

LESLIE HILL MOLECULAR SYSTEMATICS LABORATORY

Reeves, Ms G. Ph.D. Senior Agricultural Scientist (contract worker)

Balele, Ms K. Scientific Officer. DNA bank manager (contract worker)

Conrad, Ms C. M.Sc. Principal Agricultural Scientist

Khunou, Ms A. Agricultural Scientist. ALLP Manager

Mabunda, A. B.Sc.(Hons). NBI Masters student (contract worker)

INFORMATION TECHNOLOGY (SOUTH) (RRIT)

Evans, N. Network Controller. Technical IT Manager

Pekeur, Ms B.L. IT Admin. Clerk (contract worker)

SUPPORT SERVICES

Bardien-Overmeyer, Ms S. B.A.(Pharm.). Senior Provisioning Admin. Officer. Admin. Manager

Boonzaaier, I. Groundsman II. Maintenance

Bowler, Mrs M. Admin. Aid II. Cleaner. Assistant: teas and functions

De Witt, D.M. Senior Artisan (B-Group). Maintenance

PUBLICATIONS BY THE STAFF

1 April 2003 to 31 March 2004

ADAMS, T. 2003-06 .Serruria cyanoides (L.) R.Br. (Proteaceae). Internet

3 pp. http://www.plantzafrica.com/plantqrs/serruriacyan.htm.

ADAMS. T. 2003-08. Leucadendron levisanus (L.) P.J.Bergius (Protea-

ceae). Internet 3 pp.

http://www.plantzafrica.com/plantklm/leucadenlevis.htm.

ADAMS, T. 2003-10. Serruria furcellata R.Br. (Proteaceae). Internet 2

pp. http://www.plantzafrica.com/plantqrs/serrurfurcell.htm.

ADAMS, T. 2003-10. Serruria triloplia Salisb. ex Knight (Proteaceae). In-

ternet 2 pp. http://www.plantzafrica.com/plantqrs/seiTurtriloph.htm.

ANDERSON, J.M. & ANDERSON, H.M. 2003. Heyday of the gym-

nosperms: systematics and biodiversity of the Late Triassic

Molteno fructifications. Strelitzia 15.

ANON. (RUTHERFORD. M.C. & POWRIE, L.W.) 2004. Biological

diversity (contributions to). In D. Burger, South African Year-

book 2003/04: 235-239. Government Communications, Pretoria.

ANON. (RUTHERFORD, M.C. & POWRIE, L.W.) 2004. Mapping

South Africa’s floral wealth. In Anonymous, The greening of a

nation 1994-2004: 10. National Botanical Institute, Cape Town.

ARCHER, C. 2003. Agapanthaceae. In G. Germishuizen & N.L. Meyer,

Plants of southern Africa: an annotated checklist. Strelitzia 14:

954.

ARCHER, C. 2003. Alliaceae. In G. Germishuizen & N.L. Meyer, Plants

of southern Africa: an annotated checklist. Strelitzia 1 4: 956, 957.

ARCHER, C. 2003. Anthericaceae. In G. Germishuizen & N.L. Meyer,

Plants of southern Africa: an annotated checklist. Strelitzia 14:

967-97 1 .

ARCHER, C. 2003. Burmanniaceae, Cannaceae, Colchicaceae. In G.

Germishuizen & N.L. Meyer, Plants of southern Africa: an anno-

tated checklist. Strelitzia 14: 1013-1017.

ARCHER, C. 2003. Cyperaccae. In G. Germishuizen & N.L. Meyer,

Plants of southern Africa: an annotated checklist. Strelitzia 14:

1020-1047.

ARCHER, C. 2003. Eriospermaceae, Flagellariaceae, Hemerocallid-

aceac. In G. Germishuizen & N.L. Meyer, Plants of southern

Africa: an annotated checklist. Strelitzia 14: 1050-1054.

ARCHER, C. 2003. Musaceae. In G. Germishuizen & N.L. Meyer,

Plants of southern Africa: an annotated checklist. Strelitzia 14:

1120.

ARCHER, C. 2004. New typifications. In S. Cafferty & C.E. Jarvis, Typi-

fication of Linnaean plant names in Cyperaceae. Taxon 53: 179,

180.

ARCHER, C. 2003. Restionaceae, Strelitziaceae. In G. Germishuizen

& N.L. Meyer, Plants of southern Africa: an annotated check-

list. Strelitzia 14: 1195-1212.

ARCHER, C. 2003. Zingiberaceae. In G. Germishuizen & N.L. Meyer,

Plants of southern Africa: an annotated checklist. Strelitzia 14:

1215.

ARCHER, C. 2003. Review: Clivias, by Harold Koopowitz. Bothalia

33: 149, 150.

ARCHER, C. 2004. Four Schoenus names. In S. Cafferty & C.E. Jarvis,

Typification of Linnean plant names in Cyperaceae. Taxon 53:

177-181.

ARCHER, R.H. 2003. Abstract: Elaeodendron , biogeography, and the

Elaeodendron complex (Celastraceae) in Madagascar. Paper

presented at the 29th Annual Congress of the South African

Association of Botanists at the University of Pretoria, 8-11

January 2003. South African Journal of Botany 69: 225.

ARCHER, R.H. 2003. Balsaminaceae. In G. Germishuizen & N.L.

Meyer. Plants of southern Africa: an annotated checklist. Strelitzia

14: 311.

ARCHER, R.H. 2003. Burseraceae. In G. Germishuizen & N.L. Meyer,

Plants of southern Africa: an annotated checklist. Strelitzia 14:

333-335.

ARCHER, R.H. 2003. Buxaceae. In G. Germishuizen & N.L. Meyer,

Plants of southern Africa: an annotated checklist. Strelitzia 14:

335.

ARCHER, R.FI. 2003. Cornaceae. In G. Germishuizen & N.L. Meyer,

Plants of southern Africa: an annotated checklist. Strelitzia 14:

378.

ARCHER. R.H. 2003. Dichapetalaceae. In G. Germishuizen & N.L.

Meyer, Plants of southern Africa: an annotated checklist. Strelitzia

14: 418.

Bothalia 34,2 (2004)

165

ARCHER, R.H. 2003. Dioscoreaceae. In G. Germishuizen & N.L. Meyer,

Plants of southern Africa: an annotated checklist. Strelitzia 14:

1047, 1048.

ARCHER. R.H. 2003. Dracaenaceae. In G. Germishuizen & N.L. Meyer,

Plants of southern Africa: an annotated checklist. Strelitzia 14:

1048, 1049.

ARCHER, R.H. 2003. Euphorbiaceae. In G. Germishuizen & N.L.

Meyer, Plants of southern Africa: an annotated checklist. Strelitzia

14: 452-472.

ARCHER. R.H. 2003. Greyiaceae. In G. Germishuizen & N.L. Meyer,

Plants of southern Africa: an annotated checklist. Strelitzia 14:

581.

ARCHER, R.H. 2003. Haemodoraceae. In G. Germishuizen & N.L.

Meyer. Plants of southern Africa: an annotated checklist. Strelitzia

14: 1053, 1054.

ARCHER, R.H. 2003. Icacinaceae. In G. Germishuizen & N.L. Meyer,

Plants of southern Africa: an annotated checklist. Strelitzia 14:

584.

ARCHER. R.H. 2003. Kirkiaceae. In G. Germishuizen & N.L. Meyer,

Plants of southern Africa: an annotated checklist. Strelitzia 14:

584.

ARCHER, R.H. 2003. Lanariaceae. In G. Germishuizen & N.L. Meyer,

Plants of southern Africa: an annotated checklist. Strelitzia 14:

1119.

ARCHER. R.H. 2003. Malpighiaceae. In G. Germishuizen & N.L.

Meyer, Plants of southern Africa: an annotated checklist.

Strelitzia 14: 618.619.

ARCHER. R.H. 2003. Meliaceae, Melianthaceae. In G. Germishuizen

& N.L. Meyer. Plants of southern Africa: an annotated check-

list. Strelitzia 14: 629-631.

ARCHER. R.H. 2003. Ochnaceae. In G. Germishuizen & N.L. Meyer.

Plants of southern Africa: an annotated checklist. Strelitzia 14:

751.

ARCHER, R.H. 2003. Oliniaceae. In G. Germishuizen & N.L. Meyer,

Plants of southern Africa: an annotated checklist. Strelitzia 14:

753.

ARCHER. R.H. 2003. Passifloraceae. In G. Germishuizen & N.L.

Meyer. Plants of southern Africa: an annotated checklist. Strelitzia

14: 771,772.

ARCHER. R.H. 2003. Sapindaceae. In G. Germishuizen & N.L. Meyer.

Plants of southern Africa: an annotated checklist. Strelitzia 14:

857-861.

ARCHER, R.H. 2003. Tecophilaeaceae, Velloziaceae. In G. Germis-

huizen & N.L. Meyer, Plants of southern Africa: an annotated

checklist. Strelitzia 14: 1212-1214.

ARCHER. R.H. 2003. News from South Africa: reports on herbarium

internships to Natal Herbarium. SABONET News 8: 48, 49.

ARCHER. R.H. & GLEN. R.P. 2003. Callitrichaceae In G. Germis-

huizen & N.L. Meyer, Plants of southern Africa: an annotated

checklist. Strelitzia 14: 336.

ARCHER. R.H. & JORDAAN, M. 2003. Celastraceae. In G. Germis-

huizen & N.L. Meyer, Plants of southern Africa: an annotated

checklist. Strelitzia 14: 354—361.

ARCHER. R.H. & RETIEF. E. 2003. Anacardiaceae. In G. Germis-

huizen & N.L. Meyer, Plants of southern Africa: an annotated

checklist. Strelitzia 14: 115-122.

ARCHER. R.H., VICTOR. J.E. & CONDY. G. (Artist). 2003. Hoodia

pilifera subsp. pillansii. Curtis's Botanical Magazine 20: 219—

224, t. 479.

AUBREY. A. & DLAMINI. M.D. 2003-09. Nymphoides indica (L.)

Kuntze (Menyanthaceae). Internet 2 pp.

http://www.plantzafrica.com/plantnop/nymphindic.htm.

BEZUIDENHOUT, R. & BURGOYNE. P. 2004. Unraveling the secrets

of the Succulent Karoo. Farmer’s Weekly 20 February: 38.

BLOCKEEL.T.L.. ABAY, G., £ETIN. B.. BEDNAREK-OCHYRA. H..

OCHYRA, R„ LEWIS SMITH. R.I., MATTERI, C.M., FARIAS,

R.M.. NOVOTNY, I., RAO. P„ ENROTH. J„ VAN ROOY, J„

SCHIAVONE. M.M. & VANA. J. 2003. New national and re-

gional bryophyte records, 1 .Journal of Bryology 25: 141-144.

BREDENKAMP. C.L. 2003. Achariaceae. In G. Germishuizen & N.L.

Meyer, Plants of southern Africa: an annotated checklist. Strelitzia

14: 106.

BREDENKAMP. C.L. 2003. Begoniaceae. In G. Germishuizen & N.L.

Meyer, Plants of southern Africa: an annotated checklist. Strelitzia

14: 311,312.

BREDENKAMP. C.L. 2003. Canellaceae. In G. Germishuizen & N.L.

Meyer, Plants of southern Africa: an annotated checklist. Strelitzia

14: 346.

BREDENKAMP, C.L. 2003. Clusiaceae. In G. Germishuizen & N.L.

Meyer, Plants of southern Africa: an annotated checklist. Strelitzia

14: 369.

BREDENKAMP, C.L. 2003. Elatinaceae. In G. Germishuizen & N.L.

Meyer, Plants of southern Africa: an annotated checklist. Strelitzia

14: 424.

BREDENKAMP, C.L. 2003. Flacourtiaceae. Frankeniaceae, Geissolo-

mataceae. In G. Germishuizen & N.L. Meyer, Plants of southern

Africa: an annotated checklist. Strelitzia 14: 559-562.

BREDENKAMP, C.L. 2003. Gunneraceae. Haloragaceae, Hypericaceae.

In G. Germishuizen & N.L. Meyer, Plants of southern Africa:

an annotated checklist. Strelitzia 14: 581-583.

BREDENKAMP. C.L. 2003. Lecythidaceae. In G. Germishuizen &

N.L. Meyer, Plants of southern Africa: an annotated checklist.

Strelitzia 14: 601.

BREDENKAMP, C.L. 2003. Linaceae, Loasaceae. In G. Germishuizen

& N.L. Meyer, Plants of southern Africa: an annotated check-

list. Strelitzia 14: 603, 604.

BREDENKAMP. C.L. 2003. Lythraceae, Maesaceae. In G. Germis-

huizen & N.L. Meyer, Plants of southern Africa: an annotated

checklist. Strelitzia 14: 616-618.

BREDENKAMP. C.L. 2003. Melastomataceae. In G. Germishuizen &

N.L. Meyer, Plants of southern Africa: an annotated checklist.

Strelitzia 14: 628,629.

BREDENKAMP. C.L. 2003. Menyanthaceae. In G. Germishuizen &

N.L. Meyer, Plants of southern Africa: an annotated checklist.

Strelitzia 14: 633.

BREDENKAMP. C.L. 2003. Onagraceae. In G. Germishuizen & N.L.

Meyer, Plants of southern Africa: an annotated checklist. Strelitzia

14: 753.

BREDENKAMP. C.L. 2003. Plumbaginaceae, Polygalaceae. In G. Ger-

mishuizen & N.L. Meyer, Plants of southern Africa: an anno-

tated checklist. Strelitzia 14: 778-788.

BREDENKAMP. C.L. 2003. Rhizophoraceae. Rhynchocalycaceae. In

G. Germishuizen & N.L. Meyer, Plants of southern Africa: an

annotated checklist. Strelitzia 14: 817.

BREDENKAMP. C.L. 2003. Sterculiaceae, Tamaricaceae. In G. Ger-

mishuizen & N.L. Meyer, Plants of southern Africa: an anno-

tated checklist. Strelitzia 14: 918-927.

BREDENKAMP, C.L. 2003. Turneraceae. In G. Germishuizen & N.L.

Meyer, Plants of southern Africa: an annotated checklist. Strelitzia

14: 938.

BREDENKAMP. C.L. 2003. Violaceae. In G. Germishuizen & N.L.

Meyer, Plants of southern Africa: an annotated checklist. Strelitzia

14: 944.

BREDENKAMP, C.L. & BEYERS, J.B.P. 2003. Thymelaeaceae. In G.

Germishuizen & N.L. Meyer, Plants of southern Africa: an

annotated checklist. Strelitzia 14: 928-935.

BREDENKAMP. C.L. & GOVENDER, N. 2003. Gentianaceae. In G.

Germishuizen & N.L. Meyer, Plants of southern Africa: an

annotated checklist. Strelitzia 14: 562-565.

BREDENKAMP, C.L. & JORDAAN, M. 2003. Tiliaceae. In G. Ger-

mishuizen & N.L. Meyer, Plants of southern Africa: an anno-

tated checklist. Strelitzia 14: 935-937.

BREDENKAMP, C.L. & LEISTNER, O.A. 2003. Malvaceae. In G.

Germishuizen & N.L. Meyer. Plants of southern Africa: an

annotated checklist. Strelitzia 14: 619-628.

BREDENKAMP. C.L. & MANAMELA.T. 2003. Working together for

the Flora of the Eastern Cape. Veld & Flora 89: 138, 139.

BREDENKAMP. G.J., SIEBERT. F„ BROWN. L.R. & SMITH. G.F.

2003. Abstract: The vegetation of Gauteng. Paper presented at

the 29th Annual Congress of the South African Association of

Botanists at the University of Pretoria, 8-11 January 2003.

South African Journal of Botany 69: 226.

BREDENKAMP, C.L., SMITH, G.F. & BREDENKAMP, G.J. 2003.

Abstract: Flora of the Eastern Cape. Paper presented at the 29th

Annual Congress of the South African Association of Botanists

at the University of Pretoria, 8-11 January 2003. South African

Journal of Botany 69: 226.

BREDENKAMP. C.L. & VAN WYK, A.E. 2003. Taxonomy of the

genus Passerina (Thymelaeaceae). Bothalia 33: 59-98.

BREUER. L, STEYN. E.M.A. & SMITH, G.F. 2003. Haworthia. In G.

Germishuizen & N.L. Meyer, Plants of southern Africa: an

annotated checklist. Strelitzia 14: 997-1008.

BROWN, N.A.C.. VAN STADEN, J.. DAWS, M.I. & JOHNSON. T.

2003. Patterns in the seed germination response to smoke in

plants from the Cape Floristic Region, South Africa. South

African Journal of Botany 69: 514-525.

BROWN, N.A.C., VAN STADEN, J.. JOHNSON, T. & DAWS, M.I.

2003. A summary of patterns in the seed germination response

166

Bothalia 34,2 (2004)

to smoke in plants from the Cape Floral Region. In H. Pritchard

et at. Proceedings of the Kew Workshop, Seed Conservation :

Turning science into practice, Wakehurst Place, July 2001:

563-574.

BURGOYNE, P.M. 2003. Abstract: Character delimitations in the fam-

ily Mesembryanthemaceae: Delosperma, a case study. Paper

presented at the XVIIth AETFAT Congress, 21-26 September

2003, Addis Ababa, Ethiopia. In S. Demissew, E. Kelbessa & S.

Edwards , Abstracts/resumes: 112, 1 13. Addis Ababa University

Press.

BURGOYNE, P.M. 2003. Crassulaceae ( Tylecodon by P. Bruyns). In

G. Germishuizen & N.L. Meyer, Plants of southern Africa: an

annotated checklist. Strelitzia 14: 378 — 41 3 .

BURGOYNE, P.M. 2003. Santalaceae. In G. Germishuizen & N.L.

Meyer, Plants of southern Africa: an annotated checklist. Strelitzia

14: 853-859.

BURGOYNE, P.M. 2003. The Delosperma species of central southern

Africa— a call to collectors. PlantLife 29: 8-14.

BURGOYNE. P.M. 2004-01. Frithia pulchra. Internet 4 pp.

http://www.plantzafrica.com .

BURGOYNE, P.M. 2004-02. Frithia humilis. Internet 4 pp.

http ://w ww.pl antzafrica.com .

BURGOYNE. P.M. & VICTOR. I.E. 2003. Delosperma: taxonomic

notes and proposed Red List status of certain species. Aloe 40:

93-97.

BUYS, M.H.. BURGOYNE, P.M. & TIEDT, L. 2003. Abstract: Paly-

nology and the Stomatuim group in the Aizoaceae. Poster pre-

sented at the XVIIth AETFAT Congress. 21-26 September 2003,

Addis Ababa, Ethiopia. In S. Demissew, E. Kelbessa & S. Edwards,

Abstracts! resumes: 23, 24. Addis Ababa University Press.

CAROLUS, B. 2003-07. Erica perspicua J.C.Wendl (Ericaceae).

Internet 3 pp.

http://www.plantzafrica.com/plantefg/ericaperspic.htm.

CAROLUS, B. 2003-11. Satyrium carneum (Dryand.) Sims

(Orchidaceae). Internet 2 pp.

http://www.plantzafrica.com/plantqrs/satyrcarn.htm.

CHESSELET, P. 2003. Sciences: Halte a l’apartheid vegetal! Le Point

1623: 84,85.

CHESSELET, P. 2004-03. Mesembryanthemaceae. Internet 4 pp.

http://w w w.plantzafrica .com .

CHESSELET. P & ANDERSON, F. (Artist). 2003. Dorotheanthus

rourkei (Mesembryanthemaceae). Flowering Plants of Africa

58: 70-73, t. 2191.

CHESSELET, P, BURGOYNE, P.M., KLAK, C„ KURZWEIL, H„

DOLD, A.P., GRIFFIN, N.J. & SMITH, G.F. 2003.

Mesembryanthemaceae. In G. Germishuizen & N.L. Meyer,

Plants of southern Africa: an annotated checklist. Strelitzia 14:

633-738. National Botanical Institute, Pretoria.

CHESSELET, P, VAN WYK, A.E., GRIFFIN, N. & SMITH, G.F.

2003. Patterns of floristic diversity in Mesembryanthemaceae.

Aloe 40: 80-85.

CHIMPHANGO, S.B.M.. MUSIL, C.F. & DAKORA, F.D. 2003.

Response of purely symbiotic and N03-fed nodulated plants of

Lupinus luteus and Vicia atropurpurea to ultraviolet-B radia-

tion. Journal of Experimental Botany 54: 1771-1784.

CHIMPHANGO. S.B.M., MUSIL, C.F. & DAKORA, F.D. 2003.

Response to ultraviolet-B radiation by purely symbiotic and

NOi-fed nodulated tree and shrub legumes indigenous to south-

ern Africa. Tree Physiology 24: 181-192.

CLEEVELY, R.J.. NELSON, EC. & OLIVER, E.G.H. 2003. More

accurate publication dates for H.C. Andrews’ The Heatheiy,

particularly Volumes 5 and 6. Bothalia 33: 195-198.

CONRAD, F„ REEVES, G. & ROURKE, J.P. 2003. Phylogenetic rela-

tionships of the recently discovered species — Clivia mirabilis.

South African Journal of Botany 69: 204—206.

CROUCH, N.R., MULHOLLAND, D.A., POHL, T.L. & NDLOVU, E.

2003. The ethnobotany and chemistry of the genus Clivia (Ama-

ryllidaceae). South African Journal of Botany 69: 144—147.

CROUCH, N.R., NDLOVU, E„ MULHOLLAND, D.A. & POHL, T.L.

2003. The genus Clivia in ethnomedicine: usage, bioactivity

and phytochemistry. Clivia 5: 15-22.

CROUCH, N.R., POTGIETER, C„ MILLER, R.M., EDWARDS, T. &

SYMMONDS, R. 2003. Pollination of Siphonochilus aethiopi-

cus: facts and speculation. PlantLife 29: 18-21.

CROUCH, N.R., SMITH, G.F. & CONDY, G. (Artist). 2003. Siphono-

chilus aethiopicus (Zingiberaceae). Flowering Plants of Africa

58: 56-68, t. 2190.

CROUCH, N.R., SMITH, G.F., SYMMONDS, R. & CONDY, G. (Artist).

2003. Gerrardanthus tomentosus (Cucurbitaceae). Flowering

Plants of Africa 58: 134-142, t. 2200.

CUPIDO, C.N. 2003. Systematic studies in the genus Merciera (Cam-

panulaceae): a re-assessment of species boundaries. Adansonia,

ser. 3,25: 33-44.

CUPIDO, C.N. & CONRAD, F. 2003. Cyphia. A pretty perennial herb.

Veld & Flora 89: 62, 63.

DE BEER. Z.W., GLEN, H.F., WINGFIELD, B.D. & WINGFIELD,

M. J. 2003. Ophiostoma quercus or Ophiostoma quercil My co-

taxon 86: 211-214.

DE WIT, M.J. & ANDERSON, J.M. 2003. Gondwana Alive corridors:

extending Gondwana research to incorporate stemming the

sixth extinction. Gondwana Research 6,3: 369-408.

DLAMINI, M.D. (assisted by T.M. Mphaphuli). 2003-08. Crocosmia

aurea (Pappe ex Hook.) Planch. (Iridaceae). Internet 2 pp.

http://www.plantzafrica.com/plantcd/croscosaurea.htm.

DLAMINI, M.D. 2003-12. Kniphofia linearifolia Baker (Asphodela-

ceae). Internet 2 pp.

http://www.plantzafrica.com/plantklm/kniphoflin.htm.

DOLD, A.P & VICTOR. J.E. 2004. Notes on Brachystelma comptum

N. E.Br., an endangered species from the Eastern Cape Pro-

vince, South Africa. Aloe 41: 22, 23.

DOLD. A.P. & VICTOR, J.E. 2003. Fire lilies and hotspots. Veld &

Flora 89: 66,67.

DOLD. A.P. & VICTOR. J.E. 2003. Threatened plants at Coega: is ex

situ conservation a solution? African Wildlife 54: 22, 23.

DREYER, L.L. & MAKGAKGA, M.C. 2003. Oxalidaceae. In G.

Germishuizen & N.L. Meyer, Plants of southern Africa: an

annotated checklist. Strelitzia 14: 762-770.

DUNCAN, G.D. 2003. Ammocharis nerinoides (Amaryllidaceae).

Curtis's Botanical Magazine 20: 196-201.

DUNCAN, G.D. 2003. Cyrtanthus guthrieae (Amaryllidaceae).

Curtis's Botanical Magazine 20: 190-195.

DUNCAN, G.D. 2003. Endangered geophytes of the Cape Floral

Kingdom. Curtis’s Botanical Magazine 20: 245-250.

DUNCAN, G.D. 2003. Lachenalia peersii (Hyacinthaceae). Curtis's

Botanical Magazine 20: 202-207.

DUNCAN, G.D. 2003. Lachenalia salteri (Hyacinthaceae). Curtis’s

Botanical Magazine 20: 208-212.

DUNCAN, G.D. 2003. Lachenalia valeriae — an exciting new species

from Namaqualand, South Africa. Bulbs 5,1 : 3—4.

DUNCAN, G.D. 2003. Rhodohypoxis — carpets of alpine jewels. Veld

& Flora 89: 140-144.

DUNCAN, G.D. 2003. Some notes on Agapanthus . The Plantsman

New Series 2,3: 174-178.

DUNCAN, G.D. 2003. Strumaria barbarae— elegance in abundance.

Indigenous Bulb Growers Association of South Africa Bulletin

No. 52: 30-33.

DUNCAN, G.D. 2003-04. Polyxena corymbosa (L.) Jessop (Hyacinth-

aceae). Internet 3 pp.

http://www.plantzafrica.com/plantnop/polycorym.htm.

DUNCAN, G.D. 2003-05. Eulophia horsfallii (Batem.) Summerh.

(Orchidaceae). Internet 3 pp.

http://www.plantzafrica.com/plantefg/eulophhors.htm.

DUNCAN, G.D. 2003-06. Lachenalia viridiflora W.F.Barker (Hyacinth-

aceae). Internet 3 pp.

http://www.plantzafrica.com/plantklm/Iachvirid.htm.

DUNCAN, G.D. 2004-01 . Watsonia hysterantha J .W.Mathews & L.Bolus.

Internet 3 pp.

http://www.plantzafrica.com/plant/watsonia.htm.

DUNCAN, G.D. & ANDERSON, F. (Artist). 2003. Lachenalia abides

var. vanzyliae (Hyacinthaceae). Flowering Plants of Africa 58:

30-36, t. 2187.

DUNCAN. G.D. & ANDERSON, F. (Artist). 2003. Lachenalia valeriae

(Hyacinthaceae). Flowering Plants of Africa 58: 38-42, t. 2186.

FISH, L. 2003. Poaceae. In G. Germishuizen & N.L. Meyer, Plants of

southern Africa: an annotated checklist. Strelitzia 14: 1 152- 1194.

FISH, L. 2003. Review: AusGrass— grasses of Australia CD-ROM, by

D. Sharp & B. Simon, 2002. South African Journal of Botany

69: 452.

FISH, L. 2003. Abstract: Grass collection at the National Herbarium (PRE),

Pretoria. African Journal of Range & Forage Science 20: 182.

FORRESTER. J. 2003-11. Haemanthus coccineus L. (Amaryllida-

ceae). Internet 3 pp.

http://www.plantzafrica.com/planthij/haemancoc.htm.

GAUTIER, L„ SMITH, G.F., SPICHIGER, R, KLOPPER, R.R., SIEBERT,

S.J., CHATELAIN, C. 2003. Abstract: Merging tropical and

southern African flowering plant data: towards an African plant

database. Poster presented at the XVIIth AETFAT Congress,

21-26 September 2003, Addis Ababa, Ethiopia. In S. Demis-

Bothalia 34,2 (2004)

167

sew, E. Kelbessa & S. Edwards, Abstracts! resumes'. 130. 131.

Addis Ababa University Press.

GERMISHU1ZEN, G. 2003. Loranthaceae. In G. Germishuizen & N.L.

Meyer, Plants of southern Africa: an annotated checklist. Strelitzia

14: 612-616.

GERMISHUIZEN, G. 2003. Rhynchosia (Fabaceae). In G. Germis-

huizen & N.L. Meyer, Plants of southern Africa: an annotated

checklist. Strelitzia 14: 545-548.

GERMISHUIZEN, G. 2003. Viscaceae. In G. Germishuizen & N.L.

Meyer. Plants of southern Africa: an annotated checklist. Strelitzia

14: 945, 946.

GERMISHUIZEN, G. & CLARKE, B. (Artist). 2003. Illustrated guide

to the wildflowers of northern South Africa. Briza Publications

Pretoria.

GERMISHUIZEN, G. & GLEN, R.P. 2003. Polygonaceae. In G. Germis-

huizen & N.L. Meyer, Plants of southern Africa: an annotated

checklist. Strelitzia 14: 788-791.

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

Africa: an annotated checklist. Stelitzia 14. National Botanical

Institute, Pretoria.

GERMISHUIZEN. G. & RETIEF. E. 2003. 2003 South African Associa-

tion of Botanists Silver Medal Recipient. Dr Otto Leistner.

South African Journal of Botany 69: vi.

GLEN, H.F. 2003. Arecaceae. In G. Germishuizen & N.L. Meyer. Plants

of southern Africa: an annotated checklist. Strelitzia 14: 973.

GLEN, H.F. 2003. Cactaceae. In G. Germishuizen & N.L. Meyer. Plants

of southern Africa: an annotated checklist. Strelitzia 14: 335.

GLEN, H.F. 2003. Cupressaceae. Pinaceae, Podocarpaceae, Stangeri-

aceae, Welwitschiaceae. In G. Germishuizen & N.L. Meyer,

Plants of southern Africa: an annotated checklist. Strelitzia 14:

88,89.

GLEN. H.F. 2003. Fagaceae. In G. Germishuizen & N.L. Meyer. Plants

of southern Africa: an annotated checklist. Strelitzia 14: 559.

GLEN, H.F. 2003. Menispermaceae. In G. Germishuizen & N.L. Meyer,

Plants of southern Africa: an annotated checklist. Strelitzia 14:

631,632.

GLEN, H.F. 2003. Myricaceae. In G. Germishuizen & N.L. Meyer,

Plants of southern Africa: an annotated checklist. Strelitzia 14:

745,746.

GLEN. H.F. 2003. Neuradaceae. In G. Germishuizen & N.L. Meyer. Plants

of southern Africa: an annotated checklist. Strelitzia 14: 748, 749.

GLEN, H.F. 2003. News from South Africa: Pretoria visits Mozam-

bique. SABONET News 8: 50.

GLEN, H.F. 2003. Papaveraceae. In G. Germishuizen & N.L. Meyer,

Plants of southern Africa: an annotated checklist. Strelitzia 14: 770.

GLEN, H.F. 2003. Portulacaceae, Primulaceae. In G. Germishuizen &

N.L. Meyer, Plants of southern Africa: an annotated checklist.

Strelitzia 14: 792-795.

GLEN. H.F. 2003. Ranunculaceae. In G. Germishuizen & N.L Meyer,

Plants of southern Africa: an annotated checklist. Strelitzia 14:

809, 810.

GLEN, H.F. 2003. Review: Trees and shrubs of Mpumalanga and Kruger

National Park, by E. Schmidt, M. Lotter & W. McCleland, 2002.

SABONET News 8: 44, 45.

GLEN. H.F. 2003. Review: Generic tree flora of Madagascar, by G.E.

Schatz, 2001 . Plant Systematics and Evolution 241 : 158-160.

GLEN. H.F. 2003. Review: EUCLID: Eucalypts of southern Australia,

edn 2, by M.I.H. Brooker et al., 2002. Plant Systematics and

Evolution 241: 126-128.

GLEN. H.F. 2003.Theophrastaceae. In G. Germishuizen & N.L. Meyer,

Plants of southern Africa: an annotated checklist. Strelitzia 14:

928.

GLEN, H.F. 2003. What plants do we grow in southern Africa? In G.H.

Schmelzer & E.A. Omino, Plant resources of tropical Africa.

Proceedings of the First PROTA International Workshop,

23-25 September 2002, Nairobi, Kenya : 293, 294. PROTA

Foundation, Wageningen, the Netherlands.

GLEN. H.F. 2003. What’s in a name? Woody eyes and universal

lusticia. The Gardener Nov./Dec. 2003: 8. Garden Centre Asso-

ciation of South Africa.

GLEN. H.F., ARCHER, C. & COOK, C.D.K. 2003. Publication and

orthography of Pycreus mundii (Cyperaceae). Taxon 52: 601,

602.

GLEN, H.F. & HURTER. P.J.H. 2003. Zamiaceae. In G. Germishuizen

& N.L. Meyer. Plants of southern Africa: an annotated check-

list. Strelitzia 14: 89, 90.

GLEN. H.F. & IORDAAN, M. 2003. Brassicaceae. In G. Germis-

huizen & N.L. Meyer, Plants of southern Africa: an annotated

checklist. Strelitzia 14: 321-329.

GLEN, H.F. & JORDAAN, M. 2003. Myrtaceae. In G. Germishuizen

& N.L. Meyer, Plants of southern Africa: an annotated check-

list. Strelitzia 14: 747, 748.

GLEN. H.F. & NKONKI. T. 2003. Review: The Leguminosae of

Madagascar, by DJ. Du Puy et al., 2002. Plant Systematics and

Evolution 236: 232-234.

GLEN. H.F. & NKONKI. T. 2003. Review: Advances in legume sys-

tematics, by P.S. Herendeen & A. Bruneau (eds), 2000. Plant

Systematics and Evolution 237: 240-242.

GLEN, H.F. & NKONKI. T. 2003. Review: Flora Malesiana: Legumi-

nosae, Caesalpinoideae of south-east Asia, by J.W.A. Ridder-

Numan & I. de Kort (eds), 2002. Plant Systematics and Evolu-

tion 239: 297-299.

GLEN, H.F. & SIEBERT. S. The paper chase. SABONET News 8: 36-42.

GLEN, H.F. & SMITH, G.F. 2003. Abstract: Some perspectives on

tropical African resource plants: ornamentals. Paper presented

at the 29th Annual Congress of the South African Association

of Botanists at the University of Pretoria, 8-11 January 2003.

South African Journal of Botany 69: 231.

GLEN, H.F. & SMITH, G.F. 2003. PROTA 4: Ornamentals. In G.H.

Schmelzer & E.A. Omino, Plant resources of tropical Africa.

Proceedings of the First PROTA International Workshop,

23-25 September 2002, Nairobi. Kenya: 132-139. PROTA

Foundation, Wageningen, the Netherlands.

GLEN. H.F., SMITH. G.F.. BREUER, I., STEYN, E.M.A., ARCHER.

C„ MANNING, J.C.. WILLIAMSON, G„ VAN JAARSVELD.

E.J. & MEYER. N.L. 2003. Asphodelaceae. In G. Germis-

huizen & N.L. Meyer, Plants of southern Africa: an annotated

checklist. Strelitzia 14: 978-1012. National Botanical Institute,

Pretoria.

GLEN, R.P. 2003. Review: Wetland plants: biology and ecology, by

J.K. Kronk & M.S. Fennessy, 2001. Plant Systematics and

Evolution 241: 267 , 268.

GLEN. R.P. 2003. Behniaceae. In G. Germishuizen & N.L. Meyer, Plants

of southern Africa: an annotated checklist. Strelitzia 14: 1012.

GLEN. R.P. 2003. Smilacaceae. In G. Germishuizen & N.L. Meyer,

Plants of southern Africa: an annotated checklist. Strelitzia 14:

1212.

GLEN. R.P. & ARCHER, C. 2003. Liliaceae. In G. Germishuizen &

N.L. Meyer, Plants of southern Africa: an annotated checklist.

Strelitzia 14: 1120.

GLEN. R.P. & BREDENKAMP, C.L. 2003. Trapaceae. In G. Germis-

huizen & N.L. Meyer, Plants of southern Africa: an annotated

checklist. Strelitzia 14: 937.

GLEN. R.P. & COOK, C.D.K. 2003. Alismataceae. In G. Germis-

huizen & N.L. Meyer, Plants of southern Africa: an annotated

checklist. Strelitzia 14: 955.956.

GLEN. R.P. & COOK, C.D.K. 2003. Cabombaceae. In G. Germis-

huizen & N.L. Meyer, Plants of southern Africa: an annotated

checklist. Strelitzia 14: 335.

GLEN, R.P. & COOK. C.D.K. 2003. Ceratophyllaceae. In G. Germis-

huizen & N.L. Meyer, Plants of southern Africa: an annotated

checklist. Strelitzia 14: 362.

GLEN. R.P. & COOK, C.D.K. 2003. Cymodoceaceae. In G. Germis-

huizen & N.L. Meyer, Plants of southern Africa: an annotated

checklist. Strelitzia 14: 1019.

GLEN, R.P. & COOK, C.D.K. 2003. Eriocaulaceae. In G. Germis-

huizen & N.L. Meyer, Plants of southern Africa: an annotated

checklist. Strelitzia 14: 1049, 1050.

GLEN, R.P. & COOK, C.D.K. 2003. Hydrocharitaceae. In G. Germis-

huizen & N.L. Meyer, Plants of southern Africa: an annotated

checklist. Strelitzia 14: 1071.

GLEN, R.P. & COOK. C.D.K. 2003. Hydrostachyaceae. In G. Germis-

huizen & N.L. Meyer, Plants of southern Africa: an annotated

checklist. Strelitzia 14: 583.

GLEN, R.P. & COOK, C.D.K. 2003. Juncaceae, Juncaginaceae.Lemna-

ceae, Limnocharitaceae, Najadaceae. In G. Germishuizen &