Bothalia

A JOURNAL OF BOTANICAL RESEARCH

Vol. 32,1

May 2002

TECHNICAL PUBLICATIONS OF THE NATIONAL BOTANICAL INSTITUTE,

PRETORIA

Obtainable from the National Botanical Institute, Private Bag XI 01, Pretoria 0001, Republic of

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

BOTHALIA

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

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

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

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

published annually.

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

STRELITZIA

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

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

MEMOIRS OF THE BOTANICAL SURVEY OF SOUTH AFRICA

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

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

ANNALS OF KIRSTENBOSCH BOTANIC GARDENS

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

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

South African Botany). Discontinued after Vol. 19.

FLOWERING PLANTS OF AFRICA (FPA)

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

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

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

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

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

or of economic value for illustration.

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

An index to Vols 1-49 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. I. Introduction. Dicroidium, 1983, by .I.M. & H.M.

Anderson.

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

& H.M. Anderson.

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

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

RSA.

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

by J.M. Anderson (ed.)

BOTHALIA

A JOURNAL OF BOTANICAL RESEARCH

Volume 32,1

Scientific Editor: G. Germishuizen

Technical Editor: B.A. Momberg

NATIONAL

Botanical

INSTITUTE

2 Cussonia Avenue, Brummeria, Pretoria

Private Bag XI 01, Pretoria 0001

ISSN 0006 8241

May 2002

Editorial Board

D.F. Cutler

B.J. Huntley

P.H. Raven

J.P. Rourke

M.J. Werger

Royal Botanic Gardens, Kew, UK

National Botanical Institute, Cape Town, RSA

Missouri Botanical Garden, St Louis, USA

Compton Herbarium, NBI, Cape Town, RSA

University of Utrecht, Utrecht, Netherlands

Acknowledgements to referees

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

Barker, Dr N.P. Rhodes University, Grahamstown, RSA.

Bayer, M.B. 16 Hope St, Gardens, 8001 Cape Town, RSA.

Beyers, Dr J.B.P. National Botanical Institute, Cape Town, RSA.

Braggins, Dr J.E. University of Auckland, New Zealand.

Brandham, Dr P.E. Royal Botanic Gardens, Kew, UK.

Bredenkamp, Mrs C.L. National Botanical Institute, Pretoria, RSA.

Bredenkamp, Prof. G. University of Pretoria, RSA.

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

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

Burrows, J.E. Buffelskloof Private Nature Reserve, Lydenburg, RSA.

De Kok, Dr R. CSIRO Plant Industry, Canberra, Australia.

De Silva, Dr P.H.A.U. University of Ruhuna, Matara, Sri Lanka.

Dreyer, Dr L. University of Stellenbosch, RSA.

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

Ellis, Dr R.P Grassland Research Centre, Agricultural Council, Pretoria, RSA.

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

Glen, Mrs R.P. National Botanical Institute, Pretoria, RSA.

Goldblatt, Dr P. Missouri Botanical Garden, St Louis, USA.

Hansen, Dr H.V. University of Copenhagen, Denmark.

Henderson, Ms L. Plant Protection Research Institute, Agricultural Research Council, Pretoria, RSA.

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

Hilu, Dr K.W. Polytechnic & State University, Blacksburg, Virginia, USA.

Killick, Dr D.J.B. 465 Sappers Contour, Lynnwood, 0081 Pretoria, RSA.

Lassen, Dr P. Botanical Museum, Lund, Sweden.

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

Lowrey, Dr T.K. University of New Mexico, Albuquerque, USA.

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

Midgley, Dr G.F. National Botanical Institute, Cape Town, RSA.

Moll, Prof. E.J. University of Queensland, Australia.

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

Nevling, Dr L. Illinois Natural History Survey, Champaign, USA.

Nilsson, Prof. Dr S. Swedish Museum of Natural History, Stockholm, Sweden.

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

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

Peckover, R. Pyramid, RSA.

Phillipson, Dr PB. Rhodes University, Grahamstown, RSA.

Poes, Prof. T. Esterhazy College, Eger, Hungary.

Rourke, Dr J.P. National Botanical Institute, Cape Town, RSA.

Schutte-Vlok, Dr A. P.O. Box 1512, 6620 Oudtshoorn, RSA.

Scott-Shaw, R. Biodiversity Publications, Cascades, KwaZulu-Natal, RSA.

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

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

Volleson, Dr K. Royal Botanic Gardens, Kew, UK.

Wclman, Ms W.E. National Botanical Institute, Pretoria, RSA.

CONTENTS

Volume 32,1

1. Clivia mirabilis (Amaryllidaceae: Haemantheae) a new species from Northern Cape, South Africa. J.P.

ROURKE 1

2. The genus Buglossoides (Boraginaceae) in southern Africa. E. RETIEF and A.E. VAN WYK 9

3. Studies in the liverwort family Aneuraceae (Metzgeriales) from southern Africa. 3. Riccardia compacta.

S.M. PEROLD 15

4. Revision of the Tarchonanthus camphoratus complex (Asteraceae-Tarchonantheae) in southern Africa.

P.PJ. HERMAN 21

5. Taxonomy of the Passerina fdifonnis L. complex (Thymelaeaceae). C.L. BREDENKAMP and A.E.

VAN WYK 29

6. The genus Erica (Ericaceae) in southern Africa: taxonomic notes 1. E.G.H. OLIVER and I.M. OLIVER 37

7. Notes on African plants:

Apocynaceae. A new species of Brachystelma from Eastern Cape, South Africa. A.P DOLD .... 71

Campanulaceae. A new species of Merciera from Western Cape. South Africa. C.N. CUPIDO . . 74

Eriospermaceae. Validation of two sectional names in Eriospermum. J. THIEDE 76

Hyacinthaceae. A new combination in Daubenya. J.C. MANNING and A.M. VAN DER MERWE 63

Lamiaceae. New combinations in the genus Rotheca in southern Africa. P.P.J. HERMAN and E.

RETIEF 81

Marsileaceae-Pteropsida. First report of the genus Pilularia from continental Africa. J.P. ROUX 82

Thymelaeaceae. Systematics of Passerina truncata and a new subspecies monticola. C.L. BRE-

DENKAMP and A.E. VAN WYK 65

Thymelaeaceae. A new species of Passerina from Western Cape, South Africa. C.L. BREDEN-

KAMP and A.E. VAN WYK 76

Thymelaeaceae. A new species of Gnidia from the Knersvlakte, Western Cape, South Africa.

J.B.P. BEYERS 79

8. The taxonomic significance of trichome type and distribution in Melolobium (Fabaceae). A. MOTEE-

TEE, B-E. VAN WYK and PM. TILNEY 85

9. Pollen morphology and biometry of the genus Androcymbium (Colchicaceae) in southern Africa: taxono-

mic and biogeographic considerations. N. MEMBRIVES, J. MARTIN, J. CAUJAPE-CASTELLS

and J. PEDROLA-MONFORT 91

10. Biogeography of Oxalis (Oxalidaceae) in South Africa: a preliminary study. K.C. OBERLANDER, L.L.

DREYER and K.J. ESLER 97

11. A checklist of the plants of Mahwaqa Mountain, KwaZulu-Natal. E.B. METER, T.J. EDWARDS, M.A.

RENNIE and J.E. GRANGER 101

12. Preliminary DNA fingerprinting of the turf grass Cynodon dactylon (Poaceae: Chloridoideae). R.

ROODT. J.J. SPIES and T.H. BURGER 117

13. Miscellaneous notes:

Poaceae. Chromosome studies on African plants. 16. Polyploidy in the genus Ehrharta. J.J. SPIES,

M.M. VAN DYK and S.M.C. VAN WYK 123

14. Obituary: Anna Amelia Obermeyer-Mauve (1907-2001). H.F. GLEN 127

15. Book review 131

New species, subspecies, sections, name, status and combinations in

Bothalia 32,1 (2002)

Brachystelma cummingii A.P.Dold, sp. nov., 71

Clivia mirabilis Rourke, sp. nov.. 1

Daubenya inarginata ( Willd . ex Kunth) J.C. Manning & A.M.van cler Merwe, comb, nov., 65

Erica abietina subsp. atrorosea E.G.H.Oliv. & I.M.Oliv ., subsp. nov., 50

Erica abietina subsp. aurantiaca E.G.H.Oliv. & I.M.Oliv., subsp. nov., 51

Erica abietina subsp. constantiana E.G.H.Oliv. & I.M.Oliv., subsp. nov., 51

Erica abietina subsp. diabolis E.G.H.Oliv. & I.M.Oliv ., subsp. nov., 51

Erica abietina subsp. perfoliosa E.G.H.Oliv. & I.M.Oliv., subsp. nov., 52

Erica abietina subsp. petraea E.G.H.Oliv. & I.M.Oliv., subsp. nov., 53

Erica banksii subsp. comptonii ( Salter ) E.G.H.Oliv. & I.M.Oliv., stat. et comb, nov., 44

Erica banksii subsp. purpurea (Andrews) E.G.H.Oliv. & I.M.Oliv., stat. et comb, nov., 43

Erica baueri subsp. gouriquae E.G.H.Oliv. & I.M.Oliv., subsp. nov., 46

Erica coccinea subsp. uniflora E.G.H.Oliv. & I.M.Oliv., subsp. nov., 39

Erica filipendula subsp. parva E.G.H.Oliv. & I.M.Oliv., subsp. nov., 47

Erica intermedia subsp. albiflora E.G.H.Oliv. & I.M.Oliv., subsp. nov., 41

Erica melastonia subsp. minor E.G.H.Oliv. & I.M.Oliv., subsp. nov., 40

Erica petrusiana E.G.H.Oliv. & I.M.Oliv., sp. nov., 58

Erica plukenetii subsp. bredensis E.G.H.Oliv. & I.M.Oliv., subsp. nov., 41

Erica plukenetii subsp. breviflora ( Dulfer ) E.G.H.Oliv. & I.M.Oliv. , stat. et comb, nov., 43

Erica plukenetii subsp. lineata (Benth.) E.G.H.Oliv. & I.M.Oliv, stat. et comb, nov., 43

Erica plukenetii subsp. penicellata (Andrews) E.G.H.Oliv. & I.M.Oliv, stat. et comb, nov., 42

Erica regia subsp. mariae ( Guthrie & Bolus) E.G.H.Oliv & I.M.Oliv, stat. et comb, nov., 54

Erica viridiflora subsp. primulina (Bolus) E.G.H.Oliv & I.M.Oliv, stat. et comb, nov., 44

Erica viridiflora subsp. redacta E.G.H.Oliv. & I.M.Oliv, subsp. nov., 45

Erica viscaria subsp. gallorum (L.Bolus) E.G.H.Oliv. & I.M.Oliv, stat. et comb, nov., 56

Erica viscaria subsp. longifolia (Bauer) E.G.H.Oliv & I.M.Oliv, comb, et stat. nov., 56

Erica viscaria subsp. macrosepala E.G.H.Oliv. & I.M.Oliv, stat. et nom. nov., 56

Erica viscaria subsp. pendula E.G.H.Oliv. & I.M.Oliv, subsp. nov., 58

Erica viscaria subsp. pustulata (H. A. Baker) E.G.H.Oliv. & I.M.Oliv, stat. et comb, nov., 58

Eriospermum [subgenus Cyathiflorum] section Cyathiflorum P.L. Perry ex Thiede, sect, nov., 76

Eriospermum [subgenus Ligulatum] section Ligulatum PL.Perry ex Thiede, sect, nov., 76

Gnidia pedunculata Beyers , sp. nov., 79

Merciera tetraloba C.N.Cupido, sp. nov., 74

Passerina filiformis subsp. glutinosa ( Thoday ) Bredenkamp & A.E.van Wyk, stat. nov., 33

Passerina montivagus Bredenkamp & A.E.van Wyk, sp. nov., 34

Passerina nivicola Bredenkamp & A.E.van Wyk, sp. nov., 77

Passerina truncata (Meisn.) Bredenkamp & A.E.van Wyk, stat. nov., 66

Passerina truncata subsp. monticola Bredenkamp & A.E.van Wyk, subsp. nov., 69

Rotheca caerulea (N.E.Br.) P.P..I. Herman & Retief, comb, nov., 81

Rotheca cuneiformis (Moldenke) PP.J. Herman & Retief, comb, nov., 81

Rotheca louwalbertsii (PP.J. Herman) PP.J. Herman & Retief, comb, nov., 81

Rotheca pilosa (H. Pearson) PP.J. Herman & Retief, comb, nov., 81

Rotheca uncinata ( Schinz ) PP.J. Herman & Retief, comb, nov., 81

Tarchonanthus littoralis PP.J. Herman, sp. nov., 22

Tarchonanthus parvicapitulatus PP.J. Herman, sp. nov., 26

iv

Bothalia 32,1: 1-7 (2002)

Clivia mirabilis (Amaryllidaceae: Haemantheae) a new species from

Northern Cape, South Africa

J.P. ROURKE*

Keywords: Amaryllidaceae, arid Mediterranean climate, Clivia Lindl., new species. Northern Cape

ABSTRACT

Clivia mirabilis Rourke is a new pendulous tubular-flowered species from Oorlogskloof Nature Reserve in Northern

Cape. Its distribution area is some 800 km outside the previously accepted range of the genus Clivia. This sun-tolerant

species is adapted to an arid Mediterranean climate, producing vegetative growth in winter and maturing its seeds rapidly

in late summer/early autumn to synchronize with the arrival of winter rains.

INTRODUCTION

The genus Clivia Lindl., consisting of four currently

recognized species, C. nobilis Lindl. (1828), C. miniata

(Lindl.) Regel, C. gardenii Hook. (1856) and C. caulescens

R. A. Dyer (1943), is presently considered to be endemic

to southern Africa (Vorster & Smith 1994; Snijman 2000).

These species occur in coastal and inland Afromontane

forest from Eastern Cape through KwaZulu-Natal,

Swaziland and Mpumalanga to the Soutpansberg in

Northern Province. Rumours of the occurrence of Clivia

in Mozambique have not yet been confirmed by accu-

rately localized herbarium collections.

Clivia is an evergreen, rhizomatous genus in the

Amaryllidaceae, characterized by distichous strap-

shaped leaves, umbellate solid scapes and red subglobose

berries containing one to few cartilaginous, pearly-white

seeds embedded in soft yellow pulp.

In February 2001 material of a further species was

submitted to the Compton Herbarium for identification

in a batch of herbarium specimens from the Oorlogskloof

Nature Reserve near Nieuwoudtville, in Northern Cape,

collected by the nature conservation officer in charge, Mr

Wessel Pretorius. The author confirmed this astonishing

discovery during a site visit to Oorlogskloof on 22

February 2001 when two fruiting populations were

examined. The new species is here described as Clivia

mirabilis. Rarely can such an extravagant epithet as

mirabilis be confidently applied, yet in the case of this

extraordinary Clivia , so unusual in its distribution and

characters, its usage seems entirely appropriate.

Clivia mirabilis Rourke , sp. nov., a speciebus

affinibus, corollis actinomorphis rectis tubularibus bicol-

oribus (miniatis/luteis); pedicellis cemuis, 22^40 mm

longis, miniatis per anthesin (demum viridibus post

anthesin); foliis lineamentis medianis albis et apicibus

acutis, distinguitur.

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

7735 Claremont.

MS. received: 2002-01-25.

Rhizoma perenne solitarium, erectum; caespite folio-

rum 0.6-1. 2 m alto. Folia disticha, rigida, erecta,

0.6-1. 2 m longa, 30-50 mm lata, glabra, coriacea, linea-

mentis medianis albis; margines cartilagineae,

plerumque laeves, interdum leviter scabra; apices acuti.

Scapus 300-800 mm longus, carmineus, late ancipitius.

Inflorescentia umbellata, 20-48 floribus; spathae 5-7,

cymbiformi-acutae, papyraceae, 35-50 mm longae,

10-15 mm latae. Pedicelli cernui, graciles, 25-40 mm

longi, miniati per anthesin, abrupte viridescentes post

anthesin. Perianthium rectum, tubulosum, 35-50 mm

longum, dilatatum versus orem; miniatum sed viride ad

apicem per anthesin, tandem flavescens distale. Antherae

6, basifixae, leviter exsertae; filamenta 30 mm longa,

basaliter adpressa circa stylum. Stylus 40-45 mm longus,

trilobatus; apices subtiliter penicillati. Ovarium

ovoideum, nitidum, miniatum per anthesin; viride post

anthesin; loculi 3-4-ovulati. Baccae pendentes, irregu-

lariter ovoideae, glebulosae; ( 1 )2— 5(— 7) semina conti-

nentes, pericarpium maturum scarlatinum. Semina irreg-

ulariter ovoidea, ± 10 mm in diam., alba.

TYPE. — Northern Cape, 3119 (Calvinia): Nieuwoudt-

ville, Oorlogskloof Nature Reserve, eastern margin of

Oorlogskloof Canyon at Agterstevlei Farm, (-AC), ±

900 m, 18-10-2001, J.P. Rourke 2220 (NBG, holo.; BOL,

K, MO, NSW, PRE, iso.).

Stout, rhizomatous, solitary, evergreen perennial,

0.6-1. 2 m tall; stem reduced to a vertical rhizome, up to

250 mm long terminating in a tuft of leaves. Root system

massive, up to 0.7 m diam., horizontally spreading; roots

perennial, very stout, fleshy, up to 20 mm diam., outer

surface covered in a corky, velamen-like layer. Leaf

sheath prominent, flushed deep carmine (RHS 183A

greyed-purple). Leaves long-lived, stiffly erect, disti-

chous, strap-shaped, 0.6-1. 2 m long, 30-50 mm wide,

flattened to slightly V-shaped with a distinct pale whitish

grey striation in the midrib area on upper surface, stria-

tion becoming less distinct in older leaves, coriaceous,

glabrous, deep dull green, flushed carmine-maroon at

base, apex obtuse-acute; margins entire, cartilaginous,

usually smooth, occasionally irregularly scabrous,

becoming slightly revolute in old, mature leaves. Scape

2

Bothalia32,l (2002)

Bothalia 32,1 (2002)

3

300-800 mm long, broadly ancipitous, longitudinally

ridged, 10-14 mm wide, glabrous, carmine-flushed.

Inflorescence umbel-like, 20-48-flowered, subtended by

5-7 brownish/carmine, papyraceous spathe valves, nar-

rowly cymbiform-acute, 35-50 x 10-15 mm; pedicels droop-

ing, slender, 25-40 x 1 .2 mm, orange-red, abruptly turn-

ing green in post-pollination phase. Perianth straight to

imperceptibly curved, tubular, becoming progressively

flared towards apex, 35-50 x 5 mm below ovary, 10-12 mm

diam. at mouth, orange-red (RHS 32B) proximally,

green-tipped (RHS 145A) distally on opening, the green

changing to yellow (RHS 22B) distally, entire perianth

becoming deep orange-red (RHS 33A) after anthesis;

tepals fused distally to form a tube 10-15 mm long, outer

tepals narrowly oblong, apices acute, inner tepals slight-

ly wider, apices obtuse, outwardly flared. Stamens 6,

dorsifixed, 2 mm long, very slightly exserted at anthesis;

filaments 30 mm long, attached to tepals 10 mm above

ovary; inwardly bowed proximally, adpressed around

style at point of attachment forming a 10 mm long nec-

tar well above ovary. Ovary ovoid, shiny, greenish yel-

low in bud, becoming orange-red at anthesis, changing to

bright green in post-pollination phase; ovules 3 or 4 in

each locule; style 40-45 mm long, terete, glabrous,

tapering distally, included at anthesis, later elongating

and becoming exserted 5-8 mm in post-pollination

phase; stigma trilobed, lobes 0.4— 1.0 mm long, penicil-

late at apex. Fruiting heads with 25-35 pendent berries.

Berries irregularly oblong to ovoid, 10-30 x 10-15 mm,

glebulose to submoniliform, often narrowed to a distinct

neck above pedicel, apex often tapering to an eccentri-

cally angled beak, containing ( 1 )2 — 4(— 7) gongyloid

seeds projecting prominently and irregularly through

thin pericarp; pericarp glossy, pale apple green, maturing

through yellow, orange to pinkish red; mature berries red

(RHS 40B eventually becoming RHS 45B). Seeds some-

what ovoid, slightly faceted, ± 10 mm diam., pearly

white; embryo green. (Colour references according to

Royal Horticultural Society colour chart.) Figure 1; Plates

1 & 2.

Diagnostic characters

Clivia mirabilis is distinguished by its straight, actin-

omorphic, bicolored (orange/yellow) tubular corolla,

long drooping pedicels, 25^-0 mm long, that are orange-

red at anthesis and green when fruiting; the distinctive

single median white striation on the upper surface of the

leaves with smooth cartilaginous margins; and irregular-

ly shaped glebulose-gongyloid berries. The basal part of

the leaves forming the leaf sheath is flushed a deep

carmine maroon, unlike any other Clivia except

C. nobilis, which occasionally produces similarly

coloured leaf bases. The orange-red coloration of the

pedicels in this species during anthesis is a unique char-

acter in the genus Clivia.

Distribution and habitat

Apparently confined to the Oorlogskloof Nature

Reserve in Northern Cape (Figure 2), Clivia mirabilis is

restricted to a small area on the eastern margin of the

Oorlogskloof Canyon. Populations are known to occur

just north of Eland se Kliphuis adjacent to Agterstevlei

Farm and a little further south around the Driefontein

Waterfall. The distance between these sites is ± 5 km.

The species also occurs at a few sites between these two

localities.

The margins of the Oorlogskloof Canyon are capped

with 30 m cliffs of Peninsula Formation Sandstone. This

has eroded to form coarse sandstone talus screes below

the cliffs that are partly covered in a light woodland of

relictual Afromontane evergreen forest elements, princi-

pally Olea europaea subsp. africana , Maytenus acumi-

nata, M. oleoides, Cassine schinoides, Halleria lucida

and Podocarpus elongatus with additional shade provid-

ed by outsize, (4 m tall) specimens of Phylica oleaefolia.

Small groups of C. mirabilis grow rooted in humus

between cracks in the sandstone talus of the rock scree,

either as solitary individuals or in small groups.

Occasionally some clumps occur in full sun but these

tend to have shorter leaves and often show signs of water

stress (dried leaf tips). However, the remaining leaves

show no signs of sunburn, despite the intense insolation

experienced for several months each year. The main pop-

ulation extends over several hectares and probably con-

sists of well over 1 000 individuals. Due to the position

of these two sites under the eastern cliffs of Oorlogskloof

Canyon, most plants experience shade until about mid-

morning after which they are in direct sun.

The area is characterized by a semi-arid Mediter-

ranean climate with a strictly winter rainfall regime —

exactly the opposite climatic conditions experienced by

the other four species in this genus. The mean annual

rainfall for Oorlogskloof is 414 mm, falling mainly

between May and September. Vegetative growth is thus

restricted to a brief winter growing period. Situated at

850-900 m, some 100 km inland from the coast, these

populations are subject to brief but light frost in winter.

MORPHOLOGY AND BIOLOGY

Root system

On excavating several plants in the habitat for culti-

vation at Kirstenbosch, the enormous root system char-

acteristic of this species was revealed. Large adult plants

have a mass of fleshy, succulent roots radiating between

V 2 — 3/ 4 m from the base of the rhizome, each root ± 20 mm

in diameter. This disproportionally large volume of sub-

PLATE 1. — Clivia mirabilis. A, inflorescence in habitat showing orange/red pedicels; B, plants in habitat between Peninsula Formation Sandstone

boulders; C, leaves showing prominent median white stripe on upper surface and variation in dimensions. D, E, fruiting head: D, immature

berries; E, mature berries. F, seedlings five months after germination, note large radicle and prominent carmine pigmentation on cotyledon

and at base of first pair of leaves. Photos; C. Paterson-Jones.

4

Bothalia 32,1 (2002)

FIGURE 1 . — Clivia mirabilis: A,

mature inflorescence; B,

fruiting head with irregularly

shaped berries; C, longitudi-

nal section through flower;

note filament bases

adpressed to style forming

nectar reservoir; D, detail of

filament bases. Scale bars: 10

mm. Artist; John Manning.

terranean biomass gives mature plants an extensive water

storage capacity, allowing them to survive the prolonged

rainless summers of the Oorlogskloof environment.

Flower colour, development and pollination

The general impression of a fully open scape is of

bicoloured perianths, orange-red at the base, yellow

towards the mouth and with orange-red pedicels. During

the development of the flower, both perianth and ovary

progress through a series of well-marked colour changes.

The unopened bud is yellowish, but prominently green-

tipped, and the ovary is also pale green. At anthesis the

green coloration slowly disappears from the tips of the

tepals which take on the same yellow tones as the basal

half of the perianth. The pedicels and upper half of the

perianth are deep orange-red at this stage. After pollina-

tion the yellow coloration disappears and the whole peri-

anth and ovary take on a uniform orange/red colour. As

the perianth begins to wither, the ovary swells and under-

goes an abrupt colour change from orange to bright

green, as do the pedicels. No other Clivia has pedicels

the same colour as the perianth when the flower is fully

open. The pedicels abruptly change to green as the peri-

anth abscisses and the ovary swells in the post-pollina-

tion phase.

The purpose of these colour changes is not yet under-

stood, but is probably related to pollinator cues.

Pollination appears to be by sunbirds. A single sighting

of a malachite sunbird probing the perianths was made at

Oorlogskloof on 18 October 2001 suggesting that sun-

birds could be involved in pollen transfer. However, like

the other three tubular-flowered species, C. mirabilis

may also be a selfer as between 80 and 90% of the flow-

ers in each umbel are pollinated and produce viable

berries. Flowering time : ± six weeks, from October to

mid-November.

Bothalia 32,1 (2002)

5

FIGURE 2. — Distribution of C. mirabilis, 9. in relation to the distrib-

ution of the remainder of the genus Clivia (black shaded area).

Fruiting

The berries mature more rapidly than in the other

Clivia species. By the end of February, four months after

flowering, the fully developed berries turn from yellow

and orange to pinkish and later red by the end of March

and are shed shortly thereafter prior to the onset of the

first winter rains in April/May. This rapid autumn matu-

ration of berries is in sharp contrast to the summer rain-

fall area clivias which mature slowly, usually 12 months

for C. miniata and C. gardenii, about nine months for

C. caulescens and C. nobilis (Duncan 1999) to coincide

with the commencement of October/November summer

rains.

Seed dispersal and germination

Berries commence falling from late February to early

April. Germination appears to be rapid in response to the

onset of autumn/early winter rains. At Kirstenbosch

seeds sowed on 18 March 2001 had already developed a

10 mm radicle by 10 April 2001.

On germinating, the primary root develops into a

swollen, white, succulent cylinder up to 50 x 5-6 mm.

During the moist winter months (May-September), it

swells, accumulating water in its succulent tissue. By

October, two short (5-10 mm long) leaves have been

produced, whereafter further vegetative growth of the

seedling slows or largely ceases with the onset of sum-

mer dormancy (November-April). During the rainless

phase of ± six months the seedling survives on water

reserves stored in the greatly enlarged primary root.

Vegetative growth commences again in autumn. Thus the

biology of a C. mirabilis seedling in its first year is much

akin to a winter rainfall area geophyte with the swollen

primary root being functionally equivalent to a corm or

bulb.

The phenology of the germinating seed described

above is clearly an adaptation to a semi-arid Mediter-

ranean climatic regime — exactly the reverse of the sum-

mer rainfall region Clivia species.

Within a few months of germinating, the plumular

bud (cotyledon plus first true leaf) (Boyd 1932),

becomes densely pigmented with anthocyanins (Plate

IF). This prominent development of anthocyanins at the

base of the leaves is later evident in the leaf sheaths of

adult plants which are heavily suffused with purple-

carmine pigments. Why the seedlings of C. mirabilis are

so densely pigmented with anthocyanins is not clear, but

it may be a response to the intense levels of sunlight

experienced in the natural habitat, thereby providing

effective screening during the seedlings’ critical estab-

lishment phase.

Relationships

The distribution ranges of all four previously known

Clivia species are contiguous or overlap, while at many

localities different pairs of species occur sympatrically,

C. nobilis with C. miniata , C. gardenii with C. miniata ,

and C. caulescens with C. miniata. Geographically, pop-

ulations of C. nobilis in Eastern Cape, though more than

800 km distant, are the closest spatially to C. mirabilis.

C. nobilis also appears to be the closest relative to

C. mirabilis on morphological grounds: tough stiffly

erect coriaceous leaves with a median pale striation on

the upper surface (some populations of C. nobilis occa-

sionally have a faint median striation), and the small

seeds.

Phytogeographic implications

Palynological evidence indicates that in Western Cape

and southern Namaqualand, subtropical forests were pre-

sent during Miocene and Pliocene times (± 5.3 million

years BP) (Scott etal. 1997). Since then, apart from more

recent cyclical changes in the Quaternary, there has been

a progressive eastward retreat of these forest elements.

Assuming that the genus Clivia has not changed its

dependence on a forest environment significantly since

pre-Quaternary times, it can be argued that the

Nieuwoudtville species is relictual and that its survival in

the Oorlogskloof Canyon is partly fortuitous and partly

due to its adaptation to a different climate. The berry

maturation period, seedling and germination biology are

so perfectly in harmony with an arid Mediterranean cli-

matic regime that Clivia mirabilis is able to survive envi-

ronmental conditions inimical to all other Clivia species.

It is currently believed that the late Miocene also saw

the development of a Mediterranean climate in the west-

ern part of the Cape (Axelrod & Raven 1978). This

would have interrupted the further spread of an essen-

tially summer rainfall genus like Clivia into the forests of

the southern and western part of the Cape. It would also

have left the precursors of Clivia mirabilis to adapt to

increasing aridification and the onset of a pronounced

Mediterranean type climate. Thus if C. mirabilis evolved

from forms with an essentially summer rainfall phenolo-

gy, the adaptation to an arid Mediterranean type climate

is a derived condition dating from late Miocene times.

Long distance dispersal should also be considered as

a possible explanation for this bizarre distribution pat-

tern, but this seems highly unlikely as no living vectors

6

Bothalia 32,1 (2002)

for the long-distance dispersal of Clivia seed have yet

been identified nor has biotic dispersal been recorded for

any other species of Amaryllidaceae (Meerow & Snijman

1998). Birds are probably the main seed dispersal vec-

tors. Clivia miniata, C. gardenii and C. caidescens have

all been observed by the author growing epiphytically in

forest trees, five or more metres above ground level. It is

probable that frugivorous birds deposited seeds in these

positions, leading one to postulate that forest dwelling

birds are responsible for the dispersal of large scarlet

Clivia berries. While the dispersal of Clivia seed by birds

between closely adjacent forest patches is a strong possi-

bility, dispersal over distances of 800 km of arid country

seems highly unlikely.

Conservation status

No populations are known outside the Oorlogskloof

Nature Reserve where the species currently enjoys max-

imum protection. Yet there is no reason why C. mirabilis

should not occur further down the Oorlogskloof Canyon,

outside the reserve, as numerous suitable habitats occur

there. If this proves to be the case, special efforts will

have to be made to protect these populations as the

species’ horticultural potential will render it vulnerable to

exploitation.

Other material examined

NORTHERN CAPE.— 3119 (Calvinia): Oorlogskloof Nature

Reserve, (-AC). 10-1 1-2000, W. Pretorius 651 (NBG).

ACKNOWLEDGEMENTS

I am most grateful to the Northern Cape Department

of Nature and Environmental Conservation for granting

permission to collect the type material of this species;

also to Wessel Pretorius, officer in charge of the

Oorlogskloof Nature Reserve for his generous assistance

in the field. Colin Paterson Jones patiently photographed

various stages in the life history of C. mirabilis and John

Manning skilfully prepared the line drawings reproduced

here. Dee Snijman and John Manning provided critical

comments on early drafts of the manuscript. I thank all of

them for their invaluable assistance.

Auriol Batten’s fine painting of C. mirabilis was pre-

pared from the type material. I am especially grateful to

her for loaning this plate to the National Botanical

Institute for reproduction in this paper.

REFERENCES

AXELROD, D.I. & RAVEN, P.H. 1978. Late Cretaceous and Tertiary

vegetation history of Africa. In M.J.A. Werger, Biogeography

and ecology of southern Africa. Junk, The Hague.

BOYD, L. 1932. Monocotyledonous seedlings. Morphological studies

in the post-seminal development of the embryo. Transactions

and Proceedings of the Botanical Society of Edinburgh 31 : 57,

58.

DUNCAN, G. 1999. Grow clivias. Kirstenbosch Gardening Series,

National Botanical Institute, Cape Town.

DYER, R.A. 1943. Clivia caulescens. The Flowering Plants of South

Africa 23: t. 891 .

HOOKER, W.J. 1856. Clivia gardenii. Curtis’s Botanical Magazine', t.

4895.

LINDLEY, J. 1828. Clivia nobilis. Edward's Botanical Register, t.

1182.

MEEROW, A.W. & SNIJMAN, D.A. 1998. Amaryllidaceae. In K.

Kubitzki, The families and genera of vascular plants 3: 83-1 10.

Springer, Berlin.

SCOTT, L„ ANDERSON, H.M. & ANDERSON, J.M. 1997. Vegetation

history. In R.M. Cowling et at. Vegetation of southern Africa.

Cambridge University Press, Cambridge.

SNIJMAN, D.A. 2000. Amaryllidaceae. In O.A. Leistner, Seed plants

of southern Africa: families and genera. Strelitzia 10: 570-576.

National Botanical Institute, Pretoria.

VORSTER, P. & SMITH, C. 1994. Clivia nobilis. The Flowering

Plants of Africa 53: t. 2094.

PLATE 2. — Clivia mirabilis, x 0.75, showing flowering and fruiting scapes against a background depicting Oorlogskloof Canyon. A fully open

inflorescence on left shows the red/yellow colour pattern. A younger inflorescence with green tipped perianths is on its right. From the type

collection, Rourke 2220. Artist: Auriol Batten.

Bothalia 32,1 (2002)

7

Bothalia 32.1 : 9-13 (2002)

The genus Buglossoides (Boraginaceae) in southern Africa

E. RETIEF* and A.E. VAN WYK**

Keywords: Boraginaceae. Boraginoideae. Buglossoides Moench. Lithospermeae, nutlets, pollen, southern Africa, taxonomy

ABSTRACT

The genus Buglossoides Moench, a member of the family Boraginaceae, subfamily Boraginoideae. tribe Lithospermeae,

is revised for the Flora of southern Africa (FSA) region. Buglossoides comprises about seven species of annual, biennial or

perennial herbs and subshrubs native to Asia, southern Europe and northern Africa. B. arvensis (L.) I.M.Johnst., a natural-

ized weed in many parts of the world, is the only member of the genus represented in the flora of southern Africa, presum-

ably introduced with imported cereal seed. Although the similar polyaperturate pollen grains of Buglossoides and

Lithospermum support the view that they are congeneric, they are retained as separate entities here. Diagnostic characters,

a full description, various illustrations and a distribution map of B. arvensis in southern Africa are given.

INTRODUCTION

In 1794 Moench established the genus Buglossoides

(Boraginaceae/Boraginoideae/Lithospermeae), separat-

ing it from the closely related genus Lithospermum L.

Ever since, the generic status of Buglossoides has been a

matter of controversy. De Candolle (1846), Boissier

(1879), Giirke (1897), Wright (1904), Riedl (1967) and

Mabberley (1997) regarded the genus as congeneric with

Lithospermum , not accepting or unaware of Moench's

view. In 1954 Johnston published a taxonomic revision

of Buglossoides. He used the corolla throat, decorated

inside with five well-developed vertical lines of hairs, to

distinguish Buglossoides from Lithospermum , which is

characterized by a corolla throat lacking these hairs but

bearing localized faucal appendages or groups of stipi-

tate glands. Buglossoides was accepted by various

authors like Ingram (1958), Fernandes (1972),

Edmondson (1978), Qaiser (1979). Brummitt ( 1992), Al-

Shehbaz (1991), Verdcourt (1991), Herman (1993),

Retief & Herman (1997), Lebrun & Stork (1997) and

Retief (2000), and this view is also taken here.

Buglossoides is a genus of about seven species, dis-

tributed primarily in the European portion of the

Mediterranean region and in adjacent southeastern Asia,

with a single species indigenous to China, Korea and

Japan (Al-Shehbaz 1991). B. arvensis (L.) I.M.Johnst. is

the only member of the genus represented in the flora of

southern Africa, scattered across the region. B. arvensis

most probably reached southern Africa with imported

cereal seed. It is widespread as a naturalized weed in

many parts of the world. As yet it is not known from the

Flora zambesiaca region, but is recorded in Tanzania and

North Africa. Fernandes (1971, 1972) reduced three

Buglossoides species to subspecies of B. arvensis. How-

* National Herbarium, National Botanical Institute, Private Bag X101.

0001 Pretoria.

** H.G.W.J. Schweickerdt Herbarium, Department of Botany,

University of Pretoria, 0002 Pretoria.

MS. received: 2001-06-18.

ever, typically B. arvensis differs from these taxa in its

infundibuliform corolla (not hypocrateriform), a calyx

that usually equals or exceeds the corolla tube in flower

(not with calyx distinctly shorter) and in its distribution

(not confined to the Mediterranean region). It is here re-

cognized as a separate species. One of the three species

concerned, B. sibthorpianum Griseb., is regarded as con-

specific with B. arvensis by Edmondson (1978) and

Qaiser (1979).

De Candolle ( 1846) already cited Burchell 1686 under

Lithospermum arvense L. Burchell collected his speci-

men at Kloof village in the Asbestos Mountains,

Northern Cape in 1811. Wright (1904) in his revision of

Lithospermum in Flora capensis cited several localities

in South Africa where 'L. arvense' had been collected.

Bolus & Wolley-Dod (1904) also mentioned the occur-

rence of Buglossoides arvensis because Wolley-Dod had

collected it at a few places on the Cape Peninsula.

Although Wright (1904) and Ross (1972) cited Gerrard

230 for KwaZulu-Natal, B. arvensis is not recently

recorded from the region.

In this paper, diagnostic characters, a full description,

various illustrations and a distribution map of Buglos-

soides arvensis in southern Africa are given. The descrip-

tion is based exclusively on local collections. This paper

forms part of a revision of tha Boraginaceae for the Flora

of southern Africa (FSA) which is currently in progress.

MATERIALS AND METHODS

Herbarium specimens of Buglossoides arvensis

housed in BM, BOL. E, GRA. K. NBG. NH, NMB. NU,

PRE, PRU and SAM were studied to assess morpholog-

ical variation as well as phenological and distributional

attributes. Acetolysis of pollen followed the standard

method of Erdtman (1960). For scanning electron micro-

scope studies, samples were coated with gold and stud-

ied with an 1SI-SX-25 SEM. Measurements of pollen

grains were taken with the light microscope from acetol-

ysed grains mounted in glycerine jelly.

10

Bothalia 32.1 (2002)

FIGURE 1. — Buglossoides arvensis. A-C, rigid setae: A, on calyx lobes; B. on upper leaf surface; C, basal part. D, E: inner corolla; D, lines of

hairs; E, close up of hairs and pollen grains. F, part of annulus and pollen grains; G, nutlet; H, I, rugose, tuberculate nutlet surface. A-E,

H. I. Acocks 17740 ; F. Acocks 19035', G, Acocks 17805. Scale bars: 10 mm. A, 122 pm: B, 190 pm: C, 492 pm; D. 130 pm; E, 36 pm; F,

71 pm; G, 528 pm; H. 269 pm; I. 59 pm.

MORPHOLOGICAL CHARACTERS OF TAXONOMIC

SIGNIFICANCE

Macromorphology

Species of Buglossoides are divided into two groups:

perennials with corollas 12-20 mm long, nutlets smooth

or punctate-reticulate, white or yellowish and shiny, and

annuals with corollas up to 10 mm long, nutlets tubercu-

late, brownish or greyish and not shiny. B. arvensis

belongs to the latter group. The species is distinguished

from others in the genus by its calyx with a whitish indu-

mentum and by the calyx lobes which are strongly

accrescent in fruit, narrow, acute and covered with long,

rigid setae (Figure I A) and hairs (Leistner 2000: 759).

The indumentum of the leaves is appressed-setulose

with rigid setae, and the multicellular bases of the setae

are 1 -layered (Figure IB, C). B. arvensis is distinguished

from all species of Lithospermum, and from all other

southern African representatives of Boraginaceae, by a

combination of the following characters: five longitudi-

nal bands of hairs extending from the base of the corolla

lobe downwards to the tips of the anthers, as is typical for

the genus (Figure ID, E); an annulus of scale-like lobes

present near the base of the corolla tube (Figure IF); and

nutlets that are rugose and tuberculate (Figure 1G-I).

Pollen

Pollen of Buglossoides arvensis is isopolar and pro-

late, with P = 12.5-18.8 pm and E = 8.3-12.6 pm. In

equatorial view, the grains are rectangular to elliptic,

with the long sides more or less straight and the poles

convex, but frequently collapsing inwards (Figure 2A,

B). Ectoapertures are positioned at the equator, rhombic

in outline, regularly granular and the margins are not

thickened. Endoapertures are situated on or near the

equator and are ± oval and lalongate. The tectum is psi-

late. Orbicules are present.

Bothalia 32,1 (2002)

FIGURE 2. — Buglossoides cirvensis. A. pollen grains; B, pollen grain with collapsed poles; C, granular aperture and psilate tectum. All grains ace-

tolysed. A. Acocks 17805: B, C. Acocks 17740. Scale bars: 10 mm. A, 4 pm; B. 2 pm; C, 1 pm.

The polyaperturate pollen grains of Buglossoides,

similar to those of Lithospermum, support the merging of

these genera. In their studies on pollen. Clarke (1977)

and Diez et al. (1986) accepted Buglossoides as a sepa-

rate genus. Ahn & Lee (1986), in a palynotaxonomical

study of the Korean Boraginaceae, did not accept

Buglossoides. Johnston (1954) divided the genus into

two sections and concluded that the pollen of section

Eubuglossoides (e.g. B. arvensis) is isopolar, whereas

pollen of section Margarospermum (e.g. B. purpuro-

caerulea (L.) I.M.Johnst. ) is heteropolar and shows simi-

larity to pollen of Lithospermum. Pollen morphology

thus suggests that whereas the section Margarospermum

can perhaps be included in Lithospermum, members of

the section Eubuglossoides are from Lithospermum and

best retained in Buglossoides. The white, smooth, shiny

nutlets of Buglossoides purpurocaerulea similar to those

of Lithospermum afromontanum Weim., for example, as

opposed to the rugose, tuberculate ones of B. arvensis,

support Ahn & Lee's (1986) view. Heterostyly is absent

in B. an’ensis and a vast amount of pollen is released on

to the corolla and between the lines of hairs (Figure

1D-F).

Pollen of southern African Boraginaceae is classified

into seven pollen types (Retief & Van Wyk 1999). Two

genera Lobostemon and Echiostachys, mainly confined

to Western Cape of South Africa, and the two species of

Echium introduced into southern Africa, are often

regarded as belonging to the tribe Lithospermeae in the

subfamily Boraginoideae (Johnston 1954). The triapertu-

rate, heterocolpate pollen grains with a reticulate tectum

found in these genera are very different from those of

Buglossoides and Lithospermum. which are polyapertu-

rate with a psilate tectum. Palynology supports the recog-

nition of a separate tribe, Echieae, for Lobostemon,

Echiostachys and Echium.

Buglossoides Moench, Methodus plantas horti

botanici et agri marburgensis: 418 (1794); R.Fern.: 87

(1972); Al-Shehbaz: 129 (1991); Verde.: 79 (1991);

Retief: 180 (2000). Type: B. tenuiflora (L.f.) I.M.Johnst.

Lithospermum L.: 132 (1753), pro parte.

The genus name is derived from Buglossum Miller,

the name of a genus now reduced to a section of Anchusa

L., and eidos = appearance, alluding to the superficial

resemblance of the two genera; the name Buglossum is

derived from the Greek bous, an ox and glossa, a tongue,

in reference to the broad, rough leaves (Al-Shehbaz

1991).

Buglossoides arvensis (L.) I.M.Johnst. in Journal

of the Arnold Arboretum 35: 42 ( 1954); Edmondson: 316

(1978); Qaiser: 82 (1979); Meikle: 1148 (1985); Tolken:

1152 (1986); Verde.: 79 (1991); Gibbons & Brough: 204

(1992); Retief & Herman: 350 (1997). B. an’ensis (L.)

I.M.Johnst. subsp. arvensis sensu R.Fern.: 87 (1972).

Type: Europe, Herb. Linn. 181/4 (LINN, syn.-PRE,

microfiche!).

Lithospermum arvense L.: 132 (1753); A. DC: 74 (1846); Boiss.:

216 ( 1879); C.H.Wright: 24(1904); I.M.Johnst.: 7 ( 1927); Levyns: 688

(1950); Levyns: 234 (1966); Jacot Guill.: 235 (1971); J.H.Ross: 298

(1972).

Illustrations: Qaiser: 83 ( 1979); Tolken: 1153 (1986); Verdcourt: 80

(1991); Gibbons & Brough: 205 ( 1992).

Annual herb up to 0.6 m high, greyish green, hispidu-

lous-strigillose. Roots purplish red. Stems erect or

decumbent, densely covered with appressed setae,

0.7-1 .0 mm long. Leaves sessile, narrowly ovate or nar-

rowly obovate, 15^45 x 3-8 mm, somewhat clasping at

base, apex acute or slightly obtuse, margin entire, dense-

ly setulose; setae rigid, with multicellular, 1 -layered

bases. Inflorescences terminal; cymes scorpioid,

bracteate. Calyx divided almost to base into 5 narrow

lobes; lobes with apices acute, strongly strigose, accres-

cent, up to 12 mm long in fruit. Corolla white, infundibu-

liform; tube cylindric, 5-7 mm long, throat with 5 well-

differentiated, longitudinal bands of hairs extending from

bases of corolla lobes downwards to tips of anthers;

lobes 5, rounded or truncate, ±2x1 mm; annulus near

base of corolla tube of scale-like lobes. Stamens 5,

included, arising from corolla below middle of tube; fil-

aments equal, shorter than anthers; anthers oblong, with

short apical appendage. Ovary 4-lobed, 4-locular,

gynobase disc-like; stigmas 2, subterminal. Fruit of 4

nutlets; nutlets erect, ovoid, flattened on dorsal side,

keeled on ventral side, with ± straight beak, 2. 5-3.0 mm

long, rugose, tuberculate, attachment scar basal or nearly

so; fruiting pedicel ± 2.5 mm long. Flowering time :

August to December. Figure 3.

12

Bothalia 32.1 (2002)

Distinguishing characters : annual herb, branching

from base, appressed hairy; corolla infundibuliform,

white, hairy inside; stigmas 2, subterminal; nutlets

rugose, tuberculate.

Distribution'. North-West, Gauteng, Mpumalanga,

Free State, Lesotho, Northern Cape, Western Cape,

Eastern Cape (Figure 4); a native of Eurasia, naturalized

as a weed in many parts of the world.

Habitat', weed of waste places; also in disturbed areas

such as in and along edges of cultivated lands.

Common names: sheepweed, white ironweed,

gromwell, corn gromwell, gromwell-corncockle, puc-

coon; naelbossie, naaldjiebossie, nagelbossie (Afrikaans)

(Smith 1966; Wells et al. 1986).

SPECIMENS EXAMINED (southern Africa only)

Acocks 17740 K. PRE; 17805 PRE; 19035 K. PRE; 40897 PRE. Acocks

& Roux 15948 K, PRE.

FIGURE 3. — Buglossoides arvensis.

A, habit, x 0.65; B, stigma, x

13.6; C, tuberculate nutlet, x

12; D, corolla laid open, x

3.3; E, setae on upper leaf

surface, x 34.5. A, C-E, Acocks

19035', B, Acocks 17740. Illu-

strations by Gillian Condy.

Bolus 101 BOL; 122 K; 274 GRA; 9201 BOL. Bradfield 200 PRE.

Burchell 1686 K.

Dieterlen 1290 PRE.

Flanagan 1629 K, PRE. Fourcade 941 BOL. GRA.

Gerrard 230 K.

Holub s.n. K.

Jenkins TRV7232 PRE.

Leendertz TRV9748 PRE. Lovemore GRA-A1642 GRA.

MacOwan 1926 GRA. M a doth 7216. 9670. PRE40898 PRE. Mogg

15292 PRE. Muir 790 PRE.

Schlecliter 3550 GRA. Sharpe 9145 PRE. Silk 52 PRE. Smith 884 PRE.

Tyson 378 GRA. Trollope GRA-A1640 GRA.

Wardle FRE40893 PRE. Wilman 19817. PRE59952 PRE. Wilms 1006

BM. Wolley-Dod 1551 K; 2794 BOL; 3611 BM. BOL, K.

ACKNOWLEDGEMENTS

Dr O.A. Leistner, Miss G. Condy, Mines E. du Plessis

Bothalia 32.1 (2002)

13

FIGURE 4. — Known distribution of Buglossoides arvensis in the

Flora of southern Africa region.

and J. Ready are thanked for assistance with the prepara-

tion of the manuscript. The directors, curators and staff

of cited herbaria are thanked for allowing the authors to

study their material and for sending specimens on loan.

REFERENCES

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Boraginaceae. Korean Journal of Plant Taxonomy 16: 199-215.

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eastern United States. Journal of the Arnold Arboretum , sup-

plementary series 1: 1-169.

BOISSIER. E. 1879. Lithospennum (Borragineae). Flora orientalis 4:

216-220. Georg, Geneva.

BOLUS. H. & WOLLEY-DOD. A.H. 1904. A list of the flowering

plants and ferns of the Cape Peninsula with notes on some of

the critical species. Transactions of the South African

Philosophical Society 14: 207-373.

BRUMMITT. R.K. 1992. Vascular plant families and genera. Royal

Botanic Gardens. Kew.

CLARKE. G.C.S. 1977. The northwest European pollen flora. 10

(Boraginaceae). Review of Pal aeobot am and Palynology 24:

59-101.

DE CANDOLLE. A. 1846. Lithospennum (Borragineae). Prodromus

10: 73-86. Masson. Paris.

DIEZ. M.J.. VALDES. B. & FERNANDEZ. 1. 1986. Pollen morpholo-

gy of Spanish Lithospennum s.l. (Boraginaceae) and its taxo-

nomic significance. Grana 25: 171-176.

EDMONDSON. J.R. 1978. Buglossoides. In PH. Davis. Flora of

Turkey 6: 315-317.

ERDTMAN, G. 1960. The acetolysis method, a revised description.

Svensk Botanisk Tidskrift 54: 561-564.

FERNANDES, R. 1971. Boraginaceae. Flora europaea — notulae sys-

tematicae 1 1 . Botanical Journal of the Linnean Society 64: 379.

FERNANDES. R. 1972. Buglossoides. Flora europaea 3: 87. 88.

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flowers of Britain and northern Europe. Hamlyn. London.

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■

Bothalia 32,1 : 15-19 (2002)

Studies in the liverwort family Aneuraceae (Metzgeriales) from south-

ern Africa. 3. Riccardia compcicta

S.M. PEROLD*

Keywords: Aneuraceae, Riccardia Gray, R. compacta (Steph.) S.W.Arnell. southern Africa, Tanzania

ABSTRACT

Riccardia compacta (Steph.) S.W.Arnell is described and illustrated. Its presently known distribution in southern Africa

is indicated on an accompanying map. Some differences between local plants of the species and those from Tanzania are

discussed.

INTRODUCTION

Riccardia compacta (Steph.) S.W.Arnell is the third

species to be treated in the current revision of southern

African taxa of this family. It was originally collected on

Table Mountain by Jelinek, while on a Novara

Expedition, and described as Aneura compacta by

Stephani in 1893. Sim (1926) referred to it briefly, stat-

ing that it frequently occurred in forest districts through-

out South Africa, which is doubtful, however, as it is

apparently rare. Later on. two more specimens were col-

lected on Table Mountain: one by Pillans, and the second

by Arnell, who also collected it at Knysna. Arnell (1952)

then transferred it to the genus Riccardia. During the

1950s Esterhuysen collected more specimens on high

altitude peaks in Western Cape, but, regrettably, there

have been no local collections since then. Arnell (1959)

described an Esterhuysen specimen from Tanganyika

(Tanzania) as R. kilimandjaricci S.W.Arnell, which was

placed in synonymy under R. compacta by Meenks &

Poes (1985). Poes, either alone or with Ochyra, collected

more material on Mt Kilimanjaro and Mt Meru during

the 1980s. It was concluded by Meenks & Poes that there

were a few differences between Tanzanian plants of the

species and the type specimen collected by Jelinek on

Table Mountain. They also state that R. compacta is an

Afro-alpine species which occurs at higher altitudes in

East Africa than in South Africa.

Riccardia compacta (Steph.) S. W.Amell in Botaniska

Notiser 1952: 141 (1952); S.W.Arnell: 88 (1963); Meenks

& Poes: 84 ( 1985). Type: Cap. Tafelberg, Jelinek , Novara

Expedition, ( 19560G . holo.!; EGR).

Aneura compacta Steph.: 19 (1893): Steph.: 755 (1901-1905): Sim:

29 (1926).

Thalli prostrate, in compact cushions or patches, up to

3 or 4 mm thick, in several overlying layers of dense and

intricately intertwined, slightly rounded, narrow bands,

quite fleshy and firm, but rather brittle, bright green, with

an ill-defined, somewhat darker, narrow strip occupying

median V3 or more of branches; when dry, brown to dark

brown. Main axis up to 15 mm long, not markedly dif-

* National Botanical Institute, Private Bag X101, 0001 Pretoria.

MS. received: 2002-01-14.

ferentiated, though often retaining its dominance,

branching generally crowded and irregular (Figure 1A),

sometimes bifurcate (Figure IB), branches narrowing

somewhat toward tips or maintaining same width

throughout, rarely slightly dilated distally, apically ±

truncate, shallowly notched, margins obtuse, not winged.

Primary branches/pinnae arising laterally from axis,

very close together or at intervals of 0.30-0.95 mm

between successive ones, opposite or subopposite,

obliquely spreading at angles up to 30° with axis, occa-

sionally suberect, often relatively well developed,

2. 0- 3. 5 mm long, up to 0.5 mm wide, usually linear,

some bifurcate, rarely trifurcate, others poorly devel-

oped, almost rudimentary. Secondary’ branches/pinnules

quite rare, if present, mostly remaining small. Stolons

unbranched, generally arising on basal part of axis, occa-

sionally developing more distally from apices of some

primary branches. Dorsal epidermal cells in median part

of apical segment of main axis (Figure 1C) from above

4-6-sided, thick-walled. 40-50 x 27.5-37.5 pm, subdor-

sal cells larger, 75-130 x 30.0-42.5 pm, subventral cells

50.0- 87.5 x 15-20 pm, ventral epidermal cells 35-60 x

30^)0 pm. Oil bodies not seen in material studied.

Margins of apical segment of main axis (Figure ID)

entire, from above outer cells 25.0-37.5 x 27.5-35.0 pm,

subquadrate to rectangular, intramarginal cells 5- or 6-

sided, 32.5-50.0 x 27.5-37.5 pm, walls somewhat thick-

ened. Cross sections of main axis at ± middle of ultimate

segment (Figure IE) and toward base (Figure IF) bicon-

vex, ± 180 pm, or 6/7 cell layers up to 240 pm, or 8 cell

layers thick medianly, and up to 460 pm wide, margins

tapering to obtuse, dorsal epidermal cells 27.5-30.0 pm

high, medullary cells ovoid to almost spherical, sizes

variable, 25-45 x 22.5-35.0 pm, walls thickened, ventral

epidermal cells 30-35 pm high: cross section of primary

branch at ± its middle (Figure IG) plano-convex, ± 200

pm or 5/6 cell layers thick medianly, up to 475 pm wide,

medullary cell walls slightly thickened; cross section of

stolon (Figure 1H) oval, ± 160 pm or 6 cell layers thick

medianly, ± 210 pm wide, cell walls somewhat thick-

ened. Mucilage papillae ventral, clustered at emarginate

apex of pinnae and then in two, spaced, acropetal rows,

one on either side of midline, reddish brown, club-

shaped, 62.5-75.0 x 20-25 pm above, narrower below,

rarely persistent. Rhizoids in widely scattered, irregular

patches along ventral surface of pinnae, hyaline, up to 15

pm wide. Asexual reproduction by gemmae not seen.

2 mm n 2 mm z mm

16

Bothalia 32,1 (2002)

FIGURE 1 . — Riccardia compacta. A, sterile thallus with axis and irregular lateral branches crowded below; B. axis with primary branches; C.

median dorsal epidermal cells (solid lines) of ultimate segment of axis, and large subdorsal cells (stippled lines); D, margin of ultimate seg-

ment of axis; E, c/s axis at ± middle of ultimate segment; F, c/s axis below; G, c/s primary branch; H, c/s stolon; I, male thallus with 3

antheridial branches. J-L, antheridial branch: .1, from above; K, from side; L, c/s. M, female thallus with calyptra, seta and dehisced cap-

sule valves; N, gynoecial branch with very young calyptra and paraphyses. O, calyptra: Oi, corona; CF, capsule; CF, seta; CF. basal portion.

P, c/s calyptra wall; Q. c/s seta: R, c/s part of bistratose wall of capsule valve showing different cell walls: abr, abaxial radial; adr, adaxial

radial; it. inner tangential; mw, median; ot, outer tangential. S, elater; Ti_j, spores. A-F, H-T, Arnell 963; G, Jelinek I9560G , holo.

Bothalia 32,1 (2002)

17

FIGURE 2. — A, cells of epidermal

layer of wall of valve in

external longitudinal view. B.

cells of inner layer of wall of

valve in internal longitudinal

view. A, B, Arnell 963. A, B,

x 446.

Dioicous. Antheridial branches solitary or occasional-

ly in sympodial pairs, lateral on main axis opposite or

subopposite primary branch, otherwise on primary

branch close to its base (Figure II), stipitate, ± oblong

(Figure 1J), up to 600 pm long, ± 320 pm wide, mostly

bearing 3 or 4, rarely up to 8 pairs of antheridia, cavities

70-100 pm diam., encircled by 2 or 3 rows of cells,

25-30 x 30 pm; in cross section ± 250 pm high (Figure

1L), margins winged (Figure IK), with single, incurved

row of large, obliquely orientated cells, 75.0-112.5 x

40-45 pm, apically rounded and free, otherwise adjoin-

ing; ventral papillae up to 30 pm long, in lateral pairs, but

rare. Gynoecial branches (Figure IN) short and oblique-

ly ventro-lateral on main axis, opposite or at base of pri-

mary branch, ± sessile, not seen with archegonia, only

with very young calyptra, up to 500 pm long, including

corona, the latter with apical row of radiating cells, width

across widest part below 480-525 pm, including sur-

rounding paraphyses, which are scale-like and dentate,

with cells 40.0-52.5 x 27.5-30.0 pm. Calyptra (Figure

1M, O) clavate, up to 2.4 x 0.7 mm, cross section of wall

± 180 pm or 5/6 cell layers thick (Figure IP), some out-

ermost ones with low protrusions, at apex corona

smoothly rounded above, cells thick-walled, dislodged at

maturity by emerging capsule. Seta ± 6.5 mm long, ± 280

pm wide, with 4 central and an outer row of 12 cells, i.e.

4 cells diam. (Figure IQ). Capsule ± ellipsoidal,

790-850 pm long, elaterophore internally attached to

apex; valves 280-350 pm or 21-24 cell rows wide, cells

of epidermal layer in external longitudinal view (Figure

2A), 55-110 x 10.0-12.5 pm, with strong vertical (radi-

al) thickenings, evident as nodular brown bulges; in cross

section (Figure 1R) cells rectangular, ± 7.5 pm thick,

thickenings on adaxial radial and abaxial radial walls

extending slightly onto outer and inner tangential walls,

resulting in a crescentic ring of thickening, bands on one

side of median wall alternating with those on the other

side in a mirror image; inner cells in internal longitudinal

view (Figure 2B) 67.5-85.0 x 12.5-17.5 (-20.0) pm,

with nodular thickenings and with faint semiannular

bands; in cross section (Figure 1R) cells ± 10 pm thick,

thickened on adaxial radial and abaxial radial walls,

extending slightly onto outer tangential walls and further

across inner tangential walls. Spores (Figure 1 T i_3) ±

orbicular, 12.5-15.0 pm diam., translucent, scabrate.

Elaters (Figure IS) 97.5-317.5 x 7.5-10.0 pm, with sin-

gle spiral band, tapering to unspiralled apical portions.

Chromosome no.: n = unknown. [It was suggested by

Hewson ( 1970), that the ‘normal’ number of n = 10 may

be associated with the dioicous state and the monoicous

state with double that number, i.e. 20],

DISCUSSION

The occurrence of Riccardia compacta in southern

Africa appears to be restricted to Western Cape (Figure

3), generally at altitudes between 1 085 and 1 220 m, but

it has also been collected close to sea level. It usually

grows in wet, shady places, with low light intensity, on

rotten wood, soil banks or on Table Mountain Sandstone.

The oil bodies were initially and briefly described by

Arnell (1952) as ‘rare, 3-8 x 8-22 pm, brownish’ and he

also illustrated a cell containing one (Arnell 1952: fig.

2h). In a much more detailed account, Meenks & Poes

(1985) observed that the oil bodies were ‘rare or even

absent in the epidermal cells and never in the cells of the

wing’. They were, however, ‘frequently present in the

inner cells of the thallus’, and were described by the

above authors as ‘light brown, 1 or 2 per cell’, as well as

‘globose to ellipsoid or bean-like, 8 x 8-30 x 12 pm’.

18

Bothalia 32.1 (2002)

FIGURE 3. — Distribution of Riccardia compacta in southern Africa.

During a visit to Table Mountain (and other areas in

Western Cape) in October 2000 by Perold & Koekemoer,

we were unable to find fresh material in order to study

the oil bodies. All the collections that I examined, had

long ago lost their oil bodies. The colour of fresh plants

could also not be observed, but Pillans had noted on the

label of his collection from Table Mountain, that they

were bright green and Meenks & Poes (1985) stated that

they were emerald green to dark brown.

In a comparison between plants from Tanzania and

those from South Africa (with information reported by

Arnell 1963), Meenks & Poes (1985) found that the for-

mer specimens were generally longer at 20 mm versus 7

mm; in transverse section 6 or 7 cell layers thick, versus

4 or 5 cell layers; spore diameter 12-22 pm versus 12-14

pm; wing of main axis 1 cell wide, cells elongated, up to

75 x 40 pm, versus wing absent.

In my findings the thalli of our plants are up to 15 mm

long and in cross section the ultimate segment of the

main axis is 6 or 7 cell layers thick, the spore diameter is

1 2.5- 15.0 pm, and the male branches are shorter at ± 600

pm long, with 3 or 4, rarely up to 8 pairs of antheridia;

the wing is formed by a row of elongated, apically round-

ed cells that are laterally adjoined, but apically free. Male

branches in the Tanzanian specimen that I studied are

mostly longer than ours, i.e. up to 1600 pm long, with

4-14 pairs of antheridia, the wing consisting of a row of

quadrate to rectangular cells, 42.5-45.0 pm high,

37.5- 42.5 pm wide, their sides joined together right up to

the edge; ventral ly there are 2 rows of conspicuous red-

brown mucilage papillae, up to 60 pm long, rarely seen

in South African specimens. As Meenks & Poes

observed, there are a few differences between Tanzanian

and South African plants. The general facies of the plants

from the two areas is, however, closely similar and there

is no doubt that they belong to the same species.

Meenks & Poes expressed the opinion that all conti-

nental African records of R. caespitans (Steph.)

E.W.Jones (nom. inval.) probably belong to R. compacta.

Apparently, Jones ( 1 956) had identified Hedberg speci-

mens from Mt Muhawara, Uganda, as ‘possibly Aneura

caespitans ’, but had added that R. caespitans is ‘known

with certainty only from the Mascareignes’ [Stephani

(1892) Bourbon, leg. Rodriguez |. I have examined

Dusen GO 1 0680 from Cameroon, incorrectly labelled as

‘TYPUS' with a red sticker, and also figure no. 236 in

Stephani’s leones hepaticarum (1985), which was prob-

ably drawn from the aforesaid Dusen specimen (accord-

ing to a note with it). This plant is decidedly different

from R. compacta specimens: in cross section the ulti-

mate segments of the main axis are 4 cell layers thick and

winged, with 3 unistratose cell rows at both margins. The

lower axis in cross section is 6 cell layers thick median-

ly and not 9 cells as in Stephani’s leones hepaticarum.

Whether Dusen’s specimen belongs to the same species

as Rodriguez’s type collection from Reunion, is very

doubtful, as the latter species was reported by Stephani to

be 10 cell layers thick medianly.

For several reasons I also hesitate to accept R. kili-

mandjarica as a synonym of R. compacta, as was done

by Meenks & Poes. In R. kilimandjarica the thalli appear

to be lighter-coloured, lacking the central dark line, and

more delicate than in R. compacta. Cross sections of the

ultimate segment of the main axis are ± 500 pm wide and

5 or 6 cell layers rows (or 150-170 pm) thick, with the

medullary cells clearly thin-walled, not thick-walled as

in R. compacta.

Schuster (1963) suggested that a number of species,

among them R. compacta, may be assigned to the sub-

genus Phycaneura, which is, however, characterized by

relatively large, thin-walled dorsal epidermal cells.

Although they had not studied specimens and illustra-

tions of R. compacta , Brown & Braggins (1989) think

that the species that Schuster (1963) grouped together,

appear to be members of section Alcicornia , subgenus

Riccardia, rather than of subgenus Phycaneura. Stephani

(1899) had placed together some of these species, e.g. A.

alcicornis, A. compacta, A. diminuta, A. subsimplex, A.

sumatrana and others, as ‘Plantae filiformes, subteretes’.

R. compacta appears to bear some similarities to R. sub-

alpina Furuki ( 1991) from Japan, as it also grows (most-

ly) at higher altitudes, and has small thalli and thick-

walled cells.

R. compacta specimens are distinguished by the fol-

lowing characters: 1, the plants grow in dense, thick

cushions of several layers of intricately intertwining

branches, mostly at high altitudes; 2, the thalli are narrow

and small, only up to ± 15 mm long, with irregularly

branched, lateral branches, their margins generally paral-

lel and unwinged; 3, from above the dorsal cells of the

ultimate segment of the main axis are thick-walled, and

in cross section, so are the medullary cells; 4, the

antheridia] branches in South African plants are usually

short and have conspicuous marginal cells; and 5, the

corona of the calyptra is smooth and the cells thick-

walled.

SPECIMENS EXAMINED

R. compacta

Arnell 96.1, Table Mountain (S): 1623. near Park Station. Knysna (BOL).

Bothalia 32,1 (2002)

19

Esterhuysen 24253 A , 24255, N side of Tsitsikama Mts (BOL); 25497.

Apollo Peak. Cederberg (BOL. PRE).

Jelinek 19560G (holotype). Table Mountain (G).

Pillans 4247, Table Mountain (BOL). Poes & Ochyra 88152/R, Mt

Meru. Tanzania (PRE).

Quite a few specimens (held at BOL. PRE & S) have been incorrectly

referred to R. compacta , probably because of the small size of some of

these plants. Arnell 1689, Gouna Forest. Knysna (PRE) is, in fact, R.

multifida (Perold 2001).

R. kilimandjarica

Esterhuysen 27249 (holotype of R. kilimandjarica). Mt Kilimanjaro,

Tanzania (S).

ACKNOWLEDGEMENTS

The curators of BOL. G and S are thanked for the loan

of specimens. I also express my most sincere gratitude to

Dr M. Koekemoer. curator of PRE. for all her help with

fieldwork, and to the referees for their helpful sugges-

tions and advice. A special word of thanks to Mr M.

Wigginton (UK) for very kindly procuring and sending a

photocopy of the Renauld & Cardot paper. The artist.

Mrs M. Steyn. the photographer, Mrs A. Romanowski

and the typist, Ms D. Maree are warmly thanked for their

valued contributions.

REFERENCES

ARNELL. S.W. 1952. South African species of Riccardia. Botaniska

Notiser 1952: 138-156.

ARNELL, S.W. 1959. A new species of Riccardia from Kilimanjaro.

Svensk Botanisk Tidskrift 53: 543. 544.

ARNELL, S.W. 1963. Hepaticae of South Africa. Swedish Natural

Science Council, Stockholm.

BROWN, E.A. & BRAGGINS, J.E. 1989. A revision of the genus

Riccardia S.F.Gray in New Zealand with notes on the genus

Aneura Dum. Journal of the Hattori Botanical Laboratory 66:

1-132.

FURUKI, T. 1991. A taxonomical revision of the Aneuraceae

(Hepaticae) of Japan. Journal of the Hattori Botanical

Laboratory 70: 293-397.

HEWSON, H.J. 1970. The family Aneuraceae in Australia and New

Guinea: II. The genus Riccardia. Proceedings of the Linnean

Society of New South Wales 95: 60-121.

JONES. E.W. 1956. African hepatics XL The genus Riccardia in trop-

ical Africa. Transactions of the British Bryological Society 3:

74-84.

MEENKS. J.L.D. & POCS, T. 1985. East African bryophytes IX

Aneuraceae. Abstracta Botanica 9: 79-98.

PEROLD, S.M. 2001. Studies in the liverwort family Aneuraceae

(Metzgeriales) from southern Africa. 2. The genus Riccardia

and its type species R. multifida, with confirmation of its pres-

ence in the region. Bothalia 31: 183-187.

SCHUSTER. R.M. 1963. Studies on antipodal Hepaticae. I. Annotated

keys to the genera of antipodal Hepaticae with special reference

to New Zealand and Tasmania. Journal of the Hattori Botanical

Laboratory 26: 185-309.

SIM. T.R. 1926. The Bryophyta of South Africa. Transactions of the

Royal Society of South Africa 15: 1—475. Cape Town.

STEPHANI, F. 1892. Hepaticae. In F. Renauld & J Cardot. Musci

exotici novi vel minus cogniti. Bulletin de la Societe Royale

Botanique de Belgique 30: 194-207.

STEPHANI. F. 1893. Hepaticae species novae. 1. Hedwigia 32: 17-29.

STEPHANI. F. 1899. Species hepaticarum 1: 97-304.

STEPHANI. F. 1901-1905. Species hepaticarum II: 727-764.

STEPHANI, F. 1985. leones hepaticarum. Microfiche, Inter Documen-

tation Company bv. Leiden.

Bothalia 32.1: 21-28 (2002)

Revision of the Tarchonanthus camphoratus complex (Asteraceae-

Tarchonantheae) in southern Africa

P.P.J. HERMAN*

Keywords: Asteraceae, new species, resurrected names, revision, southern Africa, Tarchonanthus camphoratus L.

ABSTRACT

The Tarchonanthus camphoratus L. complex in southern Africa is revised. Five species are recognized on the grounds

of differences in synflorescences, flowering times, leaf shape and margin, and distribution. Two names, T. minor Less, and

T. obovatus DC., are resurrected and two new species, T. littoralis RRJ. Herman and T. parvicapitulatus P.P.J. Herman, are

described. A key, full descriptions of each taxon and distribution data are presented.

INTRODUCTION

Plants belonging to the Tarchonanthus camphoratus

L. complex are well known in southern Africa and are

commonly known as camphor bush Ikanferbos (Afri-

kaans) (Smith 1966; De Winter et al. 1978) or African

fleabane, camphor wood, sagewood, wild cotton, wild

sage, basterolien, bastervaalbos, bergvaalbos, kanfer-

boom, kanferhout, kapokboom , kleinvaalbos, salie ,

saliehout , vaalbos, veldvaalbos, waaibos, wildesalie and

witbos (Afrikaans) (Wells et al. 1986). or camphor tree,

siriehout (Afrikaans), isiDuli selindle (Xhosa), isiDuli-

sehlathi, iGqebe-elimhlophe (Zulu), mofahlana (South

Sotho), mohatlha (Tswana), sefahla (North Sotho),

moologa (Venda) and omutea (Herero) (Palmer &

Pitman 1972). Various parts of the plants are used medi-

cinally (Watt & Breyer-Brandwijk 1962; Smith 1966;

Palmer & Pitman 1972). The wood was used for turnery,

boat-building, musical instruments, cabinet work, fence

posts, shafts of spears and walking sticks (Smith 1966;

Palmer & Pitman 1972; Coates Palgrave 1977).

Tarchonanthus camphoratus is used as a fodder plant in

dry areas (Smith 1966; Palmer & Pitman 1972; Tree

Society of Southern Africa 1974). It is a very common

element in some areas and some veldtypes have been

named after it, e.g. the subdivisions of the Vryburg Shrub

Bushveld by Acocks (1988) into the Tarchonanthus Veld

of the Ghaap Plateau, the Mixed Tarchonanthus Veld of

the Asbestos and Kuruman Hills, the Mixed Tcirchonan-

thus-Rhus-Croton Veld of the Langeberg and the Mixed

Tarchonanthu$-Thomve\d of the Kimberley plains and

koppies. In all these cases, Tarchonanthus camphoratus

L. sens. str. is involved.

A number of different names have been published for

various Tarchonanthus taxa, but they were all put into

synonymy under T. camphoratus by Paiva (1972) and

subsequently by Hilliard (1977), Pope (1992) and

Beentje (1999). Several workers maintained that there

were different taxa under T. camphoratus (Acocks 1988;

A. Gubb pers. comm.), with differences in palatability

* National Botanical Institute, Private Bag X101, 0001 Pretoria.

MS. received: 2001-03-19.

(Burchell 1824; A. Gubb pers. comm.). While I was

preparing a manuscript for Flowering Plants of Africa

(Herman & Condy 2001 ), it became clear that there were

in fact a number of different taxa grouped under T. cam-

phoratus. Differences in synflorescences, flowering

times, leaf shape and margin and distribution, led to the

recognition of five species. Two names are herewith res-

urrected and two new species described.

The septate hairs present in the capitula, which can be

considered a generic characteristic for the genus, are

described by Herman (2001 ). The descriptions of the leaf

shapes are based on that proposed by Radford et al.

(1974) and Radford (1986).

Key to species of Tarchonanthus

For the sake of completeness, Tarchonanthus trilobus

(also occurring in southern Africa) is included in the key.

la Involucral bracts of male and female capitula linear, free;

adaxial leaf surfaces bullate (Figure 1A) . . . T. trilobus

lb Involucral bracts of male and female capitula lanceolate,

elliptic, ovate or obovate, free, or of male capitula fused

to ± halfway: adaxial leaf surfaces reticulate (Figure

IB):

2a Capitula in dense, spicate, axillary or terminal clusters

(Figure 2A. C); involucral bracts of male and female

capitula free; peak flowering time September to

December (spring to early summer) 1. T. minor

2b Capitula in lax, open panicles (Figure 2B. D); involucral

bracts of male capitula fused to halfway, of female

capitula free; peak flowering time February to August

(late summer, autumn and winter):

3a Leaf margins always entire:

4a Leaves narrowly elliptic to slightly oblanceolate. acute

or obtuse; cottony hairs enveloping cypselas

creamy or yellowish; widespread . 3. T. camphoratus

4b Leaves obovate or elliptic, obtuse or acute; cottony

hairs enveloping cypselas pure white; restricted to

Gordonia area (Northern Cape) mostly on iron

stone or limestone base in sandy soils . . . 4.T. obovatus

3b Leaf margins often faintly denticulate in upper part or entire:

5a Leaves large, up to 140 x 45 mm; petioles up to 12 mm

long; fruiting capitula large, up to 15 mm diam.;

growing along coast from southern KwaZulu-

Natal to Western Cape 2. T. littoralis

5b Leaves smaller, up to 60 x 16 mm; petioles up to 5 mm

long; fruiting capitula small, up to 10 mm diam.;

Bothalia 32,1 (2002)

22

FIGURE 1. — Scanning electron micrographs of adaxial leaf surfaces of Tarchonanthus species. A, bullate surface of T. trilobus var. trilobus,

Strey 9016 (PRE); B, reticulate surface of T. minor, Leistner 495 (PRE). Scale bars: 100 pm.

growing inland, occurring in forests, valleys or

bushveld on mountain slopes, hills and river

banks; Northern Province, North-West, Gauteng,

Mpumalanga, Swaziland and KwaZulu-Natal . . .

5. T. pan’icapitulatus

1. Tarchonanthus minor Less., Synopsis generum

Compositarum: 208 (1832); DC.: 431 (1836); Harv.: 118

(1865). Type: South Africa, [Orange] Free State, 2925

Jagersfontein, Fauresmith, under edge of plateau of

Langeberg, (-CB), NW slope, 21 October 1925, Smith

942 (PRE!, neotype, here designated).

T. angustissimus DC.: 431 (1836). Syntype: South Africa,

[Northern] Cape, Philipstown, on the Table Mountain, near Horse’s

Grave (= Paardeberg), ??Karrapoort, Burchell 2691 [G-DC; K!, lecto.

chosen by Beentje: Kew Bulletin 54,1: 83 (1999)-PRE photo.!].

Dioecious shrub or small tree up to 5 m high. Leaves

aromatic, alternate, small, narrowly elliptic (Figure 3A),

1 0 — 40(— 52) x 2.5-15.0 mm, discolorous, upper surface

bright to dark green, pubescent when young, becoming

glabrous except for hairs in lower half of furrow caused

by sunken main vein, reticulate, with golden glands on

margins of reticulations (Figure IB), lower surface

densely whitish tomentose; apex acute, often mucronate

subapically; base cuneate; margin entire, rarely denticu-

late. Petiole l-3(rarely up to 5) mm long. Synflorescences

dense, spicate, axillary and terminal clusters (Figure 2A,

C). Male plants : capitula homogamous discoid, 7-12

mm diam., 15-80-flowered, sessile. Involucral bracts

densely hairy, free, in ± 3 rows, imbricate; outer ovate,

acute to acuminate, up to 7 x 5 mm; middle row ovate,

acute or obovate, 4x3 mm; inner row linear, 5 x 0.5

mm. Corolla infundibuliform, hairy and glandular; tube

5-6 mm long; lobes 5, 1. 5-2.0 mm long, recurved, papil-

late. Anthers exserted, with ovate apical appendage, cal-

carate, caudate, tails ± I mm long, branched; filaments

1 .5 mm long. Style up to 6 mm long, well exserted, undi-

vided or with 2 very small lobes, papillate. Ovary nar-

rowly oblong, sterile, with long, septate hairs. Female

plants : capitula homogamous discoid, 6-10 mm diam.,

1-1 1-flowered, sessile or peduncle up to 3 mm long.

Involucral bracts densely hairy, free, in 2 or 3 rows;

outer ovate, acute or obovate, obtuse, 4-7 mm long; mid-

dle row obovate, obtuse or acute, 5-7 mm long; inner

row oblanceolate, 4-7 mm long. Corolla infundibuli-

form, hairy and glandular; tube 1 .5-2.0 mm long; lobes

5, 1.0-1. 5 mm long, papillate, recurved. Style well

exserted, 4 mm long, with 2 broad, 0.5- 1.0 mm long

lobes. Ovary elliptic, 3^4 mm long, glandular and sep-

tate-hairy. Cypsela enveloped by long, white, silky, sep-

tate hairs. Pappus absent. Flowering time: August to

December with a peak from September to December

(spring to early summer).

Distribution and habitat: Tarchonanthus minor

occurs in the Free State, Lesotho, Northern, Western and

Eastern Cape on hillslopes, mountainsides, rocky ridges

and hills (Figure 4).

Vernacular names: small-leaf camphor bush, klein-

blaarkanferbos (Afrikaans) are here proposed.

Notes : 1 ) Lessing described T. minor in 1832 but cited

no specimen (see also Pope 1992); neither did De

Candolle ( 1836), but only referred to Lessing’s descrip-

tion. Beentje (1999) stated that the plants Lessing saw,

were destroyed at B. Harvey (1865) cited a few speci-

mens. I chose Smith 942 as neotype as Smith made a note

on his specimen: ‘Agrees in shape and size of leaves with

Cooper 708 cited by Harvey in Flora capensis 3: 118

under T. minor Less., but Cooper specimen is barren'.

2) The name ‘minor’ has been misapplied to almost

all species of Tarchonanthus recognized here. Tarcho-

nanthus minor is distinguished from the other species by

the small, narrowly elliptic, entire leaves, dense, spicate

synflorescences and flowering time in spring and sum-

mer. The leaves in some of the Lesotho specimens reach

the upper limit of the range in leaf size.

2. Tarchonanthus littoralis P.PJ. Herman, sp.

nov., T. camphoratus L. affinis sed folia magna, oblan-

ceolata, elliptica ad anguste elliptica, raro obovata,

( 3 2— )40— 1 40 x 10-45 mm, bicoloria, supra atrovirentia.

Bothalia 32,1 (2002)

23

FIGURE 2. — Synflorescences of Tarchonanthus species. A, dense, spicate male synflorescences of T. minor, Galpin 13954 (PRE); B, paniculate

male synflorescences of T. camphoratus, Story 1057 (PRE); C, dense, spicate female synflorescences of T. minor Liebenberg 7317 (PRE);

D. paniculate female synflorescences of T. parvicapitulatus, Botha 2660 (PRE). Scale bars; 10 mm.

infra albida; margine integra vel plerumque apicem ver-

sus leviter denticulato; petiolus 5— 8(— 1 2) mm longus;

cypsela pilis involvens albidis; plerumque littora

incolens.

TYPE. — South Africa, [KwaZulu-]Natal, 3030 Port

Shepstone, Uvongo, Deppe’s road, (-CB), 10 March

1970, Strey 9713 (PRE, holo.!).

Dioecious shrubs or trees, 1-8 m high. Leaves aro-

matic, alternate, large, oblanceolate, elliptic to narrowly

elliptic, rarely obovate (Figure 3B, C), (32-)40-140 x

24

Bothalia 32,1 (2002)

FIGURE 3. — Leaf shapes of Tarclionanthus species. A, small, narrow-

ly elliptic, entire leaf of T. minor. Braack 22 (PRE). B, C,

Tarchonanthus littoralis : B, large, narrowly elliptic leaf,

O’Callaghan, Fellingham & Van Wyk 186 (PRE); C, oblanceo-

late leaf with denticulate margins, Osborne 18 (PRE). D. nar-

rowly elliptic, entire leaf of T. camplioratus, Germishuizen 369

(PRE). E, F, Tarchonanthus obovatus: E, obovate, entire leaf,

Gubb 1562 (KMG); F, elliptic, entire leaf, Gubb 1519 (KMG).

G, H, Tarchonanthus parvicapitulatus: G, oblanceolate, obtuse

leaf with entire margins, Mogg PRE43479 (PRE); H. oblanceo-

late, obtuse-mucronate leaf with denticulate margins, Compton

27888 (PRE). Scale bar: 10 mm. Drawn by G. Condy.

10-45 mm, discolorous, upper surface bright or dark

green, hairy when young, becoming glabrous, reticulate,

with glands on margins of reticulations, main vein

sunken and hairy, especially in lower half, lower surface

densely while-hairy, main and secondary veins conspic-

uous; apex obtuse to acute, sometimes with curved

mucro subapically; base cuneate; margin very often

faintly denticulate in upper part, rarely entire. Petiole

5— 8(— 12) mm long. Synflorescences terminal, paniculate.

Male plants’, capitula homogamous discoid, 5-10 mm

diam., 1 3-47-flowered; peduncle 2-12 mm long.

Involucral bracts densely hairy, fused halfway, 5-lobed;

tube 1. 5-3.0 mm long; lobes 1 .5-2.0 mm long; some-

times with few free, inner bracts. Corolla infundibuli-

form, glandular and hairy; tube 2. 0-3. 5 mm long; lobes

5, 1 .0-1.5 mm long, papillate, recurved. Anthers 1 .0-1 .5

mm, exserted, with ovate-triangular, apical appendage,

calcarate, caudate, tails ± 0.5 mm long, branched; fila-

ments 1. 5-3.0 mm long. Style well exserted, 5-8 mm

long, undivided or with 2 small lobes, papillate. Ovary

rod-like, 0. 5-1.0 mm long, glandular, sterile, septate-

hairy. Female plants’, capitula homogamous discoid, ± 5

mm diam., mostly 3(-6)-flowered; peduncle 0-4 mm

long. Involucral bracts free, 3-6 mm long, in 2 or 3

rows; outer bracts ovate or narrowly lanceolate-ovate,

acute; inner elliptic. Corolla infundibuliform, glandular

and hairy; tube 1 .5-2.0 mm long; lobes 5(6), 0. 5-1.0 mm

long, papillate, recurved. Style exserted, 2. 5-3. 5 mm

FIGURE 4. — Distribution of T. minor in southern Africa based on

material at PRE.

long, with 2 short branches, ± 1 mm long. Ovary elliptic

to obovate, 2. 5-4.0 mm long, densely septate-hairy and

glandular. Cypsela obovate to elliptic, ribbed, 3-5 mm

long, densely, white, septate-hairy and glandular. Pappus

absent. Flowering time : December to July with a peak

from February to March (July) (late summer).

Distribution and habitat: Tarchonanthus littoralis

occurs from southern KwaZulu-Natal to Eastern and

Western Cape along the coast, on hillsides, littoral dunes

and river banks (Figure 5).

Vernacular names: coastal camphor bush, kuskanfer-

bos (Afrikaans), are here proposed.

Tarchonanthus littoralis is distinguished by the usual-

ly large, oblanceolate to elliptic or narrowly elliptic, dark

green leaves with mostly denticulate upper margins and

long petiole. The cypsela-containing capitula are rather

FIGURE 5. — Distribution in southern Africa of T. littoralis. •. based

on material at PRE; and T. camplioratus. ■, based on material

at KMG and PRE.

Bothalia 32,1 (2002)

25

large. This is probably the species described and illus-

trated by Von Breitenbach (1974) under T. camphoratus.

It has often been confused with Brachylaena species.

3. Tarehonanthus camphoratus L., sens, str..

Species plantarum: 842 (1753); Less.: 208 (1832); DC.:

431 (1836); Harv.: 118 (1865); Merxm.: 176 (1967);

Paiva: 360 (1972); Compton: 622 (1976); Hilliard: 111

(1977); Pope: 9 (1992); Beentje: 82 (1999). Type:

'Aethiopia’, in this case South Africa; Hort. Cliff, has

'Cap. Bon. Sp.' (Herb. Cliff. 398 , Tarehonanthus no. I ,

BM, lectotype chosen by Anderberg in Jarvis et al. 1993:

92, see Beentje 1999-PRE, photo.!).

T. litakunensis DC.: 431 (1836); excluding lectotype chosen by

Beentje (1999), here placed under T. littoralis. Lectotype: Burchell

2202 (G-DC, here designated-PRE. photo.!).

T. camphoratus L. var. litakunensis (DC.) Harv.: 118 (1865).

Mostly a multi-stemmed, rounded dioecious shrub,

rarely a tree, 1-8 m high. Leaves aromatic, alternate, nar-

rowly elliptic to slightly oblanceolate (Figure 3D),

(20— )26— 80 x 7-20 mm, discolorous, upper surface grey-

green or khaki-green, hairy when young, becoming

glabrous, reticulate, with glands on margins of reticula-

tions, main vein sunken, hairy in lower part, lower sur-

face densely grey- or greenish hairy, main and secondary

veins prominent, tertiary veins reticulate; apex acute,

often mucronate, sometimes obtuse; base cuneate; mar-

gin entire. Petiole up to 5 mm long. Synflorescences ter-

minal. paniculate (Figure 2B). Male plants : capitula

homogamous discoid. 3.0-11.5 mm diam., 10-60-flow-

ered; peduncle 0-10 mm long. Involucral bracts fused

halfway, 5(-7)-lobed; tube 1. 5-5.0 mm long; lobes

1.0-2. 5 mm long, sometimes with a few free bracts on

inside. Corolla infundibuliform. glandular and hairy;

tube 1.5-3. 5 mm long; lobes 5, 0. 5-1.0 mm long, papil-

late, recurved. Anthers well exserted, 1.0-1. 5 mm long,

with small, ovate, apical appendage, calcarate, caudate,

tails up to 0.5 mm long, branched; filaments 2-3 mm

long. Style well exserted, 4-8 mm long, undivided or

rarely with 2 small lobes, papillate. Ovary rod-like,

0. 5-1.0 mm. sterile, septate-hairy. Female plants: capitu-

la homogamous discoid, 3. 5-7.0 mm diam., ( 1— )3(— 5 )-

flowered; peduncle 0-6 mm long. Involucral bracts

densely hairy, in 2 or 3 rows, free, imbricate, 3-6 mm

long; outer bracts linear-lanceolate to linear-obovate:

middle and inner row ovate. Corolla infundibuliform,

hairy and glandular; tube 1.0-1. 5 mm long; lobes 5, 0.5

mm long, papillate. Staminocles sometimes present. Sf’le

exserted. 1. 5-3.0 mm long, with 2 broad, 0.5 mm long

lobes. Ovary elliptic to obovate, 1^1 mm long, densely

septate-hairy and glandular. Cypsela elliptic, ribbed. 3^f

mm long, creamy or yellowish, densely septate-hairy and

glandular. Pappus absent. Flowering time : March to

August with a peak from March to July (autumn to win-

ter).

Distribution and habitat: Tarehonanthus camphoratus

is widely distributed in the northern part of southern

Africa from Namibia, Botswana, the Northern Province,

North-West, Gauteng, Free State and Northern Cape

(Figure 5). It also occurs in tropical and North Africa and

the Arabian Peninsula. It occurs in a variety of habitats

and soil types, e.g. savanna, bushveld. woodland, grass-

FIGURE 6. — Distribution in southern Africa of T. obovatus. ■.based

on material at KMG and PRE; and T. parvicapitulatus. •. based

on material at PRE.

land, on flats, rocky hills, mountain slopes and hillsides,

riverbanks on sandy, loamy, gravelly, calcrete, quartzite

and dolomitic soils.

Vernacular names: camphor bush, kanferbos

(Afrikaans) as listed by De Winter et al. (1978) is pro-

posed for T. camphoratus sens. str.

Notes: 1) It is very unfortunate that Beentje (1999)

chose Drege 5041 as lectotype for T. litakunensis.

'Fitakun' (Takoon 2723BB) is a small settlement near

Kuruman (Burchell 1824; Leistner & Morris 1976) in the

North-West. However. T. littoralis always grows near the

sea and the specimen chosen by Beentje (Drege 5041)

was collected near the sea. This specimen (Drege 5041)

also conforms to the description of T. littoralis presented

here. It is therefore proposed that Drege 5041 be reject-

ed as lectotype of T. litakunensis.

2) This is the most common taxon of the genus in the

northen parts of southern Africa. It can be distinguished

by the narrowly elliptic to slightly oblanceolate, entire

leaves with a grey-green or khaki-green colour and the

cypselas enveloped by yellowish cottony hairs. Not

browsed by stock (Burchell 1824) except as a last resort

in times of drought [A. Gubb pers. comm.. Speedy 11/22

(PRE)].

4. Tarehonanthus obovatus DC., Prodromus 5:

431 (1836), Herman & Condy: 108, t. 2180 (2001 ). Type:

South Africa, Bechuanaland Division (?Northern Cape),

Klipfontein, Burchell 2155 (G-DC. holo.; K!-PRE, photo.!).

Mostly a single-stemmed dioecious tree or rarely a

shrub, up to 2 m high. Leaves aromatic, alternate, obo-

vate or elliptic (Figure 3E, F), 1 2— 32(— 37 ) x 7-17 mm,

discolorous, upper surface bright green, hairy when

young, becoming glabrous, reticulate, glandular in reticu-

lations, main vein sunken and hairy in lower part, lower

surface densely whitish pubescent, main and secondary

veins prominent, minor veins forming a reticulation;

apex obtuse or acute; base cuneate; margin entire. Petiole

26

Bothalia 32,1 (2002)

1. 0- 4.0 mm long. Synflorescences terminal, paniculate.

Male plants: capitula homogamous discoid, 8-1 1 mm

diam., 20-40-flowered; peduncle 0-7 mm long.

Involucral bracts fused halfway. 5-lobed; tube 2.0-2. 5

mm long; lobes 2. 0-2. 5 mm long. Corolla infundibuli-

form, hairy and glandular; tube 1.5-2. 5 mm long; lobes

5(6), 1.0-1. 5 mm long, papillate. Anthers 5(6), exserted,

1.0- 1. 5 mm long, calcarate, caudate, tails ± 0.5 mm long,

branched; filaments 2-3 mm long. Style well exserted,

6-7 mm long, undivided or with 2 small lobes, ± 0.5 mm

long, papillate. Ovary rod-shaped, 0.5-1. 0 mm long,

densely septate-hairy and glandular, sterile. Female

plants: capitula homogamous discoid, 3.5— 4.5 mm diam.,

2-5-flowered; peduncle 0-7 mm long. Involucral bracts

free, in 2 or 3 rows, 3-6 mm long; outer bracts obovate,

acute; inner spathulate or narrowly obovate, acute.

Corolla infundibuliform, glandular and hairy; tube

0.5- 1.0 mm long; lobes 5(6), ± 0.5 mm long, apex papil-

late. Style exserted, 1. 0-2.0 mm long; style branches 0.5

mm long. Ovary obovate, 2-3 mm long, glandular and

septate-hairy. Cypsela brown, obovate, ribbed, 2. 5-5.0

mm long, glandular and pure white, septate-hairy.

Pappus absent. Flowering time: March to May (autumn

to early winter).

Distribution and habitat: Tarchonanthus obovatus is

restricted to Northern Cape where it occurs on hillsides,

rocky outcrops or flats often on an ironstone or limestone

base in sandy soils (Figure 6).

Vernacular names: Gordonia camphor tree, Gordonia-

kanferboom (Afrikaans), was proposed for this taxon (Herman

& Condy 2001). It is known by fanners in that region as

olienvaalbos (Afrikaans) (A. Gubb pers. comm.), but vaal-

bos refers to Brachylaena species (De Winter et al. 1978).

Note: it is interesting that, since the description, the

name T. obovatus has never been used. This species is

recognized by the obovate or elliptic, entire leaves and

the cypselas that are enveloped by pure white, cottony

hairs in contrast to T. camphoratus, occurring in the same

area, which have yellowish hairs enveloping the cypse-

las. Readily browsed by both game and domestic stock

all year round (A. Gubb pers. comm.).

5. Tarchonanthus parvicapitulatus P.P.J. Herman,

sp. nov., T. camphoratus L. affinis sed folia oblanceola-

ta, raro obovata, (22— )25— 60(— 70) mm, margine integra

vel plerumque apicem versus leviter denticulato; capitu-

la parva; synflorescentia laxa.

TYPE. — South Africa, Transvaal [Mpumalanga],

2531 Komatipoort, Barberton, (-CC), lower hill slopes,

April 1890, Galpin 952 (female plant) (PRE, holo.!).

Dioecious shrub or small, multi-stemmed tree, up to 8

m high. Leaves aromatic, alternate, oblanceolate or rarely

obovate (Figure 3G, H), (22-)25-60(-70) x 7-16 mm,

discolorous, upper surface pale, dark green, hairy when

young, becoming glabrous, glandular in reticulations,

midrib sunken and hairy in lower part, lower surface

densely whitish hairy, main and secondary veins promi-

nent; apex obtuse to obtuse-mucronate, sometimes acute;

base cuneate; margin often faintly denticulate in upper

part, rarely entire. Petiole 1. 5-5.0 mm long. Synflores-

cences terminal, paniculate (Figure 2D). Male plants:

capitula homogamous discoid, 5-9 mm diam., 10-30-

flowered; peduncle 0-7 mm long. Involucral bracts

fused halfway, 5-lobed; tube 1-2 mm long; lobes 1.0-1. 5

mm long. Corolla infundibuliform, glandular and hairy;

tube 1. 5-2.0 mm long; lobes 5, 1.0-1. 5 mm long, apex

papillate. Anthers 5, exserted, 1.0-1. 5 mm long, cal-

carate, caudate, tails 0.5 mm long, branched; filaments

2-3 mm long. Style well exserted, 4.0-5. 5 mm long,

entire or with 2 small lobes. Ovary rod-shaped, 0.5 mm

long, sterile, septate-hairy. Female plants: capitula

homogamous discoid, 3-4 mm diam., 1-3-flowered;

peduncle mostly absent or up to 5 mm long. Involucral

bracts free, 2-5 mm long, in 2 or 3 rows, narrowly lanc-

eolate, narrowly ovate or elliptic, acute. Corolla

infundibuliform, glandular and hairy; tube 0. 5-1.0 mm

long; lobes (4)5, 0. 5-1.0 mm long, papillate. Style exsert-

ed, 1-2 mm long, lobes 2, 0.5 mm long. Ovary obovate

to elliptic, 1.5-3. 5 mm long, glandular and septate-hairy.

Cypsela pale or dark brown, obovate, ribbed, 1. 5-3.0

mm long, glandular and pure white, septate-hairy.

Pappus absent. Flowering time: March to October with a

peak from April to June (autumn to winter).

Distribution and habitat: Tarchonanthus parvicapitu-

latus occurs in Namibia(?), Northern Province, the North-

West, Gauteng, Mpumalanga, Swaziland, KwaZulu-

Natal and Eastern Cape (only 2 records) (Figure 6). It is

also found in Zimbabwe (cf. Borle 157, Sim 19129,

Miller 1776, Ngoni 370 , Wild 1048). It grows in forest,

valleys and bushveld on mountain slopes, hills and river

banks.

Vernacular names: small-head camphor bush, klein-

hofiekanferbos (Afrikaans) are here proposed.

Note: this species is recognized by the oblanceolate,

obtuse-mucronate leaves mostly with denticulate mar-

gins towards the apex. The synflorescences tend to be

more open and the capitula are smaller than in the other

species. This is probably the species referred to by Moll

(1992) under T. camphoratus.

SPECIMENS EXAMINED

f, female; m, male.

Acocks 535 (3) (f & m), 543 (3) (f) PRE; 2109 (4) (f), 2117 (3) (m),

2344, 2351, 2352 (3) (f) KMG, PRE; 8617, 8627 (2) (f), 13010 (5) (m)

PRE; KMG 11730 (2) (f) KMG. Alexander PRE43490 (2) (f & m) PRE.

Allen 131 (3) (m) PRE. E.R. Anderson J.I4 (3) (m) PRE. J. Anderson

58 (1) (f & m) PRE. J.R. Anderson ORFS233 ( I ) (in) PRE. Archibald

3595 (2) (f) PRE.

M. Badenhorst 509 (3) (m) KMG. />./. Badenhorst 40, 104 (3) (f)

KMG, PRE. Balsinhas & Kersberg 1948 (3) (f) PRE. N. Barker 886 (5)

(m) PRE. N.P. Barker 621 (3) (m) PRE. Capt. Barrett- Hamilton

TRV6408 (3) (f) PRE. Basson 14 (3) (f) PRE. Bayliss BR1.B.351 (3),

BRI.B.6249 (2). BS8416 (2) (m) PRE. Bengis 447 (1) (f) PRE. Biggs

222 (5) (m) PRE. Boddam-Whethain 60 (1) (f) PRE. Bosch 154 (4) (f)

KMG. Botha 2559 (3) (f), 2660 (5) (f), 3067 (2) (m) PRE. Braack 22

(1) (m) PRE. Bradfield 250 (3) (m) PRE. Britten 33, 736 (2) (f) PRE.

4. Brueckner 845 (3) (m) KMG, PRE. Burchell 2155 (4) (f & m) K,

2691 (I) (sterile) K. Burrows 2102 (I) (m), 2286 (1) (f) PRE. Burn

Davy 148, 10741, 13933 (3) (f), 317(5) (m), 7865 (2) (f), 9537, 10375,

13820 (3) (m) PRE.

Bothalia 32,1 (2002)

27

Codd 1166 (5) (f), 1169 , 1935 (5) (m), 1283 (4) (f), 1283 (3) (m) PRE.

Collett 510 (3) (m) PRE. E. Collins PRE43484 (5) (m) PRE. .S’. Collins

12 (3) (m) PRE. Comins 1053 (2) (f) PRE. Compton 26015 , 27888,

28810, 32132 (5) (f). 28947 (5) (m) PRE. E.O. Cooke 6271 (3) (m)

KMG. O.L. Cooke 4265 (4) (m) KMG, 6266 collected 4/1940 (4) (m)

PRE. 6266 collected 4/1942 (4) (m). 6267 collected 4/1942 (4) (f)

KMG. Culverwell 23 (5) (m) PRE.

Dahlstrand 420 (2) (f) PRE. Davies, Thompson & Miller 8, 89 (3) (f)

PRE. De Souza 441 (5) (m) PRE. De Villiers PRE43498 (3) (f & m)

PRE. De Winter 2834, 3520 (5) (m), 9256 (2) (f) PRE. De Winter &

Leistner 5636 (3) ( f) PRE. Dieterlen 565 (1) (m) PRE. Dinter 4723 ( 3)

(m) PRE. Dix 211 (2) (m) PRE. Downing 560 (5) (m) PRE. Drege s.n.

(5041) (2) (f) K; PRE9727 (2) (f) PRE. Du Preez & Steenkamp 114 (3)

(f) PRE.

Ecklon & Zeyher 112.11 (1) (m) PRE. D. Edwards 2062 (5) (f), 2505

(5) (m), 4413 (3) (f) PRE. H.W.G. Edwards 52 (5) (f) PRE. C. Evrard

9276 (3) (f) PRE.

Farquharson A1483 (2) (f) PRE. Feely 32 (5) (m) PRE. Fellingham 963

(2) (f) PRE. Flanagan 192 (2) (f & m), 350 (2) (f). 1551 (1) (f), 2330

(5) (m) PRE. Fourie ORFS174 (1) (m) PRE.

Galpin 952 (5) (f & m), 7008 (3) (f). 13954 (1) (m), 133127 (5) (f),

M184 (3) (m) PRE. Gone 268 (2) (f) PRE. Germishuizen 369 (3) (f &

m), 387 (5) (f) PRE. Gerrard 1022 (2) (m) K (mounted with type).

Gerstner 593 (5) (m), 659 (5) (f) PRE. Giess 11693 (3) ( f) PRE.

Gilfillan Herb. Galpin 5533 (1) (m) PRE. Gillett 3434 (2) (m) PRE.

Glen 2761 (5) (f) PRE. Goldblatt 1424 (2) (f & m), 8013 (2) (f) PRE.

Goossens 710 (1) (f). 1673 (3) (f) PRE. Gubb 16 (4) (f), 18 (3) (m)

KMG; 168-1, 219/1, 260-84, 272-2 (4) (f) PRE; 341 (3) (m & f), 342,

343 (3) (f), 344 (4) (f), 345, 346 (3) (f). 347 (3) (m), 349 (3) (m & f).

350 (3) (0, 351 (4) (m), 352, 353, 355 (4) (f), 1088 (3) (m). 1097 (4)

(f), 1328, 1439 (4) (m), 1507 (4) (f), 1513 (4) (m), 1519, 1530 (4) (f),

1562, 1703 (4) (m), 7727(3) (m), 2105 (4) (m), 2221, 2222 (3) (f), 2243

(4) (f), 3603 (3) (m), 4361, 6002, 6516 (4) (f), 7442 (3) (m). 8118,

13680, 14303, 16125, 16127 (4) (f). 16128 (4) (m), 16130 (3) (m),

16131 (3) (f) KMG.

Hansen 3141 (3) (f) PRE. Henrici 4708 (1) (m) PRE. Herbert 29 (4)

(m) KMG. Herman 654 (3) (m). 1523 (4) (m). 1524 (4) (f) PRE.

Humbert 9537 (2) (f) PRE.

Jacobsen 2929 (3) (m) PRE. Jacot-Guillarmod 592 (1) (f), 9920 (2) (m)

PRE. Jarman 82 (1) (m) PRE. Joffe 831, 942 (2) (f), J1 (3) (f) PRE.

Jooste 104 (3) (m), 123 (3) (f), 216 (1) (f) PRE. M. Jordaan 1135 (5)

(f) PRE. P. Jordaan CBK9 (3) (f) PRE

Killick4304 (1) (f) PRE. King 69 (5) (f) PRE. Kotze43 (5) (m). 44 (5) (f) PRE.

Lang TRV31709 (3) (f) PRE. Leach & Bayliss 12920, 12962 (3) (f)

PRE. Leendertz 1136, 2142, TRV11245 (3) (f) PRE. Leighton 3112 (2)

(m) PRE. Leistner 495 (1) (m) PRE. Le Roux 366 (3) (f) PRE. Letty 223

(2) (f) PRE. L.C.C. Liebenberg 4480 (3) (f), 7377 ( 1) (f), 8078 (2) (m)

PRE. Liebenberg S.78, S.84 (3) (m) PRE. Louw 234 (3) (m) PRE.

MacDevette 252 (2) (f) PRE. Macdonald 76/24, 77/148 ( 3) (f) KMG,

PRE. MacMurray 5942 (3) (m), 5943 (3) (f) KMG; 5980 (4) (f) KMG,

PRE. Marais 1137 (2) (f) PRE. Marloth 557 (2) (f & m), 1009, 1329 (3)

(m), 5056, 12750 (3) (f) PRE. Mbedzi 1561 (5) (f) PRE. McClean 265

(2) (f) PRE. McDonald 77/121 (4) (f), 77/122 (4) (m) KMG. PRE.

McGregor Museum 5980 (4) (f) PRE. McMurtry 2432 (5) (m) PRE.

Miller B/202 (3) (f) PRE. Moffett 605 ( 1) (f) PRE. Mogg 4788, 13209

(2) (f), 7925 (3) (f & m), 8517 (3) (m), 16922 (3) (f). PRE43479 (5) (f)

PRE. Moore 11 (3) (f) PRE. Morze 2005 (2) (f) PRE. Mott 985 (3) (f)

PRE. Muir 132 (2) (f) PRE. Muller 1206, 1505 (3) (f) PRE. Munro

PS. 11 (2) (f), PRE2909 (5) (m) PRE. Murray 634 (3) (f) PRE.

Nelson 29 TRV11384 (3) (f) PRE.

Oates 312 (3) (m) PRE. O'Callaghan, Fellingham & Van Wyk 31 , 186

(2) (f) PRE. Osborne 18 (2) (m) PRE. Owens 99, 104 (3) (f) PRE.

Pagan 1729a (3) (f) KMG. Paterson 1001. TRV12337 (2) (f) PRE.

Pearson 1610 (3) (m) PRE. Peeters, Gericke & Burelli 4 (3) (f) PRE.

Phalatse 17 (3) (m) PRE. Phelan 1048 (3) (f) PRE. J. Phillips 1175,

1370 (3) (m) PRE. J.F.V. Phillips For.Herb.350 (2) (f) PRE. Pillans

3492 (2) (m) PRE. Pole Evans 1260. 2226 (3) (m), 2495, 2495(42) (4)

(f). 2496, 2496(43) (4) (m), 3594 (5) (f), H. 15695 ( 3) (f), H. 18041 (2)

(m) PRE. Pole Evans & Smith 1859 ( 1 ) (f) PRE. Potter ex Henrici 2030

(1) (m), 2031 (1) (f) PRE. Poynton For.Herb. 11990 (2) (f) PRE. Prior

PRE43495 (2) (f & m) PRE. Prosser 1844 (3) (f) PRE.

Rauh & Schlieben 9644 (3) (f) PRE. Rehm PRE43513 (1) (f) PRE.

Repton 616 (3) (f & m), 4525, 4562 (5) (f) PRE. Roberts 5379 (1) (m)

PRE. Rodin 2122 (3) (f), 3062 (2) (m) PRE. Rogers 24097 (5) (f), 26851

(2) (f), TRV22185 (5) (f) PRE. Romanowski 6 (2) (f) PRE. Rose-Innes 86

(5) (f) PRE. Ross 2117 (5) (f) PRE. Rossouw TRV26142 (3) (f) PRE.

Rowland, Sealt, Steyn PRE26464 (4) (f) PRE. Rudatis 1646 (2) (f) PRE.

Scharf 1140 (2) (m), 1393, 1419 (2) (f) PRE. Scheepers 1596 (3) (f)

PRE. Schlieben 8749 (3) (f) PRE. B. Schmidt 15 (3) (f) PRE. E. Schmidt

330 (3) (m) PRE. Schmitz 6850, 6851 (1) (m), 9262 (1) (f) PRE.

Schonland 1628 (3) (f) PRE. Shearing 1204 (1) (f) PRE. Silk 264 (3) (f)

KMG. Sim 2101, 20429 (2) (f) PRE. C.A. Smith 440A, 440B, 942 (1)

(f), 4417 (3) (f & m), 4502 (3) (f), 5452 (1) (f) PRE. PA. Smith 2591

(3) (f) PRE. Smuts 1417 (3) (f) PRE. (Mrs) J.C. Smuts PRE43492 (2)

(f) PRE. Smyth PRE43528 (2) (f) KMG, PRE. South 671 (2) (m) PRE.

Speedy 11/22 (3) (m) PRE. Stalmans 621 (5) (f) PRE. Story 101, 137

(1) (f), 1057 (3) (m), 1190 (4) (f), 4585, 4635, 4877, 4981, 5277 (3) (f)

PRE. Strev 3288 (5) (f). 6536 (2) (m). 7660a, 8825, 9713 (2) (f), 9709

(2) (m), 9757 (5) (m) PRE. Sutton 880 (5) (f), 1293 (3) (f) PRE.

Taylor 7111 (3) (m), 10219 (2) (f) PRE. Theiler TRV12362 (5) (f) PRE.

G.C. Theron 345 (1) (m), 815 (3) (f) PRE. G.K. Tlteron 1429 (5) (f)

PRE. J.J. Theron 636 (3) (m) PRE. Thode 2801 (5) (f), A527 (1) (f),

A861 (2) (m), A2479 (2) (f) PRE. Thorne PRE45409 (3) (f) PRE. Tyson

PRE3102, Herb.Marl.8541 (2) (m) PRE.

Van Breda 1147 (2) (f) PRE. Van Dam TRV18856 (2) (f) PRE. Van der

Schijff 6733, 8026 (3) (f) PRE. Van der Spuy 59 (3) (f) PRE. Van Eck 1

(3) (m), 2 (4) (f) PRE. Van Hoepen 1846 (4) (f) PRE. Van Son

TRV28805 (3) (m), TRV28806 (3) (f). TRV31769 (3) (m) PRE. A.E. van

Wyk 1842 (2) (f) PRE. P. van Wyk BSA1I8 (3) (f), BSA992 (5) (f) PRE.

Van Zinderen-Bakker 97 (3) (f), 997 (3) (m), 1063 (1) (f) PRE. F. Venter

933 (2) (f), 2006 (3) (m) PRE. H.J.T Venter 3749 (5) (m), 3750 (5) (f)

PRE. Verdoorn 1002 (1) (f), 1590 (1) (m), 1761 (3) (f). 1762 (3) (m),

PRE30042 (5) (f) PRE. Victor 347 (2) (f), 1452 (1) (f) PRE. A. Viljoen

69 (2) (f) PRE. G. Viljoen 56 (5) (m) PRE. Viviers 167 (2) (m) PRE.

Ward 1402, 2612, 7719 (5) (f), 2611, 2613, 7718 (5) (m) PRE. Walsh

PRE61373 (2) (f) PRE. Werdermann & Oberdieck 1578 (1) (m) PRE.

West 1230 (5) (f), 1804 (5) (m) PRE. White 5101 (2) (f), 10509 (5) (m)

PRE. Williamson 274 (5) (f) PRE. Wilman 1244, PRE43493 (3) (m)

PRE; 1728, KMG11733 (4) (m). 3319, 5990b, 6527 (3) (f) KMG; 5979,

5990, 6268, 6269 (3) (f) KMG. PRE. Wirminghaus 289 (2) (f) PRE.

Wood Herb.Galpin3229 (2) (m) PRE.

Zambatis 669 (5) (f) PRE. Zeyher 819 (2) (m) PRE.

ACKNOWLEDGEMENTS

A. Gubb, previously of the McGregor Museum,

Kimberley (KMG), is thanked for his field observations

and for bringing the different taxa in Northern Cape to

our attention. I am grateful to the curators of K, BM and

KMG for sending me material on loan and/or photo

images by e-mail. Dr O.A. Leistner is thanked for the

Latin translations, Ms G. Condy for the line drawings,

Ms A. Romanowski for photographic prints and Mrs S.

Turck for the scanning of micrographs and synflores-

cences from herbarium specimens.

REFERENCES

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the Botanical Survey of South Africa No. 57.

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BURCHELL, W.J. 1824. Travels in the interior of southern Africa, vols

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CANDOLLE, A.P. DE. 1836. Compositae. Prodromus 5: 431. Treuttel

& Wiirtz, Paris.

COATES PALGRAVE. K. 1977. Trees of southern Africa. Struik

Publishers, Cape Town.

COMPTON, R.H. 1976. The flora of Swaziland. Journal of South

African Botany, Suppl. vol. 11: 622.

DE WINTER, B„ VAHRME1JER. J. & VON BREITENBACH. F.

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28

Bothalia 32,1 (2002)

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HERMAN, PPL & CONDY, G. 2001. Tarchonanthus obovatus.

Flowering Plants of Africa 57: 108-112, t. 2180.

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LEISTNER. O.A. & MORRIS. J.W. 1976. Southern African place

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LINNAEUS, C. 1753. Species plantarum edn I: 842. Laurentius

Salvius, Stockholm.

MERXMULLER. H. 1967. Asteraceae. Prodromus einer Flora von

Siidwestafrika 139: 176.

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PALMER. E. & PITMAN, N. 1972. Trees of southern Africa , vol. 3.

Balkema, Cape Town.

POPE, G.V. 1992. 97. Compositae. Flora zambesiaca 6,1: 9-11.

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RADFORD, A.E.. DICKISON, W.C.. MASSEY, J.R. & BELL. C.R.

1974. Vascular plant systematics. Harper & Row, New York.

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

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the Witwatersrand. Witwatersrand University Press. Johan-nes-

burg.

VON BREITENBACH, F. 1974. Southern Cape forests and trees. The

Government Printer, Pretoria.

WATT, J.M. & BREYER-BRANDWIJK. M.G. 1962. The medicinal

and poisonous plants of southern and eastern Africa. Living-

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problem plants in southern Africa. Memoirs of the Botanical

Survey of South Africa No. 53: 509.

Bothalia 32,1: 29-36(2002)

Taxonomy of the Passerina filiformis complex (Thymelaeaceae)

C.L. BREDENKAMP* and A.E. VAN WYK**

Keywords: anatomy, new species, Passerina , southern Africa, taxonomy. Thymelaeaceae

ABSTRACT

Revision of the genus Passerina L. indicated a new delimitation of taxonomic entities within the Passerina filiformis L.

complex. Evidence from leaf anatomy greatly assisted in the recognition of taxa. P. filiformis is here divided into two sub-

species, namely P. filiformis subsp .filiformis and P. filiformis subsp. glutinosa (Thoday ) Bredenkamp & A.E. van Wyk, and

a new species. P. montivagus Bredenkamp & A.E. van Wyk. which is also described. The new taxa are geographically sep-

arated: subsp. filiformis ranges from Piquetberg in the north through the Cape Peninsula in the south, where it is quite com-

mon, to Attaquaskloof in the southwestern Cape; subsp. glutinosa occurs along the coast between Vredendal and St Helena

Bay; and P. montivagus has a wide distribution from Mossel Bay and Oudtshoorn in the south through Eastern Cape and

along the Great Escarpment northwards to Zimbabwe, with outliers in Tanzania.

INTRODUCTION

Passerina L., a genus of woody shrublets or shrubs,

comprises about 20 species and four subspecies, all con-

fined to southern and eastern Africa (Thoday 1924:

Goldblatt & Manning 2000). With the exception of a few

species growing along the Great Escarpment, most mem-

bers are endemic to the Cape Floristic Region with its

Mediterranean or semi-Mediterranean climate. In the

most recent taxonomic revision of the genus, Thoday

( 1924) considered P . filiformis L. a variable species with

a wide range, noting that plants from KwaZulu-Natal are

more robust and luxuriant in growth. The purpose of the

present paper is to present a taxonomic re-assessment of

the P. filiformis complex based on evidence from epider-

mal structure (Bredenkamp & Van Wyk 1999, 2000), leaf

anatomy (Bredenkamp & Van Wyk 2001a) and floral

morphology (Bredenkamp & Van Wyk 2001b). We pro-

pose the subdivision of P. filiformis L. into two sub-

species and describe a new species; all the new taxa are

geographically separated (allopatric).

MATERIAL AND METHODS

All collections of Passerina in the following herbaria

(acronyms according to Holmgren et al. 1990) were

studied for taxonomy and external morphology: BM.

BOL. BREM, C, GRA. K, LINN, M, MO, NBG, PR,

PRE. PRU, S. SBT. TCD. UPS.

For leaf anatomy, both fresh and herbarium material

were studied. Names of taxa and voucher specimens

used in anatomical studies are listed in Table 1 .

Light microscopy (LM) was used for general leaf

anatomical as well as epidermal studies. Methods for

preparation of transverse sections and for the study of cuti-

cles are described by Bredenkamp & Van Wyk (2000).

* National Botanical Institute, Private Bag X101, 0001 Pretoria.

** H.G.W.J. Schweickerdt Herbarium, Department of Botany,

University of Pretoria, 0002 Pretoria.

MS. received: 2001-08-31.

TABLE 1. — Passerina specimens examined anatomically and housed

at PRE

Scanning electron microscopy (SEM) was used to study

epidermal surface features (including epicuticular waxes)

and to elucidate the structure of the cuticle (Bredenkamp

& Van Wyk 2000). Transmission electron microscopy

(TEM) was used to establish the structure of mucilaginous

epidermal cell walls (Bredenkamp & Van Wyk 1999).

I . Passerina filiformis L., Species plantarum: 559

(1753); Thunb.: 75 (1794); J.C.Wendl.: 18 (1798);

Wikstr.: 324 (1818); Thunb.: 374 (1825); Meisn.: 562

(1857); C.H. Wright: 10 (1915); Thoday: 159 (1924);

Bond & Goldblatt: 432 (1984); Hilliard & B.L.Burtt: 1 82

(1987); Goldblatt & J.C.Manning: 683 (2000). Type:

Passerina filiformis , Linnean Herbarium 504.1 (LINN,

lecto.!, here designated).

Passerina phyliformis Hort. Bot. Bonarensis in Herb Zuccarinii (K.

Ml), err. typogr.

P. cupressina J.C.Wendl. nom. nud. Meisn.: 404 (1840); Meisn.:

563 (1857); Thoday: 159 (1924). P. cupressoides Steud.: 273 (1841).

P. pectinata Lodd.: 18 (1816) nom. nud. Wikstr.: 347 (1818);

Meisn.: 404 ( 1840); Meisn.: 562 (1857): Thoday: 159 ( 1924).

Large rounded shrubs up to 2 m high, often lax in shade.

Stems initially greyish tomentose, cork finely grey-brown

tessellate, becoming glabrous, with conspicuous leaf scars

and hair-like, whitish fibres protruding between bark fis-

sures. Leaves sessile, sometimes glutinous, closely adher-

ing to stem or spreading from stem at an angle of ± 30°;

lamina narrow, almost terete, acerose or linear, transverse-

ly elliptic or cordiform in c/s, length x V2 width (because

30

Bothalia 32,1 (2002)

FIGURE 1. — A-C. Passerina fili-

formis subsp. filifonnis ,

Bredenkamp 1039: A, inflor-

escence; B, leaf; C, bract.

D-F, P. filifonnis subsp.

glutinosa , Schlechter 5125:

D. inflorescence; E, leaf; F,

bract. G-M, P. montivagus,

Bredenkamp 1327: G, inflor-

escence; H, leaf; I. bract; J,

flower clasped by bract; K,

fruit enveloped by persistent

hypanthium, fragmented at

neck base; L, membranous

pericarp enveloping seed,

remnant of lateral style api-

cally present; M, seed with

black tegmen. Scale bars: 2.0

mm.

lamina is rolled) (4.0— )5.5— 8.0(— 1 0.0) x (0.4 — )0.6— 1 .0 mm.

tapering towards rounded apex, slightly widening or

widening towards base, coriaceous, smooth, dark green to

greyish green, often drying brown; adaxial groove tomen-

tose; abaxial surface convex, glabrous; margin involute.

Inflorescences subterminal, 10-20- flowered, composed of

proliferating spikes. Bracts cymbiform, outside glabrous,

inside setose from base to central part, ovate-acuminate to

widely obovate, length x 'A width ± 4. 6-7. 3 x 1 .5-2.0 mm,

gradually narrowing to point or narrowing abruptly into fil-

iform point, base cuneate to widely cuneate, main vein

strongly developed, often keeled, shortly extended or

extending to form leaf-like point; lamina coriaceous or

chartaceous; wings glabrous, membranous with distinct

venation, margins glabrous or ciliate, often with few tri-

chomes at apex adjacent to filiform point. Flowers gluti-

nous or not. Floral envelope ± 6.0-6. 5 mm long, yellow-

pink during pollination, sparsely tomentose or tomentose at

ovary, neck 1.5-1. 7 mm long, sparsely tomentose or

tomentose; outer and inner sepals with adaxial surface

glabrous, apex abaxially setose; outer sepals concave obo-

vate, inner sepals concave obovate or obovate, ± 2.2-2. 5 x

1 .4—1 .5 mm. Androecium : filament of antipetalous whorl ±

1 . 1-1 .2 mm, antisepalous whorl ± 2.2 mm; anthers ± 0.8 x

0.3 mm. Ovaty ± 2. 3-2. 5 x 0. 5-0.6 mm. Fruit enveloped

by persistent, loosely arranged hypanthium fragmenting at

neck base; pericarp membranous and dry. Flowering time :

in spring when pollen is wafted away in clouds, cause of

hay fever in sensitive persons (Marloth 1925). Figure

1 A-F.

Leaf anatomy

Leaf structural type B: bundle sheath completely

enveloping main vascular bundle, extraxylary scle-

renchyma fibres enclosed in bundle sheath (Bredenkamp

& Van Wyk 2001a).

Leaf outline in transverse section (t/s) cordiform to

transversely elliptic. Adaxial epidermis', cuticular mem-

brane (CM) ± 2 pm thick, periclinal x anticlinal cell

diam. in t/s 15 x 10 pm. Abaxial epidermis , in surface

view, glutinous or not, glabrous, CM mostly exhibiting a

striate pattern, cuticle with epicuticular waxes, wax

plates scarce, oblong, raised 90°, with sharp edges, usu-

ally arranged perpendicular to cell rows, epidermal cells

arranged in rows, oblong, pentagonal to heptagonal in

outline, 45-50 x 30 pm; CM 15-20 pm thick in t/s, pro-

Bothalia 32.1 (2002)

31

nounced at junctions of epidermal cell walls, grooved in

midline of joining walls, concavities and convexities pre-

sent. periclinal x anticlinal cell diam. in t/s (35— )40( —45 )

x 45-50(-80) pm, mucilagination of mainly inner tan-

gential cell walls often resulting in mucilage-filled cavi-

ties between remains of epidermal cells and adjacent

mesophyll. Palisade parenchyma horseshoe-shaped, 1-

or 2-layered, density 5 cells per 50 pm. Spongy

parenchyma aerenchymatic. Main vascular bundle

350-420 pm thick. 780-860 pm wide, widely ovate, var-

iously orientated in relation to mesophyll: touching pali-

sade parenchyma abaxially (type B3), sunken into pali-

sade parenchyma (type B4) (Figure 2A) or bordering on

abaxial epidermis (type B6) (Figure 2B). Bundle sheath

completely enveloping vascular bundle, consisting of

13— 15( — 27) cells. Secondary vascular bundles 2-A on

each side of main bundle. Sclerenchymatous hypodermal

sheath absent. Figure 2 A. B.

Two of the four Passerina specimens in the Linnean

Herbarium are named P. filifortnis in the handwriting of

Linnaeus; these specimens are numbered 504. 1 and 504.2

in Savage (1945). Number 504.2 is undoubtedly P.

paleacea Wikstr. Thoday (1924) maintains that number

504.1 is a Clifford specimen and probably the one

Linnaeus saw when he wrote the first edition of Species

plantarum (1753). Savage (1945) added the inscription [?

ex herb. Cliff.], indicating doubt as to the origin of this

specimen. The first author has seen this specimen and

agrees with Thoday ( 1924) that it matches the concept of

P. filiformis perfectly, as it is known in the Cape

Peninsula. The phrase 'Passerina foliis linearibus' in the

Species plantarum (1753), has clearly been copied by

Linnaeus from Hortus Cliffortianus ( 1738) and from Van

Royen (1740). The leaves in the illustration in Hortus

Cliffortianus are ± lanceolate and the bracts are very simi-

lar to those of P. filiformis subsp. glutinosa (Thoday)

Bredenkamp & A.E.van Wyk stat. nov.. which has acerose

or filiform leaves. The specimen labelled Passerina fili-

formis L. in Clifford’s Herbarium is sterile, lacks charac-

teristic bracts or flowers and could possibly be P. vulgaris

Thoday (= P. filiformis L. subsp. vulgaris Meisn.). P. vul-

garis is the dominant Passerina species in the southern

and southwestern Cape and is constantly confused with P.

filiformis. Specimen 504.1 in the Linnean Herbarium,

which is named P.filifomis by Linnaeus, is here designat-

ed as the lectotype. Thymelaea aethiopica, in Plukenet

(1700: 180), is cited in synonymy under P. filiformis by

Linnaeus (1753). The illustrated synonym of Breyne

(1678) most probably belongs to the genus Phylica L.

(Rhamnaceae) and that of Burman (1739) is clearly a

species of Struthiola L. (Thymelaeaceae).

Etymology, the Latin specific epithet filiformis (=

thread-like) obviously refers to the narrow leaves of this

species. The vernacular name 'sparrow-wort' was sug-

gested by Miller (1768) for all Passerina species, indi-

cating P. filiformis as ‘sparrow- wort with linear convex

leaves’. Wendland (1798) used the German equivalent

fadenformige Vogelkopf Marloth (1925) mentioned the

names kannabcis and kaalgaar. The following Afrikaans

names appear in Smith (1966) and some of them also in

Palmer & Pitman (1972) and Coates Palgrave (1977):

bcikbossie, bakkersbossie, bruingonna, fyntaaibos,

gannabas, gonnabas, kaalgaarbos, kaalgaring , kabel-

garing , kannabas, koordehciar , taaibos, windmakers-

bessie, windmakersbossie .

Key to subspecies

Branchlets and inflorescences dry; bracts widely obovate, nar-

rowing abruptly into filiform point la. subsp. filiformis

Branchlets and inflorescences glutinous; bracts ovate-acumi-

nate, gradually narrowing to point lb. subsp. glutinosa

la. subsp. filiformis

Leaves spreading from stem at angle of ± 30°; lamina

almost terete, cordiform in c/s, length x 'A width

(4.0— )5.5— 8.0(— 1 0.0) x 0. 6-1.0 mm, slightly widening

towards base, dark green to greyish green. Inflorescences

not glutinous. Bracts widely obovate, length x 'A width ±

7.3 x 2.0 mm, base widely cuneate, main vein strongly

developed, often keeled, extending to form a leaf-like

point; lamina chartaceous; margins glabrous, often with

a few trichomes at apex adjacent to filiform point.

Flowers not glutinous. Floral envelope ± 6.0 mm long,

tomentose at ovary, neck 1.7 mm long, tomentose, outer

sepals concave, obovate, inner sepals obovate. Figure

1 A-C.

Leaf anatomy

Leaf outline in t/s transversely elliptic to cordiform.

Adaxial epidermis: CM ± 2 pm thick; periclinal x anti-

clinal cell diam. in t/s 15 x 10 pm. Abaxial epidermis, in

surface view, not glutinous, cells oblong, 45-50 x 30

pm; CM 15-20 pm thick in t/s, periclinal x anticlinal cell

diam. (35— )40( — 45 ) x 45— 50(— 80 ) pm. Palisade paren-

chyma in 1 or 2 layers of elongated cells. Main vascular

bundle 350—420 pm thick. 780-860 pm wide, touching

palisade parenchyma abaxially (type B3) or sunken into

palisade parenchyma (type B4) (Figure 2A). Bundle

sheath consisting of 13-15 cells. Secondary vascular

bundles 3 or 4 on each side of main bundle.

Diagnostic characters and relationships: subsp. fili-

formis is morphologically distinguished by the almost

terete, adaxially grooved, acerose or linear leaves, the

cymbiform, widely obovate floral bracts, which abruptly

narrow to a filiform point (Figure 1A-C), and by the

long (± 1.7 mm) hypanthium neck. As both subsp. fili-

formis and P. vulgaris occur in the Cape Peninsula, they

are often confused. Diagnostic characters of P. vulgaris

include linear to narrowly lanceolate leaves, the dia-

mond-shaped bracts and leaves with a hypodermal scle-

renchymatous sheath (Bredenkamp & Van Wyk 2001a).

Some specimens of subsp. filiformis with incurved,

tapering leaves and the necks of the hypanthium exsert-

ed from the clasping, veined bracts, could be mistaken

for P. falcifolia. The apical beard on the young leaves

and outer sepals and the glabrous inner sides of the bracts

are reliable diagnostic characters, distinguishing the

subsp .filiformis from P. falcifolia.

Etymology and uses: according to Van Wyk & Gericke

(2000) the name bakkerbos commemorates an era in the

Cape when the official licensed bakers used the branches

32

Bothalia 32,1 (2002)

FIGURE 2. — LM photographs and SEM micrographs showing leaf anatomy and epidermal structure of selected species in Passerina: A, P. fili-

formis subsp. filiformis, Bredenkamp 1039 , leaf in t/s, illustrating leaf structural type B4. B, C, P.filiformis subsp. glutinosa , Schlechter

5125: B, leaf in t/s, from rehydrated herbarium material, illustrating leaf structural type B6; C, glutinous substance sticking pollen grain

to leaf surface. D, P. montivagus , Bredenkamp 1016 , leaf in t/s illustrating leaf structural type C. E-H, abaxial epidermis in P. montiva-

gus: E, in surface view, with cells arranged in rows, Bredenkamp 1012 ; F, epidermal cells oblong, pentagonal to heptagonal, Bredenkamp

1016 ; G, upright epicuticular wax platelets, Bredenkamp 1016. H, mucilagination of inner tangential cell walls, resulting in mucilage-

filled cavities, Killick 238. ad, adaxial epidermis; ab, abaxial epidermis; bs, bundle sheath; m, mucilage; mb, median vascular bundle; pi,

epicuticular wax plates; pp, palisade parenchyma; sp, spongy parenchyma. Scale bars: A, B, D-F, H, 100 pm; C, G, 10 pm.

Bothalia 32. 1 (2002)

33

17° 18c 19° 20° 21° 22° 23°

FIGURE 3. — Known distribution of

P. filiformis subsp. filiformis.

and leaves of this plant to heat their ovens. The plants

used at that time were clearly the subsp. filiformis. When

ignited, plants of subsp. filiformis disappear in a blaze of

hot flame owing to a waxy secretion on the leaves (Smith

1966). The plants were formerly used for heating up

stoves. Today it is quite scarce in the vicinity of Cape

Town, because of the commercial use of this once abun-

dant resource. At maturity, these plants are quite orna-

mental and have been cultivated in Britain and Europe

since the time of Linnaeus. Plants of the subsp .filiformis

are soboliferous and vigorous resprouters. They are well

adapted to the Cape climate and would be suitable for

reclamation plantings in areas where alien invasive vege-

tation has been cleared. The tough bark was used by

indigenous peoples instead of twine (Marloth 1925).

According to Laidler (1928) a decoction of this plant is

used by the rural people of the Cape for shooting pains.

Distribution and ecology. P. filiformis subsp. fili-

formis is common in the Cape Peninsula and is distrib-

uted from Piquetberg, across the Hex River Mountains,

to Attaquaskloof in the southwestern Cape. It grows in

rocky areas, mostly on south-facing mountain slopes and

on sandy plains, such as the Rietvallei and Stellenbosch

Flats. Figure 3.

Conservation status : Least Concern (LC) (IUCN

Species Survival Commission 2000).

lb. subsp. glutinosa (Thoday) Bredenkamp & A.E.

van Wyk, stat. nov.

TYPE. — Malmesbury Div., around Langeenheid Sta-

tion. Thoday 215 (BOL!; NBG. lecto.!. here designated).

P. filiformis L. var. glutinosa Thoday in Kew Bulletin 4: 1 60 (1924).

Leaves glutinous, closely adhering to stem or spread-

ing at angle of ± 30°; lamina narrow, acerose or linear,

transversely elliptic in c/s, length x ‘/2 width ± 7.0 x 0.4

mm, tapering towards rounded apex, widening towards

base, dark green, drying brown. Inflorescences glutinous,

somewhat longer than in typical subspecies. Bracts ovate-

acuminate, gradually narrowing to point, length x '/2

width ± 4.6 x 1 .5 mm. base cuneate. main vein strongly

developed: lamina coriaceous; wings membranous, with

distinct venation; margins often ciliate. Flowers gluti-

nous. Floral envelope ± 6.5 mm long, sparsely tomentose

at ovary, neck 1 .5 mm long, sparsely tomentose, outer and

inner sepal lobes concave-obovate. Figure 1D-F.

Thoday 215 in NBG was chosen as lectotype because

of the longer inflorescences and the conspicuously gluti-

nous, narrow leaves. Duplicates of the syntype of var.

glutinosa. Schlechter 5125. from BM. C, K. MO. PRE

and S were seen. Although these specimens agree close-

ly with the concept of var. glutinosa (Thoday 1924). the

glutinous character is not evident in the dried specimens.

Leaf anatomy

Leaf outline in t/s transversely elliptic. Adaxial epi-

dermis with CM ± 2 pm thick; periclinal x anticlinal cell

diam. in t/s 15 x 10 pm. Abaxial epidermis , in surface

view, glutinous (Figure 2C), cells slightly oblong. 50 x

30 pm; CM 15 pm thick in t/s. periclinal x anticlinal cell

diam. 40 x 55 pm. Palisade parenchyma in 2 layers of

elongated cells. Main vascular bundle ± 400 pm thick. ±

820 pm wide, sunken into palisade parenchyma abaxial-

ly (type B4) or bordering on abaxial epidermis (type B6)

(Figure 2B). Bundle sheath consisting of ± 27 cells,

adaxially radiating outwards, abaxially tanniniferous,

specializing into collenchyma in contact with abaxial

epidermis. Secondary vascular bundles 2 or 3 on each

side of main bundle.

Distribution and ecology, subsp. glutinosa occurs in

the Strandveld (Acocks 1988). from Doring Bay in the

north to St Helena Bay in the south. The vegetation sur-

rounding Doring Bay is described as Strandveld

Succulent Karoo by Hoffman ( 1998). The area is charac-

terized by deep, calcareous, coastal Quaternary sands

and generally low rainfall. St Helena Bay is situated in

the Sand Plain Fynbos (Rebelo 1998). This part of the

range has a Mediterranean-type climate with summer

drought and deep acid sands. Sand Plain Fynbos is a

highly endangered vegetation type because of urbaniza-

tion and the impact of alien invasive plant species

(Rebelo 1998). Figure 4.

Conservation status: Near Threatened (NT) (IUCN

Species Survival Commission 2000).

34

Bothalia 32,1 (2002)

17° 18° 19° 20°

FIGURE 4. — Known distribution of P.filiformis subsp. glutinosa.

2. Passerina montivagus Bredenkamp & A.E.vcm

Wyk , sp. nov., a P.filiforme L. habitu robustiore et luxu-

riantiore distinguenda. Folia cymbiformia, secus nervum

medium plicata, lanceolata, ad apicem rotundatum

decrescentia, basi expansa. Bracteae cymbiformes,

ovatae ad obovatae, extra glabrae, intra basaliter setosae,

in carinam longam, fere cylindricam, adaxialiter sul-

catam, leviter incurvam gradatim decrescentes, basi

cuneatae, coriaceae, sed lateris vel alis chartaceis, plus

minusve tricostatis, marginibus trichomatibus conspicuis

crassis serialibus secus dimidium distalem obsitis.

TYPE. — KwaZulu-Natal, 2930 (Pietermaritzburg):

hills above Pinetown, 2400 ft, (-DD), 3 December 1891 .

J.M. Wood in PRE49409 (PRE!, holo.; MO!, iso.).

Passerina fdiformis L.: 559 (1753) pro parte, excluding type;

Thunb.: 75 (1794); Wikstr.: 324 (1818); Thunb.: 374 (1825); Meisn.:

562 (1857); C.H. Wright: 10 (1915); Thoday: 159 (1924); Bond &

Goldblatt: 432 (1984); Hilliard & B.L.Burtt: 182 (1987); Goldblatt &

J.C. Manning: 683 (2000).

Low, spreading shrub l(-2) m high, vigorous re-

sprouter. Stems initially greyish tomentose, cork Assur-

ing lengthwise into fine, dark grey, tomentose strips,

older branchlets glabrous, with conspicuous leaf scars.

Leaves sessile, spreading from stem at angle of ± 45°;

main vein sturdy; lamina cymbiform, folded along main

vein, lanceolate, often slightly falcate, length x '/2 width

5.6-7.0(-8.0) x 0. 7-1.0 mm, tapering towards rounded

apex, expanded at base, ciliate, coriaceous, smooth,

greyish green; adaxial surface tomentose; abaxial surface

glabrous; margin involute. Inflorescences subterminal,

1 0-20-flowered, composed of proliferating spikes, com-

mon. Brads cymbiform, outside glabrous, inside basally

setose; lamina coriaceous, ovate to obovate, length x '/2

width ± 6.3 x 1.6 mm, narrowing gradually into sturdy,

leaf-like point of extended main vein, base cuneate;

wings chartaceous ± 3-ribbed, obscurely veined, greyish

green, margins conspicuously lined with strong white tri-

chotnes along distal half, often reaching up lo apex.

Floral envelope ± 6.6 mm long, yellow-pink during pol-

lination, tomentose at upper half of ovary, neck tomen-

tose ± 2.3 mm long; adaxial surface of outer and inner

sepals glabrous, apex setose abaxially; outer sepals

cymbiform, inner sepals obovate, ± 2.1 x 1.3 mm.

Androecium: filament of antipetalous whorl ± 0.8 mm

long, antisepalous whorl ± 1 .7 mm long; anthers ± 0.8 x

0.3 mm. Ovaiy ± 2.2 x 0.7 mm. Fruit enveloped by per-

sistent, loosely arranged hypanthium fragmented at neck

base; pericarp membranous and dry. Figure 1G-M.

Thoday (1924) noted that plants named P. filiformis in

KwaZulu-Natal are more robust and luxuriant than those

from Western Cape. Wood s.n. from Pinetown was cho-

sen as the holotype of P. montivagus as it is a good rep-

resentation of the new taxon and was determined and

cited by Thoday ( 1924).

Leaf anatomy

Leaf structural type C: bundle sheath capping main

vascular bundle adaxially, ± absent abaxially, extraxylary

sclerenchyma fibres not enclosed in bundle sheath, main

vascular bundle bordering on palisade parenchyma,

extraxylary sclerenchyma fibres fitting into V-shaped

palisade parenchyma (Bredenkamp & Van Wyk 2001a).

Leaf outline in t/s carinate. Adaxial epidermis: CM ±

2 pm thick, periclinal x anticlinal cell diam. in t/s 25 x

15 pm. Abaxial epidermis , in surface view: cuticle with

epicuticular waxes, wax plates scarce, ± perpendicular to

cell rows, oblong, raised 90°, with sharp edges (Figure

2G), CM mostly exhibiting a striate pattern, epidermal

cells arranged in rows (Figure 2E), oblong, pentagonal to

heptagonal (Figure 2F), 30-55 x 25-35 pm; CM ± 20

pm thick in t/s, pronounced at junctions of epidermal cell

walls, grooved in midline of joining walls, concavities

and convexities present, periclinal x anticlinal cell diam.

(22.5— )30.0(— 35.0) x (40— )55(— 90) pm, mucilagination

of mainly inner tangential cell walls often resulting in

mucilage-filled cavities between remains of epidermal

cells and adjacent mesophyll (Figure 2H). Palisade

parenchyma V-shaped, in 2 layers of elongated cells,

density 3 or 4 cells per 50 pm. Spongy parenchyma

aerenchymatic. Bundle sheath an adaxial cap of ± 21

cells, rounded. Main vascular bundle (400-)560-630

(-700) pm thick, (820— )980— 1 1 80(— 1 280) pm wide,

obovate, bordering on and fitting into the V-shaped pali-

sade parenchyma (type C) (Figure 2D). Extraxylary scle-

renchyma fibres not enclosed in bundle sheath.

Secondary vascular bundles 3 or 4 on each side of main

bundle. Sclerenchymatous hypodermal sheath absent.

Diagnostic characters and relationships: Passerina

montivagus is easily distinguished from P. filiformis by

its more robust and luxuriant habit. Furthermore, for

some distance below the inflorescences, the foliage

leaves are expanded at the base and the bracts are ovate

to obovate, narrowing gradually into a sturdy, leaf-like

point, with margins along their distal half conspicuously

fringed by strong white trichomes. P. montivagus could

also be confused with P . falcifolia, but it is distinguished

from the latter by the apical beard on the young leaves

Bothalia 32.1 (2002)

35

10' 15* 20* 25* 30" 35* 40-

FIGURE 5. — Known distribution of P. montivagus.

and outer sepals and by the adaxial surfaces of the bracts,

which are basally setose with glabrous wings.

Etymology : the specific epithet is a compound of the

Latin montanus (= pertaining to mountains) and vagus (=

in several directions), referring to the distribution of this

species. Von Breitenbach et al. (2001) uses the names

brown gonna (English), bruingonna (Afrikaans) and

unwele oluncane (Zulu) for P. filiformis in the wide

sense, but these names are most appropriate for P. mon-

tivagus because of its wide distribution.

Distribution and ecology : Passerina montivagus has a

wide distribution, from Mossel Bay and Oudtshoorn in

Western Cape northwards mainly along the Great

Escarpment to KwaZulu-Natal. Swaziland. Mpuma-

langa, Northern Province, Mozambique and Zimbabwe.

Polhill & Paolo 2372, from Tanzania, represents an

extreme form of this taxon, as the bracts are smaller and

almost diamond-shaped. The most southwesterly distri-

bution of this species is in the southern Cape, a region

transitional between winter and summer rainfall.

However, over most of its range, the species receives

summer rainfall. Because of its wide distribution, espe-

cially along the Great Escarpment, P. montivagus is

adapted to a variety of habitats, with relatively high rain-

fall. It is often found along forest margins in the ecoton-

al zone between Afromontane forest and grassland. It has

been recorded from rocky mountain peaks and slopes,

river valleys, gorges and among riverside rocks. In

coastal regions, it grows on hills and often borders small

tributaries of streams flowing to the sea. Figure 5.

Story (1952) reported that P. montivagus (= P. fili-

formis) dominated the western half of a small plateau

north of the Mount McDonald beacon in the

Keiskammahoek District. The plants were not browsed

by stock although the plateau was heavily grazed. He

regarded the species as ‘useless’ and advised that it

should be eradicated by hand, as it was not dense enough

to burn without additional fuel. This Fynbos species, dis-

tributed along the Great Escarpment has not been report-

ed as undesirable or invasive and is currently not regard-

ed as a threat, although it might be a dominant species in

restricted areas.

Conservation status: Least Concern (LC) (IUCN

Species Survival Commission 2000).

SPECIMENS EXAMINED

Abbott 43, 308 (2) PRE. Acocks 11549 (2) PRE; 19671 (lb) K, M,

NBG. PRE; 24062 (lb) PRE; 890, 5174, 5754 (la) S. Andraea 1165

( la) NBG, PRE. Anferweg 850 ( la) PR.

Barker 5795, 8095 (la) MO. NBG. Bolus 2440 (2) BOL, K; 2925 (la)

BOL. K. Botha 1445 (2) PRE. Boucher 2833 (lb) PRE. Bredenkamp

1012, 1015 (2) PRE; 1016, 1017. 1327, 1359, 1360 (2) PRE; 1039 ( la)

PRE. Bremer 317 (2) PRE. S. Buchenau s.n. ( la) BREM. Buitendag 712

(2) NBG. PRE. Burchel 66, 276. 6544 (la) K. Burmeister s.n. (la) SBT.

Cheadle & Welts 668 (2) M. PRE. Compton 11762, 14827 ( la) NBG;

15888 (lb) NBG; 25157 (2) NBG; 26157, 29473 (2) NBG. PRE.

Dlamini s.n. (2) NBG, PRE. Dr'ege s.n. (la) K.

Ecklon s.n. (la) PR. Edwards 820 (2) PRE. Esterliuysen 26437, 34096.

35680 (la) BOL. MO. M. S.

Forbes & Obermeyer 58 (2) PRE. Fourcade 3473 (2) NBG; 4809 (2)

BOL.

Galpin 11016 (2) PRE. Garthside 17 (la) K. Germishuizen 1705, 9089

(2) PRE. Gerrard 1478(\a) BM. TCD, K. Gill s.n. (la) K. Gillett4537

(2) BOL. PRE. Goodier & Phipps 270 (2) MO. Goodier 637 (2) BM.

M. PRE. Grant 1 (la) C. Hardy s.n. (la) K, S. Herb. Linairiaro (la)

PR. Herb. Regium Monacense s.n. ( la) M. Herb. Rofski (la) PR. Herb

Zuccarinii s.n. (la) M. Hilliard & Burtt 14463 (2) PRE; 14654 (2) K;

Hilliard & Burn 15671 (2) N. PRE, S; Hilliard & Burtt 18772A (2) K,

S. Hugo 2019 (2) PRE; 2086 (2) NBG, PRE. Huntley 612 (2) MO.

PRE. Hutchinson 258 ( la) PRE; 1298 (2) PRE.

Joffe 576 (2) TCD. PRE.

Keet s.n. (2) NBG. PRE. Killick 238, 3469 (2) PRE. Krebs 282 (la)

MO. Kutzelman’s Herbarium, Prague (la) PR.

Lehman 1891 ( la) C. Lewis 4064, s.n. (2) NBG. Luidberg s.n. (la) S.

MacOwen 16388 (2) BM; s.n. (2) MO. Mantel! & Vassilatos 32 (2) PRE.

McKinnon 32 (la) NBG. Medical Soc. Univ. (la) K. Meebold 15156,

15157 (2) M. Mogg 13350 (2) K. PRE; 5206, 13397, 38026, (2) PRE.

Morris 441 (2) NBG. Moss 5643, T10 (la) BM. Mund & Maire s.n. ( la)

K. Museum Bot. Hauniense (la) C. Niven & Laubert s.n. (la) S.

Pappe s.n. (la) NBG. Pedro & Pedergao 7310 (2) BOL. Pegler 1273

(2) BOL. GRA, NBG. PRE. Penther 1912, s.n. (la) S. WU; 1919 (2)

M. S. Phillipson 1200 (2) MO, PRE. Polhill & Paolo 2372 (2) PRE.

Prior s.n. (la) K.

Rob & Fries 3393. 3396-3399 (la) UPS, S. Roberts 66, s.n. (2) S.

Rogers 16701 B (la) PRE. Rudatis 1204 (2) BM. PR, S.

Schlechter 5125 (lb) BM. C. K. MO. PRE. S. Schmidt 568 (la) M.

Sidey 3862 (2) PRE, S. Sikhakhane 524 (2) PRE. Sparrman s.n. ( la) S.

Story 3765 (2) GRA. PRE. Strev 7112, 11363, 11364 (2) PRE; Strey

9321 (2) PRE. S.

Talbot s.n. (la) K. Taylor 1542 (lb) NBG. PRE. Thoday 215 (,1b) BOL.

NBG. Thode 4657 (2) NBG. Thom 553. 577 (la) K. Thompson 41 ( lb)

NBG, S; 801 (lb) NBG PRE. Trinity College s.n. (la) TCD. Tyson

1287 (2) MO. NBG; 2621 (2) NBG.

Van derMerwe 1101 (la) PRE. Van Wvk 2622 (2) PRE. PRU, M; 5332,

BSA2586 (2) PRE, PRU. Van Wyk & Bredenkamp 1 (2) PRE, PRU. Van

Wvk & Matthews 77719 (2) K. PRU. Victor 556 (2) PRE. Vincent &

Wearne 4 Cl) PRE.

36

Bothalia 32,1 (2002)

Wall 1060 (la) S. Ward 8832 (2) N. PRE. 10669 (2) PRE Wells 1809

(2) MO. Wolley-Dod , 3103 (la) K. Wood 1182 (2) BOL; 8938 (2) MO:

s.n. (2) MO, PRE. Worsdell s.n. (la) K. Wright s.n. ( la) MO. Without

collector. K692 (la).

Cultivated: Hort. Herb. Pallas (la) BM; Hort. Prague (la) PR: Vinaf

s.n. (la) PR.

ACKNOWLEDGEMENTS

The authors would like to thank Ms G. Condy for the

line drawings, Dr O.A. Leistner for translating the diag-

nosis into Latin and Ms A. Romanowski for printing the

photographs.

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Bothalia 32,1: 37-61 (2002)

The genus Erica (Ericaceae) in southern Africa: taxonomic notes 1

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

Keywords: Erica L„ nomenclature, southern Africa, taxonomy

ABSTRACT

This is the first in a series of notes on the southern African species of Erica L. which are currently recognized. Brief

synonymy citing only the important revisions and synonyms is given, as well as comments on the status and diagnostic char-

acters of each species. One new species. E. petrusiana E.G.H.Oliv. & I.M.Oliv. and 14 new subspecies are described, and

11 species are reduced to subspecific status. Where necessary drawings of important features and variations are given. Part

1 covers the first 35 species (Sections 1-3).

INTRODUCTION

The last revision of the genus Erica L. covering the

species in southern Africa was undertaken by Dulfer

(1965) and published as a brief conspectus — excluding

descriptions and with very little or no discussion about

problems of delimitation and the decisions he took. All

of this was based almost exclusively on the very small

collection of Erica housed in the Natural History

Museum in Vienna (W), in many cases with species rep-

resented by only one or two specimens. He had only a

few small loans from the Bolus Herbarium (BOL) and

the Royal Botanic Gardens, Kew (K), having been

refused large loans. His work was based on that of

Guthrie & Bolus in Flora capensis (1905) and included

the 220 species described since that publication. We esti-

mate that he had far less material on which to base his

decisions than did Guthrie & Bolus, who also consulted

the large collections at Kew and the Natural History

Museum, London (BM) and also at Trinity College,

Dublin (TCD). Bolus also consulted the important

Ericaceae collections in Berlin, which, except for the

Willdenow herbarium, were totally destroyed during

World War II. Fortunately he was allowed to remove a few

flowers from several of the types and these are in BOL.

Dulfer’s work is valuable in that he spent much time

checking the synonymy and old references that Guthrie

& Bolus had put together. This remains a much used

source of reference in our work on Erica. Since Dulfer’s

revision, another 63 species have been added, and all the

species formerly included under the 23 minor genera

have also been added, with the relegation of these genera

to synonymy under Erica (Oliver 1988, 1993a, 1993b.

2000). The number of species in the genus currently

stands at 765 in the area covered by the Flora of south-

ern Africa. The number of specimens located in the two

Cape herbaria. Bolus Herbarium (BOL) and the

Compton Herbarium (NBG), which now includes the old

South African Museum Herbarium (SAM) and

Government Herbarium. Stellenbosch (STE), greatly

increases the amount of material that exhibits variation,

compared to that which was available for Guthrie &

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

7735 Claremont, Cape Town.

MS. received: 2001-11-21.

Bolus and considerably more so in the case of Dulfer.

Having studied the genus in the herbarium and in the

field for 42 years (EGHO) and 15 years (IMO) respec-

tively, we are in a position to update the species concepts

currently recognized in the literature (Dulfer 1965;

Oliver & Van Wyk 1993), and to publish our concepts

which are incorporated as curatorial practice in the

Compton Herbarium (NBG).

It has been decided to produce a series of taxonomic

notes on the species following the short format used by

Dulfer, but with short to detailed explanations of varia-

tions, of problems with delimitation and of new delimi-

tation of some taxa. Where necessary illustrations of the

variable taxa have been included. We have followed the

numbered order started by Guthrie & Bolus (1905) and

adapted by Dulfer (1965) and then by ourselves in the

Compton Herbarium. This system follows the sectional

arrangement used by Guthrie & Bolus in which the genus

is divided into 43 sections based on the earlier work of

Bentham (1839). These sections were an attempt to

group the species into supposedly related assemblages.

In some cases this is clearly the case, but in others there

are no close relationships and species are. in our opinion,

sometimes widely separated from their nearest relatives.

An example is a species which was described twice by

Guthrie & Bolus (1905), firstly as E. auriculata , which is

no. 135, and secondly as E. greyii, which is no. 338.

This sectional/species arrangement is used here for

want of a better, natural system which is as yet not forth-

coming. The available morphological characters are not

sufficiently adequate for this puipose due to the high

degree of homoplasy present within the genus. To

attempt to resolve this impasse, a study is to be under-

taken into the molecular relationships of some 300

species in collaboration with overseas researchers to try

to ascertain the basic clades within the genus and to cou-

ple this with an analysis of morphological characters.

These results should give a good indication as to which

the important morphological characters are in a reassess-

ment of the genus at the level of subgenus and section.

In a few cases the species have not been fully resolved

and we are forced to list these in complexes under the

oldest name with pointers towards the problems that

need to be resolved through more detailed fieldwork and

molecular studies.

The genus Erica is by far the largest represented in the

Cape Flora and it is, in our opinion, clearly still evolving

38

Bothalia 32,1 (2002)

actively. There are many species which are isolated in

their relationships, and these one may regard as

palaeoendemics, whereas there are many that form

extremely complicated complexes that currently, and

perhaps always will, defy satisfactory resolution.

To date, the only subspecific category that has been

used in the main revisions is that of variety (varietas).

With the current trend to drop that category, we have

resorted to recognizing worthy subspecific groupings as

subspecies, especially when there is, in addition to some

morphological disjunction, also a disjunction in distribu-

tion, ecology or postulated pollination biology. In other

cases we have refrained from a proliferation of sub-

species and have resorted to listing unnamed variants

which reflect what we believe are low level groupings

that have been noted by us.

The concentration of so many species in the Cape

Floral Region is shown in counts we have done. Often

we have found four species growing in one square metre

with the highest count thus far obtained being seven, or,

in one case, four species in a quarter square metre — all not

closely related species. In a number of cases the disjunc-

tions between taxa can occur over very short distances

with variants being confined to relatively small areas.

With such a large genus — currently 760 species in

southern Africa, we are forced to publish this review in a

series of parts covering a varying number of species per

part depending on the complexity of the species and the

species complexes in order to provide data on species

delimitation soon, rather than wait many years for a com-

plete coverage of all 680 capsular species in one publi-

cation. The 84 indehiscent fruited species have recently

been dealt with in detail (Oliver 2000).

NOTE 1 : in the numbering system the whole numbers

for the species are those that were originally allocated by

Guthrie & Bolus (1905). The species described since

then until 1965 were placed by Dulfer nearest their sup-

posedly nearest ally as a & b numbers. These we have

altered to decimal notation since we use the a/b notation

for subspecies. Species described since Dulfer’s 1965

revision have been placed by us in their postulated

alliances. Species we recognize are printed in bold

Roman type. Those that are reduced to synonymy, or that

we regard as cultivated forms, or dubious, are printed in

bold italics.

NOTE 2: the inclusion under the synonymy of cita-

tions of publications additional to the original one, is

restricted to the most important revisions of the genus.

For all other citations and more detailed synonymy, ref-

erence should be made to Guthrie & Bolus ( 1905) and to

Dulfer (1965).

NOTE 3: in the case of citations involving the two

works of Andrews, Coloured engravings of heaths

(1794-71830) and The Heathery (1804-1812), the rele-

vant work has to be cited in full in each instance, due to

the considerable overlapping of publication dates (see

Cleevely & Oliver 2002), and cannot be cross-referenced

in the references.

ERICA COCC1NEA COMPLEX

This complex, consisting of E. coccinea, E. melas-

toma, E. intermedia and E. monadelpliia, is character-

ized by an enlarged petaloid bract and bracteoles

adpressed to the calyx, long, attenuated, well-exserted

anthers with rounded apices and basal attachment, the

often flap-like placenta with a naked abaxial zone, shiny

smooth seeds (not alveolate or reticulate) and leaves with

a large sclerenchyma bundle on either side of the sul-

cus— these visible as white stripes in fresh or pressed

material.

The inflorescence consists of 3-nate flowers, except in

E. melastoma where they are borne singly.

The ‘ coccinea ’ complex’s nearest relatives are the

species belonging to the small-flowered E. imbricata

complex (no. 369) and not to any of the other large-flow-

ered species, e.g. E. plukenetii , where similarities are the

result of convergent evolution in pollination syndromes.

It is at times difficult to differentiate between the various

species, especially in the dmbricata-placentiflora ’ com-

plex which has evolved numerous variants and in some

cases these are sympatric, thus adding to the confusion.

There could very well be a case for considering

hybridization between some of these variants.

These two complexes cover a wide range of flower

sizes from a large tubular corolla ± 20 mm long, to a

small globose one ± 2 mm long. The pollination syn-

dromes vary from bird pollination in the large-flowered

species to insect and wind pollination in the small-flow-

ered species.

Erica banksii is very similar to members of this com-

plex, but lacks the distinctive sclerenchyma bundles in

the leaves, and the anthers have the filaments attached

more or less dorsally.

1. E. coccinea L., Species plantarum edn 1,1: 355

(1753); Salter: 633 (1951) p.p.; Dulfer: 28 (1965) p.p.

Lectotype: Seba: t. 21, fig. 4 (1735), selected here by

Oliver, Jarvis & Cafferty.

Note: a paper covering the typification of all Linnaean names of

Cape Erica spp. is in preparation by Oliver, Jarvis & Cafferty.

Diagnostic features : corolla tubular, ± 16-22 mm

long; bract, bracteoles and sepals with long narrow sul-

cus; anthers far exserted; leaves recurved.

la. subsp. coccinea

E. petiveri L.: 10, t. 50 (1770); Benth.: 621 (1839); Guthrie &

Bolus: 47 (1905). Lectotype: L., t. 50, selected here by Oliver. Jarvis &

Cafferty, in prep.

E. sebana Donn: 45 (1796) nom. nud., et auct. mult.

E. petiveri var. pubescens Bolus: 47 ( 1 905 ); E. coccinea var. pubes-

cens (Bolus) Dulfer: 29 (1965). Type: Stellenbosch Div., Lowry's Pass,

Burchell 8246 (Kl); Hottentots Holland Niven 129 (K!); Caledon Div..

Houw Hoek. Burchell 8130 (K!); Div.?, Bolus 8036 (BOL!).

Illustrations: Baker & Oliver: t. 1, 1-7 (1967); Schumann & Kirsten:

35, t. 2 & 37, t. 2 (1992); Oliver & Oliver: t. 2 (2000).

Diagnostic features: leaves small, closely packed and

recurved; flowers 3-nate on ends of short, leafy, side

branches, red, orange, yellow or green and usually uni-

coloured (Figure 1 A i , A2).

A yellow-flowered variant with long narrow sepals

which has a dark corolla mouth, occurs in the Bredas-

dorp to Gansbaai area.

The typical subspecies is the common and widespread

taxon in this species occurring from the Cederberg to the

Cape Peninsula and eastwards as far as the Kammanassie

Mountains.

Vouchers: Balfour s.n. (MO!, NBG!); Bolus 6758 (BOL!, NBG!);

Compton 23683 (NBG!, S!); Johnson 304 (K!, NBG!); Lewis 5300

Bothalia 32,1 (2002)

39

flowers . 4mm

anthers. 2mm

FIGURE 1. — Erica coccinea complex: flowers and anthers. Ai, A;, E. coccinea subsp. coccinea: Ai, Baker 42, Constantiaberg; Aj, Baker 2014,

Elim. B|-B4, E. melastoma subsp. melastoma : B,, variant D, Oliver 10676: B2, variant E, Kirsten 811, De Tronk; B ,, variant A, Oliver

8013, Caledon Swartberg; B4, variant B. Oliver 11287. Klein River Mtns. B5, E. melastoma subsp. minor. Baker 1250, Baardskeerdersbos.

C, E. monadeiphia, De Vos 648, Kleinmond. Di, D;, E. intermedia: Di, subsp. intermedia, from type, Niven 127', D2, subsp. albiflora,

Oliver 9105, Ruitersberg. E, c/s leaf. Scale bars: flowers, 4 mm; anthers, 2 mm; E, 250 pm.

(BM!, NBG!, PRE); MacOwan sub Herb. Norm. 5 (BM!, K!, SAM!);

Middlemost 1556 (NBG!, NY!); Oliver STE30141 (BM!. BOLL NBG!,

PRE), STE30142 (NBG!, PL PRE); Zeyher 3185 (BOLL KL PL PRE,

SAM!, W).

lb. subsp. uniflora E.G.H.Oliv. & I.M.Oliv.,

subsp. nov., flore uno foliis suberectis ad recurvatis dis-

tinguitur.

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

(-DB ), 12 January 1958, Rycroft2104 (NBG, holo.; K, PRE).

Diagnostic features: flowers borne singly at ends of

short, leafy lateral branches, usually yellow, sometimes

orange-red; leaves semi-erect to reflexed.

This is a lowland taxon occurring on hills and flats not

far from the coast from the Cape Peninsula eastwards to

near Mossel Bay.

Paratype material (selection from numerous collections): WEST-

ERN CAPE. — 3418 (Simonstown): Karbonkelberg. 1500 ft [450 m],

(-AB), 21-07-1974, Esterhuysen 33586 (BOLL NBG!); Sir Lowry’s

Pass, 1000 ft [300 m], (-BB), 11-07-1890, Guthrie 2001 (NBG!). 3419

40

Bothalia 32,1 (2002)

(Caledon): Baviaansfontein, Gansbaai, 800 ft [250 m], (-CB), 14-03-

1977, Hugo 806 (NBG!, PRE); Kleinbos, Die Kelders, 50 ft [15 m],

(-DA), 4-11-1969, Taylor HC 7420 (NBG!. PRE); Bredasdorp Forest

Reserve, 60 ft [18 m], (-DC), 12-03-1979, Thompson MF 3974 (NBG!,

PRE). 3420 (Bredasdorp): Potberg. 500-600 ft [150-180 m], (-BC),

19- 06-1984, Oliver 8512 (NBG!). 3421 (Riversdale): Canca, (-BC),

20- 03-1975, Oliver 5739 (NBG!); Ystervarkpunt, 80 m, (-BD), 15-05-

1987, Willemse 185 (NBG!). 3422 (Mossel Bay): Mossel Bay, dunes,

(-AA), 08-1962, Liebenberg 6376 (BM!. NBG!, PRE).

1.1. E. melastoma Andrews , Coloured engravings

of heaths: t. 37 (1799). E. petiveri var. melastoma

(Andrews) Benth.: 622 (1839); Guthrie & Bolus: 47

(1905). E. follicularis Salisb. var. melastoma (Andrews)

Dulfer: 29 (1965). Iconotype: Andrews: t. 37 (1799).

Note: Dulfer overlooked the problem of the dates of publication of

Andrews' plates and selected Salisbury's name of 1802 for the species,

thereby relegating Andrews’ E. melastoma to varietal status. Dulfer

(1965) cited 1802 as the date of publication for E. melastoma which is

the date of the bound volume and not the date which is printed on the

plate. A paper detailing the publication dates of all Andrews' species is

in preparation (Cleevely & Oliver 2002).

Diagnostic features', leaves erect to semi-erect,

straight; flowers single on short, leafy side branches; sul-

cus on sepals, bract and bracteoles very short, apical,

broadly open, V-shaped (not narrow, slit-like and form-

ing a keel) or absent.

This is a very variable species in the size of the flow-

ers, size and shape of the sepals, shape and stickiness of

the corolla and colour of the corolla apex. Within all this

variation there is only one distinct discontinuity, namely

in the size of the corolla, and we propose to recognize

this at subspecific level.

1.1a. subsp. melastoma

E. vestiflua Salisb.: 346 (1802); Guthrie & Bolus: 48 (1905);

Dulfer: 30 (1965). Type: sine loc.. Roxburgh s.n. [Roxburgh 71 det.

Salisbury] (K!).

E. follicularis Salisb.: 348 (1802): Dulfer: 29 (1965). Iconotype:

Andrews, Coloured engravings of heaths: t. 44 (1797).

E. petiveri var. willdenovii Bolus: 48 (1905). E. coccinea var.

willdenovii (Bolus) H. A. Baker: 184 (1967). Type: Thunberg s.n.

(UPS).

E. coccinea var. inflata H. A. Baker 75 (1958). E. follicularis var.

inflata (H. A. Baker) Dulfer: 30 (1965). Type: Caledon Div., Koude

Mountains, between Gaansbaai and Elim. Baker 1154 (BOL, holo.;

NBG!).

Illustration: Baker & Oliver: t. 1, 8 (1967).

Diagnostic features: corolla ± 18 mm long (Figure

1B4).

Within the subspecies there is a considerable amount

of variation in several characters, mainly the corolla

shape, colour and stickiness. Despite there being some

collections which appear to be very distinct, we are

unable to recognize any formal categories and thus list

them here as unnamed variants to show possible groups

that need to be worked on in detail in the field and

through DNA analyses.

VARIANT A: the corolla is yellow with a dark, almost

black, distal end. It may, however, be greenish yellow

with the distal end fading to brown soon after anthesis.

The typical variant, shown in Andrews’ painting, has

large sepals, i.e. broad and more than half the length of

the corolla tube, the corolla slightly to much inflated

towards the base and mostly not sticky although small

sessile glands may occur on the margins of the sepals

(Figure I B.fl. The leaves can be short to very long. The

distribution is from Stellenbosch to Bredasdorp.

VARIANT B: this variant has flowers similar to

Variant A, but they are very sticky and yellowish with a

black mouth (Figure 1B4). It includes the var. inflata and

occurs in the Bredasdorp District.

VARIANT C: with similar flowers to A, but not

sticky, usually light orange-yellow, mostly with no tuft-

ed small side branches typical of the other variants. It

occurs on limestone flats of the Bredasdorp District.

VARIANT D: flowers yellow-orange with no black

mouth, with long narrow tube and a short calyx less than

half the length of the corolla tube, which is not sticky

(Figure IB,). This occurs in the region from Tulbagh to

the Cederberg.

VARIANT E: flowers the same as those in D above,

but very sticky. The stickiness is produced by a remark-

able array of sessile glands in several zones around the

margins of the sepals (Figure 1 B2). This variant is found

in the Porterville Mountains, but a few collections near

Wemmershoek and Baviaansberg, Ceres, have a reduced

amount of glands.

Vouchers: Andreae 686 (NBG!); Guthrie 2005, 2498 (NBG!);

Hanekom 2941 (NBG!); Oliver 10676 (NBG!), STE30143 (BOL!,

NBG!. PRE).

1.1b. subsp. minor E.G.H.Oliv. & l.M.Oliv. subsp.

nov., floribus minoribus, 6 ad 7 mm longis, distinguitur.

Figura 1B5.

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

SE-facing slope above Remhoogte, ± 250 m, (-DA), 27

October 2001, R.C. Turner 401 (NBG, holo.; BM, BOL,

K, NY, PRE).

Diagnostic features : corolla 6-7 mm long, yellow

with dark mouth (Figure 1B5).

The subspecies occurs in the Hermanus and Bredas-

dorp Districts.

Paratype material: WESTERN CAPE. — 3419 (Caledon): Herma-

nus Mtns, (-AD), 10-1924. Marloth 6225 (NBG!. PRE); Baard-

skeerdersbos, (-DA), 8-9-1957, Baker 1250 (NBG!): Remhoogte,

(-DA), 25 October 1967, Thomas in NBG86471 (NBG). 3420 (Swell-

endam): Bredasdorp Mtn, (-CA), 15-10-1951, Esterhuysen 19169

(BOL!. NBG!. PRE). Without locality: Caledon Wildflower Show. 09-

1965, Oliver STE32109 (NBG!).

1.2. E. intermedia Klotzsch ex Benth. in DC.

Prodromus 7: 621 (1839). E. petiveri var. intermedia

(Klotzsch ex Benth.) Bolus: 47 (1905). E. coccinea var.

intermedia (Klotzsch ex Benth.) Dulfer: 29 (1965).

Syntypes: Swellendam, Masson s.n. (K), Niven s.n.

(BOL!, K), Ecklon s.n. (Bt). Lectotype to be selected

from the material used by Bentham in K.

Diagnostic features: anther apices acuminate; corolla

distinctly longer than calyx, ±6-11 mm long.

1 ,2a. subsp. intermedia

Diagnostic features: flowers green to yellow-green,

8-11 mm long; corolla tubular-ovoid; bract, bracteoles

and sepals with small sessile glands and short hairs on

margins (Figure lDfl.

This occurs along the Langeberg and Outeniqua

Ranges from Swellendam to George.

Vouchers: Fourcade 5693 (NBG!, PRE), 4706 (BOL!, PRE);

McDonald 1274 (NBG!); Rourke 294 (NBG!); Taylor LE 3014

(NBG!).

Bothalia 32.1 (2002)

41

1.2b. subsp. albiflora E.G.H.OIiv. & I.M.Oliv.,

subsp. nov., floribus albis minoribus corolla 5. 5-6.0 mm

longa late ovoidea, glandibus in marginibus bracteae,

bracteoleae sepalorumque distinguitur. Figura 1D2.

TYPE. — Western Cape. 3322 (Oudtshoorn):

Ruitersberg, N slopes at W end, 3000 ft [910 m], (-CC),

11 March 1988. Oliver 9105 (NBG).

Illustration: Schumann & Kirsten: 35. t. 1 (1992).

Diagnostic features : flowers white; corolla broadly

ovoid, 5. 5-6.0 mm long; bract, bracteoles and sepals

with only small sessile glands on margins and no hairs

admixed (Figure 1D2).

The subspecies occurs in the Robinson Pass/Ruiters-

berg to Jonkersberg area of the Outeniqua Mountains.

Paratype material : WESTERN CAPE. — 3322 (Oudtshoorn):

Outeniqua Mtns, Ruytersbosch, (-CC), 19-09-1951, G. van Niekerk 79

(BOL!. NBG!); Robinson Pass, 1850 ft [560 m], (-CC), 2-04-1979,

Campbell 13342 (NBG!); ibid.. 30-06-1947, Compton 19595 (NBG!);

ibid., 2-05-1974, Goldblatt 1781 (MO, NBG!); Ruitersberg, 3000 ft

[914 m], (-CC), 11-03-1988, Oliver 9105 (NBG!); ibid., 765 m, 5-04-

1994, Oliver 10429 (NBG!); Jonkersberg, (-CC), 29-08-1978, W. Bond

1462 (NBG!); ibid., 790 m, 4-02-1986. Brusse 4794 (NBG!, PRE);

ibid., 09-1912, Rogers 4273 (NBG); ibid.. 2-06-1951, L.E. Taylor 3006

(NBG!, PRE); ibid.. Van Breda & Admiraal 2105 (NBG!, PRE); ibid.,

2600 ft [790 m], 6-02-1978, Williams 2433 (NBG!); Engelsberg, 1800

ft [549 m], (-CC), 23-10-1984, VI ok 858 (NBG!. PRE).

2. E. vestiflua Salisb. — see E. melastoma subsp.

melastoma (1.1a).

6. E. monadelphia Andrews , Coloured engravings

of heaths; t. 38 (1797); Benth.: 622 (1839); Guthrie &

Bolus: 50 (1905); Duller: 32 (1965) cum auct. Willd.

Iconotype: Andrews: t. 38 (1797).

Note: Andrews published the name as ‘monodelphia’ in the proto-

logue (text and plate), but later changed it to 'monadelphia' in the index

which was published some five years later in 1802 when the complet-

ed volume was bound. He also used the spelling ‘Monadelphia’. This

latter spelling was taken up by all subsequent authors who did not

query the ‘-ia’ ending. There is no indication why Andrews used this

substantival epithet in apposition. This could be assumed to be

Linnaeus’ major Class name. Monadelphia (stamens united by their fil-

aments into one body) (Steam 1957: 31). The grammatically correct

adjectival epithet would be 'monodelpha’.

Illustrations: Baker & Oliver: t. 6 (1967); Schumann & Kirsten: 37,

t. 1 (1992); Oliver & Oliver: t. 3 (2000).

Diagnostic features: anthers with a distinct boundary

between thecae and filament, and a slight basal adaxial

‘nose’; corolla tube straight. ± 11 mm long, bright red

(Figure 1C).

This is a distinct species within the ‘ coccinea' com-

plex and not with E. banksii as its number would suggest.

There are two branching forms within the species — one

with numerous very short side branches as in E. coc-

cinea, the other without these. The species is a resprouter

after fires.

Vouchers: Baker 1886 (NBG!); Oliver 3360 (NBG!. PRE); Oliver

& Palser 78 (K!, NBG!, PRE): Salter 4278 (BM!, BOL!. K!, PRE);

Schlechter 9750 (BM!, BOL!, K!, PRE).

ERICA PLUKENETII COMPLEX

Three separate species used to be recognized in this

complex — E. plukenetii, E. linecita and E. breviflora.

3. E. lineata Benth. — see E. plukenetii subsp. lineata

(5d).

4. E. breviflora Dulfer — see E. plukenetii subsp. bre-

viflora (5e).

5. E. plukenetii L., Species plantarum, edn 1, 1:

356 (1753); Benth.: 622 (1839); Guthrie & Bolus: 50

(1905); Dulfer: 31 (1965). Lectotype: Plukenet, Alma-

gesti Mantissa botanici 45: t. 344, fig. 3 (1700) selected

here by Oliver, Jarvis & Cafferty, in prep.

Note: the iconotype accurately reflects Plukenet's specimen. Herb.

Sloane 89: 11 (BM! photo.-NBG).

Diagnostic features: bract and bracteoles very

reduced, remote from calyx; anthers exserted. long atten-

uated with rounded apices; placenta covered entirely by

ovules abaxially; seeds reticulate.

The main variation in this species is in the length and

shape of the corolla and sepals, size and texture of the

leaves, and in its habit. The flowers are 7—1 6(— 28) mm

long with a tubular or inflated tubular corolla with far-

exserted stamens (Figure 2). Flowers change shape as

they mature, often being rather slender in bud and quite

inflated at maturity. The sepals range from 2-12 mm

long, lanceolate to ovate in shape with the sulcus as long

as the sepal. The plants are mostly reseeders, but can be

resprouters in some areas, and can vary from woody,

rounded shrubs 0.75 m tall near the coast, to tall delicate

erect shrubs in the Bredasdorp District. In the Karnies-

berg the multi-stemmed resprouters can be up to 3 m tall.

This species is often confused with species in the E.

coccinea complex. The position and size of the bract and

bracteoles is the most visible character to distinguish it

from that group.

5a. subsp. plukenetii

E. plukenetii var. densa Bartl.: 630 (1832); Dulfer: 31 (1965). Type:

Caledon und Hottentotsholland, Ecklon s.n. (?).

E. plukenetii var. brachysepala Bartl.: 631 (1832); Dulfer: 31

(1965). Type: Zwischen Strauchen am Fusse des Teufelsbergs [Table

Mtn] in zweiter Hbhe. Ecklon s.n. ( P! ).

E. plukenetii var. brevifolia Bolus: 50 (1905); Dulfer: 31 (1965).

Type: Piquetberg Div.; Piquetberg Range, Schlechter 5208 (BOL!).

Illustrations: Baker & Oliver: t. 2 (1967); Schumann & Kirsten: 36.

t. 7, 8, 9 (1992); Oliver & Oliver: t. 1 (2000).

Diagnostic features: sepals 2-8 mm long, short and

ovate to longer and lanceolate (Figure 2A, B).

There is considerable overlapping variation in the

length of the leaves and in the width and length of the

sepals. This subspecies also exhibits considerable range

in the size of its flowers.

Vouchers: Barker 10423 (MO!, NBG!); Maguire 1129 (NBG!.

NY!): Oliver 3764 (K!, NBG!, PRE), 4918 (NBG!, PRE); Oliver &

Palser 22 (K!, NBG!. PRE); Phillips 7587 (K!, NBG!, NY!, PRE);

Rycroft 1949 (NBG!. P!); Schlechter 5208 (BM!, BOL!, K!, NBG!).

5b. subsp. bredensis E.G.H.OIiv. & I.M.Oliv,

subsp. nov., sepalis latibus ± 5-6 x 3 mm foliis

7— 10(— 1 2) mm longis dignoscenda. Figura 2E.

42

Bothalia 32,1 (2002)

FIGURE 2. — Erica plukenetii complex. A. B. subsp. plukenetii'. A, Oliver 5953, Kamiesberg; B. Rycroft 1949, Bainskloof. C. subsp. penicellata,

Oliver 4226, Stanford: D, subsp. lineata. Oliver 8749, Pearly Beach. Carruther’s Hill; E, subsp. bredensis. Marsh 1468, Cape Infanta. F.

G. subsp. breviflora: F. Taylor 1 1670, Cederberg; G, Rourke 653, Boboskloof. Scale bar: A-G, 4 mm.

TYPE. — Western Cape, 3420 (Bredasdorp): De Hoop

Nature Reserve, 100 ft [30 m], (-AD), 28 July 1970,

Marsh 1468 (NBG).

Diagnostic features: sepals broad (elliptic to ovate), ±

5-6 x 3 mm, leathery; leaves 7— 1 0(— 1 2) mm long; corol-

la ± 15 mm long, white with a pink mouth, red or occa-

sionally yellowish (Figure 2E). Flowers look very much

like those of subsp. lineata , but leaves are broader and

shorter.

The taxon occurs on limestone hills and lateritic Bats

from Heuningrug south of Bredasdorp, eastwards as far

as Cape Infanta and also on the sandstone of Potberg.

There are numerous collections from this area.

Paratype material (selection from many specimens): WESTERN

CAPE. — 3419 (Caledon): Bredasdorp Dist., Heuningrug, 70 m, (-DB),

1-04-1995. Paterson-Jones 218 (NBG!); De Hoop area, (-AD), 8-04-

1957, Barker 8681 (NBG!); ibid., 200 m, 14-06-1979, Burgers 1865

(NBG!); ibid.. Hardevlakte, 10-03-1985, Fellingham 947 (NBG!):

Albcrtsdal, (-BC), 31-05-47, Barker 4553 (NBG!); Potberg,

Elandspad, 600 ft 1 182 ml, (-BC), 1-05-1985, Leiih 51 (NBG!, PRE);

Cape Infanta, 150 ft [45 nt], (-BD), 7-04-1984, Oliver 8428 (NBG!);

Die Poort, (-CA), 10-08-1949. Steyn 341 (NBG!); Moerasfontein. 50 ft

[15 m], (-CB), 15-03-1977, Thompson 3430 (NBG!, PRE).

5c. subsp. penicellata (Andrews) E.G.H.Oliv. &

I.M.Oliv. stat. et comb. nov.

E. penicillata Andrews, Coloured engravings of heaths 2: t. 116

(1802). Iconotype: Andrews: t. 1 16 ( 1802).

E. revolvens Barth: 631 (1832). E. plukenetii var. bicarinata Bolus:

51 ( 1905); Dulfer: 31 ( 1965). Type: berge bei Gnadenthal, Ecklon s.n.

(?holo.; ?P). Note: the collection in P is this species, but not yet veri-

fied as this subspecies.

Diagnostic features: sepals up to 12 mm long, thick

and fleshy, with distinctive non-revolute thick margins

forming two longitudinal keels like a catamaran (Figure

2C), bases of these keels produced into small lobes

beyond point of attachment to pedicel. These features are

well displayed in Andrews' drawing. Bartling's descrip-

tion fits this taxon — ‘sepalis base productis solutis, mar-

gine revolutis’.

Vouchers: Oliver 4226 (NBG!); Schlechter 7743 ( BML BOLL KL PRE).

Bothalia 32,1 (2002)

43

5d. subsp. lineata ( Benth.) E.G.H.OIiv. &

I.M.OIiv., stat. et comb. nov.

E. lineata Benth. in DC.. Prodromus 7: 622 (1839); Guthrie &

Bolus: 48 (1905); Duller: 30 (1965). Type: in colonia Capensi [Cape

Colony], Mund s.n. (Kl).

Illustrations: Baker & Oliver: t. 3 (1967); Schumann & Kirsten: 35,

t. 3,4 (1992).

Diagnostic features: leaves very long, narrow; sepals

(Figure 2D) broad and flat sepals like those in subsp. bre-

densis.

The subspecies occurs on sandy soils associated with

coastal limestone deposits from Gansbaai to Zoetanys-

berg.

Vouchers: Kerr STE30029 (BM!, Kl, NBG!. PRE); Oliver 3365

(NBG!. PRE): Schlechter 9711 (BM!, BOL!. Kl. PI. PRE. SAM!, W).

5e. subsp. breviflora (Dulfer) E.G.H.OIiv. &

I.M.OIiv., stat. et comb. nov.

E. breviflora Dulfer in Annalen des Natiirhistorischen Museums,

Wien 68: 30 (1965). Type: Tliunberg s.n. (UPS).

E. scariosa Thunb.: 350 (1823); Guthrie & Bolus: 49 (1905); non

P.J.Bergius (1767). Type: as for£. breviflora above.

E. penicillata Benth.: 622 (1839), non Andrews, Coloured engrav-

ings of heaths 2: t. 116 (1802) (see subsp. 5c).

Illustrations: Schumann & Kirsten: 36, t. 5. 6 (1992).

Diagnostic features : corolla short, ovoid to almost

globose, mostly ± 7 mm but less than 12 mm, white,

occasionally pink: synflorescence typically very long

spike-like up to 250 mm long (Figure 2F, G).

There is an increase in flower size in some collections

from the Cederberg together with a reduction in the

length of the synflorescences. These can merge with

some short variants of subsp. plukenetii in the

Franschhoek Mountains. We have noted that populations

in the Porterville Mountains have flowers that are sweet-

ly scented. This coupled with the smaller flowers would

indicate that the subspecies is pollinated by insects as

opposed to the postulated bird-pollinated, longer flowers

of all the other subspecies. The plants are single-

stemmed reseeders.

Guthrie & Bolus (1905) noted under E. scariosa 'we

admit this species with doubt' and kept it as distinct ‘with

some reluctance’.

The subspecies occurs in the Cederberg to Porterville

and the southern Cold Bokkeveld area.

Vouchers: Bolus 5114 (BM!. BOL!. K!, PRE, SAM!); Drege 7694

(BM!. K!, NBG!, PRE, W); Oliver 4075 (NBG!. PRE); Rourke 653

(K!, NBG!): Schlechter 9084 (BM!, K!, NBG!. PRE).

6. E. monadelphia Andrews — this has been placed in

the E. coccinea group after E. intermedia ( 1.2).

7. E. banksii Andrews, Coloured engravings of

heaths 1: t. 5 (1797) as E. banksia: Willd.: 395 (1799);

Bauer: t. 29 (1803); Benth.: 624 (1839); Dulfer: 32

(1965); as banksia Guthrie & Bolus: 51 (1905). Icono-

type: Andrews: t. 5 (1797).

Note: as with several other epithets coined by Andrews, the ‘-ia’

ending was thought to reflect the use of a generic name used in appo-

sition, in this case Banksia (Australian Proteaceae), and should not be

corrected. However, there are some cases where a generic name did not

exist or was coined later than the publication of the species name.

These -ia endings are being replaced by the relevant genitive ending (-

i, -ii, -ae or -iae) for all Andrews’ names commemorating persons. The

date of the Bauer plate is actually 1 Jan. 1801 and not the date of the

first published fascicle in the bound publication, i.e. 1796, as used by

Dulfer to antedate Bauer’s name over that of Andrews. However, fas-

cicle 3 containing this plate only appeared in 1803 (Britten 1899).

Diagnostic features: anthers well exserted with fila-

ments held in a tight bundle, bilobed with basal attach-

ment and distinct basal ‘chin’; corolla lobes spreading to

reflexed; tube straight, white or greenish with lobes

green or purple; leaves slightly serrated with sharp, short

to long mucro and longish petiole; bract and bracteoles

large and approximate to calyx; sepals with terminal

sharp mucro. Inflorescences numerous, pendulous, con-

sisting of 3-nate flowers at ends of main branches.

7a. subsp. banksii

Diagnostic features : leaves short, ± 5-6 mm long;

corolla yellowish green; ovary hairy (Figure 3A).

The typical subspecies forms a small, compact,

woody shrublet growing on rock ledges and occurs in the

mountains around the Elgin Basin.

Vouchers: Esterhuysen 2695 (BOL!. NBG!. PRE): Schlechter 7572

(BOL!, K!, PRE).

7b. subsp. purpurea (Andrews) E.G.H.OIiv. &

I.M.OIiv., stat. et comb. nov.

E. banksia var. purpurea Andrews, The Heathery 3: t. 106 (1806);

Guthrie & Bolus: 51 (1905). E. banksia var. purpurea (Andrews)

Dulfer: 32 (1905). Iconotype: Andrews: t. 106 (1806). Note: Andrews’

painting in Coloured engravings of heaths: t. 151 is not dated, so the

volume date has to be used in this case, namely 1809.

Illustrations: Baker & Oliver: t. 5 (1967); Schumann & Kirsten: 38,

t. 3,4(1992).

Diagnostic features: corolla white with purple lobes;

leaves short, ± 5-9 mm long; ovary glabrous.

This subspecies grows on the mountains from

Babylon’s Tower southeast to Quoin Point.

Vouchers: Oliver STE30033 (BOL!. K!, NBG!. PRE); Zeylier 3189

(BOL1. P!, W).

FIGURE 3. — Erica banksii. A, subsp. banksii, Esterhuysen 2695,

Rooskraalberg: Bi. B2, subsp. comptonii, holotype, Compton

6066, Hangklip. Scale bars: A, Bi, 4 mm; B?, 2 mm.

44

Bothalia 32,1 (2002)

7c. subsp. comptonii ( Salter ) E.G.H.Oliv. &

I.M.Oliv., stat. et comb. nov.

E. comptonii Salter in Journal of South African Botany 2: 60

(1936): Dulfer: 32 (1965). Type: Caledon Div., among rocks, northeast

slope of Hangklip, 1400 ft [426 m], Jan, Compton 6066 (BOL!, K!,

NBG1).

Illustrations: Baker & Oliver: t. 4 (1967); Schumann & Kirsten: 38,

t. 5, 6 (1992).

Diagnostic features : corolla greenish white; leaves

long, 1 2— 20(— 26) mm long; ovary glabrous (Figure 3B).

This subspecies is confined to the higher mountain

peaks from Kogelberg to Hangklip. In the Kogelberg

complex the subsp. banksii occurs at lower altitudes at

the northeastern end.

Subsp. comptonii differs from the other two sub-

species in the larger, more open habit (up to 500 mm tall)

in open ground between rocks and in the longer leaves

(up to 26 mm) with longer mucros on the leaves and

sepals (Figure 3Bi). The sepals can vary from about half

to the full length of the corolla tube.

The variation in this species occurs in the flower

colours, the habit, habitat, length of leaves and length of

the mucro on the leaves and sepals. Disjunctions occur

regionally, but warrant recognition only at subspecific

level.

Vouchers: Boucher 1332 (NBG!, PRE); Compton 6066 (BOL!, K!,

NBG!); Pillans 8183 (BOL!, K!).

8. E. primulina Bolus — see E. viridiflora subsp.

primulina (9b).

8.1. E. leucosiphon L. Bolus in Kew Bulletin 1933:

186 (1934); Dulfer: 32 (1965). Type: Cape Town Wild

Flower Show, BOL19253 (BOL!).

Illustrations: Schumann & Kirsten: 39, t. 9, 10 (1992).

Diagnostic features: pedicel long, with dense plumose

hairs; calyx white; anther appendages broad; corolla

white.

This distinct species has no alliances with any other

long-tubed species, but has, rather, an alliance with sev-

eral species with much shorter flowers, such as E. mon-

soniana L.f. (no. 402) and E. goatcheriana L. Bolus (no.

405.1 ). The possession of plumose hairs on the pedicel

are shared by all of them.

Vouchers: Kellerman 157 (PRE); Kirsten 434 (NBG!); Oliver 4101

(NBG!).

9. E. viridiflora Andrews , The Heathery: t. 299

(1812); Guthrie & Bolus: 52 (1905); Dulfer 32 (1965).

Iconotype: Andrews: t. 299 (1812).

Note: Andrews’ much fuller painting in Coloured engravings of

heaths: t. 287, must have appeared after 1820, which date Andrews

mentioned in the text accompanying the plate, even though the volume

is dated ‘1809’.

Diagnostic features: corolla lobes large and erect;

anther appendages decurrent along filament, pendulous

to very reduced and ear-like.

9a. subsp. viridiflora

Illustrations: Baker* Oliver: t. 7 (1967); Schumann & Kirsten: 39,

t. 11, 12 (1992).

Diagnostic features: corolla lime-green, sticky, with a

patch of sessile, sticky glands in centre of upper half of

adaxial surface of sepals. Inflorescence consists of three

flowers; it is terminal on main and occasionally sec-

ondary branches; bract and bracteoles approximate to

calyx; anthers bilobed with thecae often divergent; ovary

ovoid and glabrous (Figure 4A).

The typical subspecies forms an erect shrub, 0. 5-1.0

m tall, growing in open ground or between rocks. It

occurs on the coastal mountains from George through to

Humansdorp.

Erica clavata Andrews (Coloured engravings of

heaths: t. 159, 1809) was included by Guthrie & Bolus

(1905) and Dulfer (1965) in synonymy. Careful exami-

nation of his painting leads to the conclusion that E.

clavata could be a hybrid with E. viridiflora as a possi-

ble parent. Characters which do not fit the current taxon

are the erect imbricate leaves, the broadly ovate sepals

with long attenuate apices, and the muticous anthers.

Vouchers: Oliver STE30030 (BM!, BOL!, K!, NBG!, PRE!);

Schlechter 5801 (BM!, BOL!, K, PRE, W).

9b. subsp. primulina (Bolus) E.G.H.Oliv. & l.M.

Oliv., stat. et comb. nov.

E. primulina Bolus in Flora capensis 4: 51 (1905); Dulfer: 32

(1965). Type: Ladismith Div., on rocky mountain slopes of the Klein

Zwartbergen, near Seven Weeks Poort, 3250 ft [990 m], Marloth 2937

(BOL!).

Illustrations: Schumann & Kirsten: 39, t. 7, 8 (1992).

Diagnostic features: corolla cream to greenish, non-

sticky; shrubs small, woody shrublets, erect to 300 mm

FIGURE 4. — Erica viridiflora, with flower, sepal in adaxial view

showing zone of sessile glands, and anther. A, subsp. viridiflo-

ra, Visser 29, Ruitersberg; B, subsp. primulina. Oliver 10819,

Meiringspoort; C, subsp. redacta, Vlok s.n., Meiringspoort.

Scale bars: anthers, sepals, 2 mm; flowers, 4 mm.

Bothalia 32.1 (2002)

45

high but sometimes prostrate (Figure 4B).

Subsp. primulina occurs only in the Swartberg Range

from the Klein Swartberg to near Willowmore. A collec-

tion from Gamkaberg, which is further to the south and

west than the rest of the collections, is an intermediate

variant having the flowers of this subspecies, but the

growth of subsp. viridiflora — it is recorded as a woody

erect shrub 500 mm tall.

In the protologue. Bolus mentioned the affinity with

E. viridiflora , but surprisingly stressed a closer likeness

to E. banksii and gave the differences from that species.

Vouchers: Esterhuysen 6286 (BOL!, K!, NBG!. PRE); Oliver 3436

(NBG!).

9c. subsp. redacta E.G.H.Oliv. & I.M.Oliv., subsp.

nov., a subspecie typica floribus multo minoribus differt.

Figura 4C.

TYPE. — Western Cape, 3322 (Oudtshoorn):

Meiring's Poort, (-BC), 14-10-2001, Vlok & Schutte 458

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

Diagnostic features : differs from subsp. primulina

only in size of flowers and their parts, corolla 4. 5-5.0

mm long (Figure 4C).

This small-flowered form is restricted to a single popu-

lation of some 100 plants occurring in crevices on a large

steep slab of quartzite rock. The plants were about 200

mm tall and bore creamy white to sometimes green-

tinged flowers. Subsp. primulina grows in similar habi-

tats in the vicinity.

This is clearly a case of a short-tubed variant adapted

to a different pollination syndrome (insect versus bird).

Paratype material. WESTERN CAPE. — 3322 (Oudtshoorn):

Meiring's Poort, (-BC). 01-2001. fruiting, Vlok s.n. (NBG).

10. E. sphenanthera Tausch, Flora: 17: 626 (1834);

Guthrie & Bolus: 52 (1905); Duller: 33 (1965). Type:

without locality or collector (PRG!, holo.; photo.

-NBG!).

This taxon is regarded as a hybrid of garden origin in

Europe. No material matching the description and the

type has been found in the Cape. Parentage could include

E. abietina because of the similar bracteoles, anthers and

ovary.

11. E. cerviciflora Salisb. in Transactions of the

Linnean Society 6: 362 (1802); Guthrie & Bolus: 53

(1905); Dulfer: 33 (1965). Type: Hottentots Holland, 1.

Mulder s.n. ( K ! ).

No material matching the description and type has

been found in the Cape. We postulate that this collection

is of hybrid origin in the wild. Parentage could include E.

abietina because of the similar anthers and ovary. Jacob

Mulder is known only through the citation by Salisbury

of 37 species of Erica from the Cape (Gunn & Codd

1981). Only one specimen is labelled as a Mulder col-

lection by Salisbury.

11.1. E. beatricis Compton in Journal of South

African Botany 9: 128 ( 1943); Dulfer: 33 (1965). Type:

Uniondale Dist., Helpmekaar (or Thumb) Peak, Bond

892 (NBG!).

Illustration: Compton, l.c„ t. 8 (1943).

Diagnostic features: anthers exserted, with short stub-

by appendages below thecae; ovary puberulous; corolla

glabrous; stems, leaves, pedicel, bract, bracteoles and

sepals all puberulous.

This species has been collected only once. There are

no indications of its alliances. There is a possibility that

it is a naturally occurring hybrid, as found in several

other supposedly rare species that have been investigat-

ed in the wild, e.g. E. x flavisepala (Oliver 1977) and E.

x vinacea (Oliver 1986).

Vouchers: Bond 892 (NBG!); Esterhuysen 5021 (BOL!).

The remaining 24 species with whole numbers dealt

with in this paper were placed by Guthrie & Bolus

(1905) in § Evanthe which was based on ‘Inflorescence

mostly axillary, more rarely also terminal on the same

plant. Corolla tubular, mostly over 6 lin. [12 mm] long,

rarely shorter.' The inflorescence is 1 -flowered on a very

short lateral branchlet often less than 1 mm long and

sometimes bearing very reduced scarious leaflets. These

occur in the axils of leaves on the main branch (hence

axillary) and are arranged in dense spike-like synflores-

cences towards the ends of the main branches which con-

tinue their vegetative growth. This contrasts with the

often 3-nate inflorescences on leafy lateral branchlets

found in §Pleurocallis which will be dealt with in Part 2.

The inclusion of species in this section, § Evanthe , has

been rather arbitrary, being based mainly on the size of

the corolla. Species with smaller flowers, but with the

same inflorescence structure, have been included in

§ Hermes , and one of these is certainly very closely relat-

ed to some species in this section, e.g. E. axilliflora

L. Bolus.

12. E. mammosa L., Mantissa plantarum altera:

234 ( 1771); Benth.: 624 (1839); Guthrie & Bolus: 53 ( 1905);

Dulfer: 33 (1965). Lectotype: Herb. Linn. 498.33 (LINN),

selected here by Oliver, Jarvis & Cafferty in prep.

E. gilva J.C. Wendt.: 48 et fig. (1798); Benth.: 624 (1839); Guthrie

& Bolus: 54 (1905). Type: sine. coll, et loc. (?); iconotype: J.C. Wendt,

fig. (1978).

Illustrations: Baker & Oliver: t. 9 (1967); Schumann & Kirsten: 40,

t. 1-3; 42, t. 5, 6 ( 1992); Oliver & Oliver: t. 4 (2000).

Diagnostic features: corolla narrowed at base with 4

indentations or folds, glabrous; bract and bracteoles

small and remote from calyx; anthers with long thin

appendages and a basal, projecting ‘nose’; leaves erect

and green; ovary short, rounded and glabrous.

We agree with Salter’s reduction of E. gilva to syn-

onymy under E. mammosa, both having four broad to

narrow indentations at the base of the corolla. The former

occurs only on Table Mountain on the Cape Peninsula

and was separated by Guthrie & Bolus (1905) on the

longer bract, bracteoles and sepals — it has white flowers

sometimes with a green tip. There is a lot of variation in

the colour of the flowers on the Peninsula, particularly in

the southern parts and some plants in the Cape of Good

Hope Nature Reserve have very long spike-like synflor-

escences.

Erica mammosa is widespread from the Cape

Peninsula to the Cold Bokkeveld and eastwards to

Bredasdorp with flowers orange or deep purple-red,

greenish yellow to white. Variations occur in the length

of the spike- 1 ike synflorescence from short and dense to

46

Bothalia 32,1 (2002)

very long (up to 250 mm) and rather open. The pedicel

can be glabrous or hairy and the sepals 3-6 mm long. On

the mainland the plants are resprouters, whereas on the

Peninsula they may be resprouters or single-stemmed

reseeders. The most variation occurs on the mountains

above Simonstown.

Vouchers: Bolus 3354 <BM!, BOL!, K!, P!, PRE, SAM!);

Schlechter 10115 (BM!, K!. P!. PRE. W).

13. E. broadleyana Andrews, Coloured engravings of

heaths: t. 154 (1809). Type: Andrews: t. 154 (1809).

This is regarded as a hybrid of garden origin in

England. The plant looks like E. plukenetii, but has

included anthers. Andrews’ drawing shows the upper part

of the filaments zig-zagged, which condition would indi-

cate that the anthers could become exserted from the

very narrow mouth.

14. E. baueri Andrews , The Heathery: t. 252 (1812)

as bauera ; Dulfer: 33 (1965). Iconotype: Andrews: t. 252

(1812).

Note: Andrews’ epithet is corrected to the genitive case instead of the

generic name, Bauera , used in apposition (see note under E. banksii no. 7).

E. bowieana Lodd.: t. 842 (1824): Benth.: 624 (1839): Guthrie &

Bolus: 54 (1905). Iconotype: Lodd.: t. 842.

Illustrations: Baker & Oliver: t. 10 (1967): Schumann & Kirsten:

40, t. 4(1992).

Diagnostic features', corolla base inflated and rounded

with no indentations, white or shell pink; anthers with

basal nose and long thin appendages; leaves usually

greyish and recurved; long curved pedicel, small bract

and bracteoles remote from calyx; ovary short, rounded,

glabrous.

14a. subsp. baueri

Diagnostic features', leaves short, spreading to

recurved; flowers usually 1 0-20 in few short dense sub-

terminal spike-like syn florescences, white to pale pink.

This commonly cultivated and well-known sub-

species is now becoming rare in its habitat due to the

spread of agriculture and alien vegetation (mainly Acacia

cyclops) in the sandy flats which are derived from quart-

zitic sandstone and lie west of Albertinia.

Vouchers: Kerr STE30084 ( BOL!, K!, NBG!. PRE); Oliver 174 sub

Baker 1589 { BM!, NBG!).

14b. subsp. gouriquae E.G.H.Oliv. & I.M.Oliv.,

subsp. nov., a subspecie typica foliis erectis floribus plus

numerosis plus sparsis differt. Figura 5.

TYPE. — Western Cape, 3421 (Riversdale): Alber-

tinia, Rein’s Coastal Nature Reserve, main valley just N

of Trig. Beacon 64, 150 m, (-BC), E.G.H. & I.M. Oliver

11933 (NBG, holo.; BM, BOL, E, K, MO, NY, P, PRE, S).

Diagnostic features: leaves erect; Bowers 5-10 in

loose, spike-like syn florescences, more numerous and

scattered over shrub, pale to deep pink (Figure 5).

This subspecies forms a taller, denser shrub up to 2.5

m. It grows very localized in sandy areas associated with

limestone between the hills to the south of Albertinia

near the coast, an area called Gouriqua.

Paratype material: WESTERN CAPE. — 3421 (Riversdale):

Buffelshoek. N of Ystervarkpunt. 140 m, (-BC), 13-09-1983, Burgers

3152 (NBG!); Ystervarkpunt/Gouriqua, 130 m. (-BC), 9-07-1987,

Willemse 624 (NBG!); ibid.. 200 nr, 27-10-1986. T.J. van cler Merwe

146 ( NBG!).

15. E. giiva J.C.Wendl., see under E. mammosa (12).

16. E. sessiliflora L.f, Supplementum plantarum:

222 (1782); Benth.: 625 (1839); Guthrie & Bolus: 55

(1905). Type: Cap. bonae spei, sine coll. (UPS).

E. clavaeflora Salisb.: 365 (1802). E. sessiliflora var. clavaeflora

(Salisb.) Bolus: 55 ( 1905); Dulfer: 34 ( 1965). Type: comm. Hibbert s.n. (K!).

E. sceptriformis Salisb.: 365 (1802). E. sessiliflora var. sceptri-

formis (Salisb.) Bolus: 55 (1905); Dulfer: 34 (1965). Type: sine loc.

Roxburgh s.n. (K!).

E. sessiliflora var. oblanceolata Bolus: 55 (1905); Dulfer: 34

(1965). Type: sine loc., Guthrie 3795 (BOL).

Illustrations: Baker & Oliver: t. 8 ( 1967); Schumann & Kirsten: 42.

t. 7-9 ( 1992); Oliver & Oliver: t. 5 (2000).

Diagnostic features: pedicel very short; bract, bracte-

oles and sepals equally long and becoming enlarged,

Beshy and persistent in fruiting stage; corolla pale green

to yellowish green; sepals spathulate with entire to ser-

rated margins; anther appendages narrow; ovary short,

rounded and glabrous.

This is a very distinctive species being the only one in

the genus exhibiting serotiny — the fruiting synflores-

cences remain on the plants for several seasons as cone-

like structures with the fruits protected by the thickened

calyces. The species is very variable in the size of the

corolla and in the shape of the sepals from narrowly

oblanceolate to very broadly spathulate — characters used

to separate several varieties by previous authors. There

is, however, no clear-cut disjunction between these

shapes that warrant recognition. A very showy variant

occurs on the Riviersonderend Mountains with green

corolla and bright red sepals compared to the normal

green sepals.

Vouchers: Burchell 5354 (NBG!, NY!. W); Compton 14230 ( NBG);

Oliver 8995. 11236 (NBG): Parker 3798 (BOL!, NBG!); Schlechter

7583 (BM!, K!, NBG!, PRE. W).

17. E. filipendula Benth. in DC., Prodromus 7:

663 (1839); Guthrie & Bolus: 56 (1905); Dulfer: 34

(1965). Type: Cape Colony, Bowie s.n. (K!).

Diagnostic features: corolla cyathiform or slightly

closed at mouth, 5-10 mm long; sepals narrow lanceo-

late, about 73-72 length of corolla; anther appendages

narrow with very few lateral teeth, anther nose long,

erect to absent.

17a. subsp. filipendula

Illustrations: Baker & Oliver: t. 13 (1967); Schumann & Kirsten:

43, t. 12, 13 (1992).

Diagnostic features: corolla cyathiform, 8-10 mm

long, yellow or dark pink; anthers situated in middle of

corolla. Figure 6B.

The typical subspecies occurs on sandy hills between

Viljoenshof and Pearly Beach in the Bredasdorp District.

Both colour forms have been noted growing sympatrically.

Vouchers: Baker 1775 (BM!, NBG!); Bolus 8450 (BM!. BOL!,

PRE); Guthrie 3786 (BOL!, NBG!); Oliver 3414 (NBG!. PRE), 8748

(NBG!).

Bothalia 32.1 (2002)

47

FIGURE 5. — Erica baueri subsp. gouriquae , Willemse 624. A, flowering branch, x 1; B, stem with leaves removed; C, leaf; D, flower; E, anther,

side, front and back views; F, gynoecium; G. ovary, opened laterally; H, capsule; 1, seed; J, testa cells. Scale bars: B, C, E, F-H, 2 mm; I,

1 mm; D, 4 mm.

17b. subsp. parva E.G.H.Oliv. & I.M.Oliv., subsp.

nov., subspeciei typicae similis sed floribus albis ad pal-

lide roseis corolla 5-7 mm longa antheris in corolla

altioribus differt. Figurae 6C; 7.

TYPE. — Western Cape. 3419DB (Caledon): Bredas-

dorp Dist.. Soetanysberg area, west end just east of Melk-

pan, 40 m. 12 May 1999, E.G.H. & l.M. Oliver 11254

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

E. filipendula var. minor Bolus: 56 (1905) p.p. Syntypes:

Bredasdorp Div., between Elim and Ratel River, 200-300 ft [60—90 m],

Guthrie 3784 (BM!. BOL!, SAM!); ibid.. Bolus 8451 (BM!, BOL!);

sine loc.. Zeyher 1090 (K!. BOL! fragm.).

Diagnostic features: corolla white to pale pink, urceo-

late-cyathiform. 5-7 mm long; anthers positioned higher

up in corolla than in typical subspecies. Figures 6C; 7.

The type population had flowers varying from white to

pink on the same plant and grew with low plants of the

green-flowered form of E. penduliflora (17.2). The subspecies

occurs on sandy flats and hills between Elim and Zoetanys-

berg. There are numerous collections from this region.

Paratype material: WESTERN CAPE. — 3419 (Bredasdorp): Ratel

River. 150 ft [45 m], (-DA), 16-07-1895, Guthrie 3784 (BM!, BOL!

fragm., SAM!); Uintjieskuil, near Ratel River, (-DA), Van Breda 2149

(NBG!); Soetanysberg, W end above Suur-en-Soet, 350 ft [106 m],

(-DB), 29-03-1971, Oliver 3357 (NBG!, PRE); Rietfontein. near

homestead, 45 m, (-DB), Pater son- Jones 234 (NBG!); Rietfontein.

(-DB), 14-05-1993, Schumann 843 (NBG!); 1 mile [1.6 km] N of

Soetanysberg. 300 ft [90 m], (-DB), 2-06-1967, Williams 1005 (NBG!,

PRE). Without precise locality: near Elim and Ratel River, 300 ft [90

in], (-DA), 07-1895. Bolus 8451 (BM!, BOL!).

This species forms a complex with the next two

species, which were all treated as one species by Guthrie

& Bolus ( 1905). It varies in a number of characters — size

and colour of flowers, form of anther appendages, anther

shape and position in the flower (Figure 5). Bolus’ var.

major was removed as a distinct species, E. penduliflora

(17.2) and Dulfer (1965) described var. minor Bolus as E.

globulifera. The remaining medium to small-flowered

forms constitute this species.

17.1. E. globulifera Dulfer in Annalen des

Natirrhistorischen Museums, Wien 68: 35 (1965). E. fili-

pendula Benth. var. minor Bolus: 56 (1905) p.p. Type:

Bredasdorp Div., between Elim and Ratel River,

200-300 ft [90 m], Schlechter 10472 (BM!, BOL!, P!.

PRE, SAM!, W).

48

Bothalia 32,1 (2002)

FIGURE 6. — Relationship of floral

parts in A, Erica penduliflo-

ra: B, E. filipendula subsp.

filipendula ; C, E. filipendula

subsp. parva ; D, E. globulif-

era. Left, flower cut open

longitudinally to show posi-

tion of anthers; centre, sepal;

right, stamen in side view

showing variations in basal

nose. Scale bars: flower, 4

mm; sepal, stamen, 2 mm.

Diagnostic features : sepals ovate, V5-V4 as long as

corolla; anther appendages broadly based and much

toothed; corolla purplish pink, 3-5 mm long.

The larger-flowered collections southwest of Bredas-

dorp have very reduced anther noses, whereas the collec-

tions from the De Hoop area have smaller flowers and

remarkable turned-up noses, a character not found in any

other species (Figure 6D).

Dulfer only examined the single collection,

Schlechter 10472 (W), when he raised E. filipendula var.

minor to specific status and selected that as the lectotype,

unaware that the other three syntypes constituted anoth-

er distinct taxon (see 17b).

Vouchers: Schlechter 10472 (BM!, BOL!, P!, SAM!, W).

17.2. E. penduliflora E.G.H.Oliv. in E.G.H.Oliv.

& I.M.Oliv. in Yearbook of the Heather Society 2001: 31

(2001 ). Type: Western Cape, 3419DA, Bredasdorp Dist.,

hills NW of Viljoenshof, 120 m, 12 May 1999, white-

flowered, E.G.H. & I. M. Oliver 11245 (NBG, holo.; BM,

BOL, K, MO, NY, P. PRE, S).

E. filipendula var. major Bolus: 56 (1905). Types: Syntypes:

Bredasdorp Div., fairly abundant on downs between Elirn and Ratel

River, 300-600 ft [90-180 m], Guthrie 3786 (BOL!, NBG!); ibid..

Bolus 8452 (BM!, BOL. NBG!, PRE); ibid., Schlechter 7618 (BM!,

BOL!. NBG!, P!. PRE. W), Schlechter 7726 (BM!, BOL!, P!.W); ibid.,

MacOwan [Schlechter] in Herb. Aust. Aft: 1920 (BM!, BOL!, P!,

SAM!).

Illustrations: Baker & Oliver: t. 13 (1967); Schumann & Kirsten:

43, t. II (1992).

Diagnostic features: corolla 12-18 mm long; sepals

broadly ovate; anthers with projecting nose and long thin

appendages; ovary glabrous with long stipe. Figure 6A.

'Iliis was regarded as a large-flowered variety of E. fili-

pendula by Bolus (1905), but was deemed sufficiently

distinct to be recognized as a distinct species based on

the above characters. The species has two distinct colour

variants, white or yellowish green, which are allopatric.

In the Pearly Beach area a white variant has recently

been recorded with green tips.

Vouchers: Oliver 11245 (BM!. BOL1. K!, MO!, NBG!, NY!, P!,

PRE!, S ! ), 11246 (BM!, K!. NBG!. NY!, PRE!); Schlechter 7618 (BM!.

BOL!, NBG!. P!, PRE. W).

18. E. grandiflora L.f. — see E. abietina subsp. auran-

tiaca (23e).

19. E. exsurgens Andrews — see E. abietina subsp.

aurantiaca. (23e).

20. E. longisepala Guthrie & Bolus: 57 (1905);

Dulfer: 35 (1965). This is just a large-flowered form of

E. parilis Salisb. (Guthrie & Bolus 1905: sp. no. 301).

21 . E. hibbertii Andrews , Coloured engravings of

heaths: t. 172 ( 1805) as hibbertia ; Guthrie & Bolus: 58

( 1905); Dulfer: 36 (1965). Type: Andrews: t. 172 (1805).

Note: see note under E. hanksii (7) for the need to change Andrews’

personal epithets.

Illustrations: Baker & Oliver: t. 18 (1967); Schumann & Kirsten:

44, t. 16, 17 (1992).

Diagnostic features: corolla smooth (lacking any lon-

gitudinal ridges), sticky; bract and bracteoles nearly as

long as sepals and approximate to them; inflorescences

umbel-like.

The inflorescence is umbel-like on the main stems and

can have, in addition, below the umbel, short leafy

branchlets ending with a 3- or 4-nate inflorescence. This

latter situation is shown in Andrews’ painting.

The main axis can sometimes continue growth.

Vouchers: Bolus 5168 (BOL!. PRE); Oliver STE30035 (BM!. K!.

NBG!, PRE!). 11952 (K, NBG, NY, PRE); Salter 4952 (BOL!, K!,

SAM!).

21.1 E. tenax L. Bolus — see E. thomae (2 1 .2).

Bothalia 32.1 (2002)

49

FIGURE 7. — Erica filipendida subsp. parva , type, Oliver 11254. A, flowering branch, x 1; B, stem with leaves removed; C, leaf; D, vestigial lat-

eral branch in axil of vegetative leaf, showing vestigial leaves at base and flower with corolla removed; E, flower; F, anther, side, back and

front views. Scale bars: B-G, 2 mm.

21.2. E. thomae L. Bolus in Annals of the Bolus

Herbarium 4: 17, t. 1A (1925); Duller: 36 (1965). Syn-

types: Cape Province, South-Western Region; Caledon

Div., fl. Jan., Rooi Els near Hangklip, T.P. Stokoe in

BOL17571 (BOL! K!); Palmiet River Valley, Stokoe in

BOL17572 (BOL!). Lectotype selected here: Stokoe in

BOL17571 (BOL).

E. thomae L. Bolus var. brevisepata L. Bolus: 18 (1925). Type: Cape

Town Wildflower Show, sine coll. BOL16233 (BOL!).

E. tenax L. Bolus: 17. t. IB ( 1925): Dulfer: 36 (1965). Type: Cale-

don Div., Palmiet River Valley, fl. Jan., T.P.Stokoe in BOL17623

(BOL!, K!).

E. porteri Compton: 125. t.7 (1953): Dulfer: 36 (1965). Syntypes:

Caledon Dist.. Buffels Kloof, near Pringle Bay, 500 ft. alt.. 25 March

1953, [specimen figured]. Porter s.n. (NBG!); ibid., 25 March 1953.

Porter s.n. (NBG!); ibid., 17 August 1953. Porter s.n. (NBG!).

Illustrations: L. Bolus: t. 1 A (1925); Schumann & Kirsten: 44, t. 18;

45, t. 19-21 (1992).

Diagnostic features', corolla sticky, with longitudinal

ridges, densely to sparsely covered with hair-like

spicules or pustules; flowers in a dense spike-like syn-

florescence, with 1 -flowered florescences on vestigial

lateral branchlets bearing small scarious leaves; bract

and bracteoles small, tough and situated in middle of

pedicel; anthers included with ear-like appendages below

thecae; filaments with spicule-like hairs; ovary rounded

and glabrous.

We retain the epithet thomae in recognition of

Thomas Stokoe’s considerable collections of fynbos

plants made from the 1920s to the early 1950s — both L.

Bolus’ species were described on the same page.

The species shows much variation in the size, thick-

ness and degree of spreading of the leaves, the length of

pedicels (from 4-16 mm long), and the colour and size of

the Bowers. There are no clear-cut discontinuities to war-

rant formal taxonomic rank and the three species have

been grouped here as variants of a single species. All

occur in the mountains from Kogelberg to Hangklip to

Kleinmond. Some detailed molecular studies may help to

elucidate the relationships between the variants.

VARIANT A (formerly E. thomae sensu stricto ):

medium-long pedicels, 6-10 mm long; corolla 22-30 mm

50

Bothalia 32, 1 (2002)

long, rose pink to dark reddish pink with or without paler

tips. It occurs in the southwestern parts of the distribution

range.

VARIANT B (formerly E. tenax ): long pedicels,

13-16 mm long; corolla 22-30 mm long, green to white.

This is the variant from the northern and eastern parts of

the distribution range.

VARIANT C (formerly E. porteri ): short pedicels, 4-7

mm long; corolla 20-25 mm long, more delicate and dark

reddish pink with white mouth; leaves more spreading up

to 90°. This is restricted to a single small population in

the Buffelsrivier Valley near Pringle Bay where it grows

with Variant A and appears to produce hybrids. However,

some collections of Variant A from other areas possess

similar small flowers, but not the bicoloured corolla. The

Buffelsrivier population has plants more delicate than the

other two variants.

Vouchers: Oliver 92 (NBG!, PRE!); Porter s.n. (K!, NBG!). sub

PRE28592 (K!. PRE); Stokoe 8347 (BOL!. NBG!. PRE).

21.3. E. porteri Compton — see E. thomae (21.2).

22. E. phylicifolia Salisb. — see E. abietina subsp.

atro rosea (23b).

22.1. E. nevillei L.Bolus in Annals of the Bolus

Herbarium 3: 172, t. 7B (1923); Dulfer: 37 (1965). Type:

Cape Province: South-Western Region: Cape Peninsula,

southwestern slopes of Noord Hoek Mountain, fl.

Jan-Feb., N.S. Pillans 4124 (BOL!, Kl, NBG!).

Illustrations: Schumann & Kirsten: 46., t. 24 (1992); Oliver &

Oliver: t. 12 (2000).

Diagnostic features: corolla tube with basal restriction

zone; inflorescence spike-like and/or umbel-like and with

no continuing apical growth; ovary emarginate, glabrous.

With the bipartite anthers and emarginate turbinate

ovary, this species clearly belongs in the E. abietina

group. It is a restricted endemic on the Cape Peninsula.

Vouchers: Baker 612 (BM!, NBG!, PRE!); Galpin 12531 (K!. P!.

PRE); Pillans 4124 (BOL!. K!, NBG!).

22.2. E. quadrisulcata L.Bolus in Annals of the

Bolus Herbarium 3: 1 72, t. 7D (1923); Dulfer: 37 (1965).

Type: Cape Province: South-Western Region: Cape

Peninsula, neck between Signal Mountain and Zwartkop,

Klaver Valley, near Simonstown, 11. Jan., Pillans 3944

(BOL!, NBG!).

Illustrations: L.Bolus: t. 7D (1923); Schumann & Kirsten: 46, t. 25

(1992); Oliver & Oliver: t. 14 (2000).

Diagnostic features: ovary acute, glabrous; inflores-

cence umbel-like, not continuing with vegetative elonga-

tion.

The species is a very restricted endemic on the Cape

Peninsula.

Vouchers: Baker 852 (BM!. NBG!); Galpin 12798 (K!. PRE);

Pillans 3944 (BOL!, NBG!).

23. E. abietina L., Species plantarum edn 1,1: 355

(1753); Salter: 634 (1951); Dulfer: 37 (1965). Type: this

will be dealt with in a paper on Linnaean typification

(Oliver, Jarvis & Cafferty, in prep.).

Diagnostic features: ovary emarginate, obovoid, cov-

ered with dense, short, retrorse hairs (Figure 8).

The considerable range of variation and lack of clear

disjunctions have resulted in the reduction of several

well-known species to subspecific rank and with two new

taxa warranting only subspecific rank.

23a. subsp. abietina

E. coccinea sensu P.J.Bergius: 92 (1767), non L. (1753); Benth.:

627 (1839); Guthrie & Bolus: 59 ( 1905). Type: without locality or col-

lector (SBT).

Illustrations: Schumann & Kirsten: 46, t. 26 (1992); Oliver &

Oliver: t. 9 (2000).

Diagnostic features: corolla dark red, tubular, 18-26

mm long, spiculed to sparsely puberulous and slightly

viscid; sepals subovate, sparsely pilose with adaxial ses-

sile glands; anthers included to exserted (Figure 8B).

The subspecies is confined to the upper rocky slopes

and plateau of Table Mountain.

Vouchers: Bolus 3366 (BM!. BOL!. K!. PRE), sub Herb. Norm. 189

(BM!. BOL!. P!); Oliver 110 (NBG!).

23b. subsp. atrorosea E.G.H.Oliv. & I.M.Oliv.,

subsp. nov., corolla rosea ad atrorosea, tubulosa, 18-22

mm longa, ± glabra aliquantum viscida. sepalis late

lanceolatis, sparse puberulis glandibus adaxialibus,

antheris inclusis interdum manifestis. Figura 8C.

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

Pond, (-AB), 14 June 1949, Barker 5355 (NBG).

E. purpurea Andrews, Coloured engravings of heaths: t. 50 ( 1795);

Benth.: 627 (1839); Guthrie & Bolus: 58 (1905). Iconotype: Andrews:

t. 50(1795).

E. phylicifolia Salisb.: 364 (1802); Salter: 636 (1951); Dulfer: 36

(1965). Type: Sponte nascentem in Hottentots Holland, 1. Mulder s.n.

( K ! ) . Note: the type has a finely puberulous calyx.

E. hesseana J.C.Wendl. ex Klotzsch: 634 (1835): Guthrie & Bolus:

61 (1905): Dulfer: 38 (1965). Type: Prom. b. sp.. Hesse s.n. (MEL!).

Note: there are two specimens in MEL — one the original from

Wendland’s herbarium determined by Klotzsch. the other a branch bro-

ken off by Klotzsch and retained in B and seen by Bentham. Both were

borrowed by Sonder who was working on the genus for his Flora

capensis when he died, and were never returned to the original herbaria.

His herbarium was sold in two parts with the Ericaceae going to Baron

Ferdinand von Mueller in MEL.

Illustration: Schumann & Kirsten: 45, t. 23 (1992).

Diagnostic features: corolla rose to deep rose, tubular,

1 8-22 mm long, ± glabrous and somewhat sticky; sepals

broadly lanceolate, sparsely puberulous with adaxial ses-

sile glands; anthers included, occasionally manifest

(Figure 8C).

The taxon is confined to the Cape Peninsula and

occurs from the lower slopes of Table Mountain at

Kirstenbosch along the mountains southwards to Cape

Point. It is not sympatric with subsp. abietina.

Paratype material (selection from numerous collections): WEST-

ERN CAPE — 3318 (Cape Town): Kirstenbosch, 150 ft [45 m], (-CD).

27-08-1997. McGrath s.n. (NBG!). 3418 (Simonstown): Muizenberg.

1400 ft (426 m], (-AD). 02-1878. Bolus 4475 (BM!. BOL!. K!. PRE!);

Muizenberg/Kalk Bay Mtns, 300 ft [90 m], (-AD), 03-1880, Bolus

4516 (BM!. BOL!, K!, NBG!); St James, 900 ft [274 m], (-AD). 12-02-

1959, Oliver 50 (NBG!); Kalk Bay Mtn, 800 ft [243 m], (-AD), 12-02-

1959. Oliver 54 (NBG!); Swartkop, Klaver Valley, 1100 ft [335 m|,

(-AD), 14-02-1985. Oliver 8675 (NBG!); Vasco da Gama Peak, (-CB ),

12-12-1959. Oliver 353 (NBG!); Cape Point, (-CB). 10-02-1929, J.B.

Gillett 3465 (NBG!); ibid., 06-1967, Williams 1008 (NBG!).

Bothalia 32.1 (2002)

51

FIGURE 8. — Erica abietina com-

plex. A, subsp. aurantiaca ,

Van Wyk 481. Rocklands,

Cold Bokkeveld; B. subsp.

abietina. Taylor 5777, Table

Mtn. Platteklip; C. subsp.

atrorosea, Oliver 8675, Sim-

onstown: D. subsp. constan-

tiana, Oliver 11335. Constan-

tiaberg; E. subsp. diabolis.

Kirsten 422, Devil’s Peak

Saddle. Scale bars: anthers,

sepals, 2 mm: flowers, 4 mm.

23c. subsp. diabolis E.G.H.Oliv. & I.M.OIiv.,

subsp. nov., corolla breviter obconica, rosea, 11-14 mm

longa, subglabra, sepalis ovatis pilosis glandibus ses-

silibus adaxialibus dignoscenda. Figura 8E.

TYPE. — Western "Cape. 3318 (Cape Town): saddle

between Devil's Peak and Table Mountain, 2100 ft [640

m], (-CD). 25 August 1973. Kirsten 422 (NBG).

E. coccinea L. var. echiiflora sensu Bolus: 60 (1905) non E. eclii-

iflora Andrews. E. abietina var. echiiflora (Bolus) Salter: 643 (1951);

Dulfer: 37 (1965).

Illustration: Schumann & Kirsten: 46. t. 27 (1992).

Diagnostic features: corolla shortly obconical, rose-

pink, 11-14 mm long, subglabrous, subviscid; sepals

ovate, pilose with adaxial sessile glands (Figure 8E).

The subspecies is a very restricted endemic occurring

only on the saddle between Devil’s Peak and Table

Mountain.

Andrews’ E. echiiflora is placed under E. viscaria

subsp. gallorum (34d). His paintings show a ridged

corolla with nipped-in mouth and spreading corolla lobes

and an ovary with longish erect hairs — all typical of that

species.

Vouchers: Bolus 3772 (BM1. BOL!, K!); Kirsten 422 (NBG!).

23d. subsp. constantiana E.G.H.Oliv. & I.M.OIiv .,

subsp. nov., floribus obconicis, roseis, sepalis sparse

puberulis vel glabris, antheris inclusis dignoscenda.

Figura 8D.

TYPE. — Western Cape. 3418 (Simonstown):

Constantiaberg, middle N slopes, 620 m. (-AB), 21-09-

1999, E.G.H. & I. M. Oliver 11335 (NBG).

E. cotiica Lodd.: t. 1179 (1824); Benth.: 664 (1839); Guthrie &

Bolus: 60 (1905); Salter: 649 (1951); Dulfer: 37 (1965). Iconotype:

Lodd.: t. 1179 (1824).

Illustrations: Schumann & Kirsten: 47, t. 28, 29 (1992); Oliver &

Oliver: t. 10 (2000).

Diagnostic features: corolla pale to deeper rose-pink.

obconic, 8-11 mm long, glabrous, subviscid; sepals

lanceolate-ovate, with adaxial sessile glands; anthers

always included, situated about 2/3 way up tube (Figure 8D).

This subspecies is confined to the Cape Peninsula

occurring on the mountains from Constantia Neck to

Chapman’s Peak.

Guthrie & Bolus (1905) noted that there is little to

separate E. conica from subsp diabolis (quoted by them

as E. abietina var. echiiflora). We retain it as a subspecie's

in this complex.

Paratype material (selection from numerous collections): 3318

(Cape Town): Kasteelspoort, 1200 ft [365 m], 08-1877. Bolus 3715

(BOL!, K! ); no precise locality, mtns near Cape Town, 1894, Bolus

7949 (BOL!, NBG!, PRE). 3418 (Simonstown): Constantiaberg, 2000

ft [609 m], (-AB), 22-05-1941, Compton 10842 (NBG!); ibid.. 2900 ft

[883 m], 5-06-1985. Oliver 8742 (NBG!); ibid., 29-08-1963. Stauffer

5063 (K!, NBG!. PRE); Orangekloof, (-AB), 28-08-1954. Esterliuysen

23085 (BOL!); Chapman’s Peak/Noordhoek. 900-1000 ft [2744304

m], (-AB), 1-08-1972, Kirsten 304 (NBG!); Karbonkelberg, (-AB),

16-08-1972. Oliver s.n. (NBG!); Vlakkenberg, (-AB). 13-09-1936.

Salter 6283 (BOL!. K!); Noordhoek Peak, (-AB), 1400 ft [426 m], 20-

07-1967. Taylor 6395 (NBG!. PRE).

23e. subsp. aurantiaca E.G.H.Oliv. & I.M.OIiv.,

subsp. nov., corolla ( 1 0— )25— 30[— 34] mm longa, glabra,

viscida vel non viscida, sepalis longe acuminata ex base

ovata, antheris inclusis ad perexertis. Figura 8A.

TYPE. — Western Cape, 3319 (Worcester): Fransch

Hoek Pass, mtn slopes NE of top of pass, 2500 ft [760

mj, (-CC), February 1966. Chafer in STE30037 (NBG,

holo.; BM. BOL, K, PRE).

E. grandiflora L.f.: 223 (1782); Benth.: 628 (1839); Guthrie &

Bolus: 57 (1905). Type: Caput bonae spei. Thunberg s.n. (UPS).

E. exsurgens Andrews, Coloured engravings of heaths: t. 22 (1796);

Benth.: 627 (1839); Guthrie & Bolus: 57 (1905): Dulfer: 35 (1965); E.

grandiflora var. exsurgens E.G.H.Oliv.: 204 (1967). Iconotype:

Andrews: t. 22 (1796). Note: Andrews' paintings bearing the above

names all seem to be of the Franschhoek/Wemmershoek form with

longer leaves and pale to dark orange flowers, but not pure yellow.

Illustrations: Baker & Oliver: t. 14 (1967); Schumann & Kirsten:

44, t. 14 (1992).

52

Bothalia 32.1 (2002)

Diagnostic features', corolla tubular, ( 1 0— )25— 30[— 34]

mm long, glabrous, sometimes with few hairs on lobes,

orange to orange-red, sticky to non-sticky; sepals long

acuminate from ovate base, with large area of adaxial ses-

sile glands; anthers included to far exserted (Figure 8A).

This is the most widespread, common and variable of

the subspecies, occurring from the hills just northeast

and east of Cape Town to as far inland as the Witteberg

at Matjiesfontein and southeast to near Ashton, but

absent from the Cape Peninsula. The flowers can be pale

to dark orange with zones of yellow below, to complete-

ly deep orange-red.

Paratype material (selection from numerous specimens): WEST-

ERN CAPE. — 3318 (Cape Town): Klein Dassenberg, Kanonkop,

(-DA), 3-05-1986, Fellingham 1077 (NBG!); Kuils River hills, 600 ft

[182 nr], (-DC), 19-06-1972, Oliver 3767a (K!. NBG!. PRE). 3319

(Worcester): Agter Witsenberg, (-AB), 10-03-1959, Barker 8875

(MO!. NBG!): MichelTs Pass, 1 400 m, (-AD), 01-1892, Guthrie 2285

(NBG!): ibid., 1200 ft [365 m], 15-01-1896, Schlechter 9958 (BOL!,

K!, NBG!. PRE!. W); Ceres, Waboomsberg, 1 320 m, (-AD). 12-11-

1989, Oliver 9272 (NBG!); Bokkerivier, (-BD), 9-11-1963,

Middlemost 2241 (NBG!. NY!); Du Toit's Kloof, foot Paarl side,

(-CA), 15-10-1949. Barker 6076 (NBG!, P!): Paarl, Donkerkloof,

(-CC), 12-09-1948, Esterliuysen 14560 (BOL!. NBG!); French Hoek.

(-CC), 1894. Fair sub Bolus 6321 (BOL!. NBG!, PRE); Fransch Hoek

Pass, 920 m. (-CC), 8-01-1970, Oliver & Palser 12 (NBG!. PRE).

3320 (Montagu): Touwsrivier. Pienaarspoort, 1 000 m. (-AA), 5-05-

1994, Oliver 10464 (NBG!): Remhoogte SE of Ashton, 425 m, (-CC),

21-03-1986. Oliver 8814 (NBG!).

23f. subsp. perfoliosa E.G.H.Oliv. & I.M.Oliv.,

subsp. nov., foliis longioribus 20-30(-42) mm, corolla

flava dense puberula non viscida dignoscenda. Figura 9.

TYPE. — Western Cape. 3318 (Cape Town):

Stellenbosch, Jonkershoek Twins, SW slopes, 600 m,

(-DD), 24 May 2001, E.G.H. & I.M.Oliver 11912 (NBG,

holo.; BM, BOL. K, MO. NY, P, PRE, S).

Illustration: Schumann & Kirsten: 44, t. 15 (1992).

Diagnostic features: corolla pure yellow, 20-25 mm

long, densely, finely hairy, non-sticky; sepals broadly

elliptic and long acuminate, with adaxial non-sticky ses-

sile glands; anthers included to manifest; leaves

20— 30( — 42 ) mm long (Figure 9).

The material of this taxon was at one stage classified

as E. exsurgens Andrews. The taxon does not match any

of Andrews’ plates and is a distinct, very localized entity

occurring only in the Jonkershoek Valley near Stellen-

bosch where it is occasional to locally common on the

moister granitic slopes facing south and southwest.

Paratype material : WESTERN CAPE — 3318 (Cape Town): all

from Stellenbosch. Jonkershoek Valley, (-DD), 29-03-1943,

Esterhuysen 8789 (BOL!); 3-04-1949, Esterliuysen 15230 (BOL!); 25-

01-1975, Esterhuysen 33758 (BOL!, K!); 19-05-1950, Parker 4477

(BOL!, K!, NBG!); 2300 ft [700 m], 14-06-1962. Taylor 3399 (BM!.

NBG!. PRE); 3000 ft [914 m], 30-01-1963, Taylor 4628 (NBG!, PRE);

13-04-1961, Van Rensburg 465 (NBG!, PRE).

FIGURE 9. — Erica abietina subsp. perfoliosa, type Oliver 11912. A, flowering branch, x 1 ; B, leaf; C, flower; D, flower cut in half with sepals

removed; E, bract; F, bracteole; G, sepal; FI, sepal, adaxial surface showing sessile glands; 1, anther, side, back and front views; J, anther

variant, side and front views; K, gynoecium seed; M, testa cell. Scale bars: B, E-K. 2 mm; C, D, 4 mm; L, 1 mm; M, 50 pm.

Bothalia 32.1 (2002)

53

23g. subsp. petraea E.G.H.Oliv. & I.M.Oliv.,

subsp. nov., corolla flava dense puberula non viscida,

sepalis sine glandibus sessilibus adaxialibus, habitu

petrensis dignoscenda. Figura 10.

TYPE. — Western Cape. 3319 (Worcester): Porterville

area, Groot Winterhoek Mtns, Kliphuisvlakte, road to

Groot Kliphuis, rock crevices in rocky outcrop. 1 140 m,

(-AA), 23 November 1999, E.G.H. & I.M. Oliver 11440

(NBG. holo.; K, PRE).

Diagnostic features: corolla pure yellow, ± 20 mm

long, densely, finely hairy, non-sticky; sepals narrow-

lanceolate with no adaxial sessile glands; anthers mani-

fest to exserted (Figure 10).

The subspecies is restricted to rocky outcrops on the

mountains above Porterville where the common and

widespread subsp. aurantiaca is not known to occur.

Paratype material: WESTERN CAPE. — 3219 (Wupperthal): Groot

Winterhoek Wilderness area, mtns above and ENE of Porterville, 3200

ft [975 m], (-CC), 23-11-1999, E.G.H. & I.M. Oliver 11449 (BMI, NBG!).

Erica abietina is highly variable in flower size and

colour, indumentum of calyx and corolla, stickiness of

the corolla, degree of inclusion/exsertion of the stamens,

anther shape, and leaf length and habitat preferences. It

used to consist of four separate species that were long

established in the literature — E. abietina. E. phylicifolia,

E. grandiflora and E. conica. There are no clear disjunc-

tions in the ranges of characters that were formerly used

to separate them, but there are some slight discontinuities

which warrant only subspecific recognition.

The whole complex is held together by flowers borne

1 -nate or occasionally 2-nate on vestigial lateral branch-

lets arranged in compact, spike-like synflorescences

towards the ends of main branches. The leaves are all

apiculate and vary from 10-30(^4-2) mm long, the se-

pals vary from very narrow, long-lanceolate to lanceo-

late, to broadly lanceolate, to almost ovate and long-

acuminate and have a flattened, raised area below the

sulcus. They may be villous to pilose or glabrous and

often bear numerous sessile glands over their inner sur-

face, thus rendering the corolla viscid. The sepals all

have sessile glands on the margins and, except for subsp.

petraea , numerous sessile glands adaxially in the middle

zone next to the margins (Figure 9H) — these may be

sticky or non-sticky.

The corolla is mostly long-tubular varying from 18 to

30(-34) mm in length, but in two subspecies, obconical

and only 8-14 mm long. It varies from glabrous to sub-

glabrous to densely and finely hairy and may be red,

orange-red, orange, deep pink, pink or yellow. In the

fresh state these colours are very distinctive, and would

clearly lead one to use them as specific characters, but in

dried material without colour notes, identification is nigh

impossible and one has to resort to a few morphological

characters. The anthers can be distinctly bipartite, even

splitting in the apex of the filament, to having the thecae

closely adpressed — sometimes varying in a single col-

lection. They are mostly attached basally to slightly sub-

basally. The filaments are often sparsely strigulose with

the style sparsely hairy.

The Peninsula taxa tend to form a group having the

apex of the corolla lobes a little more rounded, whereas

the taxa from the mainland have more acute apices to the

corolla lobes.

FIGURE 10. — Erica abietina subsp. petraea, type Oliver 11440. A, flowering branch, x 1; B. stem with leaves removed; C, leaf; D, flower; E,

bract; F, bracteole; G, sepal; H, anther, back, side and front views; I, ovary; J, 1/s ovary. Scale bars: B, C, E-J, 2 mm; D, 4 mm.

54

Bothalia 32,1 (2002)

24. E. conica Lodd. — see E. abietina subsp. constan-

tiana (23d).

25. E. pinea Thunb., Dissertatio botanica de Erica :

23 (1785); Guthrie & Bolus: 60 (1905); Duller: 38

(1965). Type: sine loc., Thunberg s.n. (UPS; BM!, K!).

E. aurea Andrews, Coloured engravings of heaths: t. 76 (1803);

Benth.: 628 (1839). Iconotype: Andrews: t. 76 (1803).

E. argentiflora Andrews, The Heathery: t. 202 (1809). E. pinea var.

argentiflora (Andrews) Bolus: 61 (1905); Dulfer: 38 (1965). Iconotype:

Andrews: t. 202 (1809).

E. aurea Andrews var. viscosissima Benth.: 628 (1839). E. pinea

var. viscosissima (Benth.) Bolus: 61 (1905); Dulfer: 38 (1965). Type:

Fransche Hoek Kloof, Masson s.n. (K!).

Illustrations: Schumann & Kirsten: 47. t. 30. 31 (1992).

Diagnostic features', corolla glabrous, smooth, dry;

sepals with no glands on adaxial surface; ovary globose,

substipitate, glabrous, 6-8-locular.

This species is characterized by the 6-8-locular ovary

which is stipitate and glabrous, the glabrous dry smooth

corolla and no glands on the adaxial surface of sepals.

The flowers may be yellow with a white tip or pure

white. It can easily be mistaken for white-flowered forms

of E. viscaria subsp. longifolia.

Vouchers: Esterhuysen 1769 . 19987 (BOL!, NBG!. PRE); Oliver

1136 (NBG); Schlechter 10254 (BM1, K!, NBG!, PRE); Thompson

3834 (K!, NBG!, PRE).

26. E. hesseana J.C.Wendl. ex Klotzsch — see E. abi-

etina subsp. atrorosea (23b).

27. E. annectens Guthrie & Bolus in Flora capen-

sis 4: 61 (1905); Salter: 637 (1951); Dulfer: 38 (1965).

Type: Cape Div., mountains near Kalk Bay, Guthrie 1002

(BOL!, PRE).

Illustrations: Baker & Oliver: t. 16 (1967); Schumann & Kirsten:

48, t. 32, 33 (1992).

Diagnostic features : flowers 4-nate, terminal on

branches; corolla glabrous, smooth, non sticky; anthers

dorsally attached; ovary globose, glabrous, 8-locular.

This is a rare, highly restricted endemic on the Cape

Peninsula. It is related to E. pinea and E. verticillata (64),

which also have 8-locular ovaries.

Vouchers: Oliver STE30036 (BM!, BOL!, K!, NBG!, PRE!).

28. E. regia Bartl. in Linnaea 7: 630 (1832);

Benth.: 626 (1839); Guthrie & Bolus: 62 (1905); Dulfer:

38 (1965). Type: Zoutcndaelsvalley im Distr. Caledon

[Zoetendalsvlei near Agulhas], Miss Joubert s.n.

(GOET?).

Diagnostic features', corolla glabrous and smooth, tex-

ture stiff and cartilaginous; anthers with vestigial

appendages on apex of filaments; ovary hairy at top with

erect hairs.

With only a slight disjunction in one morphological

character coupled with a clear difference in habitat pref-

erences, we are reducing this complex of three species

and three varieties to one species with two subspecies.

28a. subsp. regia

E. regia var. variegata Bolus in Flora capensis 4: 62 (1905); Dulfer:

38 (1965). Syntypes: Bredasdorp Div., hills near Elim, 300—400 ft

[90-120 m], Pappe 60 (?); ibid.. Bolus 6754 (BM!. BOL!. K!, PRE);

ibid.. Guthrie 2362 (BOL!); ibid.. Will sub MacOwan in Herb. Austr.-

Afr. 1718 (BM!, BOL. K!.P!).

E. regia var. williana Bolus: 62 (1905): Dulfer: 38 (1965).

Syntypes: Bredasdorp Div., hills near Elim. 300—400 ft [90-120 m],

Guthrie 3788 (NBG!); ibid.. Bolus 8448 (BM!, BOL!); ibid., Schlechter

7680 (BM!. BOL!. PRE); ibid. Will sub MacOwan Herb. Austr. Apr.

1719 (BM!. BOL!, K!, P!, SAM!). Lectotype selected here: Will sub

MacOwan Herb. Norm. Austr. Afr. 1719 (BOL!).

E. casta Guthrie & Bolus: 62 (1905); Dulfer: 38 (1965). Syntypes:

Bredasdorp Div., maritime downs and hills near Elim, about 300 ft [90

m], Guthrie 3719 (7): ibid. Bolus 6752 (BM!. BOL!). 6762 (BOL!. K!)

& 8446 (BM!. BOL!, K!). Note: Bolus 6762 = E. vestita (31).

E. casta var. breviflora Guthrie & Bolus: 63 (1905); Dulfer: 38

(1965). Syntypes: Bredasdorp Div., hills near Elim, 300 ft [90 m],

Guthrie 3790 (BOL. SAM!): ibid. Bolus 8449 (BOL!. K!). 8460 (BOL!

K!). Note: Guthrie’s 3790 in SAM = E. axilliflora also Bolus 8449 in

BM. Bolus determined his 8449 in BOL as E. axilliflora and cites it as

such in Flora capensis. He noted on the label of 8460 that it grew with

his 8446 and 8449. Guthrie 3789 is E. axilliflora and cited as such in

Flora capensis.

Illustrations: Baker & Oliver: t. 11 (1967); Schumann & Kirsten:

48, t. 34-37 (1992).

Diagnostic features: corolla nipped in at apex just

below spreading lobes.

Guthrie & Bolus (1905) noted under E. casta — ^close-

ly allied to E. regia we propose the species with some

doubt’. We concur with them.

The variant with variegated flowers (basal white zone,

middle mauve/purple zone and bright red upper zone) is

very striking and well known as the Elim Heath, but has no

morphological differences from the unicoloured forms, some

of which can have a slightly paler whitish basal portion.

The var. williana described by Bolus is a problem in

that it was considered by us as a short-tubed form (flow-

ers 6-12 mm long versus 14-20 mm in the typical sub-

species) with a postulated different pollination syndrome

and worthy of subspecific status. However, we have

recorded it as a few plants growing together with the

orange-red long-tubed typical subspecies and the dark

pink E. axilliflora Bartl. just northwest of Zoetanysberg

(see Note above under synonymy). The flowers varied

from pale to darker purple. At the time we concluded that

the plants were possible hybrids between the two species.

Further ‘populations’ of this form need to be located and

investigated thoroughly before any definite conclusions

can be made as to its identity.

Vouchers: Bolus 8448 (BM!. BOL!. K!, PRE); Oliver 3371 (K!.

NBG!. PRE), 3415 (K!, NBG!. PRE!). STE30138 (K!. NBG! NY!, PRE!).

28b. subsp. mariae (Guthrie & Bolus) E.G.H.Oliv.

& I.M.Oliv ., stat. et comb. nov.

E. mariae Guthrie & Bolus in Flora capensis 4: 63 (1905); Dulfer:

39 ( 1965). Type: Riversdale Div., at Milkwood Fontein, 600 ft [180 m],

Galpin 3565 (BOL!, K!, PRE).

Illustrations: Schumann & Kirsten: 50, t. 38—40 (1992).

Diagnostic features: corolla not nipped in at apex.

This subspecies occurs only on limestone ridges from

southwest of Bredasdorp to Stilbaai. In the Mierkraal

area this taxon grows on the limestone ridge, whereas the

variegated variant of subsp. regia grows on the nearby

lateritic/sandy flats.

Guthrie & Bolus separated the species in this variable

complex on the colour and shape of the corolla and the

Bothalia 32.1 (2002)

55

degree of protrusion of the nose at the base of the

anthers.

The protologue for the species described the type with

‘flores purpurei’. We have not seen a purple-flowered

long-tubed plant in the wild, but have seen fresh material

of the purple-flowered short-tubed form (var. willicina ).

The Elim hills and Zoetanysberg possess a form with

bright orange-red flowers and further south towards the

sea the white-flowered (sometimes tinged pinkish) form,

formerly E. casta , occurs, both on sandy substrates. On

the lateritic flats southeast and east of Elim the form with

variegated flowers occurs. On the limestone ridges from

Mierkraal right through to Stilbaai, there occur plants

with dark red flowers. The colours are very distinctive

and recognizable, but have no major taxonomic signifi-

cance to warrant subspecific ranking. Herbarium materi-

al in which colour has not been recorded cannot be iden-

tified with any certainty.

The corolla shape in E. regia and E. casta is almost

identical, both having a restriction below the spreading

corolla lobes and sometimes a tapering towards the base.

The flowers of E. marine do not have the distinct restric-

tion. There is no clear-cut boundary between the more

extreme anther noses in some material of E. regia to the

almost lack of a nose in E. casta and E. mariae.

Vouchers: Compton 23202 (NBG!. W): Oliver 5788 (NBG!. PRE):

Taylor 8951 (K!. NBG!, PRE).

29. E. casta Guthrie & Bolus — see E. regia subsp.

regia (28a).

30. E. mariae Guthrie & Bolus — see E. regia subsp.

mariae (28b).

31. E. vestita Thunb., Dissertatio botanica de

Erica : 22 (1785); Benth.: 626 (1839): Guthrie & Bolus:

63 (1905): Dulfer: 39 (1965). Type: sine loc., Thunberg

s.n. (UPS. K!, fragm.).

E. vestita var. coccinea Curtis: t. 402 (1798). E. longifolia Bauer

var. amplicata Bolus: 66 (1905); Dulfer: 40 (1965). Type: Curtis: t. 402

(1798).

E. vestita var. fnlgida Andrews: t. 137 (1804); Bolus: 64 (1905);

Dulfer: 39 (1965). Iconotype: Andrews: t. 137 (1804).

E. longifolia Bauer var. maritime Bolus: 66 (1905); Dulfer: 40

(1965). Type: Bredasdorp Div., hills near Cape Agulhas, Schlechter

10556 (BML BOLL PRE!, W).

Illustrations: Baker & Oliver: t. 12 (1967); Schumann & Kirsten:

50, t. 41,42 (1992).

Diagnostic features: long leaves with long petiole (i.e.

versatile leaves); corolla trumpet-shaped, 16-24 mm

long, finely hairy, not ridged: ovary hairy on top. hairs

erect.

The variation in this species occurs in the shape of the

sepals from narrow-lanceolate to those with a broader

base, the corolla indumentum from very finely downy to

pilose and flower colour — white, pink, purple to red. The

northern groups of populations have distinctly geographi-

cal colours — white at Ezeljacht hills, red on the

Riviersonderend Mountains and purplish pink on the

Langeberg near Swellendam. They all have finely downy

corollas, whereas the southern populations from the

Klein River Mountains to Cape Agulhas have more

pilose corollas. The population north of Stanford has

white flowers, whereas there is a mixture of purple-

pink to reddish in the colour of the flowers from the hills

east of Stanford to Agulhas. The type is described as

white-flowered and would probably match the Ezeljacht

material.

Under E. longifolia Bolus commented on the close

similarity between his var. maritima and E. vestita. We

find it inseparable from the various forms of E. vestita

found in the Bredasdoip District and transfer it as a syno-

nym under this species — the smooth corolla (without

ridges) being sparsely pilose and the long petioles mak-

ing the leaves versatile, are the defining characters.

Vouchers: Burchell 7949 <KL PL PRE); Oliver 7536. 8982. 10970

(NBG!); Schlechter 7634 (BM!, KL PL PRE. W).

32. E. nematophvlla Guthrie & Bolus in Flora

capensis 4: 64 (1905); Dulfer: 39 (1965). Syntypes: Ri-

versdale Div., 1000 ft [300 m]; slopes of the Langeberg

Range near Riversdale, Schlechter 1728 (BM!, BOL!,

K!. P!. Z); roadside, Garcia’s Pass, Galpin 3643 (BOL!,

K!. PRE). Lectotype selected here: Galpin 3643 (BOL).

E. filamentosa Andrews var. longiflora Bolus: 65 (1905); Dulfer: 40

(1965). Syntypes: Swellendam Div., mountain ridges along lower part

of River Zondereinde. Zeyher 3171 (BOL!. KL PL PRE, W); Caledon

Div., without collector's name or number. Cape Govt. Herb. (?).

Illustrations: Schumann & Kirsten: 51, t. 43, 44 (1992).

Diagnostic features: corolla tubular, 10-12 mm long,

narrowed slightly towards apex, finely hairy to sub-

glabrous, white or pink.

The long-flowered variety of E. filamentosa has the

closed corolla mouth of this species, and even though the

corolla is pink, it is transferred to this species. There is a

clear disjunction in the distribution range with the pink

variant on the eastern Riviersonderend Range and the

white variant on the Langeberg at Grootvadersbos and

Garcia’s Pass.

Vouchers: Kirsten 677 (BOL!, KL NBG!); Oliver 8633 (NBG!);

Schlechter 1728 (BM!. BOL!, KL PL Z); Zeyher 3171 (BOL!. KL PL

PRE, W).

33. E. filamentosa Andrews , Coloured engravings

of heaths: t. 91 (1804); Benth.: 664 (1839); Guthrie &

Bolus: 65 (1905); Dulfer: 39 (1965). Iconotype: l.c., t. 91

(1804).

Note: Dulfer incorrectly dated Andrews' plates of this species —

Coloured engravings of heaths: t. 91 is dated Feb. 1804 on the full

painting, with the reduced version. The Heathery: t. 22, dated Feb. 1805

on the plate.

Illustration: Schumann & Kirsten: 51, t. 45 ( 1992).

Diagnostic features: corolla broadly obconic, 8-9 mm

long, sparsely and very finely hairy, pink.

The species is confined to the gravelly flats southeast

of Swellendam in the Bontebok National Park. It clearly

has a different pollinator from the pink-flowered form of

E. nematophylla as the anthers are situated lower down

in the very open-mouthed corolla.

E. filamentosa var. longiflora Bolus is transferred to

E. nematophylla (32).

Vouchers: Kirsten 744 (BM!, NBG!); MacOwan 1494 (KL NBG!,

PRE, W).

56

Bothalia 32.1 (2002)

34. E. viscaria L., Dissertatio botanica de Erica : 10

(1770); Andrews: t. 71 (1800); Benth.: 664 (1839); Guth-

rie & Bolus: 210 (1905); Salter: 649 (1950); Dulfer: 102

(1965). Lectotype: Herb. Linn. no. 498.74 , lectotype

selected here by Oliver, Jarvis & Cafferty, in prep.

Diagnostic features', corolla longitudinally ridged with

very short bristle-like hairs or pustules; tube slightly con-

stricted below spreading lobes, not or rarely obconical to

broadly so; ovary obconical, not stipitate, covered with

erect dense fairly long, white hairs (Figure 1 1 ).

This is a very variable species which occurs from the

Cape Peninsula to Franschhoek and to the Bredasdorp

coastal flats. It is probably the most variable species of

Erica with respect to flower colour — white, green, yellow-

ish, pink, purple, red, or combinations of pink with a white

mouth or red with a yellow mouth. Some of these colour

variants are very striking and showy. The corolla varies in

shape and size over a wide range. The indumentum may be

almost absent, finely puberulous, strigose, spiculate,

strigose from pustules, or markedly pustulate. The flowers

may be very viscid, partially so or non-viscid and the plants

may be reseeders or resprouters with long to short leaves.

Colour is an unsuitable character since this is not

retained by older herbarium material and can therefore

not be used as a sole distinguishing feature.

Corolla length we regard as important in distinguish-

ing subspecies, since the open corolla shape of these taxa

coupled with a lower position of the stamens in their

flowers would suggest a different pollination syndrome

from the bird-pollinated long-flowered variants.

34a. subsp. viscaria

E. decora Andrews, Coloured engravings of heaths: t. 162 (1807);

Benth.: 664 (1839); Salter: 649 (1950); Baker & Oliver: 119 (1967). E.

viscaria L. var. decora (Andrews) Bolus: 211 (1905); Dulfer: 103

(1965). Iconotype: Andrews: t. 162 (1807).

Illustrations: Baker & Oliver: t. 114 (1967); Schumann & Kirsten:

169, t. 19 (1992).

Diagnostic features: inflorescence mostly lax; corolla

short (5-9 mm long), urceolate to campanulate-obconic,

soft and more transparent with slightly softer and shorter

hairs; anthers often adhering laterally; thecae with more

prominent basal bulges towards filament (Figure 1 1G).

Bolus (1905) noted the slight difference between

subsp. viscaria and subsp. gallorum by describing E. vis-

caria var. hispida from the Sir Lowry’s Pass area (see

below under 34d). The inflorescence is mostly laxer and

longer and the corolla soft and more transparent in the

typical subspecies.

This subspecies occurs on the Cape Peninsula moun-

tains and also the surrounding flats from where it is now

mostly extinct except in the northern parts. Two variants

occur — the commoner one with narrower, hairy sepals on

the mountains and the rarer with broader, subglabrous

sepals, as represented by the lectotype selected above,

mostly on the flats.

Vouchers: Baker 1228, (BM!, NBG!), 1773 (NBG!. PRE); Bolus

4610 (BOLL K!, NBG!, PRE); Oliver 8683 (NBG!); Steyn 88 (BOL!,

NBG!); Taylor 6404 (NBG!).

34b. subsp. longifolia (Bauer) E.G.H.Oliv. &

I.M.Oliv., comb, et stat. nov.

E. longifolia Bauer, Delineations of exotick plants cultivated in the

royal garden at Kew: t. 4 (1796); Benth.: 625 (1839); Guthrie & Bolus:

65 (1905); Dulfer: 40 (1965). Iconotype: Bauer: t. 4 (1796). Note: the

plate was dated 1 Jan 1793 by the artist/engraver, but published in the

fascicle, part 1, dated April 1796 (see Britten 1899).

E. pinea J.C.Wendl.: l.t. II (1798), non Thunb. (1785). E. longifo-

lia var. contracta Bolus: 66 (1905); Dulfer: 40 (1965). Type:

J.C.Wendl.: t. 11 (1798).

E. longifolia var. squarrosa Bolus: 66 (1905); Dulfer: 41 (1965).

Syntypes: sine loc.. Bolus 8039 (BOL!) & Schlechter 4789 (BM!,

BOL!, NBG!. W).

E. longifolia var. stricta Dulfer: 41 (1965). Type: Caledon Distr.:

Palmiet Rivier, Schlechter 7326 (BM!. BOL!, W).

Illustrations: Baker & Oliver: t. 15 (1967); Schumann & Kirsten:

52, t. 47 (1992).

Diagnostic features: corolla ± 12-20 mm long, tubu-

lar, non-viscid, hairy or pustulate, red, pink, purple,

white, yellowish or green or, in some cases, bicoloured —

pink with a white mouth or red with a yellow mouth;

sepals, bract and bracteoles long, linear-lanceolate

(Figure 1 1 A, B).

This is the most variable subspecies, which it may, on

more detailed population studies coupled with molecular

analyses, be possible to divide into more subspecific

taxa. All plants appear to be single-stemmed reseeders.

There are problems in distinguishing material of the

pink/purple-flowered form in the Bredasdorp District

from similarly coloured forms of E. vestita. It is very pos-

sible that the two species hybridize where they are sym-

patric. The flowers of E. viscaria subsp. longifolia have a

longitudinally ridged corolla and do not possess the

longer, soft hairs of E. vestita.

E. longifolia var. maritima Bolus has been transferred

to E. vestita (31).

Vouchers: Burchell 8039 ( K!, NBG!, P!, W); Oliver 4175 (BM!. K!,

NBG!, PRE!), STE30038 (BOL!. K!. NBG!, PRE!); Schlechter 4789

(BM!. BOL!, NBG!. W); Zeyher 3172 (BOL!, NBG!, P!, PRE, W).

34c. subsp. macrosepala E.G.H.Oliv. & I.M.Oliv .,

stat. et nom. nov.

E. glutinosa Andrews, Coloured engravings of heaths: t. 25 (1798),

non P.J.Bergius (1767). Iconotype: Andrews: t. 25 ( 1798).

E. onosmiflora Salisb.: 363 (1802); Benth.: 626 (1839); Guthrie &

Bolus: 66 ( 1905); Dulfer: 41 (1965). Type: as for E. glutinosa Andrews.

E. viridis Andrews: t. 140 (1805). E. longifolia var. viridis

(Andrews) Bolus: 66 (1905); Dulfer: 41 (1965). Type: Andrews: t. 140

(1805).

Diagnostic features: sepals lanceolate to ovate-lan-

ceolate, often with a broader base and attentuated apex,

corolla ± 15-20 mm long, tubular (Figure 1 ID).

The corolla is yellowish to green, occasionally white

and the plants are mostly resprouters. Some variants may

have very viscid flowers. The subspecies occurs in the

region from Betty’s Bay to Elim.

Vouchers: Compton 6135. 6149 (NBG!); Esterhuysen 4949 (BOL!,

NBG!); Hugo 1541 (NBG!, PRE); /. Kruger 1026 (NBG!); Oliver 3351.

4202 , 7423. 8700 (NBG!).

34d. subsp. gallorum (L.Bolus) E.G.H.Oliv. &

I.M.Oliv ., stat. et comb. nov.

E. gallorum L.Bolus in Annals of Bolus Herbarium 1: 155, t. 1 1A

(1918); Dulfer: 41 (1965). Type: Cape Province: South-Western

Region, French Hoek, fl. Oct., Anon, in BOL14029 (BOL!, K!, PRE).

E. echiiflora Andrews: t. 164 (1805). Iconotype: Andrews: t. 164

(1805).

E. echiiflora Andrews var. purpurea Andrews: t. 260 (71812).

Iconotype: Andrews: t. 260 (1812).

E. viscaria L. var. hispida Bolus: 211 (1906). Syntypes:

Stellenbosch Dist., mountains near Sir Lowry's Pass, 900 ft [274 m],

Bolus 5548 (BOL, K!, NBG!, SAM!); ibid., Guthrie 3524 (BOL).

Bothalia 32.1 (2002)

57

Illustrations: L. Bolus, t. 11A (1918); Schumann & Kirsten: 52, t. 49

(1992).

Diagnostic features', corolla short 5— 10(— 1 2) mm

long, elongate-campanulate to obconic with pink lower

half and white mouth or sometimes pinkish throughout;

hairs short and stout (Figure 1 IF).

The subspecies is confined to the Nuweberg Reserve,

Elgin, on the eastern and southeastern side of the

Hottentots-Holland Mountains where the red-flowered

form of subsp. longifolia occurs at higher altitudes.

Andrews’ E. echiiflora is included as a synonym

based on the ridged corolla with spreading to reflexed

lobes and the ovary with erect, fine hairs, which are all

characteristic of E. viscaria and not of E. abietina.

Vouchers: Anon, in BOL14029 (BOL!. K!, PRE); Kirsten 546

(NBG!); Stokoe 7851 (BOL!, NBG!. PRE).

FIGURE 11. — Erica viscaria complex with flowers, sepal in adaxial view showing zones of sessile glands, anther and ovary. A, B, subsp. longifo-

lia: A, Oliver 11473, Steenbras; B, Gillett 999. Hermanus. C, subsp. pendula, Oliver 7610. Highlands; D. subsp. macrosepala. Porter s.n.,

Hangklip; E, subsp. pustulata, Williams 1552, Hermanus; F, subsp. gallorum, Kirsten 546, Nuweberg; G, subsp. viscaria, Oliver 8683,

Simonstown. Scale bars: anthers, ovaries, sepals, 2 mm; flowers. 4 mm.

58

Bothalia 32,1 (2002)

34e. subsp. pustulata (H. A. Baker) E.G.H.OIiv. &

I.M.Oliv., stat. et comb. nov.

E. pustulata H. A. Baker in Journal of South African Botany 39: 207

(1973). Type: Cape, 3419 (Caledon): above the dams on Mossel River

at Hermanus, (-AC), at about 609 m, 11-10-1971, Williams 1552

(BOL!, holo.: NBG!).

E. patersonii L. Bolus: 134 (1928). non E. patersonia Andrews. E.

longifolia var. breviflora Dulfer: 40 (1965). Type: Caledon Div.,

Hermanus, fl. Sept. 1926, Paterson in BOL18548 (BOL!).

Diagnostic features: corolla markedly pustulate, yel-

low, ± 7 mm long, ovoid-urceolate (Figure HE).

Subsp. pustulata is closely related to the long-tubed

form of subsp. longifolia with yellowish to green flowers

from the same locality.

Vouchers: Paterson in BOL18548 (BOL!, K!); Williams 1552

(BOL!, NBG!).

34f. subsp. pendula E.G.H.OIiv. & I.M.Oliv .,

subsp. nov., floribus pendulis albis non viscidis tubo

corollae sub ore constricto ± 12-18 mm longo dignos-

cenda. Figura 1 1C.

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

south of Highlands Forest Reserve, (-BC), 550 ft [150

m], 26 February 1980, Oliver 7610 (NBG).

Illustration: Schumann & Kirsten: 52, t. 50 (1992) sub E. onosmi-

flora.

Diagnostic features: flowers pendulous (not spreading

to semi-erect); corolla ± 12-18 mm long, tubular, white,

sometimes tinged pink, non-sticky, with short hairs only

along longitudinal ridges and around base; plants sparse-

ly branched and up to 1.5 m tall, with relatively short

grey-green leaves (Figure 11C).

This subspecies is confined to the eastern end of the

Paardeberg Range west of Bot River Village.

A few collections from the region of Shaw’s Mountain

(e.g. Oliver 8022) have semi-pendulous flowers with the

corolla red and a yellow mouth, but sticky and strigose.

These could possibly be included under this subspecies,

but further investigations need to be done to assess their

identity within this highly variable species.

Paratype material : WESTERN CAPE. — 3419 (Caledon):

Arieskraal, (-AA), 17-11-1944, Compton 9021 (NBG!); ibid., 255 ft

[78m], 13-02-1993, Rhode & Boucher 22 (NBG!); Paardeberg

/Highlands, 800 ft [240 m], (-AC), 25-02-1970, Boucher 1151 (K!,

NBG!); ibid., 1000 ft, 22-03-1971, F.J. Kruger 1176 (NBG!); ibid., 550

ft [167 m], 12-01-1970, Oliver & Falser 67. 69 (NBG!); ibid., 150 m,

3-1985, Schumann 307 (NBG!); ibid., 3-1949, Stokoe s.n. (SAM!);

Houtech, SW of Houhoek, 1700 ft [578 m], (-AC), 17-09-1987, Oliver

9021 (NBG!); Shaw’s Mtn, above Langhuis, 300 m, (-AD), 2-08-1983,

Oliver 8022 (NBG!). Without precise locality (3418BB/3419AA):

Palmiet/Grabouw, 800 ft [240 m], 20-07-1895, Guthrie 4971 (NBG!);

ibid., foothills, 10-1948, Stokoe s.n. (SAM!); road from Sir Lowry’s

Pass to Houwhoek, 6-04-1892, Guthrie 2294 (NBG!).

The var. amplicata of E. longifolia described by Bolus

(1905: 66) and upheld by Dulfer (1965: 40), was based

on a coloured painting of E. vestita var. coccinea Curtis:

t. 402 (1798). This plate does not depict any long-tubed

subspecies of E. viscaria and is clearly the red-coloured

form of E. vestita (31).

34.1. E. gallorum L. Bolus — see E. viscaria subsp.

gallorum (34d).

34.2. E. petrusiana E.G.H.OIiv. & I.M.Oliv., sp.

nov., Ericae viscariae L. affinis sed corolla breve

infundibuliformi-obconica, ± 8.5 x 5.0 mm, porcis longi-

tudinalibus, subglabra, subviscida, obscure flava, lobis

erectis et Ericae latiflorae L. Bolus sed lobis corollae bre-

vioribus 1.2 mm longis, coloris corollae flava non pur-

purea distinguitur. Figura 12.

TYPE. — Western Cape, 3418 (Simonstown):

Kogelberg area, between Steenbras River and Kogelberg,

± 1000 ft [± 300 m], (-BB), 16 March 1969, Esterhuysen

32128 (BOL, holo.; NBG, K, PRE).

Erect low woody shrubs. Branches: numerous main

branches ± 30-50 mm long with continuous apical

growth, secondary branches few, very reduced, bearing a

subterminal flower; internodes ± 1 mm long; stems cov-

ered with very short, dense, stiff, spreading hairs and

occasional short-stalked glands. Leaves 4-6-nate, 8-12

mm long, erect, incurved, shortly acute, finely hairy to

FIGURE 12. — Erica petrusiana. type Esterhuysen 32128. A, flowering branch, x I; B. leaf; C, flower; D, flower opened laterally; E, bract; F,

bracteole: G, sepal, abaxial view (left), adaxial view (right); H, stamen, side, front and back views; I. gynoecium. Scale bars: B, E-I, 2 mm;

C, D, 4 mm.

Bothalia 32,1 (2002)

59

glabrous on both sides, sulcus narrow, closed at base;

petiole ± 18 mm long, puberulous. Inflorescence : 1 flower

subterminal on very short, secondary branches at each

node, all aggregated into a spike-like synflorescence, ±

15-20 mm long, towards ends of main branches, flowers

suberect to spreading; pedicel ± 6 mm long, indumentum

like stem; bract partially recaulescent ± V4 way up pedi-

cel, lanceolate, ± 1 mm long, glabrous, ciliate with ses-

sile glands, esulcate; bracteoles about '/2 way up pedicel,

linear-lanceolate, ± 3 mm long, finely hairy and ciliate

with sessile glands, green, sulcate for V3 their length.

Calyx 4-partite; segments adpressed to corolla, lanceo-

late. ± 4.0 x 0.8 mm, abaxially finely hairy, adaxially

with a basal, central zone of sessile sticky glands, sulcate

for 3/4 their length. Corolla 4-lobed, funnel-shaped-

obconic, widest at mouth, longitudinally ridged, sparsely

strigulose, dull yellow; lobes ± 1.2 x 3.5 mm, erect, mar-

gins entire. Stamens 8, free, included just below mouth;

filaments straight, glabrous, with vestigial appendages

just below thecae; anthers bilobed. oblong in adaxial

view, thecae oblong in lateral view, ± 1.5 x 0.5 mm,

smooth, pore V2 length of theca; pollen in tetrads. Ovary

4-locular, broadly obovoid. ± 1.8 x 1.8 mm, emarginate,

densely hairy with thick erect hairs, with basal nectaries;

style exserted, glabrous; stigma capitate. Figure 12.

Diagnostic features', corolla shortly funnel-

shaped/obconic with an open mouth, ± 8. 5-5.0 mm long,

longitudinally ridged, dull yellow, almost glabrous (only

a few minute stiff hairs) with no pustules, slightly sticky,

not nipped in below mouth, lobes erect, not spreading

(Figure 12).

Erica petrusiana is closely related to the short-tubed

subspecies of E. viscaria, but differs in the shape of the

corolla with its open mouth. It shares this character with

E. latiflora L. Bolus (303.1) which has an even more

open corolla with longer lobes. This latter relationship

points to the problem in the genus of the long-tubed and

short-tubed species being closely related, but placed very

far apart in the current sectional system.

The name is derived from the generic name of the

fish, Petrus rupestris, the ‘red steenbras' (red rock-bream),

after the locality, the Steenbras area which is the type and

only known locality. Esterhuysen records the plants as

very local, but common, in stony shaly soil and also on

sandstone ridge. She also records ‘corolla was yellow,

not very bright, but certainly yellow, slightly sticky’.

35. E. onosmiflora Salisb. — see E. viscaria subsp.

macrosepala (34c).

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INDEX TO SPECIES

Erica L., 37

abietina L., 50

subsp. abietina, 50

subsp. atrorosea E.G.H.Oliv. & l.M.Oliv., subsp. nov., 50. 54

subsp. aurantiaca E.G.H.Oliv. & l.M.Oliv., subsp. nov., 51

subsp. constantiana E.G.H.Oliv. & l.M.Oliv., subsp. nov., 51, 54

subsp. diabolis E.G.H.Oliv. & l.M.Oliv, subsp. nov., 51

subsp. perfoliosa E.G.H.Oliv & l.M.Oliv., subsp. nov., 52

subsp. petraea E.G.H.Oliv. & l.M.Oliv., subsp. nov., 53

var. echiiflora (Bolus) Salter, 51

annectens Guthrie & Bolus, 54

argentiflora Andrews, 54

aurea Andrews, 54

var. viscosissima Benth., 54

banksii (banksia) Andrews, 43

subsp. banksii, 43

subsp. comptonii ( Salter ) E.G.H.Oliv. & l.M.Oliv, stat. et comb,

nov., 44

subsp. purpurea ( Andrews ) E.G.H.Oliv. & l.M.Oliv., stat. et comb,

nov., 43

var. purpurea Andrews, 43

var. purpurea (Andrews) Dulfer, 43

baueri (baueria) Andrews, 46

subsp. baueri, 46

subsp. gouriquae E.G.H.Oliv. & l.M.Oliv., subsp. nov., 46

beatricis Compton. 45

bowieana Lodd., 46

breviflora Dulfer, 41. 43

broadleyana Andrews, 46

casta Guthrie & Bolus, 54, 55

var. breviflora Guthrie & Bolus, 54

cerviciflora Salisb., 45

clavaeflora Salisb., 46

coccinea L., 38

sensu P.J.Bergius, 50

subsp. coccinea, 38

subsp. uniflora E.G.H.Oliv & l.M.Oliv, subsp. nov., 39

var. echiiflora sensu Bolus, 51

var. inflata H. A. Baker, 40

var. intermedia (Klotzsch ex Benth.) Dulfer, 40

var. pubescens (Bolus) Dulfer, 38

var. willdenovii (Bolus) H. A. Baker, 40

comptonii Salter, 44

conica Lodd., 51, 54

decora Andrews, 56

echiiflora Andrews, 51, 56

var. purpurea Andrews, 56

exsurgens Andrews, 48, 51

filamentosa Andrews, 55

var. longiflora Bolus, 55

filipendula Benth., 46

subsp. filipendula, 46

subsp. parva E.G.H.Oliv. & l.M.Oliv., subsp. nov., 47

var. major Bolus, 48

var. minor Bolus, 47

follicularis Salisb., 40

var. inflata (H. A. Baker) Dulfer, 40

var. melastoma (Andrews) Dulfer, 40

gallorum L. Bolus, 56, 58

gilva J.C.Wcndl., 45, 46

globulifera Dulfer, 47

glutinosa Andrews, 56

grandiflora L.f., 48, 51

var. exsurgens E.G.H.Oliv., 51

hesseana J.C.Wendl. ex Klotzsch, 50, 54

hibbertii (hibbertia) Andrews, 48

intermedia Klotzsch ex Benth., 40

subsp. albiflora E.G.H.Oliv & l.M.Oliv., subsp. nov., 41

subsp. intermedia, 40

leucosiphon L.Bolus, 44

lineata Benth., 41, 43

longifolia Bauer, 56

var. amplicata Bolus, 55

var. breviflora Dulfer, 58

var. contractu Bolus, 56

var. maritima Bolus, 55

var. squarrosa Bolus, 56

var. stricta Dulfer, 56

var. viridis Bolus, 56

longisepala Guthrie & Bolus, 48

mammosa L., 45

mariae Guthrie & Bolus, 54, 55

melastoma Andrews, 40

subsp. melastoma, 40, 41

subsp. minor E.G.H.Oliv. & l.M.Oliv., subsp. nov., 40

monadelphia Andrews, 41, 43

nematophylla Guthrie & Bolus, 55

nevillei L.Bolus, 50

onosmiflora Salisb., 56, 59

patersonia Andrews, 58

patersonii L.Bolus, 58

penduliflora E.G.H.Oliv., 48

penicillata Andrews, 42

penicillata Benth., 43

petiveri L.. 38

var. intermedia (Klotzsch ex Benth.) Bolus, 40

var. melastoma (Andrews) Benth.. 40

var. pubescens Bolus, 38

var. willdenovii Bolus, 40

petrusiana E.G.H.Oliv & l.M.Oliv., sp. nov., 58

phylicifolia Salisb., 50

pinea J.C.Wendl., 56

pinea Thunb., 54

var. argentiflora (Andrews) Bolus, 54

var. viscosissima (Benth.) Bolus, 54

plukenetii L., 41

subsp. bredensis E.G.H.Oliv. <& l.M.Oliv, subsp. nov., 41

subsp. breviflora ( Dulfer ) E.G.H.Oliv. & l.M.Oliv., stat. et comb,

nov., 43

subsp. lineata {Benth.) E.G.H.Oliv. & l.M.Oliv., stat. et comb, nov., 43

subsp. penicellata (Andrews) E.G.H.Oliv. & l.M.Oliv., stat. et

comb, nov., 42

subsp. plukenetii, 41

var. bicarinata Bolus, 42

var. brachysepala Barth, 41

var. brevifolia Bolus, 41

var. densa Barth, 41

porteri Compton, 49, 50

primulina Bolus, 44

purpurea Andrews, 50

pustulate t H. A. Baker, 58

quadrisulcata L.Bolus, 50

regia Barth, 54

subsp. mariae (Guthrie & Bolus) E.G.H.Oliv. & l.M.Oliv., stat. et

comb, nov., 54, 55

subsp. regia, 54, 55

var. variegata Bolus, 54

var. williana Bolus, 54

revolvens Barth, 42

scariosci Thunb., 43

sceptriformis Salisb., 46

sebema Donn, 38

Bothalia 32,1 (2002)

61

sessiliflora L.f , 46

var. clavaeflora (Salisb.) Bolus, 46

var. oblanceolata Bolus, 46

var. sceptrifonnis (Salisb.) Bolus, 46

sphenanthera Tausch, 45

tenax L. Bolus, 48, 49

thomae L.Bolus, 49

var. brevisepala L.Bolus, 49

vestiflua Salisb., 40, 41

vestita Tluinb., 55

var. coccinea Curtis, 55

var .fiilgida Andrews, 55

viridiflora Andrews, 44

subsp. primulina (Bolus) E.G.H.Oliv. & I.M.Oliv.,

subsp. redacta E.G.H.Oliv. & I.M.Oliv., subsp. nov.. 45

subsp. viridiflora, 44

viridis Andrews, 56

viscaria L., 56

subsp. gallorum (L.Bolus) E.G.H.Oliv. & I.M.Oliv., stat. et comb,

nov., 56

subsp. longifolia E.G.H.Oliv. & I.M.Oliv., comb, et stat. nov., 56

subsp. macrosepala E.G.H.Oliv. & I.M.Oliv., stat. et nom. nov., 56

subsp. pendula E.G.H.Oliv. & I.M.Oliv, subsp. nov., 58

subsp. pustulata (H. A. Baker) E.G.H.Oliv. & I.M.Oliv.. stat. et

comb, nov., 58

subsp. viscaria, 56

var. decora (Andrews) Bolus, 56

stat. et comb. nov.. 44 var. hispida Bolus, 56

1

Bothalia 32.1: 63-83 (2002)

Notes on African plants

HYACINTHACEAE

A NEW COMBINATION IN DAUBENYA

The circumscription of the genus Daubenya has

recently undergone substantial modification as a result of

phylogenetic analysis of plastid DNA sequence data

(Goldblatt & Manning 2000; Van der Merwe et al. in

prep.). The genus now includes the previously monotyp-

ic genera Amphisiphon and Androsiphon, as well as the

taxa previously recognized in the genus Neobakeria.

These various genera were distinguished from one

another primarily on the basis of floral differences that

are now understood to be related to different pollination

strategies. Despite the exaggerated effect that these floral

specializations have on the appearance of the plants, the

species accord perfectly in vegetative morphology. The

genus is characterized by the two prostrate or spreading

leaves, glabrous and rather glossy above with impressed

longitudinal striations along the main veins, with sheaths

that split vertically on drying to form a neck of papery

strips. The subcorymbose or racemose inflorescence

develops small or medium-sized bracts subtending white

to yellow or red flowers with a narrow perianth tube. In

a few species the upper bracts are larger than the lower

and form a ± conspicuous sterile coma at the tip of the

inflorescence.

Among the taxa recently transferred to Daubenya

(Goldblatt & Manning 2000) was the species known until

then as Massonia angustifolia L.f. This species was

described by the younger Linnaeus in 1 782, based on the

collections and notes of Thunberg. No specimen was des-

ignated as the holotype and although the description is too

brief to be diagnostic in any way, his account of the

species which appeared in Hortus kewensis (Aiton 1789)

a few years later, is accompanied by a fine engraving

(Figure 1). Thunberg’s own description of the species

appeared after Linnaeus’ under the name Massonia lan-

ceolata Thunb. The latter name was lectotypified by

Jessop (1976), based on Thunberg s.n. ( UPS -

THUNB7990) and the same collection was later also des-

ignated as the lectotype of Massonia angustifolia L.f. by

U. & D. Miiller-Doblies ( 1997).

As understood in recent revisions of the group (Jessop

1976; Miiller-Doblies 1997), Massonia angustifolia was

readily distinguished from other species of Massonia by

its relatively smaller bracts, cucullate tepals and orange

filaments (Figure 2). These small bracts are characteris-

tic of the group of species previously segregated as

Polyxena subgenus Astemma (Baker 1896) and later as

the genus Neobakeria (Schlechter 1924; Phillips 1951).

In contrast, all species of Massonia s. str. have uniform-

ly white or pinkish flowers that are subtended by large,

leafy bracts and the infructescences are invariably sub-

globose on account of the highly condensed, corymbose

inflorescence typical of the genus. In addition, the tepals

in true species of Massonia display a highly characteris-

tic orientation not found in other genera in the family.

Initially reflexed from the base, they then curve outwards

more or less abruptly, forming a sigmoid fold. Even a

cursory look at the engraving of M. angustifolia pub-

lished in Hortus kewensis reveals that the plant depicted

has the large bracts characteristic of true Massonia

species plus a clear indication of a sigmoid curvature in

the tepals. It is clear, therefore, that this illustration does

not coincide with the concept of Massonia angustifolia

as understood by recent authors (Jessop 1976; Miiller-

Doblies & Miiller-Doblies 1997; Goldblatt & Manning

2000).

Among the specimens in the Thunberg herbarium are

only two sheets labelled M. lanceolata , one of them the

lectotype of both M. lanceolata and M. angustifolia. This

sheet ( UPS-THUNB7990 ) bears two complete flowering

FIGURE 1. — Massonia angustifolia

L.f. from Hortus kewensis

(Aiton 1789).

64

Bothalia 32,1 (2002)

specimens plus three separate infructescences lacking

both leaves and bulbs. The other sheet (UPS-

THUNB7989) bears three flowering plants only. The

flowering plants on both sheets match the illustration of

M. angustifolia perfectly, but the fruiting stems on the

lectotype are clearly not from the same species as they

are rather more racemose with small floral bracts. It is

quite clear that both Linnaeus and Thunberg were refer-

ring to the leafing and flowering material in their con-

cept. The fruiting stems must therefore be excluded from

the lectotype material. The type collection of M. angus-

tifolia was made by Thunberg along the Roggeveld

Escarpment in November 1774. All species of Massonia

and Daubenya are winter-growing and winter-flowering,

and at that time of the year the plants would have been in

fruit. Thunberg was accompanied on this journey by the

Scottish plant collector Masson, who had been sent out

to South Africa to collect plants for the Royal Botanic

Gardens at Kew. It was Masson’s collections, received at

Kew in 1775 and later flowered there, that formed the

basis for the engraving of M. angustifolia that appeared

in Hortus kewensis in 1789, and subsequently for a later

FIGURE 2. — Daubenya marginata

(Willd. ex Kunth) J.C. Man-

ning from Hooker’s leones

Plantarum (1888) (as Potyxena

haemanthoides Baker).

illustration in Curtis's Botanical Magazine (Ker Gawler

1804). It is almost certain, therefore, that the fruiting

bulbs collected by the men in November were similarly

flowered in cultivation to provide the flowering material

for the Thunberg herbarium, probably in the garden that

they established in Cape Town for growing their collec-

tions. One can only surmise that Thunberg unwittingly

combined flowering plants of the one species with previ-

ously collected fruiting specimens of a second species.

This raises the question of the true identity of

Massonia angustifolia. There is no doubt that the species

is actually a smooth-leaved form of M. echinata L.f. This

species is common along the Roggeveld Escarpment and

is extremely variable in leaf vestiture, even within popu-

lations. It is characterized in the genus by its narrowly

tubular flowers with relatively short stamens and tepals

with a sigmoid fold (Figure 3). The sole anomaly in this

identification is the extremely long perianth tube depict-

ed in the engraving in Hortus kewensis (Figure 1 ). This

long tube is not, however, matched by the Thunberg

herbarium material and must represent either an abnor-

FIGURE 3. — Massonia echinata L.f.

from The Flowering Plants

of South A frica 1 1 : t. 429 (as

M. bolusiae W.F. Barker).

Bothalia 32.1 (2002)

65

mality resulting from etiolation of the cultivated plants

grown under low light intensity or, more probably, a mis-

representation by the artist who incorrectly identified the

junction between flower and pedicel. M. angustifolia and

M. echinata were both described contemporaneously by

Linnaeus. The latter is a well-known species and M.

angustifolia is therefore best placed into synonomy

under M. echinata. The oldest available name for the

plant currently known as Daubenya angustifolia is either

Massonia marginata Willd. ex Kunth or Massonia rugu-

losa Licht. ex Kunth. Again, both species were described

in the same publication. Enquiries suggest that the

Lichtenstein material is no longer extant (R. Vogt pers.

comm.) but the Willdenow collection is a fine specimen

that coincides with the current concept of the taxon and

is therefore chosen here to represent the species.

The confusion between M. angustifolia and the taxon

now correctly known as D. marginata can probably be

traced to Baker’s treatment in Flora capensis (Baker

1896). An earlier coloured engraving of the species (Ker

Gawler 1804), based on Masson’s original collections,

shows all of the characteristics evident in the engraving in

Hortus kewensis, thereby reaffirming the Linnean con-

cept of the species. Likewise, Baker’s interpretation of M.

angustifolia as having white, fragrant flowers with

reflexed tepals also coincides perfectly with the original

concept of the species. However, the second of the two

specimens cited by him {Zeyher s.n., K, drawing BOL!;

the other is the typical Thunberg material ) represents the

plant described as Massonia zeyheri Kunth, which was

considered by some authors (Jessop 1976) to be conspe-

cific with D. marginata. Both Jessop (1976) and MUller-

Doblies & Miiller-Doblies (1997) were apparently misled

by the fictitiously long tube in the original engraving and

by the misattributed Zeyher collection, into equating the

species with M. marginata and M. zeyheri respectively.

Daubenya marginata {Willd. ex Kunth) J.C. Man-

ning & A.M.van der Merwe, comb. nov.

Massonia marginata Willd. ex Kunth, Enumeratio plantarum 4:

299 (1843). Polyxena marginata (Willd. ex Kunth) Benth. & Hook.f.

ex T.Durand & Schinz: 366 (1893). Neobakeria marginata (Willd. ex

Kunth.) Schltr.: 150 (1924). Type: South Africa. Caput Bona Spei [B-

WILLD6373. holo.-NBG. photo.!).

Massonia rugulosa Licht. ex Kunth: 299 (1843). Polyxena rugu-

losa (Lichtenst. ex Kunth) Baker: 420 (1896). Neobakeria rugulosa

(Licht. ex Kunth) Schltr.: 150 (1924). Type: Caput Bona Spei,

Lichtenstein 224 (B, holo., ?destroyed).

Polyxena haemanthoides Baker: t. 1727 (1888). Neobakeria hae-

manthoides (Baker) Schltr.: 150 (1924). Type: South Africa. Nuweveld

Mountains near Fraserburg. Bolus 5493 (BOL!. lecto., here designated,

G. SAM).

Massonia angustifolia auct. non M. angustifolia L.f. ( = M. echina-

ta L.f)

Massonia echinata L.f: 193 (1782). Type: South

Africa, Crescit juxta margines montis, dicate Bocklands

Berg |= Bokkeveld Mountains). Thunberg s.n. ( UPS-

7992, holo.!).

Massonia angustifolia L.f.: 193 (1782). M. lanceolata Tliunb.: 40

(1794) nom. superfl. Polyxena angustifolia (L.f.) Baker: 419 (1896).

Neobakeria angustifolia (L.f.) Schltr.: 150 (1924). Daubenya angusti-

folia (L.f.) A.M.van der Merwe & J.C. Manning in Goldblatt &

Manning: 713 (2000). Type: South Africa, Crescit in summo monte

Onderste Roggeveldt, 16.11.1774 ( UPS-THUNB7990 excl. fruiting

fragments, holo.!), syn. nov.

ACKNOWLEDGEMENTS

Our thanks to Dr R. Vogt of the Botanische Museum

Berlin-Dahlem, who was extremely helpful in forward-

ing a high-resolution digital image of the type of

Massonia marginata. and to Dee Snijman for her useful

comments on the manuscript.

REFERENCES

AITON. W. 1789. Hortus kewensis. edn 1. Nicol, London.

BAKER. J.G. 1888. Polyxena haemanthoides Baker. Hooker's leones

Plantarum 18: t. 1727.

BAKER. J.G. 1896. Liliaceae. Flora capensis 6,2: 385-528.

DURAND, T. & SCHINZ, H. 1893. Conspectus florae africae, vol. 2.

Jardin Botanique de l'etat, Brussels.

GOLDBLATT, P. & MANNING, J.C. 2000. Cape Plants. A conspectus

of the Cape flora of South Africa. Strelitzia 9.

JESSOP. J.P 1976. Studies in the bulbous Liliaceae in Africa: 6. The

taxonomy of Massonia and allied genera. Journal of South

African Botany 42: 401 — 437.

KER GAWLER. J.B. 1804. Massonia angustifolia. Curtis's Botanical

Magazine 19: t. 736.

KUNTH. K.S. 1843. Enumeratio plantarum 4. Stuttgart & Tubingen.

LINNAEUS, C. fil. 1782 ('1781')- Supplementarum plantarum.

Orphantrophius, Braunschweig.

MULLER-DOBLIES. U. & D. 1997. A partial revision of the tribe

Massonieae (Hyacinthaceae). Feddes Repertorium 108: 49-96.

PHILLIPS. E. P. 1951. The genera of South African flowering plants.

Memoirs of the Botanical Sun’ey of South Africa No. 25.

SCHLECHTER. R. 1924. Drei neue Gattungen der Liliaceen aus

Siidafrika. Notizblatt des Botanischsen Gartens und Museums

z u Berlin-Dahlem 9: 145-151.

THUNBERG, C.P 1794. Prodromus plantarum capensium. Edman, Uppsala.

VAN DER MERWE, A , BELLSTEDT. D.U., MARAIS, E M. &

HARLEY, E.H. in prep. Phylogenetic relationships in subtribe

Massoniinae of tribe Massoniae (Hyacinthaceae) based on non-

coding trnL-trnF chloroplast sequences.

J.C. MANNING* and A.M. VAN DER MERWE**

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

7735 Claremont, Cape Town.

** Department of Botany. University of Stellenbosch, Private Bag XL

7602 Matieland, Stellenbosch.

MS received: 2001-02-12.

THYMELAEACEAE

SYSTEMATICS OF PASSERINA TRUNCATA AND A NEW SUBSPECIES MONTICOLA

INTRODUCTION Lachnaea conglomerata L. (Rhamnaceae) (Cafferty &

Beyers 1999; Brummitt 2000) has necessitated nomen-

Most of the Passerina L. species are endemic to the clatural changes to P. glomerata Thunb., a taxon also

Cape Floristic Region. The rejection of the name recognized in the now outdated revision of Thoday

66

Bothalia 32,1 (2002)

(1924). In the present paper the name P. glomerata is

rejected and Passerina rigida Wikstr. var. truncata

Meisn. is raised to the species level. Based on leaf anato-

my (Bredenkamp & Van Wyk 1999, 2000, 2001a) as well

as vegetative and floral morphology (Bredenkamp & Van

Wyk 2001b), a new subspecies of P. truncata is described.

MATERIALS AND METHODS

Material from the following herbaria was studied: B,

BM, BOL, BREM, C, G, GRA, HAL, K, LINN, M,

MEL, MO, NBG, NCY, P, PR, PRC, PRE, S, SAM, SBT,

SRGH, TCD, UPS, W and WU. Lor the anatomical

study, both fresh and rehydrated leaves were used. Lor

Passerina truncata subsp. truncata, vouchers of

Bredenkamp 973, 977, 984, 985, 988, 994, and 1002

were used; for P. truncata subsp. monticola, Stokoe 8040

and Esterhuysen 28587 were used.

Light microscope ( LM ) was used in general leaf

anatomical studies. Methods used to prepare transverse

sections and cuticle preparations are described by

Bredenkamp & Van Wyk (2000).

Scanning electron microscope (SEM) was used to

study the leaf surface (including epicuticular waxes) and

to verify the structure of the cuticular membrane

(Bredenkamp & Van Wyk 2000).

Transmission electron microscope (TEM) was used to

study the structure of the mucilaginous epidermal cell

walls (Bredenkamp & Van Wyk 1999).

1. Passerina truncata (Meisn.) Bredenkamp &

A.E.van Wyk, stat. nov. Type: Western Cape, near

Tulbagh Waterfall, April 1865, Zeyher 43 (K ! , lecto.,

here designated; MEL!, MO!, NBG!, S!, W!).

Passerina rigida Wikstr. var. truncata Meisn. in A.D.C., Prodromus

systematis naturalis 14 563 (1857); C.H. Wright; 13 (1915); Thoday:

153 (1924).

Passerina rigida Wikstr. var. tetragona Meisn.: 563 (1857) pro

parte quoad specimen Drege 2971\ C.H. Wright: 13 (1915); Thoday:

153 (1924). Type: Western Cape, circa Ezelsbank Drege 2971 (P!,

lecto., here designated; K!).

Lachnaea conglomerata L.: 560 (1753); L.: 374 (1784); Willd.: 434

(1799); Wikstr.: 323 (1818); Thunb.: 374 (1825); Meisn.: 562 (1857);

Cafferty & Beyers: 171, 172(1999); Brummitt: 805 (2000) nomen reji-

ciendum. Passerina conglomerata Thunb. fide Wikstr.: 322 (1818)

nom. illeg.

Passerina glomerata sphalm. quoad L. conglomerata L. sensu

Thunb.: 75 (1794); Willd.: 434 (1799); Wikstr.: 323 (1818); Thunb: 374

(1825); Meisn.: 562 (1857); Cafferty & Beyers: 171, 172 (1999).

Lachnaea glomerata sphalm. quoad L. conglomerata L. sensu Thunb.:

75 (1794). Passerina ericoides sensu Thunb.: 374 (1825) non L.: 236

( 1767, 1771 ). Passerina glomerata sensu Thoday: 153 (1924) nom. illeg.

Shrubs, shrublets or low stunted shrublets (0.2—)

0.4 — 1 .5(— 2.0) m high. Stems branching from base to grow-

ing points; branchlets ascending, straight or arcuate (subsp.

monticola), greyish brown, younger branchlets whitish

tomentose, older branchlets with whitish scales, remains of

indumentum and scales flaking off with cork on older

branchlets; cork fissured between leaf scars or tesselate,

scaberulent. Leaves sessile, decussate, imbricate on young

branchlets, diverging at 15°^15°, four-ranked, cymbiform;

lamina inversely ericoid, narrowly oblong to oblong in out-

line, length x depth ( 1 .7— )2.0— 3 .0(— 4.0) x (0.6-)0. 8-1.0

(—1.7) mm, becoming keeled in upper part; adaxial surface

concave, tomentose; abaxial surface laterally compressed

and glabrous; apex truncate or rounded (subsp. monticola)',

base dilated; margins involute. Inflorescences comprising

polytelic synflorescences, spikes reduced, resembling ter-

minal subcapitulate inflorescences, each characterized by

two terminal leaves with axillary blind-ending rudimenta-

ry flowers, enveloping minute growing point; proliferating

growth less common; subcapitulate inflorescences ± ellip-

soid, pale green or straw-coloured, 4— 8-flowered. Bracts

enveloping flowers and fruits, accrescent after anthesis,

becoming more coriaceous and rounded at fruit set, decus-

sate, imbricate, sessile, appressed or ascending, obovate to

widely obovate in outline, length x depth (2.5— )2.9— 4.0 x

1 ,4(— 2.4) mm, adaxially concave, villous, abaxially con-

vex, glabrous, coriaceous to thickly coriaceous, ribbed

(subsp. monticola) or ribbed and reticulately veined on

both sides of main vein, concolorous, pale green or greyish

green, senescing to yellowish brown; base cuneate; main

vein extending into obtuse to acute apex; margins involute.

Flowers actinomorphic, bisexual, hypogynous. Floral

envelope 4.9-6.4 mm long, comprising a hypanthium

(fused calyx and androecium) and calyx lobes, papyra-

ceous and yellow-pink during pollination, dehydrated after

shedding of pollen, turning red to brown. Pedicel very

short or absent. Receptacle very short. Disc absent.

Hypanthium a membranous cylindric tube, glabrous at

ovary or apex of ovary tomentose, neck tomentose, 0.7-0.8

mm long, abscission tissue and articulation plane absent.

Calyx petaloid; lobes 4, imbricate in bud, flexed in flower;

outer lobes concave ovate or cymbiform, adaxially sparse-

ly pubescent or apex tomentose, abaxially glabrous; inner

lobes oblong to elliptic or widely obovate, adaxially

sparsely pubescent or tomentose, abaxially glabrous.

Corolla absent. Petaloid scales absent. Androecium dimor-

phic, diplostemonous, arising from hypanthium at separa-

tion point of calyx lobes; filaments of antipetalous whorl

0.9-1. 2 mm and those of antisepalous whorl ± 1.4 mm

long; anthers (0.8— )0.9 x 0.4 mm, sub-basifixed, 2-thecous

and 4-locular. Ovary superior (2.4-)2.5 x 0.9 mm, uniloc-

ular, placentation parietal; ovule 1, pendulous, laterally

attached near top of ovary. Style situated laterally from top

of ovary, maintaining lateral position in hypanthium neck,

reaching beyond hypanthium rim. Stigma ± globose, mop-

like or penicillate. Fruit an achene with pericarp membra-

nous and dry, 2.5 x 1.2 mm, enveloped by persistent, loose-

ly arranged hypanthium, the latter fragmenting over widest

circumference of fruit, resulting in shedding of fragmented

hypanthium, calyx lobes and androecium. Seed broadly

fusiform with outgrowths at both micropylar and funicular

ends; tegmen black and shiny, often with white spots;

endosperm formation nuclear, but later becoming cellular

throughout. Conservation status: Not Threatened (IUCN

Species Survival Commission 1994).

Key to subspecies

la Shrubs or shrublets from various habitats; stems branching

from base, straight; branchlets straight; leaves diverging

at ± 15°, narrowly oblong to oblong, apex truncate,

keeled, appearing humped on the back; bracts appressed.

Bothalia 32.1 (2002)

67

thickly coriaceous, ribbed and reticulately veined on

each side of main vein, greyish green, senescing to yel-

lowish brown: main vascular bundle variously arranged.

types B2, 3, 5 (Bredenkamp & Van Wyk 2001a)

la. subsp. truncata

lb Shrubs or low shrublets from mountain habitats; stems often

stunted; branchlets lax, arcuate; leaves diverging at ±

45°, narrowly oblong, apex truncate to rounded and

keeled; bracts ascending, coriaceous, ribbed on each

side of main vein, pale green; main vascular bundle

bordering on abaxial epidermis, type B6 . . lb. subsp. monticola

Leaf anatomy

Leaf inverse-dorsiventral, epistomatic. Outline in

transverse section (t/s) transversely elliptic. Lamina

width (570— )740— 830(— 980 ) pm, midrib in t/s (290-)

330— +60(— 500) pm. Adaxial epidermis uniseriate, sto-

mata anomocytic, cuticular membrane (CM) ± 2 pm

thick; periclinal cell diam. 15 pm, anticlinal cell diam.

12.5 pm. Abaxial epidermis in surface view, cells

arranged randomly, isodiametric or slightly oblong, pen-

tagonal to heptagonal, (30-)40-55 x 30— 10 pm; CM

smooth or with several domes per cell, wax platelets

scarce, square to oblong, raised 30°-90°, rounded to

sharp edges; CM in t/s (20-)30— f0(— 50 ) pm thick, peri-

clinal cell diam. (20-)30-45 pm, anticlinal cell diam.

(25-)30-60 pm, epidermal cells large, with thick inner

periclinal cell walls, becoming mucilaginous and con-

taining abundant amounts of tanniniferous compounds;

outer periclinal wall domed. MesophylL palisade

parenchyma in 2 or 3 layers of elongated cells with abun-

dant chloroplasts and tanniniferous compounds, density

3-5 cells per 50 pm; spongy parenchyma of irregular

cells ± densely arranged with largest intercellular spaces

close to vascular bundles. Main vascular bundle central-

ly arranged (type B2) (Bredenkamp & Van Wyk 2001a),

touching palisade parenchyma abaxially (type B3), or

with palisade parenchyma abaxial of main vascular bun-

dle specializing into collenchyma, containing ample

amounts of tanniniferous substances (type B5), or bor-

dering on abaxial epidermis (type B6) (subsp. montico-

la), ovate with ample sclerenchyma, or widely to very

widely ovate with sclerenchyma bulging laterally.

Bundle sheath of (2 1— )25— 30(— 32 ) parenchymatous

cells, ± rounded, tanniniferous substances strongly pre-

sent adaxially or specializing into collenchyma abaxially.

Secondary growth present (subsp. truncata). Secondary

vascular bundles 2^1 on each side of main bundle.

Sclerenchymatous hypodermal sheath absent. Figure 4.

Nomenclatural notes

Key to tissues

Spongy parenchyma

Palisade parenchyma

Parenchyma

Xylem

Phloem

Sclerenchyma

Epidermis

?oWo< Cuticular membrane

.02,0-0*0.

Collenchyma

FIGURE 4. — Diagrammatic representation of t/s of leaves. P. trunca-

ta subsp. truncata characterized by leaf structural types B2, 3

& 5, and subsp. monticola by type B6.

9577 (UPS) (Figure 5), bears the inscription ' Passerina

glomerata a (Lin. herb.)’ and was determined by Thoday

as the type of P. glomerata Thunb. However, specimens

504.2 & 504.3 referred to in the Linnean Herbarium

(Savage 1945), are both P. paleacea Wikstr., suggesting

that Thunberg did not have a clear interpretation of

P. glomerata. The specimen Herb. Swartzii s.n. (S) bears

the name Lachnaea conglomerata L. and has been cor-

rectly identified as P. glomerata fide Thunberg (1794)

and Thoday (1924). Unfortunately, neither Thunberg nor

any of the contemporary Swedish botanists were aware

that the illustrated synonym in Breyne on which

Lachnaea conglomerata sensu Linnaeus was based, rep-

resents Phylica stipularis L. (Rhamnaceae). Thunberg

clearly intended to base his new name on the Linnean

concept, referring to 'Linn Syst. 374’ and 'Lin. herb.’.

Lachnaea conglomerata was described by Linnaeus

(1753), with synonyms from Plukenet (1700), Ray

(1704) and Breyne (1678). This name was subsequently

also used by Linnaeus in Sy sterna vegetabilium (1784).

However, there is no material of this taxon in the

Linnean Herbarium (Savage 1945). The only original

element found, is the illustration by Breyne (1678),

determined as Phylica stipularis L. by Cafferty & Beyers

(1999).

Thunberg (1794), in error, made the illegitimate com-

bination Passerina glomerata , citing L. glomerata Linn.

Syst. 374’ in synonymy. The specimen Herb. Thunberg

Willdenow (1799) once again reinstated L. conglom-

erata. sinking P. glomerata Thunb. Wikstrom (1818)

retained P. conglomerata , citing both P. glomerata and

L. conglomerata in synonymy. Wikstrom’s concept is

clearly illustrated by two specimens from the Bergius

Herbarium (SBT) bearing the inscriptions Lachnaea

conglomerata Linn, and Passerina conglomerata Thunb.

Adding to the confusion about the concept of the taxon,

he referred to 'Hautbay’, today Hout Bay, as the locality

for the species, which is the known habitat of both P.

paleacea and P. ericoides L. Thunberg (1825) accepted

Wikstrom’s concept of P. glomerata. citing P. ericoides

in synonymy. This confusion is reflected by many

68

Bothalia 32,1 (2002)

At Sta'

y.*. jx

( 7^n»t<yv 1 /if , <6 ys~

Dntormin.aviti Bdk o-eL*

i 4

I XI

t&pkft WA. ' /--T/vr/u^,, 4J.)

FIGURE 5, — The specimen Herb. Thunberg 9577 (UPS), bearing

inscription 'Passerina glomerata a (Lin. herb.)’ and determined

by Thoday as the type of P. glomerata Thunb.

herbarium specimens, e.g. Thunberg s.n. (S), presently

identified as P. glomerata Thunb., misidentified as P. eri-

coides ( = Lachnaea conglomerata ); the specimen Herb.

Thunberg 9596D (UPS), identified as P. paleacea by

Thoday in 1922, with inscriptions lP. ericoides Linn.’

and 'L. conglomerata (Linn.)’. Recognizing the continu-

ous confusion, Meisner (1857) cited both P. glomerata

and L. conglomerata in synonymy under P. ericoides.

In the interest of nomenclatural stability, the proposal

by Cafferty & Beyers (1999) to reject the name

Lachnaea conglomerata L. was recommended by the

Committee for Spermatophyta (Brummitt 2000). Article

56.1 of the International Code of Botanical Nomen-

clature (Greuter et al. 2000) states that combinations

based on nomina rejicienda should be similarly rejected.

As Thunberg clearly indicated that his concept of

P. glomerata was based on the Linnean concept of L.

conglomerata , the name P. glomerata is also rejected.

Meisner (1857) placed both P. glomerata and L. con-

glomerata in synonymy under P. ericoides. Furthermore,

he described two new varieties under P. rigida , namely

var. tetragona and var. truncata. Both these varieties

included authentic material later cited by Thoday (1924)

in his description of P. glomerata. Based on the specimen

Herb. Thunberg 9577 (UPS) (Figure 5) as the type,

Thoday ( 1924) reinstated P. glomerata Thunb. and placed

Meisner’s two varieties in synonymy, unaware of the fact

that Thunberg (1794) had made the illegitimate combina-

tion P. glomerata , in error. The rejection of L. conglomer-

ata (Cafferty & Beyers 1999; Brummitt 2000) and P.

glomerata , implied the choice of a new name for the exist-

ing species. As all the specimens included in P. rigida var.

truncata are delineated by \foliis ramulo adpressis vel

semipatulatis truncatis' and their distributions coincide

perfectly with the distribution of the previously known

P. glomerata sensu Thoday, the var. truncata was conse-

quently selected and is here raised to the species level. The

var. tetragona included Drege 2971 (K, P) from Ezelsbank

in Western Cape, presently placed in synonymy under

P. truncata , and Drege s.n. from Stormberg in the Eastern

Cape, presently identified as P. montana.

la. subsp. truncata

Diagnostic characters and relationships: the typical

subspecies may easily be distinguished by its decussate,

imbricate, four-ranked leaves, spreading at an angle of

15°. The leaves are abaxially laterally compressed and

glabrous, narrowly oblong to oblong, with a dilated base

and a truncate, keeled apex, appearing humped on the

back. Inflorescences are reduced spikes, terminally sub-

capitulate. Bracts are obovate to widely obovate. The

lamina is villous on the inside and glabrous on the out-

side. It is thickly coriaceous, ribbed and reticulately

veined on each side of the main vein, which extends into

an obtuse apex. Bracts are greyish green in colour,

senescing to yellowish brown. P. truncata could be con-

fused with P. quadrifaria Bredenkamp & A.E.van Wyk

ined., which is characterized by shortened leaf-bearing

branchlets with closely arranged nodes, very short

internodes and leaves spreading at an angle of 45°. The

abaxial surface is convex and tomentose in young leaves

and the leaf apex is rounded.

Etymology : the specific epithet is from the Latin trim-

cat us = ending very abruptly, as if cut straight across. It

describes the truncate apex of the leaves.

Distribution .and habitat: subspecies truncata is con-

fined mainly to the Fynbos (Rebelo 1998) and Succulent

Karoo Biomes (Hoffman 1998) of the Northern and

Western Cape. It occurs in the Northwestern, South-

western, Karoo Mountain and Southeastern centres of

endemism (Goldblatt & Manning 2000) within the Cape

Floral Region. In the Northern Cape, it is distributed from

Steinkopf, through Namaqualand, Nieuwoudtville and

Bothalia 32,1 (2002)

69

HEIGHT ABOVE SEA I

FIGURE 6. — Known distribution of

P. truncata subsp. truncata.

Vanrhynsdorp to Wolfberg. This subspecies is common in

the Western Cape, where it is distributed from

Vanrhynsdorp, the Koue Bokkeveld, Clanwilliam, the

Olifants River Valley, Citrusdal, along the Cederberg

Mountains, to Malmesbury, Ceres, Tulbagh, Matjiesfontein

and Seven Weeks Poort. The most southeasterly known

locality is Baviaanskloof near Patensie ( Bayliss 546).

Subsp. truncata is adapted to a wide range of habitats. It

occurs on low-lying plains as well as on several mountain

ranges, where it is common on level ground, rocky slopes

and along watercourses. Subsp. truncata is one of the most

dominant taxa between Tulbagh and Clanwilliam, where

the size of the shrubs varies from 0.4— 1 ,5(— 2.0) m on flat

areas and plains. They grow prolifically, and during flow-

ering time yellow, yellowish pink and red flowers can often

be seen on the same plant, representing various stages of

maturation. This subspecies is also common in the arid

Karoo environment of the Witteberg Mountains near

Matjiesfontein. where it grows in rocky areas. In extreme-

ly dry areas of Namaqualand, the Ceres-Karoo and on top

of the Skurweberg and Cederberg Mountains, the grow th

form of the subsp. truncata changes to a hardy, skeletal,

less-branched, sclerophyllous shrublet. with yellow, coria-

ceous leaves, closely arranged on the stem, but with each

growing point protected by a pair of conspicuous, coria-

ceous bracts. Figure 6.

lb. subsp. monticola Bredenkamp & A.E.van W\'k,

subsp. nov., a subspecie typica praecipue habitu differt.

Frutices vel fruticuli in habitu montano in fissuris rupi-

um radicantes. Ramuli adscendentes, laxi, arcuati. Folia

ad angulum 45° divergentia. anguste oblonga, carinata,

apice truncato vel rotundato. Bracteae adscendentes,

obovatae ad late obovatae, coriaceae, utrinque costae

costatae, pallide virides.

TYPUS. — Western Cape, 3319 (Worcester): Robert-

son Division, Omklaar, (-DD), 1923, Stokoe 22330

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

Shrubs or low stunted shrublets 0.2-1. 3 m high, often

rooted in rock crevices. Stems often stunted, branching

profusely on younger growth, branchlets ascending,

arcuate; younger branchlets densely whitish tomentose;

cork fissured between yellowish leaf scars. Leaves

diverging at ± 45°, narrowly oblong in outline, length x

depth (2.4— )2.6(— 3.0) x 0.6-1. 7 mm; abaxial surface lat-

erally compressed and glabrous; apex truncate to round-

ed; base dilated. Inflorescences subcapitulate, ± ellip-

soid. pale green. Bracts ascending, obovate to widely

obovate; lamina adaxially concave, villous, abaxially

convex, glabrous, coriaceous, ribbed, pale green, base

cuneate; main vein extending into obtuse to acute apex;

length x depth ( 2.5— )2.9 x 1.5 mm. Flowers : floral enve-

lope ± 4.9 mm long. Hypanthium: apex of ovary tomen-

tose, neck tomentose. ± 0.7 mm long. Calyx: outer lobes

concave ovate, apex adaxially tomentose, abaxially

glabrous; inner lobes widely obovate, adaxially tomen-

tose, abaxially glabrous. Androecium: filaments of

antipetalous whorl ± 1 .2 mm and antisepalous whorl ± 1.4

mm long; anthers 0.9 x 0.4 mm. Ovary 2.5 x 0.9 mm.

Conservation status: Not Threatened (IUCN Species

Survival Commission 1994). Figure 7.

Diagnostic characters and relationships: Passerina

truncata subsp. monticola can easily be distinguished

from the typical subspecies by its growth form. The

plants are often low shrublets rooted in rock crevices on

mountain peaks. The stems are stunted, branching pro-

fusely on younger growth, the branchlets are ascending,

lax and arcuate. The leaves spread at 45°, are narrowly

oblong and the apex is truncate to rounded and keeled.

The bracts retain the characteristic shape of subsp. trun-

cata, but are ascending, smaller and pale green. The

characteristic lax, arcuate branchlets of this subspecies

also distinguish it from P. quadrifaria.

Etymology: the subspecific epithet monticola = moun-

taineer, referring to the mountainous habitat of the subspecies.

70

Bothalia 32,1 (2002)

FIGURE 7. — P. truncata subsp.

monticola , Esterhuysen 28587.

A, flowering branch; B,

Bower clasped by bract; C,

indumentum of hypanthium

tube and glabrous abaxial sur-

faces of calyx lobes; D, fruit

in tomentum of bract; E, leaf;

F, bract. Scale bars: 2 mm.

Distribution and ecology : subspecies monticola is dis-

tributed south of 32°S latitude and between 18°E and 20°

longitude. It occurs in the Northwestern, Southwestern,

and Agulhas Plain centres of endemism (Goldblatt &

Manning 2000) of the Cape Floristic Region. It grows in

mountainous areas and is distributed from Clanwilliam,

along the Cederberg Mountains, southwards along

mountain ranges and summits to Table Mountain,

Helderberg, Kogelberg, the Hex River Mountains and

the Riviersonderend Mountains.

Plants are often found in rock crevices or in damp shel-

tered gullies. They also occur on western, southwestern

and eastern slopes and on shale bands at an altitude of

1 667-2 000 m, on Sneeukop (Goudini). The most com-

mon habitat is on mountain summits, e.g. Sneeuberg,

Slanghoek Pile, Zebra Kop, Jonaskop and Buffelshoek, as

well as on mountain peaks, e.g. Milner, Du Toits, Uitkyk

and Bailey’s Peaks, at altitudes of ± 2 000 m. Figure 8.

ACKNOWLEDGEMENTS

The authors would like to thank Ms Gill Condy for

the line drawings. Dr O.A. Feistner for translating the

diagnosis into Fatin and Ms A. Romanowski for print-

ing the photograph.

SPECIMENS EXAMINED

Acocks 1002 , 1004 , 3045 , 3214 (la) S; 17234 (la) K, PRE; 19397 (la)

K, M, PRE; 23678 (la) PRE. Almborn 475 (la) MO; 539 (la) M. F.

Archer 120 (la) NBG, PRE.

Barker 5751. 9217 . 10286 (la) NBG. Bayliss 577 (la) M, MO, PRE;

591. 1105, BRI-B591 (la) K, MO. PRE; BR1-B546 (la) PRE. Bellamy

2 (la) PRE. Bolus 9507 (la) BOL, K. Boucher 1983 (la) NBG; 2872

(la) K, NBG, PRE; 2880 (la) PRE; 3177 (la) C, K, NBG, PRE.

Bredenkamp 972 , 973. 977. 978. 983. 985, 986. 988-994. 1004-1011

( la) PRE. Bremer 264 ( la) PRE, S. Buchell 387 (la) PRE.

Compton 519 (la) NBG, S; 1404 (lb) NBG; 2800 (la) BOL, NBG;

3327 (la) BOL, K; 3743 (la) NBG; 4894 (la) BOL; 4901 . 5890 (la)

BOL, NBG; 6347 (lb) NBG; 6546. 6902. 9537 (la) NBG; 9970 (la)

MO, NBG; 11701. 11718 . 12237. 15020. 18781 (la) NBG; 19949 (la)

NBG, MO; 22821 (la) S; 24331 (la) NBG, MO.

Dahlgren & Peterson 257 (la) K. Davis s.n. (la) NBG. PRE, SAM.

Drege 2971 (la) K, P; s.n. (la) K.

Ecklon & Zeylier s.n. ( la) BREM, MO, S. Emdon 197 (la) C, K. PRE.

Esterhuysen 1327 (la) BOL; 1744 (lb) BOL; 2273 (la) BOL; 3638

(lb) BOL; 3703 (la) K; 7380. 8416. 9542. 9822 , 9921 (lb) BOL;

10886 (la) BOL; 11046 ( lb) BOL; 11255 (lb) BOL, K; 26859 (la) K;

27451 (lb) BOL; 28555 (lb) BOL, K; 28574. 28587, 30573. 30910

(lb) BOL; 32195 (la) MO. Evrard 9001 (la) MO. PRE, UPS.

Forrester 114, s.n. ( la) NBG.

Galpin 11209. 11210. 11218 ( la) BM, C, PRE, UPS. Gillett 397 ( la)

NBG; 4012 (la) BOL, K, PRE. Alice Godman 527 (la) BM. GoUlblatt

2757 ( la) M, NBG, PRE, S; 2802 (la) C, PRE, M, MO; 7264 (la) MO,

PRE, S. Greuter 21765 (la) PRE. Guthrie s.n. (lb) NBG.

Hanekom 943 (la) NBG. PRE; 1249 (la) K, PRE. Hardey 448. 449

(la) PRE, BM, K, M. Herb Swartzii s.n. (la) S. Herb Thunberg s.n.

(la) LIPS. Hiendhnayr s.n. (la) M. Howes 225 ( I a) K, PRE. Hugo 545.

679 (la) K, PRE. Hugo & Mauve 4 (la) K, MO, NBG, PRE.

Hutchinson 416 (la) BM, GRA. K, PRE; 834 (la) GRA. K. PRE.

Bothalia 32,1 (2002)

71

Johnson 519 (la) NBG. Jordaan 545 ( la) NBG.

Kerfoot K5061 (lb) NBG. Kotze 45 (la) NBG. Kruger 911 (la) NBG,

PRE: 1754 (lb) PRE.

Le Roux 2310 (la) PRE. Levyns 1232, 1335, 1370, 1377, 1808, 1941,

2165, 2382, 2415a, 4108 (la) BOL. Lewis 2730, 4061, 4062 ( la) NBG;

62671 (la) MO; SAM67057, SAM67058 (la) NBG, PRE; Lewis s.n.

(la) NBG. Linscherg s.n. (la) S. Loubser 598 (la) NBG.

Maguire 2025 (la) C, NBG. Marloth 4238 (la) K. PRE; 6626 (lb)

NBG. PRE; 6873, 6957 (la) PRE; 10668 (la) NBG; 10798, 11426,

11686, 12943 (la) PRE; 13006, 13007 (la) NBG. PRE; 13144 (la)

PRE. Merxmuller & Giess 3832 ( la) M. PRE. Metelerkamp 562, 562a

(la) BOL. Meyer 74 (la) NBG. Morley 250 (la) NBG, PRE. Museum

Botanicum Hauniense (la) C.

Nortje 77 (la) NBG.

Pappe s.n. (la) K. Pearson 5709, 6256 ( la) BOL, K; 6765 (la) K; 6773

(la) BOL; 6782 (la) K. Pearson & Pillans 5928 (la) K. Phillips 2057

(la) NBG. Pillans 7298 (la) K; 7298, 7769 ( lb) BOL; 8021 (la) BOL;

9606, 10705 (la) MO; 63799 (la) BOL, MO. PRE. Pocock 770 (la)

NBG, PRE.

Rauh 3553 (la) M. Rourke 961 ( lb) MO. NBG.

Salter 1568 ( la) K. Schlechter 10627 (la) BM. K. MO. PRE. S; 10660

(la) BM, K. PR. PRE. S. Solomon 92 ( la) NBG. Stehle 309 ( lb) NBG.

Stephens 7226. 7264 (la) K. Stirton 10148 (la) PRE. Stokoe 1460 (la)

PRE; 2520 (lb) PRE; 8040 (lb) BOL; 8041 (la) BOL; 8042 (la) BOL.

GRA, NBG. PRE; 8225 (lb) BOL; 17574 (lb) BOL. PRE; 22330 (lb)

K. PRE; 22331 (la) BOL. PRE; 64616 (lb) PRE: s.n. (la) NBG: s.n.

(lb) BOL. NBG. Stokoe & Davis s.n. (la) NBG. Story 4303 (la) K.

GRA. PRE. Strid 37832 (la) C, NBG.

Taylor 2865 (la) NBG, S; 3921 (la) PRE: 4011 (la) NBG; 6145 (lb)

PRE; 6495 (la) NBG. PRE; 8664 (la) K, NBG. PRE. S; 9293 (la)

PRE; 10483, 10618 (la) NBG. PRE. Thoday 213 (la) NBG: 1921 ( la)

BOL. Thompson 1278, 1519 (la) NBG; 2823 (la) K. MO. NBG. PRE.

Thunberg s.n. (la) S. SBT. Tyson 318 (la) NBG; 1452 (la) PRE: 2318

(la) K.

Van Breda 198 (la) PRE. Van Rooyen 2401. 2626 ( la) M. Van Rooven,

Steyn & De Villiers 618 (la) NBG. Van Son TRV36623 ( la) PRE .Van

Wyk 6485 ( la) PRE. Van Zyl 3295 (la) K. MO. PRE.

Wagener 259 ( la) NBG. Whitehead s.n. ( la) TCD.

Zeyher43 ( la) K. NBG. MO. S; s.n. ( la) M. Zietsman & Zietsman 925,

1151 (la) PRE.

REFERENCES

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BREDENKAMP. C.L. & VAN WYK. A.E. 2000. The epidermis in

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BREDENKAMP. C.L. & VAN WYK. A.E. 2001a. Leaf anatomy of the

genus Passerina (Thymelaeaceae): taxonomic and ecological

significance. Bothalia 31: 53-70.

BREDENKAMP, C.L. & VAN WYK, A.E. 2001b. Taxonomic signifi-

cance of inflorescences, floral morphology and anatomy in

Passerina (Thymelaeaceae). Bothalia 31: 213-236.

BREYNE, J. 1678. Exoticarum plantarum centuria. David-Fredericus

Rhetius, Gdansk.

BRUMMITT, R.K. 2000. Report of the Committee for Spermatophyta:

50. Taxon 49: 799-808.

CAFFERTY, S. & BEYERS, J.B. 1999. Proposal to reject the name

Lachnaea congloinerata (Rhamnaceae). Taxon 48: 171, 172.

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

the Cape flora of South Africa. Strelitzia 9: 15-19.

GREUTER, W„ MCNEIL, J, BARRIE, F.R., BURDET, H.M.,

DEMOULIN, V.. F1LGUEIRAS, T.S., NICOLSON, D.H.,

SILVA, PC., SKOG. J.E.. TREHANE. P„ TURLAND, N.J. &

HAWKSWORTH. D.L. 2000. International Code of Botanical

Nomenclature ( Saint Louis Code). Koeltz Scientific Books,

Konigstein.

HOFFMAN. T.M. 1998. Succulent Karoo Biome. In A.B. Low & A.G.

Rebelo, Vegetation of South Africa, Lesotho and Swaziland: 74.

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IUCN. Gland. Switzerland.

LINNAEUS, C. 1753. Species plantarum, edn 1. Salvius, Stockholm.

LINNAEUS, C. 1767. Mantissa plantarum. Cramer, Weinheim.

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Candolle, Prodromus systematis naturalis 14: 493-605.

Masson, Paris.

PLUKENET, L. 1700. Almagesti Botanici Mantissa. Sumptibus

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APOCYNACEAE

A NEW SPECIES OF BRACHYSTELMA FROM EASTERN CAPE. SOUTH AFRICA

Brachystelma cummingii A.P.Dold, sp. nov., B.

tenellum R.A.Dyer affinis base inflata gemmae pentago-

na (non globosa); lobis interioribus coronae rectangulatis

(non anguste linearibus); lobis interioribus coronae non

appendiculatis-dentatis; corpusculo clavato (non rectan-

ulato); pagina interiori loborum trichomis sparsis acuti-

cylindricis, albis adprementibusque, in longitudinem

ordinatis (non pilis longis albisque).

TYPE. — Eastern Cape, 3324 (Steytlerville): Klein-

poort, (-BD), 550 m, 27-01-2001, Bold 4368 (GRA,

holo.).

Perennial herb with tuber. Tuber depressed, spherical,

not exposed. 20-25 mm high, ± 30 mm diam. Stems 1-3

from single central growth point, erect, occasionally

branching above ground level, below 10 mm, 0. 5-1.0 mm

72

Bothalia 32,1 (2002)

simultaneously; pedicels spreading, 15-18 x 0.4 mm,

puberulous, purple. Bract persistent, single, extra axillary

clasping, lanceolate, acute, ladle-shaped, cuspidate,

bluntly spurred at base, 1 x 0.2 mm at base, green becom-

ing dark purple-black. Sepals 5, lanceolate, acute, 1 .6 x

0.6 mm at base, glabrous, reflexed below corolla sinus,

reaching pedicel, persistent, purple. Corolla : in bud,

inflated base pentagonal, 2 x 2.5 mm, cylindrical beak

4-6 x 0.5 mm, apex rounded; at anthesis, (4.5-)

5 .5— 6.0(— 10) x 3. 5-4.0 mm; tube 0.25-0.50 mm, sur-

rounding base of gynostegium, then reflexed up to 1 mm

to sinus; lobes deltoid at base, produced into long attenu-

ate lobes connivent at apices, 5-8 mm from apex to sinus,

1.2 mm broad at base, narrowing abruptly to 0.3 mm,

sinus between lobes deeply incised, forming an open,

pyramidal to ± globose, cage-like structure, inner surface

of lobes with sparse, cylindrical-acute, white, adpressed

trichomes, each attached to a raised papilla, arranged lon-

gitudinally, base dark purple, outer surface paler, upper

surfaces dull green to yellow-green. Gynostegium 1 mm

diam. at base, 1-2 mm high, glabrous, stipe white; outer

corona lobes 0.4 x 0.3 mm at base, erect, slightly spread-

ing, flattened, distal end bilobed, lobes spreading lateral-

ly forming a shallow V, black, tips sparsely furnished with

minute white trichomes; inner corona lobes 5, 0.4 x 0.2

mm at base, flattened, incumbent on and curving inwards

distally to cover anthers, black. Pollinium semi-ovate,

flattened, 0.22 x 0.12 mm, insertion crest along outer

edge narrowly, transparently winged, golden brown; cor-

pusculum clavate, 0. 15 mm long, orange-brown; caudicle

short, broad, orange-brown. Follicles paired, linear ± 60

x 3 mm, broadest in the middle, green, minutely and

densely mauve-speckled, apices forming minute club-

shaped tip. Flowering time : November to December.

Figures 9-11.

Specimens examined

EASTERN CAPE. — 3324 (Steytlerville): De Bordtjie, west of

Kleinpoort, (-BC), Bruyns 6913 (BOL); south of Kleinpoort, (-BD).

Brityns 4914 (BOL); Kleinpoort, (-BD), Camming 7037 (GRA); Dold

4368 (GRA).

Brachystelma cununingii resembles B. tenellum

R. A. Dyer (Dyer 1973, 1980, 1983), but differs morpho-

FIGURE 10. — Brachystelma citmmingii , Dold 4368, pollinariutn.

Scale bar: 100 pm.

FIGURE 9. — Brachystelma citmmingii, Dold 4368. A, flower bud; B,

flower at anthesis; C, D, corona, view from top and side. Scale

bars: 1 mm. Illustrations: A. Dold.

thick at base, 0.4-0. 5 mm thick above, 100-150 mm long,

internodes 4 mm at base to 10-14 mm above, dull purple

at base, new growth green, white puberulous. Leaves ses-

sile, basal 3 or 4 pairs elliptic, 14-20 x 3-5 mm, erect,

often twisted on central axis, margins tinged reddish,

upper 10-15 pairs linear, spreading, simple, (I0-)

1 8— 20(— 22) x 1 .0-1.5 mm, entire, V-shaped in cross sec-

tion, lower keel prominent, upper surface with minute

shiny water cells, dark green, lower surface leathery, yel-

low-green. Flowers single, extra-axillary at nodes, open-

ing successively at every node whilst growing, seldom

Bothalia 32. 1 (2002)

73

FIGURE 1 1. — Brachystelma cummingii, Dold 4368, section of corolla

lobe showing adpressed trichomes. Scale bar: 100 pm.

logically in several ways (Table 1). The recurved corolla

lobe margins of B. cummingii resemble those of B.

gracile E. A. Bruce (Bruce 1949: Dyer 1980, 1983), but

are more pronounced, obscuring all but the tips of the

calyx. The unusual adpressed trichomes on the inner sur-

face of the corolla lobes of B. cummingii are also found

on B. pygmaeum.

B. cummingii is only known from four specimens, one

from De Bordtjie and three from Kleinpoort (Figure 12),

at 550-800 m above sea level and is scattered in open

rocky grassland on Hat surfaces. The vegetation type of

both localities is Central Lower Nama-Karoo (Hoffman

1996). Tubers are well below the surface in shallow

sandy soil (derived from the Peninsula formation of the

Table Mountain geological group) associated with

Aristida adscensionis. Dicoma spinosa , Digitaria eri-

antlia , Eragrostis curvula , Euphorbia jansenvillensis and

Lebeckia pungens. An average annual rainfall (23 years)

of 350 mm has been recorded at Glenconnor, 30 km to

the east of Kleinpoort. In the absence of data, it is expect-

ed that Kleinpoort and De Bordtjie would have a similar

rainfall figure.

TABLE 1. — Morphological differences between Brachystelma cum-

mingii and B. tenellum

FIGURE 12. — Known distribution of Brachystelma cummingii.

The locality of B. cummingii is significant, as the

genus is poorly represented in arid regions (Peckover

1993) compared to areas of higher rainfall. In his revi-

sion of the genus. Dyer (1980) recorded only a single

species in the Karoo. In comparison. Retief & Herman

(1997) record 35 species for the northern provinces of

South Africa, whereas Bruyns (2000) records three

species of Brachystelma for the Cape flora, of which

only one is endemic to that region.

ACKNOWLEDGEMENTS

Rhodes University Joint Research Council is thanked for

financial support. Nancy Lawrence, Albuquerque, New

Mexico, is thanked for the Latin diagnosis. Dr Peter Bruyns

is thanked for his valuable advice on the manuscript.

REFERENCES

BRUCE, E.A. 1949. Brachystelma gracilis. The Flowering Plants of

Africa 27: t. 1077.

BRLTYNS. RV. 2000. Apocynaceae. In P. Goldblatt & J. Manning,

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

Strelitzia 9: 280-290.

DYER. R.A. 1973. Brachystelma tenellum. The Flowering Plants of

Africa 42: t. 1 664.

DYER, R.A. 1980. Asclepiadaceae. Flora of southern Africa 27.4: 31.

DYER, R.A. 1983. Ceropegia, Brachystelma and Riocreuxia in south-

ern Africa. Balkema, Rotterdam.

HOFFMAN. T. 1996. Central Lower Nama-Karoo. In A.B. Low & A.G.

Rebelo, Vegetation of South Africa, Lesotho and Swaziland.

Department of Environmental Affairs and Tourism, Pretoria.

PECKOVER. R. 1993. Taxonomic questions within the genus

Brachystelma : a few examples. Aloe 30: 114, 115.

RETIEF. E. & HERMAN. P.P.J. 1997. Plants of the northern provinces

of South Africa: keys and diagnostic characters. Strelitzia 6:

257-281.

A.P DOLD*

* Selmar Schonland Herbarium, P.O. Box 101, 6140 Grahamstown,

South Africa.

MS. received: 2001-04-23.

74

Bothalia 32,1 (2002)

CAMPANULACEAE

A NEW SPECIES OF MERCIERA FROM WESTERN CAPE, SOUTH AFRICA

INTRODUCTION

Merciera A. DC. is one of five Campanulaceae genera

endemic to Western Cape. It is a genus of four dwarf shrub

species, occurring in sandy, rocky or clayey habitats.

Complex morphological variation render Merciera taxo-

nomically challenging at the species level. Detailed mor-

phometric analysis in Merciera (Cupido 2000) revealed

the existence of a well-defined group of specimens, which

did not match any previously recognized taxon. This

group of specimens was identified as M. brevifolia A. DC.

The first known specimen was collected by Drege during

the mid 1 820s on the Du Toitskloof Mountains and was

considered to be a member of Roella L., a genus closely

related to Merciera. Floral characters of these plants dis-

play a clear interval in the morphological variation pattern

within M. brevifolia. In addition to the distinct morpho-

logical characters, the plants also occupy a distinct geo-

graphical range, west of the Hottentots Holland Mountains

in Western Cape. All the other species in the genus occur

east of these mountains. This discrete group of specimens

is described here as a new species.

Merciera tetraloba C.N. Cupido, sp. nov., ex

affiniate M. brevifolia A. DC. et specierum affinum folia

glabra in pagina abaxiale, calyx 4 lobis et ciliatis mar-

gines, corolla alba, interdum purpureus apice, lobis 4

ovatis, stamina 4 distinguenda.

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

Harmony Flats, Tortoise Nature Reserve, off Disa Road,

(-BB), 17 January 2000, C.N. Cupido 77 (NBG, holo.;

BM, K, MO, NY, PRE).

Subshrubs, hispidulous to hispid. Stems decumbent,

slender, branched, occasionally with groups of branches at

the end of a year’s growth. Leaves alternate, linear, less

than 8 mm long, subulate, entire, scattered, ascending, the

older spreading, sessile, glabrous on abaxial surface, mar-

gins ciliate; axillary cluster of smaller leaves present.

Inflorescence 3-flowered, I terminal, lateral 2 rudimenta-

ry, on highly reduced lateral branches with bract-like

leaves, aggregated into spike-like synflorescences towards

ends of main branches. Flowers sessile, axillary, actino-

morphic. Bract-like leaves 2, succulent, subtending each

of rudimentary flowers, absent in terminal llower, mm

long. Hypanthium obconical, 1-2 mm long, hispid with

uncinate or circinate triehomes. Calyx: lobes 4, 0.6-1. 2

mm long, fused at base to form short tube, hairs often on

hyaline tips and margins. Corolla narrowly funnel-shaped,

white, occasionally with purple tips, or very rarely pale

blue; lube 4—6 mm long; lobes 4, ovate, glabrous or hairy

on back, 2-3 mm long. Stamens 4, free, inserted at base of

corolla tube; filaments llattened, wider and pilose about

the middle, narrower towards apex, 3.0-4. 5 mm long;

anthers linear, basifixed. Ovary inferior, I -locular, con-

taining 4 erect ovules situated at base of ovary; style fili-

form, exsertcd, glabrous, swollen at base, 6-10 mm long;

stigma bifid, glabrescent, bluish purple. Fruit a hispid cap-

sule, crowned with persistent calyx, I -seeded, indehiscent.

Flowering time: November to January, flowering most

profusely in recently burnt veld (Figures 13 & 14).

Diagnostic features: the reduction in the number of calyx

lobes, corolla lobes and stamens make M. tetraloba

unique in the genus. This species differs from the other

two white-flowered species, Merciera leptoloba A. DC.

and M. brevifolia, in tetramerous flowers and calyx lobes

often with marginal hairs. Furthermore, in M. leptoloba

the corolla lobes are linear-lanceolate and the lower

leaves are more than 8 mm long, whereas in M. tetralo-

ba the corolla lobes are broadly ovate and the lower

leaves less than 8 mm long. The other species in the

genus, Merciera tenuifolia (L.f.) A. DC. and Merciera

eckloniana Buek ex Ecklon & Zeyher, differ from M.

tetraloba in having violet-blue flowers and corolla tubes

of more than 7 mm long. The species is closely related to

M. brevifolia from which it differs in floral characters

and geographical distribution. The two species occur

allopatrically. M. brevifolia is confined to mountains in

Bot River, Houwhoek and Caledon, whereas M. tetralo-

ba occurs west of the Hottentots Holland Mountains.

Distribution and habitat: this species is found in Faure,

Gordon’s Bay, Sir Dowry's Pass, Somerset West, Strand,

Dal Josaphat, Du Toitskloof and Stellenbosch (Figure

15) on flats and lower mountain slopes at altitudes

between 30 m and 350 m. It grows in open clayey soil,

often in disturbed habitats.

Farge areas of the habitat of this species in the

Helderberg and Stellenbosch areas have been destroyed

because lower mountain slopes and lowland areas are

sought after for urban development. On the Harmony

FIGURE 13.- — Merciera tetraloba. inflorescences, x 0.75. Photograph

by J.C. Manning.

Bothalia 32,1 (2002)

75

B , 4mm .

C-G 2mm

H i 1 mm

FIGURE 14. — Merciera tetraloba. A, portion of plant, life size: B, flowering branch; C, leaf; D, bract-like leaf with rudimentary flower; E, flower

with two bract-like leaves; F. stamen; G, base of flower longitudinally opened, showing ovary and basal part of style; H. seed. Drawn from

Cupido 77 by Inge Oliver. Scale bars: B. 4 mm; C-G. 2 mm; H, 1 mm.

Flats in Strand, the few existing populations are under

serious threat of extinction.

FIGURE 15. — Known distribution of Merciera tetraloba.

Etymology, the name of this species is derived from the

respective number of lobes in the corolla and calyx (tetra

= four, lobi - lobes).

Specimens examined

WESTERN CAPE.— 3318 (Cape Town): Dal Josaphat, (-DB),

Barker 8865 (NBG), on hills, Tyson 899 (SAM); between Bottelary and

Stellenbosch, (-DD), Bolus BOL98256 (BOL); near roadside

Stellenbosch, Markotter 8639 (NBG); Stellenbosch Mountain, lower

western slopes above the university farm and NE of Brandwacht sub-

urb, Oliver & Oliver 11866 (NBG). 3319 (Worcester): Du Toitskloof,

(-CC), Dr'ege SAM17297 (SAM). 3418 (Simonstown): Sir Lowry’s

Pass, foot of pass, (-BB ), Adamson 4895 ( BOL); Sir Lowry’s Pass, foot

of pass on the Somerset West side. Cupido 117 (NBG); Sir Lowry’s

Pass, Guthrie 2792 (NBG); Faure Hills, Compton 10372 (NBG);

Gordon’s Bay, mountain slopes above, Bayliss 4089 (NBG); flats

between Strand and Gordon’s Bay, Jordaan s.n. (NBG); Somerset

West, Parker 3550 (NBG); Strand, upper Harmony Flats between

Lwandle Township and Strand Foam Factory, Boucher 3447 (NBG);

Strand, Tortoise Nature Reserve, Cupido 75, 77 (NBG).

ACKNOWLEDGEMENTS

I wish to thank the following people for their assis-

tance: Prof. H.P. Linder for supervising the project;

76

Bothalia 32,1 (2002)

Mrs P. Runnalls of the Friends of Heidelberg Nature

Reserve for finding a population of the living plants

for me, after my own futile attempts; Ferozah Conrad

for assistance with the Latin diagnosis; the curator of

the Bolus Herbarium for making their specimens

available for the study; Mrs I. Oliver for doing the line

drawings and Dr J.C. Manning for taking the photo-

graph.

REFERENCE

CUP1DO, C.N. 2000. A re-assessment of the species boundaries in

Merciera A DC. (Campanulaceae). M.Sc. thesis (Systematics &

Biodiversity Science), University of Cape Town.

C.N. CUPIDO*

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

7735 Claremont, Cape Town.

MS. received: 2001-07-12.

ERIOSPERMACEAE

VALIDATION OF TWO SECTIONAL NAMES IN ERIOSPERMUM

Perry’s revision of Eriospermum (1994) included a new

infrageneric classification of the genus with new infra-

generic taxa being formally described. However, two of the

new sectional names were invalidly published due to the

lack of typification (Art. 37. 1 St Louis Code; Greuter et al.

2000). Although this omission was mentioned in the Index

kewensis (IK) as available in the CD-Rom version, or with-

in the International Plant Names Index (IPNI) internet data-

base, no action has been taken to validate the names.

Consequently, both names are validated here:

Eriospermum [subgenus Ligulatum| section Ligu-

latum P.LPerry ex Thiede, sect. nov. Type: Eriospermum

flagelliforme (Baker) J.C. Manning.

For the Latin diagnosis see Perry in Contributions

from the Bolus Herbarium 17: 77 (1994).

Eriospermum flagelliforme is the correct name for the

species previously known as Eriospermum abyssinicum

Baker (Manning 2000).

Eriospermum [subgenus Cyathiflorum] section

Cyathillorum P.L.Perry ex Thiede, sect. nov. Type:

Eriospermum cooperi Baker.

For the Latin diagnosis see Perry in Contributions

from the Bolus Herbarium 17: 120 (1994).

The ‘description’ of Eriospermum [subgenus Erio-

spermum] section Eriospermum P.L.Perry [in tom. cit. 160]

was superfluous under Art. 22.3 St Louis Code (Greuter

et al. 2000) and consequently not included in IK.

REFERENCES

GREUTER. W.. MCNEILL. J„ BARRIE, F.R., BURDET, H.M..

DEMOULIN, V., FILGUEIRAS. T.S., NICOLSON, D.H., SILVA,

PC., SKOG, J.E., TREHANE, P„ TURLAND, N.J. &

HAWKSWORTH, D.L. 2000. International Code of Botanical

Nomenclature (Saint Louis Code). Koeltz Scientific Books, Konigstein.

MANNING, J.C. 2000. A new combination in Eriospermum

(Convallariaceae). Bothalia 30: 157.

PERRY, PL. 1994. A revision of the genus Eriospermum (Eriosperm-

aceae). Contributions from the Bolus Herbarium 17: 1-320.

J. THIEDE*

* Hamburg University, Botanical Institute, Ohnhorststr. 18, D-22609

Hamburg, Germany. E-mail: thiede@botanik.uni-hamburg.de

MS. received: 2001-09-10.

THYMELAEACEAE

A NEW SPECIES OF PASSERINA FROM WESTERN CAPE, SOUTH AFRICA

INTRODUCTION

Passerina L. comprises about 20 species and four

subspecies of evergreen perennial shrublets (Thoday

1924; Goldblatt & Manning 2000). Most members are

confined to the Cape Floristic Region with its Mediterranean

or semi-Mediterranean climate. Three species extend

northwards along the Great Escarpment, with P. monti-

vagus Bredenkamp & A.E.van Wyk extending north-

wards to Tanzania. Under the heading ‘Incertae’, Thoday

( 1924) discussed specimens of doubtful identity, most of

them from mountain summits and outlying localities.

He concluded that further collecting would solve the

problem of their identification. Here we describe a new

species confined to high altitudes on some of the moun-

tains in the western sector of the Cape Floristic Region

and the adjacent Roggeveld Escarpment. The separate

specific status of the new taxon is supported by evidence

derived from the epidermal structure (Bredenkamp &

Van Wyk 1999, 2000), leaf anatomy (Bredenkamp & Van

Wyk 2001a) and vegetative and floral morphology

(Bredenkamp & Van Wyk 2001b).

MATERIAL AND METHODS

All collections of Passerina from 22 herbaria were

studied for taxonomy and external morphology.

Specimens of the new species were found in the follow-

ing herbaria (acronyms according to Holmgren et al.

1990): BOL, K, NBG, PRE.

For leaf anatomy, both fresh and herbarium material

were studied. Vouchers used in anatomical studies:

Bredenkamp 1044-1047; Goldblatt & Manning 8627;

Bothalia 32.1 (2002)

77

Oliver 9281. Methods used in the study of leaf structure

are described in Bredenkamp & Van Wyk (1999. 2000,

2001a).

Passerina nivicola Bredenkamp & A.E.van Wyk ,

sp. nov., P. comosae C.H. Wright affinis sed bracteis cir-

cumscriptione late obovatis, basi cuneatis, rubrofusis in

alabastro, lamina utrinque glabra, chartacea, alis bullatis,

membranaceis, marginibus ciliatis dimidio superiore,

costa crassa, ventraliter tomentosa in apicem subacutum

extensa, caespite apicali pilorum. Hypanthium circa

ovarium extra tomentoso. Sepala tempore pollinationis

membranacea, lutea vel luteorosea; sepalis, exterioribus

apice setosis, sepalis interioribus utrinque glabris.

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

Waboomberg, 1 760 m, (-DD), 12 November 1989, E.G.H.

Oliver 9281 (PRE, holo.; NBG. iso.).

Rounded evergreen shrublets 0.3-0. 5 m high. Stems

greyish brown, younger branchlets greyish tomentose,

cork finely fissured, grey-brown. Leaves decussate,

imbricate on young branchlets, sessile, closely adpressed

to stem, cymbiform. cylindric, often slightly dilated api-

cally, linear-lanceolate, (2.5-)3.1(^4.5) x (0.5— )0.6(— 0.7 )

mm, inversely ericoid; adaxial surface concave, tomen-

tose; abaxial surface convex, glabrous; apex rounded,

with tuft of trichonres, tinged red; base sessile; margin

sometimes ciliate. Inflorescences composed of polytelic

synflorescences; main florescences as well as co-flores-

cences spicate. Bracts tinged red in bud stage, enveloping

flowers and fruits, largest after anthesis, becoming more

coriaceous and rounded at fruit set, decussate, imbricate,

cymbiform; lamina rolled, widely obovate, length x '/2

width (3. 1— )3.5 x 1 ,4(— 1 .9) mm, adaxially (inside) con-

cave, abaxially (outside) convex and glabrous, midrib

tomentose on inside, chartaceous; base cuneate; main

vein strongly developed, extending to form subacute

apex, with apical tuft of trichomes; wings glabrous, bul-

late. membranous; margin ciliate in upper half. Pedicel

very short or absent. Receptacle very short. Flora! enve-

lope petaloid. up to 5.3 mm long, membranous and yel-

low or yellow-pink during pollination, papyraceous and

turning red after pollination; hypanthium tomentose at

ovary, neck ± 1.2 mm long, tomentose on outside, often

hairy inside, articulation at neck base absent because of

lack of abscission tissue, fragmentation of neck base

caused by dehydration and torsification of tissue after

fruiting, resulting in shedding of sepals and androecium;

sepals 4, imbricate in bud, flexed in flower, outer sepals

cymbiform, abaxially setose at apex, inner sepals widely

ovate, glabrous. Corolla absent. Petaloid scales absent.

Disc absent. Androecium dimorphic diplostemonous,

arising from hypanthium at separation point of sepals; fil-

aments of antipetalous whorl ± 0.4 mm long, those of

antisepalous whorl ± 1 .2 mm long; pollen grains shed as

monads, spheroid, pantoporate, basal reticulum of typical

crotonoid pattern no longer discernible, replaced by sec-

ondary reticulum derived from fused sexine (Breden-

kamp & Van Wyk 1996). Ovary superior, ± 1.7 x 0.5 mm,

pseudomonomerous, uniloculate, with one pendulous ovule

laterally attached near top of ovary, placentation parietal;

ovule anatropous, position ventrally epitropous, bitegmic,

with exotegmic palisade, crassinucellate, obturator of

elongate cells extending from base of style to mi-cropyle;

stigma developing at base of sepals, penicillate. Fruit a 1-

seeded berry, pericarp membranous and dry, enveloped

by persistent, loosely arranged hypanthium fragmented at

neck base. Seed broadly fusiform with outgrowths at both

micropylar and funicular ends, ± 2.2 x 1.2 mm, tegmen

black and shiny; endosperm formation nuclear, but later

becoming cellular throughout. Figure 16.

Leaf anatomy

Leaf structural type A: leaf isobilateral. Main vascu-

lar bundle central; bundle sheath completely enclosing

vascular tissue. Secondary vascular bundles close, with

bundle sheaths adhering, forming a central plate of veins.

Sclerenchymatous hypodermal sheath absent. (Breden-

kamp & Van Wyk 2001a).

Leaf isobilateral, outline in transverse section (t/s)

transversely elliptic. Adaxial epidermis: cuticular mem-

brane (CM) ± 3 pm thick; periclinal x anticlinal cell

diameters in t/s 25 x 40 pm; stomata often dispersed in

FIGURE 16. — Passerina nivicola.

Oliver 9281: A, inflores-

cence; B. leaf; C, bract; D.

flower clasped by bract; E,

fruit enveloped by persistent

floral bract. Scale bars: A. D,

E, 2 mm; B. C. 1mm.

78

Bothalia 32,1 (2002)

FIGURE 17. — Passerina nivicola . Bredenkamp 1044 : leaf structural

type A, t/s; leaf isobilateral, vascular bundles closely arranged,

bundle sheaths adhering, forming central plate of veins, ad,

adaxial epidermis; ab, abaxial epidermis; bs, bundle sheath;

mb, median vascular bundle. Scale bar: 100 pm.

two columns in adaxial epidermal folds, with ± 3-5 rows

of epidermal cells in between, sunken or arranged in

stomatal crypts. Abaxial epidermis : cuticular ornamenta-

tion comprising several domes per cell, CM ± 20 pm

thick, irregularly marked by grooves, wax platelets

scarce, geometrical, flat or raised, with rounded to sharp

edges; cells arranged in rows, oblong, pentagonal-hep-

tagonal, dimensions in surface view 45-55 x 35^40 pm,

periclinal x anticlinal cell diameter in t/s 35^40 x 40-55

pm. Mesophyll consisting of ± 3 adaxial and 4 abaxial

layers of ± homogeneous, palisade-like parenchyma cells;

density 3 or 4 cells per 50 pm; intercellular spaces sur-

rounding central veins. Main vascular bundle 41CM50

pm thick, 810-850 pm wide, widely ovate with ample

extraxylary fibres. Bundle sheath consisting of ± 19-25

parenchymatous cells, completely enclosing vascular tis-

sue, cells rounded, transversely or longitudinally elongat-

ed. Secondary vascular bundles forming central plate of 3

or 4 veins on each side of main bundle. Figure 17.

Diagnostic characters and relationships : Passerina

nivicola is superficially similar to P. comosa , but differs

from it by its glabrous leaves and red-tinged bracts,

which are glabrous outside, with bullate, membranous

wings. The flowers of the new species are membranous

and yellow or yellow-pink at pollination, with outer

sepals abaxially setose at the apex and the inner sepals

glabrous on both surfaces.

Etymology, the specific epithet is a compound of the

Latin nivalis (= pertaining to snow) and cola (= dweller),

thus nivicola = a dweller in the snow. The name alludes

to the occurrence of this species at high altitudes, where

the plants are periodically covered by snow.

Distribution and ecology, because of its confinement

to mountainous areas P. nivicola is clearly still under-col-

lected. The most northerly record is from Sneeukrans on

the Roggeveld Escarpment, where it is associated with

Escarpment Mountain Renosterveld (Rebelo 1998).

Most records are from the Ceres and Worcester Districts

(Figure 18), with plants occurring in either Mountain

Fynbos or Central Mountain Renosterveld (Rebelo

1998).

The new species was found on Waboomberg near

Ceres growing at an altitude range of 1 200-1 760 m and

although it was summer, the plants were dug out from

under a layer of snow (Figure 19). Here the plants occur

in low restioid or graminoid veld on sandy loamy soil.

They are somewhat stunted and attain a height of ± 0.3

m, possibly because of the effects of snow. The same

species is also found lower down, next to the road lead-

ing to the mountain, where the plants are more robust

and reach a height of up to 0.5 m; the floral bracts are

more coriaceous. Accessible plants are grazed by stock.

Information from Grobler 540 , indicates that this species

also occurs on shale flats at Kareevlakte in Ceres.

Conservation status : Least Concern [LC] (IUCN Spe-

cies Survival Commission 2000).

Specimens examined

NORTHERN CAPE. — 3220 (Sutherland): Roggeveld Escarpment,

Quaggasfontein Farm, on road to Uitkyk, Sneeukrans W of Sutherland,

(-AB), Goldblatt & Manning 8627 (PRE).

17° 18° 19° 20° 21° 22°

FIGURE 18. — Known distribution

of P. nivicola.

Bothalia 32.1 (2002)

79

FIGURE 19. — Typical habitat of P. nivicola: A. rounded shrublets covered in snow on level area close to summit of Waboomberg, near Ceres; B.

individual stunted plants, ± 0.3 m high; C. plants flowering lower down mountain.

WESTERN CAPE. — 3319 (Worcester): Ceres, Kareevlakte,

(-AD), Grobler 540 (PRE); Ceres, Waboomberg, Farm Merino. (-AD),

Bredenkamp 1044-1047 (PRE); vlei N of FM tower, Cillie 9 (NBG);

level area S of beacon, Oliver 9281 (NBG. PRE); Ceres, Baviaansberg,

(-BA), Compton 8718 (NBG); Worcester Dist.. Tafelberg, (-CC),

Pillans 17159 (BOL, K), s.n. (K).

ACKNOWLEDGEMENTS

The authors would like to thank Gill Condy for the

line drawings, Otto Leistner for translating the diagnosis

into Latin and Adela Romanowski for printing the pho-

tographs.

REFERENCES

BREDENKAMP. C.L. & VAN WYK. A.E. 1996. Palynology of the

genus Passerina (Thymelaeaceae): relationships, form and

function. Grand 35: 335-346.

BREDENKAMP, C.L. & VAN WYK, A.E. 1999. Structure of

mucilaginous epidermal cell walls in Passerina (Thyme-

laeaceae). Botanical Journal of the Linnean Society 129:

223-238.

BREDENKAMP. C.L. & VAN WYK, A.E. 2000. The epidermis in

Passerina (Thymelaeaceae): structure, function and taxonomic

significance. Bothalia 30: 69-86.

BREDENKAMP. C.L. & VAN WYK, A.E. 2001a. Leaf anatomy of the

genus Passerina (Thymelaeaceae): taxonomic and ecological

significance. Bothalia 31: 53-70.

BREDENKAMP. C.L. & VAN WYK. A.E. 2001b. Taxonomic signifi-

cance of inflorescences, floral morphology and anatomy in

Passerina (Thymelaeaceae). Bothalia 31: 213-236.

GOLDBLATT, P. & MANNING. J. 2000. Cape plants. A conspectus of

the Cape flora of South Africa. Strelitzia 9.

HOLMGREN, K„ HOLMGREN. N.H. & BARNETT, L.C. 1990.

Index Herbariorum. International Association for Plant

Taxonomy, New York Botanical Garden, New York.

IUCN SPECIES SURVIVAL COMMISSION 1994. IUCN Red List

Categories. ILrCN. Gland, Switzerland.

REBELO, A.G. 1998. Fynbos Biome. In A.B. Low & A.G. Rebelo,

Vegetation of South Africa. Lesotho and Swaziland: 62-74.

Department of Environmental Affairs & Tourism, Pretoria.

THODAY, D. 1924. XV III. A revision of Passerina. Kew Bulletin 4:

146-168.

C.L. BREDENKAMP* and A.E. VAN WYK**

* National Botanical Institute, Private Bag X101, 0001 Pretoria.

** H.G.W.J Schweickerdt Herbarium, Department of Botany, Uni-

versity of Pretoria, 0002 Pretoria.

MS. received: 2001-08-31.

THYMELAEACEAE

A NEW SPECIES OF GN1D1A FROM THE KNERSVLAKTE, WESTERN CAPE, SOUTH AFRICA

Gnidia pedunculata Beyers , sp. nov., Gnidia

polycephala (C.A.Mey.) Gilg similis sed ab ea peduncu-

lo plerumque longo, involucralibus bracteis coriaceis

abaxiale glabris, floribus quadrimeris squamis quarternis

camosis petaloideis differt.

TYPE. — Western Cape, 31 18 (Vanrhynsdorp): Kners-

vlakte, Olifants River Settlement 316, near Eastern bor-

der, (-BC), 6-08-1993, Le Roux & Hilton-Taylor 27

(NBG, holo.; K, PRE).

Erect dwarf shrub up to 0.25 m tall. Branches ascend-

ing, later becoming woody, glabrous, densely leafy, later

becoming naked with prominent leaf scars. Leaves alter-

nate, ascending, densely imbricate below, becoming dis-

tant above, petiolate, glabrous; blade oblanceolate,

4. 8-7. 8 x 0. 5-1.4 mm, subfleshy, enervate, ad- and abax-

ially flat, apex obtuse or subacute; petiole 0. 3-0.4 mm

long. Inflorescence terminal, indeterminate, bracteate

umbel with up to 14 mature flowers at a time; flower

development sequential — fruiting stages present with

young buds developing within; peduncle 10-30 mm

long, glabrous. Involucral bracts 5 or 6, ovate or elliptic,

7-10 x 3-6 mm, green or green with maroon tinge, cori-

aceous, faintly nervate, adaxially shortly adpressed hairy,

abaxially glabrous, apex rounded or subacute. Flowers

subactinomorphic, yellow, strongly scented. Pedicel

1. 3-2.0 mm long, adaxially sericeous, abaxially densely

covered with short, silky, spreading hairs. Hypanthium

9.5-22.0 mm long, circumscissile 3. 5-5.0 mm from

base, upper portion narrowly funnel-shaped, covered

with short crisped hairs becoming sericeous towards

apex, basal portion narrowly ovate, densely covered with

long, silky, ascending hairs. Sepals 4, narrowly elliptic,

2. 9-3. 6 x 1.0-2. 2 mm, adaxially glabrous, abaxially

sericeous, margin recurved, apex rounded. Petaloid

scales 4, inserted at rim of hypanthium alternate to sepa-

ls, fleshy, narrowly oblong to oblong, narrowly elliptic to

80

Bothalia 32,1 (2002)

FIGURE 20. — Gnidia pedunculata, Le Roux & Hilton-Taylor 27 (NBG). A. portion of plant, x 1 ; B, leaf, abaxial view and c/s; C, involucral bract,

abaxial view. D. E, flower: D. side view, circumscission zone arrowed; E, 1/s above circumscission zone with gynoecium removed. F,

gynoecium and stigma enlarged. Scale bars: B-F, 2 mm.

elliptic, obovate or spathulate, 0.9-1. 6 x 0.5-0. 6 mm,

apex rounded or obtuse, glabrous. Stamens 4 + 4, sub-

sessile; outer whorl semi-exserted, inserted just below

rim of hypanthium; inner whorl included, inserted a short

way down throat of hypanthium; filament 0.1 -0.2 mm

long; anthers introrse, ellipsoid, 0.5-0. 9 x 0.3-0. 5.

Ovary narrowly ellipsoid, attenuated at base, 1.6-3. 6 x

0.5-0. 9 mm, glabrous, unilocular with single, pendulous

ovule; style filiform, inserted laterally on ovary, includ-

ed, 4. 8-5.0 mm long, glabrous; stigma penicillate. Fruit

an achene, enclosed in persistent base of hypanthium.

Flowering time'. August. Figure 20.

This species at first glance appears similar to Gnidia

polycephala (C.A.Mey.) Gilg which also has alternate

leaves, a terminal, indeterminate, bracteate umbel with

involucral leaves much broader than the cauline leaves

and yellow sericeous flowers. It, however, differs from

G. polycephala by the usually well-developed peduncle,

the coriaceous involucral bracts which are glabrous

abaxially and the four-merous flowers with four, fleshy,

petaloid scales. In G. polycephala there is no well-devel-

oped, naked peduncular region, the papery thin involu-

cral bracts are adpressed hairy abaxially, the flower is

pentamerous and the petaloid scales are absent.

Gnidia pedunculata has only been recorded from the

Knersvlakte north of Vredendal, growing in Succulent

Karoo vegetation in well-drained, sandy, loam soil.

Figure 21.

FIGURE 21. — Geographical distribution of Gnidia pedunculata.

Bothalia 32.1 (2002)

81

The usually well-developed, naked peduncles in this

species prompted the specific epithet which is derived

from the Latin, pedunculatus.

Parcitype material

WESTERN CAPE.— 3118 (Vanrhynsdorp): Knersvlakte,

Moedverloor. near river on steep slope, (-AD), 31-08-2001 . Schmiedel

118546 (NBG).

ACKNOWLEDGEMENTS

I would like to thank Dr Ted Oliver for the Latin diag-

nosis and Mrs Inge Oliver for the line drawings.

J.B.P. BEYERS*

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

7735 Claremont, Cape Town.

MS. received: 2001-12-04.

LAMIACEAE

NEW COMBINATIONS IN THE GENUS ROTHECA IN SOUTHERN AFRICA

The genus Clerodendrum L. was divided into a num-

ber of different sections or subgenera by Gtirke (1895),

Pearson (1912) and Thomas ( 1936) on the basis of inter

alia the flower symmetry. In the past, some of these sec-

tions or subgenera were considered to be independent

genera, e.g. the genus Cyclonema Hochst. (1842). Steane

et al. (1997) showed that Clerodendrum sens. lat. is

polyphyletic and that Cyclonema should be reinstated as

a genus. Steane & Mabberley (1998) revived the genus

Rotheca Raf. for taxa under the subgenus Cyclonema

(Hochst.) Thomas and/or sections Cyclonema (Hochst.)

Giirke and Konocalyx Verde, of the genus Clerodendrum

on the grounds of the corolla (flower) symmetry, anther

attachment and stigma symmetry. More new combina-

tions for taxa from the Flora zambesiaca and Conspectus

florae angolensis areas were made by Fernandes &

Verdcourt (2000).

A number of new combinations for taxa occurring in

southern Africa, not covered by the above-mentioned

publications, are presented here:

Rotheca caerulea ( N.E.Br .) P.PJ. Herman &

Retief comb. nov.

Clerodendrum caeruleum N.E.Br. in Kew Bulletin 1895: 115

(1895). Type: [KwaZulu-JNatal, Mooi River Valley, Gerrard 1252 (K).

Rotheca cuneiformis ( Moldenke ) P.P.J. Herman &

Retief, comb. nov.

Clerodendrum cuneiforme Moldenke in Phytologia 59: 1 19 (1986).

C. cuneatum Giirke: 303 (1900), nom. illeg., non Turcz.: 221 (1863).

Type: Transvaal [Mpumalanga], Lydenburg, Wilms 160 (?B).

C. scheffleri Giirke var. ellipticum Moldenke: 417 (1940). Type:

[KwaZulu-]Natal, Durban, Evans 544a (NH).

Rotheca louwalbertsii (P.P.J .Herman) P.P.J. Herman

& Retief, comb. nov.

Clerodendrum louwalbertsii P.P.J.Herman in Bothalia 25: 100

(1995). Type: Transvaal [Northern Province], between Nylstroom and

Warmbaths, Germishuizen 3343 (PRE, holo.!).

Rotheca pilosa (H. Pearson) P.P.J.Herman &

Retief, comb. nov.

Clerodendrum pilosum H. Pearson in Transactions of the South

African Philosophical Society 15: 181 (1905). Type: [Eastern Cape],

Kentani, Pegler 164 (PRE!).

Rotheca uncinata ( Schinz ) P.P.J.Herman &

Retief, comb. nov.

Clerodendrum uncinatum Schinz in Verhandlungen des

Botanischen Vereins der Provinz Brandenburg 31: 206 (1889).

Kalaharia uncinata (Schinz) Moldenke: 132 ( 1955). Type: South West

Africa [Namibia], Goroxas, Scliinz 1890 (?Z).

C. spinescens (Oliv.) Giirke: 180 (1893), nom. illeg.

REFERENCES

BROWN, N.E. 1895. Plantarum novarum in herbario horti regii con-

servatarum. Decades XV-XIX. Kew Bulletin 1895: 102-120.

FERNANDES. R.B. & VERDCOURT. B. 2000. Rotheca (Labiatae)

revived — more new combinations. Kew Bulletin 55: 147-154.

GURKE, M. 1893. Verbenaceae africanae. Botanische Jahrbucher 18:

165-183.

GURKE, M. 1895. Verbenaceae. In A. Engler, Die Pflanzenwelt Ost-

Afrikas, Theil C: 337-342. Dietrich Reimer. Berlin.

GURKE. M. 1900. Verbenaceae africanae. II. Clerodendron L. Bota-

nische Jahrbucher 28: 291-305.

HERMAN, P.P.J. 1995. Verbenaceae. A new species in the genus

Clerodendrum. Bothalia 25: 100-102.

HOCHSTETTER, C.F. 1842. Nova genera plantarum Africae turn aus-

tralis turn tropicae borealis proponit et describit. Verbenaceae.

Flora 25: 225-228.

MOLDENKE. H.N. 1940. Novelties in the Avicenniaceae and

Verbenaceae. Phytologia 1: 409^-19.

MOLDENKE. H.N. 1955. Notes on new and noteworthy plants. XIX.

Phytologia 5: 132-134.

MOLDENKE, H.N. 1986. Notes on the genus Clerodendrum

(Verbenaceae) XII. Phytologia 59: 99-126.

PEARSON, H.H.W. 1905. South African Verbenaceae. Transactions of

the South African Philosophical Society 15: 175-182.

PEARSON. H.H.W. 1912. Verbenaceae. Flora capensis 5,1: 180-226.

SCHINZ, H. 1889. Beitrage zur Kenntnis der Flora von Deutsch-

Siidwest-Afrika und der angrenzenden Gebiete. Verhandlungen

des Botanischen Vereins der Provinz Brandenburg 31: 206,

207.

STEANE, D A. & MABBERLEY, D.J. 1998. Rotheca (Lamiaceae)

revived. Novon 8: 204—206.

STEANE. D.A., SCOTLAND, R.W.. MABBERLEY, D.J..

WAGSTAFF, S.J.. REEVES. PA. & OLMSTEAD. R.G. 1997.

Phylogenetic relationships of Clerodendrum s.l. (Lamiaceae)

inferred from chloroplast DNA. Systematic Botany 22:

229-243.

THOMAS, B. 1936. Die Gattung Clerodendrum in Afrika. Botanische

Jahrbucher 68: 1-106.

TURCZANINOW, N.S. 1863. Verbenaceae et Myoporaceae nonnullae

hucusque indescriptae. Mosquae. Bulletin de la Societe des

Naturalistes de Moscou. Section Biologique 36: 221.

P.P.J. HERMAN* and E. RETIEF*

* National Herbarium. National Botanical Institute, Private Bag X101,

0001 Pretoria.

MS. received: 2001-05-21.

82

Bothalia 32.1 (2002)

MARSILEACEAE-PTEROPSIDA

FIRST REPORT OF THE GENUS P1LULARIA FROM CONTINENTAL AFRICA

Pilularia L. is a genus of five poorly defined species.

Two species, P. globulifera L. and P. minuta A.Braun

have a European distribution, whereas P americana

A.Braun (inch P. mandoni A.Braun) occurs in North and

South America. One species each occurs in Australia (P.

novae-hollandiae A.Braun) and New Zealand (P. novae-

zelandiae Kirk). Hitherto no taxa belonging to the genus

have been recorded for the African continent.

Recently, however, I collected a species of Pilularia

in Northern Cape, South Africa. An assessment of P.

americana , P novae-hollandiae and P. novae-zelandiae

by Large & Braggins (1989) shows that these species all

have four locules in each sporocarp, rather than the two

locules per sporocatp recorded for the European taxa.

Due to a lack of suitable characters and the large overlap

in diagnostic features between these species, Large &

Braggins (1989) concluded that some of the names might

have to be placed in synonymy.

A study of the South African specimen showed that it

also has four locules in each sporocarp, thus making it

part of the P. americana group of species. Other congru-

ent features the South African specimen has with the P.

americana group of species include frond length, pedicel

length, sporocarp size, and number of megasporangia per

locule, micro- and megasporangia size as well as epi-

spore ornamentation. For this reason the oldest valid

name for the taxon, P. americana , is tentatively applied

to the South African Pilularia species.

Pilularia americana A.Braun in Monatsberichte

der Koniglichen Akademie der Wissenschaften zu Berlin

15: 435 (1864). Type: North America, Fort Smith, on the

Arkansas River, Thomas Nutt all s.n. (Holotype not locat-

ed; NY, iso.).

Plants terrestrial. Rhizome with single unbranched

stele, creeping, irregularly branched, up to 0.5 mm diam.,

set with single root or root clusters at nodes, roots

unbranched, internodes up to 11 mm long, initially

sparsely haired, glabrous with age, mostly with a few

hairs at frond base, hairs up to 5 cells long, laterally

attached. Fronds with single unbranched stele, borne

singly at nodes, erect, simple, terete, up to 19 mm long,

basally with few laterally attached, 2-celled hairs,

glabrous higher up; stomata of aperigenous type,

(34.0— )39.23(— 44.0) pm long. Sporocarp globose, up to

2.5 mm diam., laterally attached, 4-locular, densely

haired, hairs up to 5 cells long, epigeal at maturity; vena-

tion catadromous, pinnately branched, each locule has 3-

veined vascular supply, central vein bearing a sorus near

branching point containing micro- and megasporangia;

veins free or anastomose at distal end of sporocarp;

sporocarp pedicel arises at node on rhizome, with single

unbranched stele, up to 2 mm long, initially growing

downwards, twisted and burying sporocarps in substrate.

Microsporangia clavate, several per sorus, wall one cell

layer thick, hyaline, each bearing 23-32 microspores.

Microspores trilete, spherical in polar view, asymmetric

in lateral view, laesura straight or curved, perispore hya-

FIGURE 22. — Pilularia americana

A.Braun. A, habit: A,, rhi-

zome; Ai, frond; Ai, sporo-

carp and cluster of roots. B,

lamina hair; C, sporocarp

hair; D, sporocarp venation

showing position of sori.

Scale bars: A, 10 mm; B. 0.1

mm; C, 0.5 mm; D, not to

scale. Drawn by J.P. Roux.

Bothalia 32,1 (2002)

83

FIGURE 23. — Distribution of Pilularia americana in southern Africa.

line, larger than spore, epispore thicker on distal face,

densely and finely rugulate, exospore (32.0-)34.87(-36.0)

pm diam. Megasporangia globose, wall one cell layer

thick, hyaline, 8-17 per sorus, each bearing a single

megaspore. Megaspores globose in polar and lateral

view, with prominent ridged papilla up to 104 pm long,

perispore hyaline, much larger than megaspore, epispore

thicker around papilla and at distal pole than at equator,

finely and deeply folded giving it a finely rugulate appear-

ance, exospore up to 360 pm diam. Figure 22A-D.

Specimen examined

NORTHERN CAPE. — 3119 (Calvinia): Nieuwoudtville, in season-

al pan. (-AC), 10 October 2001. J.P. Roux 3156 (NBG).

Distribution and ecology

Pilularia americana was recorded from the edge of a

seasonal pan on the Bokkeveld Escarpment, an area of

low rainfall (500-650 mm per annum) which occurs dur-

ing winter and spring (June-September) (Figure 23).

Sandstone beds of the Cape System are the dominant

geological formation in the region and support fynbos

vegetation.

Until a thorough study of the distribution of the species

and population size has been done, its conservation status

should be considered as data-deficient (IUCN 1994).

ACKNOWLEDGEMENTS

My thanks to Dr J.T. Mickel who verified the pres-

ence of the isotype in the New York herbarium.

REFERENCES

BRAUN, A. 1864. Uber Marsilea und Pilularia. Monatsberichte der

Konielichen Akademie der Wissenschaften zu Berlin 15:

434-436.

IUCN SPECIES SURVIVAL COMMISSION. 1994. IUCN Red List

Categories. IUCN Gland. Switzerland.

LARGE. M.F. & BRAGGINS, J.E. 1989. An assessment of characters

of taxonomic significance in the genus Pilularia (Mar-

sileaceae): with particuar reference to P. americana , P. novae -

hollandiae , and P. novae-zelandiae. New Zealand Journal of

Botany 27: 481^186.

J.P ROUX*

* Compton Herbarium. National Botanical Institute, Private Bag XI,

7735 Claremont, Cape Town.

MS. received: 2001-12-07.

Bothalia 32.1: 85-89(2002)

The taxonomic significance of trichome type and distribution in

Melolobium (Fabaceae)

A. MOTEETEE* B-E. VAN WYK* and P.M. TILNEY*

Keywords: Fabaceae, Melolobium Eckl. & Zeyh., sessile glands, stalked glands, trichomes, uniseriate hairs

ABSTRACT

All species of Melolobium Eckl. & Zeyh. were examined for epidermal features and particularly the glands which are a

distinctive feature amongst the southern African Genisteae. For comparative purposes, three species of Argyrolobium Eckl.

& Zeyh.. all five species of Dichilus DC. and five species of Polhillia C.H. Stilt, were also examined for trichome type and

distribution. Three trichome types are recognized in Melolobium. Trichome type and distribution provide an important

insight into taxonomic relations at species level in Melolobium and sometimes even allow a distinction between regional

forms. The distribution of glands (sessile and stalked) is of considerable diagnostic value in identifying species of

Melolobium. A key to all the species of the genus based mainly on type and distribution of trichomes, is presented.

INTRODUCTION

Melolobium Eckl. & Zeyh. is a papilionoid legume

genus restricted to southern Africa. Although about 20

species have been described, we recognize only 15 of

them (a complete synonymy will be published else-

where). The genus consists of small shrubs or perennial

herbs, characterized by their usually spiny habit, auricu-

late stipules and bilabiate calyces. Some species have

glandular trichomes, referred to as glandular papillae by

Gibbs (1967), glandular tubercles by Polhill (1976) and

stipitate glands by Harvey (1862). Glands are also char-

acteristic of the Mediterranean genus Adenocarpus DC.

Melolobium and related genera were originally placed in

the tribe Genisteae (Harvey 1862), then transferred to

Crotalarieae (Bentham 1865; Polhill 1976, 1981), and

finally moved back to Genisteae by Van Wyk & Schutte

(1995), where they are now firmly placed.

In the latest available revision of the genus, Harvey

(1862) used hairs and glands as diagnostic characters,

but the full extent of the variation, especially at micro-

scopic level, has not yet been studied. The aims of this

study were: to determine the taxonomic potential of epi-

dermal features in Melolobium (at both species and

generic levels); to record the microscopic structure of

hairs and glands in this genus; and to determine the

homology of glands in Melolobium and Adenocarpus.

MATERIALS AND METHODS

Hair type and distribution were investigated in all 1 5

of the species of Melolobium that we recognize, as well

as in three species of Argyrolobium Eckl. & Zeyh., all

five species of Dichilus DC. and five of the seven species

of Polhillia C.H.Stirt. A list of voucher specimens of all

species of Melolobium and the related African genistoid

genera used in this study is given in Table 1. For light

microscope studies, material taken from formalin: acetic

* Department of Botany, Rand Afrikaans University, RO. Box 524,

2006 Auckland Park, Johannesburg.

MS. received: 2001-04-12.

acid: alcohol (FAA) and herbarium specimens was

embedded in glycol methacrylate (GMA) according to a

modification of the method of Feder & O'Brien (1968).

This modification involves infiltrating the material for a

minimum of 24 hours between the first two changes and

for a longer period (usually at least five days) before

placing in the gelatine capsules, which are then heated in

the oven at 60° C for 24 hours to polymerize. A Porter

Blum MT-1 ultramicrotome was used for sectioning and

the sections were stained according to the periodic acid-

Schiff/Toluidine Blue (PAS/TB) staining method. For

epidermal peels, pieces of leaves were treated according

to the method of Ram & Nayar (1974). To study tri-

chome distribution, several specimens of each taxon

were examined with a stereomicroscope. For SEM stud-

ies of trichomes, herbarium or washed, air-dried FAA

material was used and at least two specimens of each

taxon were examined using a JEOL JSM 5600 scanning

electron microscope.

RESULTS AND DISCUSSION

Trichome type

Trichome type and distribution in Melolobium and

related African genera are summarized in Table 2. Three

trichome types were recognized in Melolobium'. uniseri-

ate hairs with a long narrow terminal cell and two or

three short basal cells (Figure 1A, B, D, E); stalked

glands with a unicellular head and a multicellular stalk

(Figures 2A-D; 4C); and sessile glands (Figure 3). Uni-

seriate hairs occur in all species of Melolobium [except

in M. exudans Harv. and M. lampolobum (E.Mey.)

A.Moteetee & B.-E.van Wyk which are glabrous],

Dichilus and in all the examined species of Argyrolobium

and Polhillia. The two types of glands are found only in

Melolobium.

In the subfamily Papilionoideae, uniseriate hairs con-

sist of three cells: a frequently enlarged epidermal cell,

serving as a basal cell; a short stalk cell, which occa-

sionally has special contents and is suberized; and an

elongated terminal cell (Solereder 1908). In Melolobium ,

86

Bothalia 32,1 (2002)

TABLE 1. — Voucher specimens of Argyrolobium, Dichilus, Melolobium and Polhillia examined for trichome characters

Taxon Voucher

Melolobiwn

adenodes Eckl. & Zeyh.

alpinum Eckl. & Zeyh.

aethiopicum (L.) Druce

calycinum Benth.

candicans Eckl. & Zeyh.

canescens Benth.

exudans Harv.

humile Eckl. & Zeyh.

lampolobum (E.Mey.) A.Moteetee & B.-E.van Wyk

macrocalyx Dummer

microphyllum (L.f.) Eckl. & Zeyh.

obcordatum Harv.

stipulatwn Harv.

subspicatum Conrath

wilmsii Harms

Dean 756* (JRAU); Van Wyk 3070, 4036 ( JRAU)

Moteetee & Van Wvk 5 (JRAU); Sclmtte 158*, 332 (JRAU)

Van Wyk 2452*, 2685*, 4040 (JRAU)

Moteetee 10 (JRAU); Schutte 349* (JRAU); Thome 54470 (SAM)

Rourke 1739 (PRE); Schutte 499 (JRAU); Schutte 252 (JRAU)

Dean 648 (JRAU); De Castro 126* (JRAU); Van Wvk 3058* (JRAU)

Van Wyk 2468, 2692, 2702* (JRAU)

Powrie 648 (PRE); Van Wyk 2351, 2543* (JRAU)

Marshall 234 (JRAU); Van Wyk 2143 (JRAU); M. van Wyk 1081* (PRE)

Basson 105* (PRE); Moteetee 8 (JRAU); Van Wyk 3061 (JRAU)

Bredenkamp 1121 (PRE); Bolus 37 (BOL); De Winter 2601 (NBG);

Moteetee & Van Wyk 3 (JRAU)

Moteetee & Van Wyk 4 (JRAU); Schutte 147*, 394* (JRAU)

Van Wyk 2239, 2562*, 4037 (JRAU)

Moteetee & Van Wyk 2 (JRAU); Schutte 108* (JRAU); Van Wyk 1779 (JRAU)

Moteetee & Van Wyk 1 (JRAL1). Schutte 402*, Van Wyk 2624* (JRAU)

Adenocarpus

mannii Hook.f.

Teixeira & Andrade 4665 (PRE)

Argyrolobium

frutescens Burtt Davy

lanceolatum Eckl. & Zeyh.

megarrhizum Bolus

Van Wyk 1858, 2815* (JRAU)

Schutte 469*: Van Wvk 2080, 2087 (JRAU)

Van W\’k 2923, 3611 (JRAU)

Dichilus

gracilis Eckl. & Zeyh.

lebeckioides* DC.

pilosus * Conrath ex Schinz

reflexus* (N.E.Br.) A. L. Schutte

strictus E.Mey.

Polhillia

brevicalyx* (C.H.Stirt.) B.-E.van Wyk & A. L. Schutte

canescens C.H.Stirt.

involucrata* (Thunb.) B.-E.van Wyk & A. L. Schutte

obsoleta* (Harv.) B.-E.van Wyk

pollens C.H.Stirt.

Schutte 227, 241 (JRAU); M. van Wyk 2501* (JRAU)

De Castro 128 (JRAU); Schutte 118 (JRAU); Van Wvk 1538 (JRAU)

De Castro 115 (JRAU): Schutte 95. 127 (JRAU)

Sclmtte 183, 188 (JRAU); Stirton 11795 (JRAU)

Schutte 150*. 155 (JRAU); Van W\’k 1553 (JRAU)

Sclmtte 388 (JRAU); Van Wyk 2104, 2134 (JRAU)

Van Wyk 2094, 2092 (JRAU)

Schutte 398 (JRAU); M. Steenkamp sub Schutte 379 (JRAU)

Van Wyk 214, 2701 (JRAU)

Van Wyk 2095, 2129*. 2708 (JRAU)

* Species and specimens used only for SEM studies.

however, some hairs have one basal cell, whereas others

have two. In the latter case, the basal cell appears to have

divided periclinally (Figure IB). The basal cells are

structurally similar to other epidermal cells and the stalk

cell is very thick-walled (Figure IB). In Argyrolobium,

Dichilus and Polhillia on the other hand, there is always

one basal cell, which like the other epidermal cells, is

papillate (Figure 1C). In this case the stalk cell is also

markedly thickened. According to Solereder (1908) the

terminal cell is often uniformly or spirally thickened,

with verrucose or peg-shaped irregularities on the sur-

face. Scanning electron microscopy shows that in the

examined species there are two types of surface sculp-

turing of the terminal cell: striated and verrucose. In all

species of Melolobium, hairs have striated surfaces

(Figure 4A), whereas in all species of Dichilus they are

verrucose (Figure 4B). In Argyrolobium and Polhillia,

both striated and verrucose hairs are present (Schutte 1988).

Stalked glands and sessile glands occur only in

Melolobium species and not in any of the other genera.

Structural details of these two types of glands are record-

ed and illustrated for the first time. Stalked glands have

a spherical, unicellular head and a multicellular stalk

consisting of several elongated cells (Figure 2A-D).

According to Solereder ( 1908), the glands in Melolobium

are unicellular and consist of a short globular head.

Polhill ( 1976) likened them to those found in Adenocar-

pus, but Solereder (1908) described the glands of

Adenocarpus as ‘multicellular glandular shaggy hairs,

columnar in shape and broadened in a capitate manner at

their apex'. Gibbs (1967) referred to Adenocarpus glands

as glandular papillae arising ‘as outgrowths of columnar-

shaped epidermal cells'. Examination of these glands,

however, shows that they are neither shaggy hairs nor

papillae, but rather multicellular glands (Figure 2E) with

broad capitate apices unlike the unicellular-headed and

narrow-stalked glands of Melolobium.

Sessile glands (Figure 3) are not visible to the naked

eye and are barely visible under the dissecting micro-

scope. Some species such as M. alpinum Eckl. & Zeyh.

and M. candicans (E.Mey.) Eckl. & Zeyh. have previ-

ously been described as non-glandular (Harvey 1862),

possibly because these minute glands were not detected.

Stalked glands and sessile glands never co-occur, but

may be found on different parts of the same plant. For

example, two species with sessile glands on their leaves,

M. exudans and M. stipulation Harv., also have stalked

ones on their calyces and pods.

Bothalia 32,1 (2002)

87

TABLE 2. — Epidermal features of Argyrolobium, Dichilus, Melolo-

biuin and Polhillia

FIGURE 2. — LM photographs of stalked glands: A, Melolobium

adenodes, Van Wyk 3070, 1/s; B, M. obcordatum, Moteetee & Van

Wyk 4, 1/s. C. D, M. adenodes, Van Wyk 4036: C, 1/s; D. surface

view. E, Adenocarpus mannii Hook.f., Teixeira & Andrade

4665 , 1/s multicellular glands. Scale bars: A-D, 25 pm; E, 75 pm.

FIGURE 1. — LM photographs of uniseriate hairs: A, Melolobium

aethiopicum. Van Wyk 4036, 1/s with one basal cell; B. M.

calycinum , Moteetee 10, 1/s with two basal cells; C, Argyrolo-

bium megarrhizum Bolus, Van Wyk 3611, 1/s with papillate

basal cells; D, M. aethiopicum. Van Wyk 4040, surface view,

with some of terminal cells broken off; E, M. microphyllum,

Moteetee & Van Wyk 3, surface view. Scale bars; 10 pm.

Bothalia 32,1 (2002)

FIGURE 3. — LM photographs of 1/s

of sessile glands: A, Melolo-

bium alpinum, Schutte 332:

B. glandular form of M. can-

dicans, Schutte 499. Scale bars:

A, B, 5 pm.

The co-occurrence of • structures is generally indica-

tive of non-homology. Since stalked and sessile glands

never co-occur in Melolobium, a sessile gland may be a

stalked gland in which development was merely arrested

at the unicellular stage. Sessile and stalked glands have

diagnostically different distributions at the species level

with no variation at all within species. Hairs, on the other

hand, are more variable in distribution and can be used to

distinguish between different populations within some of

the species.

The function of glandular trichomes in Melolobium is

not yet clear, but many species are viscid. Examination

of epidermal peels of the leaves revealed the heads of the

glands to have dense protoplasts (Figures 2A, B, D; 3),

further suggesting that these structures might be secre-

tory in nature. In Adenocarpus, ‘the inner cells of the

glands break down at maturity to produce a viscous

secretion' (Gibbs 1967). Glandular trichomes are known

to secrete a large number of different substances, includ-

ing water, salt, nectar, mucilage, terpenes and digestive

enzymes (Esau 1977). Studies are being carried out to

determine the chemical nature of the contents of these

glands.

Hair distribution

In Melolobium the distribution of hairs on the leaflet

blade varies greatly, even within a species, but is highly

consistent within various forms or provenances. For

example, in the ‘typical' form of M. humile Eckl. &

Zeyh., hairs occur on all parts of the lamina, whereas in

the ‘Malmesbury’ form they occur only on the adaxial

midrib and are rare or absent on other parts. Hairs are

most commonly present on the leaflet margins and abax-

ial midrib, with the exception of M. adenodes Eckl. &

Zeyh. and some forms of M. microphyllum (L.f.) Eckl. &

Zeyh. where they are absent. In general, hairs are less

frequent on the adaxial surface than on the abaxial one.

Hairs can be used to distinguish many of the species of

Melolobium. M. adenodes is allied to M. humile (both

glandular), for example, but the former is subglabrous

and the latter is densely hairy. M. aethiopicum (L.) Druce

superficially resembles M. alpinum, but leaves of the for-

mer are hairy on both surfaces, whereas the latter is only

sparsely hairy on the upper surface. In all species of

Melolobium, glands (whether stalked or sessile) and

hairs co-occur. The distribution of hairs is not correlated

with the type of glands.

CONCLUSIONS

The type and distribution of glands is of diagnostic

significance at the generic level in African Genisteae and

at the species level within Melolobium. The type and dis-

tribution of hairs is of taxonomic value at both species

and population (provenance) levels. Within the Genis-

teae, the microscopic structure of hairs and glands is

unique in Melolobium. The glands in Melolobium and

Adenocarpus are not homologous.

Key to species of Melolobium based on trichome type

and distribution

la Stalked glands present:

2a Stalked glands on calyces only:

3a Plants unarmed:

4a Plants hairy M. stipulation

4b Plants almost glabrous M. exudans

FIGURE 4. — SEM micrographs: A,

striated hair surface in Melo-

lobium calycinum, Thome

54470 ; B, verrucose hair sur-

face in Dichilus gracilis

Eckl. & Zeyh., Schutte 241:

C, M. humile, Van Wyk 2351.

stalked glands. Scale bars: A,

B, 2 pm; C, 100 pm.

Bothalia 32.1 (2002)

89

3b Plants spiny:

5a Pods straight M. candicans*

5b Pods falcate M. canescens* (E.Mey. ) Benth.

2b Stalked glands on stems, leaves and calyces:

6a Plants glabrous or subglabrous M. adenodes

6b Plants sparsely to densely hairy:

7a Plants unarmed or slightly spinescent:

8a Leaflets distinctly obcordate, apex sharply emar-

ginate: bracts obliquely lanceolate to ovate ....

M. obcordatum Harv.

8b Leaflets obovate-oblong. apex mucronate; bracts

cordate to suborbicular M. luimile

7b Plants distinctly spiny M. microphyllum

lb Stalked glands absent:

9a Sessile glands present, at least on calyces (visible under 20 x

magnification):

10a Plants distinctly spiny:

11a Stems and pods subglabrous, the latter distinctly

shiny M. lampolobum

lib Stems and pods usually densely hairy, velutinous:

12a Pods straight M. candicans

12b Pods falcate M. canescens

10b Plants unarmed:

13a Sessile glands on leaves and calyces M. alpinum

13b Sessile glands on calyces only:

14a Stipules present M. wilmsii Harms.

14b Stipules absent M. subspicatum Conrath

9b Sessile glands absent:

15a Plants distinctly spiny M. calycinum Benth.

1 5b Plants unarmed:

16a Leaves and calyces densely silky: pods short, scarce-

ly exceeding calyx M. macrocalyx Dummer

16b Leaves and calyces pubescent; pods several times

longer than calyx M. aethiopicum

* Some forms of M. candicans and M. canescens are glandular as in M.

microphyllum. Since there is no strong geographical pattern in the dis-

tribution of glandular forms of M. candicans and M. canescens, we

suspect there is hybridization/ introgression between M. microphyllum

and these two species

ACKNOWLEDGEMENTS

The curators of the cited herbaria (NBG. PRE, SAM)

are thanked for the loan of specimens. Financial assis-

tance from the Government of Lesotho and the National

Research Foundation is gratefully acknowledged. We

appreciate the assistance of Cassim Petker (RAU

Photographic Department).

REFERENCES

BENTHAM. G. 1865. Leguminosae. In G. Bentham & J.D. Hooker.

Genera plantarum 1: 434-600. Reeve, London.

ESAU. K. 1977. Anatomy of seed plants, edn 2. Wiley, New York.

FEDER. N. & O'BRIEN, T.P 1968. Plant microtechnique: some prin-

ciples and new methods. American Journal of Botany 55:

123-142.

GIBBS. PE. 1967. A revision of the genus Adenocarpus. Boletim da

Sociedade Broteriana, ser. 2, 41: 67-121.

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

Flora capensis 2: 77-81. Hodges Smith, Dublin.

POLHILL, R.M. 1976. Genisteae (Adans.) Benth. and related tribes

(Leguminosae). Botanical Systematics 1: 143-368.

POLHILL. R.M. 1981. Tribe 29. Crotalaneae (Benth.) Hutch. In R. M.

Polhill & P. H. Raven, Advances in legume systematics 1:

399-402. Royal Botanic Gardens. Kew.

RAM. H.Y.M. & NAYYAR. V.L. 1974. A rapid method of obtaining

epidermal peels in plants by treatment with cupric sulphate and

hydrochloric acid. Stain Technology 49: 114-116.

SCHUTTE. A.L. 1988. 'n Taksonomiese studie van die genus Dichilus

DC. ( Fabaceae-Crotalarieae ). M.Sc. thesis. Rand Afrikaans

University. Johannesburg. Unpublished.

SOLEREDER, H. 1908. Systematic anatomy of the dicotyledons, Vols

1 & 2 (English translation by Boodle & Fritsch). Clarendon

Press, Oxford.

VAN WYK. B-E. & SCHUTTE A.L. 1995. Phylogenetic relationships

in the tribes Podalyrieae. Liparieae and Crotalarieae. In M.D.

Crisp & J.J. Doyle, Advances in legume systematics 7: 283-308.

Royal Botanic Gardens, Kew.

Bothalia 32,1:91-96 (2002)

Pollen morphology and biometry of the genus Androcymbium

(Colchicaceae) in southern Africa: taxonomic and biogeographic

considerations

N. MEMBRIVES*, J. MARTIN**, J. CAUJAPE-CASTELLS*** and J. PEDROLA-MONFORTf

Keywords: Androcymbium Willd., biogeography. Colchicaceae, pollen biometry, pollen morphology, southern Africa, taxonomy

ABSTRACT

Pollen characters in 16 southern African type specimens of Androcymbium Willd. that represent the three sections into

which the genus is presently divided ( Androcymbium , Dregeocymbium and Erythrostictus ), were examined to assess the

possibility of sectional characterization based on pollen traits and to explore the relationship between pollen traits and geo-

graphic species distribution. Differences in pollen grain size, shape, number of apertures and exine pattern, were observed.

Principal Component Analysis, using both qualitative and biometric pollen characters, distinguished four groups of species

with four different pollen types. Most of the species have a microreticulate exine pattern with either diaperturate or tria-

perturate grains. The remaining species have either a rugulate exine pattern with diaperturate grains, or a rugulate-reticu-

late exine pattern with thickened (hypertrophied) muri with diaperturate grains. While section Dregeocymbium can be well

delimited by the unique thickened muri. and by a significantly larger grain size, no diagnostic pollen traits were found for

sections Androcymbium and Erythrostictus. Similarly, although the six northern African species exhibit a microreticulate

exine pattern, the high variability observed for this trait in their southern African counterparts, does not follow a geographic

pattern.

INTRODUCTION

The genus Androcymbium Willd. (Colchicaceae) com-

prises ± 50 species (Miiller-Doblies & Miiller-Doblies

1984. 1990. 1998; Arnold & De Wet 1993; Pedrola-

Monfort et al. 1999a, b, in press) whose distribution

(Figure 1 ) embraces arid areas in western southern Africa

(with about 35 species), eastern southern Africa (with six

species), northern southern Africa and eastern Africa

(with two species), the Mediterranean area (with four

species) and the Canary Islands (with two species).

Androcymbium is the only member of the tribe

Colchiceae together with Colchicum L. sensu into (i.e.

including Bulbocodium L., Merendera Ram. and

Synsiphon Regel). At present, the genus is divided into

three sections based on tepal blade morphology ( Krause

1921): section Androcymbium , characterized by green,

curved, winged tepal blades, with representatives

throughout southern Africa; section Dregeocymbium ,

possessing Hat, unwinged tepals and very short stamens

and styles, with representatives in western southern

Africa; and section Erythrostictus , featuring white, Hat,

unwinged tepal blades, with representatives in southern

Africa, the Mediterranean area and the Canary Islands.

However, these differential morphological traits are cur-

rently considered irrelevant (Miiller-Doblies & Miiller-

* Jardf Botanic de Barcelona. C. Doctor Font i Quer s/n. 08038

Barcelona, Spain. Corresponding author: e-mail: jbotanicbcn@ inter-

com.es.

** Laboratori de Botanica. Facultad de Farmacia. Universitat de Barce-

lona. Avda. Joan XXIII s/n 08028 Barcelona. Spain.

*** Jardin Botanico Canario Viera y Clavijo. Apartado 14. Tarifa Alta,

35017. Las Palmas de Gran Canaria, Spain.

t Estacio Intemacional de Biologia Mediterrania-Jardi Botanic Mari-

murtra P.O. Box 112. 17300. Blanes. Spain.

MS. received: 2000-11-06.

20 10 0 10 20 30 40 50

FIGURE 1. — Distribution of the genus Androcymbium.

Doblies 1990, 1998; Pedrola-Monfort 1993) and, conse-

quently, of little taxonomic value. A previous taxonomic

study based on pollen morphological characters of 1 1

Androcymbium species representing the sections

Erythrostictus and Androcymbium (Martin et al. 1993),

92

Bothalia 32,1 (2002)

TABLE 1. — Herbarium specimens of Androcymbium studied in South Africa. Abbreviations of the herbariums follow Holmgren et al. (1990)

KZN. KwaZulu-Natal: NC. Northern Cape; WC, Western Cape.

TABLE 2. — Morphological and biometrical pollen characters of Androcymbium species. Values correspond to the mean ± standard deviations.

Numbers in parenthesis are the minimum and maximum values of each parameter

Ap, no. apertures; EL equatorial long axis; E2, equatorial short axis; MR, microreticulate; P. polar axis; R. rugulate; RA, rugulate-reticulate with

thickened (hypertrophied) muri.

Bothalia 32.1 (2002)

93

offered four important insights. Firstly, there is no single

pollen character that separates the two sections.

Secondly, a microreticulate exine pattern was observed

in all but the Canary Island species, A. psammophilum

Svent., which has a rugulate exine pattern. Thirdly,

whereas the pollen in section Erythrostictus is always

diaperturate, both di- and triaperturate forms co-exist

within section Androcymbium. And lastly, the pollen size

of the western southern African A. bellum, that was his-

torically ascribed to section Erythrostictus , fits the range

of values observed for the species in section Androcymbium.

Given the variability in pollen traits noted in that sur-

vey, we undertook the analysis of all the measurable

pollen characters, the number of apertures and the exine

patterning in a broader geographic representation of the

genus. The aims of this study are to assess whether there

might be traits of diagnostic value for the three sections

of the genus currently recognized and to examine the

relationship between pollen morphology and geographic

distribution.

MATERIAL AND METHODS

Sixteen type specimens of Androcymbium were exam-

ined (Table 1), representing southern African species

housed in the Museum Botanicum Berolinense (B),

Royal Botanic Gardens, Kew (K), and Natal Herbarium

(NH).

Pollen was extracted from the anthers with a drop of

alcohol (96%) and then acetolysed according to the

Avetissian micromethod (Avetissian 1950). For the bio-

metric study, pollen grains were mounted in glycerol

jelly and measured with the image analyser IMAT at the

facilities of the Serveis Cientffico-Tecnics of Barcelona

University.

The four biometric characters used (Table 2), follow

Erdtman (1969) and Reitsma (1970). The arithmetical

mean, standard deviation, and maximum and minimum

values were calculated after 15 measurements for each

parameter (Table 2) using the SPSS/PC+ program (SPSS

1984). Exine patterning and the number of apertures

were determined with the aid of a scanning electron

microscope (SEM). The pollen was first mounted onto

SEM stubs and then coated with gold in a diode sputter-

coater. The stubs were observed using a Hitachi 52300

electron microscope at 15 kv at the facilities of the

Serveis Cientffico-Tecnics of Barcelona University.

A test of comparison of means was conducted using

the program SPSS/PC+ (SPSS 1984) to assess the possi-

ble relationship between the pollen parameters consid-

ered. Principal component analysis (PCA) was conduct-

ed using the program NTSYS-PC (Rohlf 1992) to visu-

alize the relationships among the studied species based

on pollen data. For this multivariate analysis, four con-

tinuous quantitative characters (P, El, E2 and P/El

ratio), one discrete quantitative character (aperture num-

ber) and one qualitative character (exine patterning) were

included. The latter was coded as: 1 = microreticulate, 2

= rugulate, or 3 = rugulate-reticulate with thickened

(hypertrophied) muri. We used the arithmetical mean

values of quantitative characters across species to build a

6x16 data matrix that was standardized and used to cal-

culate the correlation matrix after which the principal

components were derived.

RESULTS

General morphology, the pollen of Androcymbium is

heteropolar, planoconvex in equatorial view and elliptic

in polar view, with bilateral symmetry. This pollen mor-

phology is typical of the Colchicaceae (Erdtman 1952,

1969; Diez 1987; Dfez et al. 1985; Martin et al. 1993).

The values of P. El , E2 and P/El ratio are summarized in

Table 2. The smallest pollen is produced by A. volutare,

8.60 x 14.43 x 8.93 pm, and the largest by A. exiguum

subsp. vogelii, 29.61 x 37.14 x 35.23 pm. The pollen

grains of the Androcymbium species examined are con-

siderably smaller than those of the closely related genera

Colchicum L. and Merendera Ram. (Erdtman 1952,

1969; Faegri & Iversen 1975; Dfez 1987; Dfez et al.

1985).

Among the species studied, the most frequent pollen

shape is oblate (0.5 < P/El < 0.69) and found in A. albo-

marginatum, A. burchellii subsp. pulchrum, A. ciliola-

tum , A. circinatum, A. crispum, A. irroratum, A. scabro-

marginatum, A. villosum, and A. volutare. However, the

grains of A. cruciatum tended to be peroblate (P/El =

0. 5), and those of A. dregei, A. decipiens, A. hantamense,

A. henssenianum, A. natalense, and A. exiguum subsp.

vogelii are suboblate (0.70 < P/El < 0.79).

Apertural system: Androcymbium' s pollen apertures

are circular or elliptic pores. Seven of the species exam-

ined (Table 2) are two-zonoporate, anaporate with two

pores either in zonal or in presumed equatorial position

and the third in distal position (Figure 2A). The remain-

ing nine species are two-zonoporate (two pores in zonal

position. Figure 2C, E, G). We have not studied the tetrad

of pollen grains in Androcymbium , but the position of the

pollen grains in the tetrad was well studied in the close-

ly related genus Colchicum (Beguin & Huynh 1978).

Exine patterning', three different exine patterns were

observed among the species of Androcymbium analysed:

1 , microreticulate (Figure 2A-D: Table 2), with the lumi-

na less than 1 pm diameter, observed in A. albomargina-

tum, A. burchellii subsp. pulchrum, A. ciliolatum, A. cir-

cinatum, A. crispum, A. cruciatum, A. hantamense, A.

irroratum, A. scabromarginatum, A. villosum, and A.

volutare', 2, rugulate (Figure 2E, F), observed in A.

decipiens, A. henssenianum, and A. natalense; and 3,

rugulate-reticulate with thickened (hypertrophied) muri

(Figure 2G, H) and lumina’s diameter higher than I pm,

observed in A. dregei and A. exiguum subsp. vogelii.

Pollen terminology follows Blackmore et al. (1992).

Comparison of means: the ANOVA test revealed sig-

nificant differences among species only in terms of the

number of pores and pollen grain size, with diaperturate

pollen tending to be larger than triaperturate pollen.

Furthermore, there are significant differences in the val-

ues of P, El, E2 and P/El ratio for the exine patterning,

Factor 2

94

Bothalia 32,1 (2002)

FIGURE 2. — Scanning electron micrographs showing the four pollen

types of Androcymbium observed. A, B. A. scabromarginatum,

Type 1: triaperturate microreticulate, arrow points to third aper-

ture. C, D, A. cruciatum, Type 2: diaperturate microreticulate.

E, F, A. henssenianum, Type 3: diaperturate rugulate. G, H. A.

exiguum subsp. vogelii, Type 4: diaperturate rugulate-reticulate

with thickened (hypertrophied) muri.

TABLE 3. — Values of first and second principal components from

multivariate analysis of pollen characters studied

El, equatorial long axis: E2, equatorial short axis; P. polar axis

with rugulate and rugulate-reticulate grains with thick-

ened (hypertrophied) muri, larger than microreticulate

pollen grains.

Principal component analysis : the first and second

principal components (Table 3) explain 93.5% of the

variance of the sample, 79.1% for the first component —

Factor 1; and 14.4% for the second — Factor 2. Whereas

the former is mainly determined by the biometric char-

acters and the exine pattern (P, El, exine patterning, E2

and P/E ratio, in order of importance), the latter is most

influenced by the number of apertures. The relationships

among species according to this analysis are represented

in a two-dimensional graph (Figure 3).

The results of the PCA indicate four groups of species

with different pollen types:

Type 1: triaperturate (two-zonoporate anaporate),

microreticulate (Figure 2A, B), found in A. albomar-

ginatum, A. ciliolatum, A. circinatum , A. hantamense, A.

irroratum, A. scabromarginatum , and A. villosum (sec-

tion Androcymbium)',

Type 2: diaperturate (two-zonoporate), microreticu-

late (Figure 2C, D), observed in A. crispum, A. burchel-

lii subsp. pulchrum and A. volutare (section Androcymbium)

and in A. cruciatum (section Erythrostictus)',

FIGURE 3. — Two-dimensional graph

representing species in the

PCA. Variance of sample is

93.5 % (79.1 % for factor 1,

and 14.4 % for factor 2).

Names of spp. abbreviated

with first three letters of epi-

thet. Distribution of spp.

indicates four pollen types.

Type 1 : in left lower part, A.

albomarginatum, A. ciliola-

tum, A. circinatum , A. hanta-

mense, A. irroratum, A. sca-

bromarginatum, A. villosum.

Type 2: in left upper part: A.

crispum, A. burchellii subsp.

pulchrum, A. volutare, A.

cruciatum. Type 3: central :

A. decipiens, A. hensseni-

anum, A. natalense. Type 4:

on right, A. dregei, A. exigu-

um subsp. vogelii.

Bothalia 32.1 (2002)

95

Type 3: diaperturate (two-zonoporate) rugulate

(Figure 2E. F), observed in A. decipiens, A. hensseni-

anum, and A. natalense (section Androcymbium)-,

Type 4: diaperturate (two-zonoporate), rugulate-retic-

ulate with thickened (hypertrophied) muri (Figure 2G,

H), found in A. dregei and A. exiguum subsp. vogelii

(section Dregeocymbium).

DISCUSSION

Taxonomic implications

According to our results, the southern African species

of Androcymbium contain most of the diversity of pollen

types described for the entire genus (Figure 2). The most

common exine patterning observed in this geographical

region is microreticulate (Figure 2A-D) and was found

both in species from section Androcymbium and in sec-

tion Erythrostictus (A. cruciatum in Miiller-Doblies &

Mtiller-Doblies 1984). Although all the species with tri-

aperturate pollen (Type 1 ) are in section Androcymbium,

this fact is of very limited taxonomic value, since rough-

ly half of the species within this section exhibit diapertu-

rate pollen (Types 2 and 3) and pollen Type 2 was also

observed in most of the North African Androcymbium

species (section Erythrostictus ) by Martin et al. ( 1993).

The diversity of pollen shape notwithstanding, two

additional facts hinder an unambiguous sectional charac-

terization in the genus Androcymbium. Firstly, the pollen

of the taxa within section Androcymbium (A. burcliellii

subsp. pulchrum, A. crispum and A. volutare) is indistin-

guishable from that of A. cruciatum, which belongs to

section Erythrostictus. Secondly, the rugulate exine pat-

terning (Type 3; Figure 2E, F) observed in A. decipiens,

A. henssenianum and A. natalense (section Androcymbium)

was also found in the Canary Island A. psammophilum,

which belongs to section Erythrostictus (Martin et al.

1993). However, the relevance of the latter observation is

only minor because, unlike A. decipiens, A. henssenianum,

and A. natalense, the structural elements of A. psam-

mophilum are always less than 1 pm in diameter and might

be considered as a different pollen pattern.

By contrast, the pollen of the constituents of section

Dregeocymbium, A. dregei and A. exiguum subsp. voge-

lii, can be unambiguously characterized in terms of two

traits: a significantly larger grain size and a rugulate

exine pattern (Type 4, Figure 2G, H).

Section Androcymbium embraces a diverse array of

pollen types, including either microreticulate or rugulate

exine patterning, and di- or triaperturate pollen grains.

Species from section Erythrostictus have microreticulate

exine patterning and diaperturate pollen grains, and share

these features with some of the species of section

Androcymbium.

Biogeographic and evolutionary implications

A microreticulate exine pattern is a general feature of

species in the families Liliaceae and Colchicaceae

(Erdtman 1952, 1969; Valdes 1978; Diez 1987; Diez et

al. 1985) and is considered to be the plesiomorphic con-

dition for the Liliales (Goldblatt 1990; Goldblatt &

Henrich 1991). Although the species of Androcymbium

that have a microreticulate exine pattern (Type 1 and 2)

occur in two of the three broad areas of distribution of

the genus (western southern Africa and North Africa), we

cannot assert that this feature is represented in all areas

of distribution until the study of some species from east-

ern southern Africa (mainly those within the Androcymbium

melanthioides complex) is completed.

By contrast, rugulate forms (Type 3) are considered

evolutionarily recent and have been observed both in

western southern African (A. henssenianum) and eastern

southern African species (A. decipiens and A. natalense).

Rugulate-reticulate exine pattern with thickened (hyper-

trophied) muri can also be considered evolutionarily

recent and is present only in the western southern African

species A. dregei and A. exiguum subsp. vogelii (section

Dregeocymbium). Thus, available evidence indicates that

more recent exine pattern types are exclusive to southern

African species, whereas plesiomorphic forms occur

both in this region and in northern Africa. However, our

survey failed to detect a geographic pattern in the genus,

because southern African species exhibit a high variabil-

ity of exine patterning.

The pollen of the tribe Colchiceae is generally char-

acterized by two, three or four apertures (Erdtman 1952,

1969; Diez 1987; Diez et al. 1985), although four aper-

tures have been reported much less frequently (Dahlgren

et al. 1985). Where the pollen grains are triaperturate, the

two equatorially positioned apertures are pores, whereas

the third one, situated at the distal pole, is an ulcus

(Erdtman 1952, 1969). The dichotomy observed in terms

of number of apertures in Androcymbium has evolution-

ary implications. As a rule, the monosulcate aperture is con-

sidered the most primitive in the monocotyledons (Nair

1970; Goldblatt 1990; Goldblatt et al. 1991). Some authors

argue that the porate apertural system derives from a reduc-

tion of the aperture zone, and the triaperturate form rep-

resents a transition from monosulcate to two-zonoporate

(Thanikaimoni 1986; Takhtajan 1991). Given that species

of Androcymbium having triaperturate pollen occur only in

western southern Africa, our data support the hypothesis

that this area might be the ancestral area of distribution

of the genus (Margelf et al. 1999; Caujape-Castells et al.

1999, 2001). However, this conclusion must be comple-

mented with the pollen morphological survey of the

species of Androcymbium distributed in eastern southern

Africa that could not be included in this work.

CONCLUSIONS

Although our results suggest a general trend towards

parallel evolution of pollen size and structural complex-

ity of exine pattern in Androcymbium, they do not indi-

cate a relationship between pollen morphology and geo-

graphic distribution. Species with pollen Type 2

(microreticulate exine pattern with diaperturate apertural

system) are ubiquitous and occur throughout the range of

the genus. The presence of the third aperture in the distal

96

Bothalia 32,1 (2002)

position in triaperturate pollen, which has been consid-

ered by previous authors to be more primitive than the

diaperturate condition, was observed only in western

southern African species of Androcymbium. Pending the

survey of some of the eastern southern African species

that could not be included in this work, this result

enforces the hypothesis that western southern Africa is

the ancestral area of distribution for the genus. The

pollen characteristics only reflect the currently accepted

sectional subdivision of the genus in the two species of

section Dregeocymbium. These results prompt a thor-

ough re-examination of macromorphological characters

in Androcymbium to assess whether the present sectional

classification should be modified.

ACKNOWLEDGEMENTS

We thank the curators of the Museum Botanicum

Berolinense (B), the Royal Botanic Gardens, Kew (K),

and the Natal Herbarium (NH) for allowing us to take

pollen samples from the type specimens analysed in this

work. We also thank Dra Julia Perez de Paz for the revi-

sion of the manuscript. This study was supported by the

Karl Faust Foundation.

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Bothalia 32.1: 97-100(2002)

Biogeography of Oxalis (Oxalidaceae) in South Africa: a preliminary

study

K.C. OBERLANDER.* L.L. DREYER* and K.J. ESLER*

Keywords: biogeography, diversity, endemism, Fynbos, Oxalis L., refugia. South Africa, Succulent Karoo

ABSTRACT

Oxalis L.. commonly called sorrel, is a large and cosmopolitan taxon that has undergone spectacular speciation within

southern Africa (± 270 taxa), and more specifically within the winter rainfall regions of the western Cape Region (CR). The

main objective of this study was to analyse the geographical distribution of Oxalis in South Africa in relation to currently

defined phytogeographic units. The observed patterns of biodiversity and endemism within South African members of the

genus show interesting disjunctions and concentrations of species. Oxalis is one of the few CR taxa that is shared between

the core Fynbos and Succulent Karoo Biomes, and this study therefore provides a novel insight into evolutionary trends

across, and not only within, these phytogeographic units. The major centre for diversity for Oxalis is situated on Table

Mountain and the northern areas of the Cape Peninsula (grid square 3318CD). Subsidiary centres are located in the

Clanwilliam/Niewoudtville and Kamiesberg regions. The reported patterns in Western Cape suggest that Oxalis species

richness has been generated and retained in areas which have been identified as core Fynbos (Table Mountain), Fynbos refu-

gia during interglacials (Kamiesberg). and an ecotonal region which might switch between the two biome types

(Clanwilliam/ Niewoudtville). Presumably these three types of areas would provide interesting material for DNA-based

phylogenetic work, and a test of the climate change 'species-pump' hypothesis proposed by Midgley et al. (2001).

INTRODUCTION

Oxalis L. (Oxalidaceae) is a large and cosmopolitan

genus, including more than 800 species (Chant 1993).

Most of its range is covered by a limited number of

species, such as O. pes-caprae L. and O. comiculata L.,

which are ubiquitous weeds (Salter 1944).

Oxalis has two well-developed centres of diversity,

which represent over 90% of the species. One centre is

located in South-Central America, including more than

500 species. The other is in South Africa, more specifi-

cally within the Cape Region (CR) (Cowling & Hilton-

Taylor 1997), where ± 150 species are found in an area

of less than 90 000 km2, with extensions both northwards

and eastwards along the coast. In total, southern Africa

hosts ± 270 Oxalis taxa (Dreyer 1996). The Cape

Region, as defined by Cowling & Hilton-Taylor (1997),

coincides with the Fynbos Biome, but also includes

enclaves of karoo and forest, which form part of the

Succulent Karoo and Forest Biomes respectively

(Rutherford & Westfall 1986).

There has been a tremendous degree of speciation

among the CR representatives. A number of other fami-

lies and genera with cosmopolitan distributions exhibit

similar patterns of massive speciation within the region.

Examples include the Proteaceae, with 330 species in the

region, as opposed to roughly 70 species within the rest

of Africa (Rebelo 1995), the Restionaceae with 275

species (Cowling & Richardson 1995), the Mesem-

bryanthemaceae, with 1 800 species (Smith et al. 1998)

and the genus Erica L., with 658 species (Goldblatt &

Manning 2000).

* Department of Botany, University of Stellenbosch, 7600 Matieland.

Stellenbosch.

MS. received: 2000-06-08.

Possible reasons for the dramatic speciation within

the CR have been suggested (Linder 1985; Cowling et al.

1992). A combination of topographic gradients (Linder et

al. 1992) and the regular occurrence of fire (Cowling

1987) have been suggested as the main driving factors

towards speciation. although the latter two explanations

are not entirely appropriate for the Succulent Karoo

Biome, which forms part of the CR. More recently, evi-

dence has been published supporting the ‘species pump'

hypothesis (Midgley et al. 2001) that Pleistocene climate

change played a role in determining plant species rich-

ness and phytogeographic patterns in the greater Cape

Mediterranean region (Goldblatt 1978). Although some

work has been done on the distribution of plant species

within the CR, it has focused almost exclusively on taxa

predominantly confined to the Fynbos Biome, the typical

but not exclusive vegetation type of the CR (Oliver et al.

1983). Despite some cursory observations by Salter

(1944), very little has been published on the distribution

of Oxalis within South Africa. The aim of this study was

to map the distribution of this genus within South Africa,

and to analyse the geographical distribution of Oxalis in

South Africa in relation to currently defined phytogeo-

graphic units.

MATERIALS AND METHODS

Data on the distribution of Oxalis were obtained from

Salter ( 1944) and the Pretoria Computerised Information

System (PRECIS) database. These data were provided in

16th degree grid square increments, and provided distri-

butional information for 237 of the 270 taxa. This is cur-

rently the most complete and up-to-date list of biogeo-

graphical data available for the genus, given the taxo-

nomic uncertainty within several sections of Oxalis.

Nevertheless, the total of 237 represents ± 88% of even

the most comprehensive list of Oxalis taxa, and is thus

considered adequate to represent the entire genus within

98

Bothalia 32,1 (2002)

FIGURE 1. — Oxalis diversity in

South Africa.

the country. Most of the taxa not included are taxonomi-

cally unresolved or vague.

Totals for each 16th degree grid square (which would

provide taxon numbers for that grid square i.e. richness)

and for each taxon restricted to only one, or two adjacent,

16th degree grid squares (i.e. reasonable marker for

endemism levels) were extracted.

These data were then summarized at quarter degree

grid square level, and the repeats eliminated. The figures

were transferred to the appropriate grid squares on maps

of South Africa.

RESULTS

Oxalis is distributed throughout the study area (Figure

I), with an observed increase in diversity and endemism

levels to the south and west of South Africa (with the

notable exception of the Great Karoo), culminating with-

in the CR, where the vast majority of taxa are located.

This pattern is a classic example of the distribution of

many prominent CR families and genera.

Three centres of diversity were identified. The largest

and most speciose is positioned on the Cape Peninsula,

extending to the Kogelberg/Hottentot's Holland Moun-

tain Ranges to the east. Although the two ranges are sep-

arated by the Cape Flats, they are considered as a single

unit for the purposes of this project, because of the high

degree of Oxalis species shared between them (60%).

A secondary centre of diversity is situated in the

Clanwilliam/Niewoudtville region. Although not as spe-

ciose as the Hottentots Holland epicentre, this region

contains a wide range of unique species, both within

Oxalis and in other taxonomic groups (Schumann &

Kirsten 1992; Rebelo 1995). This epicentre is higher in

altitude and further inland than the Cape Peninsula, and

thus has a somewhat different, more arid, climate.

The third main centre of Oxalis biodiversity is situat-

ed in the Kamiesberg region of the Northern Cape. This

is a region of extreme botanical interest as the unique

Namaqualand and Richtersveld areas, containing many

semi-desert paleo- and neo-endemics, surround it. In

addition, the region still harbours fynbos taxa (Cowling

& Pierce 1999), probably isolated there after the retreat

of the last glacial maximum (Midgley et al. 2001).

Other minor centres of diversity occur throughout

South Africa, but at least two can be considered some-

what biased as they are located near or on major cities,

and have therefore been extensively collected. These two

sites are located on the Witwatersrand and on Algoa Bay

grid squares, the first containing the cities of Johannes-

burg and Pretoria and the latter the city of Port Elizabeth.

The Cape Peninsula centre probably also shows some

collecting bias, as it is located on a major city, and is the

site of South Africa’s main Botanical Gardens at

Kirstenbosch. Collection records for the Cape Peninsula

are probably much better than for Namaqualand, in part

due to the brief and ephemeral appearance of Oxalis

species. Collectors in more remote areas, which are vis-

ited only occasionally and not on such a regular basis as

areas around Cape Town, may easily miss these plants.

At present no reason can be offered for the existence of

other sites, such as the one located in Swaziland and in

the East London/Umtata area. The last centre, located on

the eastern Langeberg/Outeniqua, could possibly also be

a southern Cape refugium.

Oxalis is represented throughout most of the rest of

South Africa by only six species, namely O. clepressa

Eckl. & Zeyh., O. obliquifolia Steud. ex Rich., O. obtusa

Jacq., O. semiloba Solid., O. smithiana Eckl. & Zeyh.,

and the naturalized American species O. corniculata L.

All taxa confined to one, or to two adjacent 16th

Bothalia 32.1 (2002)

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