Part 1 RUBIACEAE

Fascicle 2 Rubioideae (Second part)

Paederieae, Anthospermeae, Rubieae

by C. Puff

Botanical Research Institute

Department of Agriculture and Water Supply

Republic of South Africa

FLORA OF SOUTHERN AFRICA

All contributions should be compiled in accordance with the Guide to Contributors to the Flora of Southern Africa

(compiled by Ross, Leistner & De Winter) which is available from the Librarian, Botanical Research Institute, Private Bag

X 101 . Pretoria, 0001 .

Cryptogam volumes will in future not be numbered but will be known by the name of the group they cover. The

number assigned to the volume on Charophyta therefore becomes redundant.

Alien families are marked with an asterisk.

Published volumes and parts are shown in italics.

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INTRODUCTORY VOLUMES

The genera of Southern African flowering plants

Vol. 1: Dicotyledons (Published 1975). Price: R 1 1 ,23. Other countries: R14,00

Vol. 2: Monocotyledons (Published 1976). Price: R8,21. Other countries R10,00

Botanical exploration of Southern Africa (Published 1981). Price: R40,00 (Obtainable from booksellers)

CRYPTOGAM VOLUMES

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(Continued on inside of back cover)

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Printed by the Government Printer and obtainable from the Directorate of Agricultural Information, Department of

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REPUBLIC OF SOUTH AFRICA

REPUBLIEK VAN SUID-AFRIKA

DEPARTMENT OF AGRICULTURE AND WATER SUPPLY

DEPARTEMENT VAN LANDBOU EN WATERVOORSIENING

FLORA OF SOUTHERN AFRICA

VOLUME 31

PART 1, FASCICLE 2

ISBN 0 621 08876 5

G.P.-S.

FLORA OF SOUTHERN AFRICA

which deals with the territories of

SOUTH AFRICA, CISKEI, TRANSKEI, LESOTHO, SWAZILAND, BOPHUTHATSWANA,

SOUTH WEST AFRICA/NAMIBIA, BOTSWANA AND VENDA

VOLUME 31

PARTI RUBIACEAE

Fascicle 2 Rubioideae (Second part)

Paederieae, Anthospermeae, Rubieae

by

C. Puff

University of Vienna, Austria

Edited by

O. A. Leistner

Editorial Committee: B. de Winter, D. J. B. Killick. G. E. Gibbs Russell and O. A. Leistner

Botanical Research Institute,

Department of Agriculture and Water Supply

1986

Digitized by the Internet Archive

in 2016

https://archive.org/details/floraofsoutherna3112unse

CONTENTS

Page

New taxa and new combinations published in part 1 , fascicle 2 vi

Introduction vii

Plan of Flora of Southern Africa viii

Paederieae 1,2:1

Paederia 1 ,2: 1

Crocyllis 1,2:3

Anthospermeae 1 ,2: 7

Anthospermum 1 ,2: 8

Nenax 1,2:37

Galopina 1,2:48

Carpacoce 1 ,2: 52

Rubieae 1,2:61

Rubia 1,2:61

Galium 1,2:65

Index 1,2:77

v

NEW TAX A AND NEW COMBINATIONS PUBLISHED IN PART 1,

FASCICLE 2

Anthospermum basuticum Puff, sp. nov., p. 1 ,2: 20

Anthospermum bicorne Puff, sp. nov., p. 1 ,2: 36

Anthospermum comptonii Puff, sp. nov., p. 1 ,2: 32

Anthospermum esterhuysenianum Puff, sp. nov., p. 1 ,2: 33

Anthospermum esterhuysenianum Puff, var. hirsutum Puff, var. nov., p. 1,2: 33

Anthospermum monticola Puff, sp. nov. 1,2: 19

Anthospermum spathulatum Spreng. subsp. saxatile Puff, subsp. nov.,p. 1,2: 18

Anthospermum spathulatum Spreng. subsp. tulbaghense Puff, subsp. nov.,p. 1,2: 19

Anthospermum spathulatum Spreng. subsp. uitenhagense Puff, subsp. nov., p. 1,2: 17

Anthospermum streyi Puff, sp. nov. , p. 1 ,2: 28

Carpacoce burchellii Puff, sp. nov., p. 1 ,2: 59

Carpacoce curvifolia Puff, sp. nov., p. 1 ,2: 57

Carpacoce gigantea Puff, sp. nov., p. 1 ,2: 56

Carpacoce scabra (Thunb.) Sond. subsp. rupestris Puff, subsp. nov., p. 1 ,2: 59

Carpacoce spermacocea (Reichenb.f.) Sond. subsp. orientalis Puff, subsp. nov., p. 1 ,2: 56

Nenax arenicola Puff, sp. nov., p. 1 ,2: 47

Nenax coronata Puff, sp. nov., p. 1 ,2: 41

Nenax elsieae Puff, sp. nov. , p. 1 ,2: 43

Nenax hirta (Cruse) Salter subsp. calciphila Puff, subsp. nov.,p. 1,2:44

Nenax namaquensis Puff, sp. nov.,p. 1,2:41

Anthospermum dregei Sond. subsp. ecklonis (Sond. ) Puff, comb, etstat. nov., p. 1,2: 32

Anthospermum galioides Reichenb.f. subsp. reflexifolium (Kuntze) Puff, comb, et stat. nov., p.

1,2: 30

Anthospermum pumilum Sond. subsp. rigidum (Eckl. & Ze\h.) Puff, comb, et stat. nov., p.

1,2:26

Anthospermum spathulatum Spreng. subsp. ecklonianum (Cruse) Puff, comb, et stat. nov., p.

1,2:18

Nenax acerosa Gaertn. subsp. macrocarpa (Eckl. & Zeyh.) Puff, comb, et stat. nov. p. 1 ,2: 46

Nenax cinerea (Thunb.) Puff, comb, nov., p. 1 ,2: 40

Date of publication: April. 1986

VI

INTRODUCTION

For a key to the families and the genera not keyed out in this fascicle, the Flora should be used

in conjunction with R. A. Dyer’s Genera of Southern African Flowering Plants, Vol. 1 (1975) and

Vol. 2 (1976), which are arranged on the lines of the Engler system. The genera are numbered, as

far as possible, according to the list published by De Dalla Torre and Harms in their Genera

Siphonogamarum (1900-1907) in order to facilitate reference, though genera in the Flora are not

necessarily arranged in this sequence.

The following condensed abbreviations for literature references are used in this Flora but not

necessarily in the present fascicle:

C.F.A Conspectus Florae Angolensis

R. A. Dyer, Gen. ... The Genera of Southern African Flowering Plants by R. A. Dyer.

Vol. 1 (1975) and Vol. 2(1976)

F.C Flora Capensis

F.C.B Flore du Congo et du Rwanda-Burundi

F.S.W.A Prodromus einer Flora von Siidwestafrika

F.T.A Flora of Tropical Africa

F.T.E. A Flora of Tropical East Africa

F.W.T.A Flora of West Tropical Africa

F.Z Flora Zambesiaca

Burtt Davy, FI.

Transv Manual of the Flowering Plants and Ferns of the Transvaal and

Swaziland, Vol. 1 (1926) and Vol. 2 (1932).

Localities are sometimes referred to in terms of the degree reference system (Leistner &

Morris in Ann. Cape Prov. Mus. 12: 1-565; 1976).

Cited voucher specimens given without indication of herbarium are housed in PRE (National

Herbarium, Pretoria).

This fascicle was compiled in accordance with a Guide to Contributors to the Flora of

Southern Africa (Ross, Leistner & De Winter, 1977), which is available from the Librarian,

Botanical Research Institute, Private Bag X101 , Pretoria, 0001 .

Volume 31 of the Flora, of which the present publication is a component, will appear in parts

of which the first is divided into fascicles (see p. ix). The number of the part, which in the present

publication is ‘1’, and the number of the fascicle, namely ‘2’, precede the page number on all

pages marked with Arabic numerals. This \yas done with a view to compiling a combined index to

the entire volume.

vii

PLAN OF FLORA OF SOUTHERN AFRICA

Cryptogam volumes will in future not be numbered but will be known by the name of the group they cover. The

number assigned to the volume on Charophyta therefore becomes redundant.

Alien families are marked with an asterisk.

Published volumes and parts are shown in italics.

Please note that local prices as given below do not include GST. Prices given for other countries include postage.

INTRODUCTORY VOLUMES

The genera of Southern African flowering plants

Vol. 1: Dicotyledons (Published 1975). Price: R1 1,23. Other countries: R14,00

Vol. 2: Monocotyledons (Published 1976). Price: R8,21. Other countries R10,00

Botanical exploration of Southern Africa (Published 1981). Price: R40,00 (Obtainable from booksellers)

CRYPTOGAM VOLUMES

Charophyta (Published as Vol. 9 in 1978). Price: R4,25. Other countries: R5,30

Bryophyta:

Part 1 : Mosses: Fascicle 1 : Sphagnaceae — Grimmiaceae

(Published 1981). Price: R24,34. Other countries R30,40

Fascicle 2: Gigaspermaceae — Bartramiaceae

Fascicle 3: Erpodiaceae — Hookeriaceae

Fascicle 4: Fabroniaceae — Polytrichaceae

Pteridophyta (in press)

FLOWERING PLANTS VOLUMES

Vol. 1: Stangeriaceae , Zamiaceae, Podocarpaceae, Pinaceae*, Cupressaceae, Welwitschiaceae, Typhaceae, Zostera-

ceae, Potamogetonaceae , Ruppiaceae, Zannichelliaceae, Najadaceae, Aponogetonaceae, Juncaginaceae, Alis-

mataceae, Hydrocharitaceae (Published 1966). Price: Rl,98. Other countries: R2,60

Vol. 2: Poaceae

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Vol. 4: Part 1: Restionaceae

Part 2: Xyridaceae, Eriocaulaceae , Commelinaceae, Pontederiaceae, Juncaceae (Published 1985). Price:

R7,50. Other countries: R9,40

Vol. 5: Liliaceae, Agavaceae

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Vol. 7: Iridaceae: Part 1: Nivenioideae, Iridoideae

Part 2: Ixioideae: Fascicle 1

Fascicle 2: Syringodea, Romulea

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Vlll

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IX

Paederieae

1,2: 1

Tribe PAEDERIEAE

by C. Puff*

Paederieae DC., Prodr. 4: 343 & 470 (1830); Puff in Bot. J. Linn. Soc. 84: 371 (1982).

Shrubs, dwarf shrubs or perennial herbs, often fetid. Flowers <£ , occasionally also corolla

cylindrical to campanulate; filaments inserted above middle of corolla tube; disk present; ovary

2-5-locular, each locule with a single basally attached anatropous ovule. Fruit fleshy to dry,

indehiscent or variously dehiscent. Seed with membranous testa and copious endosperm, remain-

ing enclosed in endocarp. Chromosome numbers : x = 1 1 (2n= 22, 44 in the Flora area).

A tribe centred in the northern hemisphere; only two genera (three infrageneric native taxa) occur in Africa south of the

Sahara [see Puff in Bot. J. Linn Soc. 84; 355-377 ( 1982) for details].

Key to genera

la Climbing shrubs; leaves petiolate, elliptic to ovate, longer than 40 mm; fruits with a shiny

papery exocarp and 2 laterally compressed, winged, glabrous pyrenes Paederia (below)

lb Many-stemmed, intricately branched shrubs or dwarf shrubs; leaves subsessile, linear to ±

elliptic, shorter than 15 mm; fruits (tardily) dehiscing into 2 mericarps with long whitish

hairs Crocyllis (p. 1 ,2: 3)

8430 PAEDERIA

Paederia L., Mant. PI. 1: 7, 52 (1767), nom. conserv.; R.A. Dyer, Gen. 1: 621 (1975); Verde, in

F.T.E.A. Rubiaceae 1: 175 (1976). Type species: P.foetida L.

Hondbessen Adans., Fam. PI. 2: 158 (1763), nom. rej.

Lygodisodea Ruiz & Pav., Prodr. 32, t.5 (1794).

Lecontea A. Rich, ex DC., Prodr. 4: 470 (1830); A. Rich, in Mem. Rubiac. 1 15 (1830) & Mem. Soc. Hist, nat., Paris 5:

195(1834).

Siphomeris Boj., Hort. Maurit. 170 (1837), nom. superfl.

Shrubs, mostly climbing, fetid. Leaves usually petiolate; stipules entire, ± triangular, decidu-

ous. Flowers in axillary or terminal thyrsic inflorescences, or solitary. Calyx : lobes 4—5, persistent.

Corolla : tube cylindrical to narrowly campanulate, often hairy inside, base often splitting into 4-5

parts; lobes 4-5, shorter than tube, valvate in aestivation. Stamens 4-5, inserted near or at mouth

of corolla tube; anthers included or exserted and often at different heights; filaments short. Ovary

2-3-locular, with a solitary erect ovule in each chamber, crowned by a small disk; styles filiform,

free or joined; stigma lobes 2-3, filiform. Fruit subglobose to compressed-ovoid or -ellipsoid;

exocarp thin, papery and brittle, at length breaking open and falling off, exposing 2-3 round to

elliptic, laterally compressed and often distinctly winged 1 -seeded pyrenes which are pendulous

from a detached, persistent median vein. Seed compressed, similar in shape to unwinged part of

pyrene; testa thin; endosperm copious; embryo large, with conspicuous cordate cotyledons. Chro-

mosome number : 2n=44 (tetraploid).

A tropical to subtropical genus of c. 50 species; only two taxa are native to Africa and only one subspecies reaches

Southern Africa.

* University of Vienna, Austria.

Financial support from The Department of Agriculture and Water Supply (Research Fellowship) and the

Hochschuljubilaumsstiftung der Stadt Wien is acknowledged.

1,2:2

Paederieae

Paederieae

1,2: 3

Paederia bojeriana (A. Rich.) Drake in

Grandidier, Hist. Madag. 36, Hist. Nat. PI. t.

412/a (1897); Verde, in F.T.E.A. Rubiaceae 1:

176 (1976), as bojerana. Type: Madagascar,

Bojer (P, holo.).

Lecontea bojerana A. Rich, ex DC., Prodr. 4: 470

(1830); A. Rich, in Mem. Rubiac. 1 15, 1. 10/1 bis (1830) &

Mem. Soc. Hist, nat., Paris 5: 195, t.20/1 bis (1834).

Paederia ligun Sweet, Hort. Brit., App. 487 (1827),

nom. nud. Siphomeris ligun (Sweet) Boj., Hort. Maurit.

170 (1837), nom. superfl.

subsp. foetens (Hiern) Verde, in Kew

Bull. 30: 285 (1975). Syntypes: Mozambique,

north of Sena, N’Keza and by the Shire R.

about the cataracts, Kirk (K!) & ‘Zambesiland’,

Stewart (BM!).

Siphomeris foetens Hiern in F.T.A. 3: 229 (1877). Pae-

deria foetens (Hiern) K. Schum. in Pflanzenfam. 4, 4: 125

(1891) & in Engl. Pflanzenw. Ost-Afr. C: 393 (1895).

Shrub, climbing, fetid. Stems to several

m long, slender and flexuous, pubescent to

velvety hairy. Leaves decussate or in whorls

of 3, petiolate; blades (30— )50— 1 10(— 140) x

(30-)35-80(-l 10) mm, elliptic to ovate, acumi-

nate at apex, cordate or rounded at base,

somewhat pubescent above, densely velvety -

tomentose beneath; petioles (15— )25— 150 mm

long, (velvety) pubescent; stipules to 10 mm

long. Inflorescences mostly axillary, on stalks

c. 40-130 mm long, often ± condensed and

many-flowered, sometimes rather extensive;

pedicels to 1 mm long. Flowers 5-merous. Ca-

lyx: lobes c. 3-8 mm long, subulate, recurved

above. Corolla whitish to greenish yellow,

somewhat pubescent outside; tube 6-10 mm

long, more or less cylindrical, splitting into 5

segments near base, hairy inside at least near

throat; lobes oblong-lanceolate, 3-5 x 1-2

mm, hairy inside at least near base. Anthers

exserted. Ovary c. 2-3 mm long, ovoid, pu-

bescent; style to c. 10 mm long; stigma-lobes to

c. 5 mm long. Fruit elliptic in outline, strongly

compressed, 10-12 x 8-11 mm; exocarp

shiny, yellowish brown to brownish, bearing

persistent calyx lobes; pyrenes slightly smaller

than exocarp, conspicuously winged. Seed

black, c. 3-6 x 3-5 mm. Fig. 1 .

Subsp. foetens occurs from southern Tanzania to the

northern and north-eastern Transvaal; growing in lowland

woodland and riverine scrub. Map 1 .

Vouchers: Codd 5344; Obermeyer 676.

[Subsp. bojeriana is confined to Madagascar, Mauri-

tius and the Comoro Islands.]

MAP 1. — Paederia bojeriana subsp. foetens

8449 CROCYLLIS

Crocyllis E. Mey. [in Dr&ge in Flora 26, Bes. Beigabe 176 (1843), nom. nud.] ex Hook. f. in

Benth. & Hook, f., Gen. PI. 2: 136 (1873); R. A. Dyer, Gen. 1: 623 (1975); Puff & Mantell in

Bot. Jb. 103: 90 (1982). Type species: Anthospermum crocyllis Sond., i.e. Crocyllis anthosper-

moides E. Mey. ex K. Schum.

Gynodioecious, fetid shrub or dwarf shrub. Leaves decussate, subsessile, linear to ± elliptic,

small; stipular sheath narrow, with a small triangular appendage on either side. Flowers shortly

pedicellate, in clusters of several to many, terminal on new growth, $ (and protandrous) or 5-

merous. Stamens inserted at throat of corolla tube. Ovary bicarpellate, each locule with a solitary

erect ovule; style joined; stigma-lobes 2, as long as style. Fruit (tardily) dehiscing into 2 mericarps.

Seed remaining enclosed in mericarp.

Monotypic. Endemic to the southern Namib Desert (southern South West Africa/Namibia and adjacent parts of South

Africa) and centred in the lower Orange River valley. Without close allies in Southern Africa; related to Saharo-Sindian and

Irano-Turanian taxa.

1,2:4

Paederieae

1:IG. 2. — Crocyllis anthospermoides: 1 , flowering branch, x 1;2, node with pair of long shoot leaves and clustered

short shoot leaves, x 5; 3, bud, x 5; 4, $ flower, x 5; 5, J flower, corolla with rudimentary anthers folded open,

longitudinal section of ovary, x 5; 6, fruit, x 3 (1 & 2: Giess 14597; 3 & 4: Giess & Merxmiiller 14302; 5: Puff 78081 1-

1/1 ; 6: Leistner 2588).

Paederieae

1,2:5

Crocyllis anthospermoides E. Mey. [in

Drfege in Flora 26, Bes. Beigabe 92-93, 176

(1843), nom. nud.] ex K. Schum. in Pflanzen-

fam. 4, 4: 132 (1891); Puff & Mantell in Bot.

Jb. 103: 90 (1982). Lectotype: Cape, Drege

s.n. (E!).

Anthospermum crocyllis Sond. in F.C. 3: 32 (1865). Lec-

totype: Cape, Drige s.n. (E!).

Crocyllis intricatissima Dinter in Feddes Reprium Beih

53: 64 (1928), nom. nud.

Shrub, ± erect, many-stemmed, intri-

cately branched, c. 0,6-1, 5 m tall, occasionally

low dwarf shrub; stems and branches whitish

grey. Leaves on younger parts on long and short

shoots, on older parts tightly fasciculate on

much contracted short shoots only; blades

(2— )3,5— 12(— 14,5) x 0,5— 1(— 1 ,5) mm, often ±

round in section, fleshy, occasionally with a

few short hairs, obtuse at apex. Flowers : calyx

lobes c. (0,2— )0,4 — 1 , 1 (—1 ,3) mm long, ±

triangular, erect, usually hidden amongst ovary

hairs; corolla lobes 1— 2,8(— 3,4) mm long, ellip-

tic to lanceolate, ± recurved, pilose on both

surfaces, tube funnel-shaped to ± campanulate,

usually less than half as long as lobes; filaments

0,5-3 mm long; anthers c. 1,5 mm long in $

($ with rudimentary anthers to c. 0,5 mm

long); ovary c. 0,3-0, 8 mm long, densely

hairy; style c. 1-2 mm long. Fruit 1— 2(— 2,5) x

0,5-1, 5 mm, obovate, densely covered with

long white hairs. Chromosome number. 2n—22

(diploid). Fig. 2.

Found on sandy flats, along ephemeral watercourses or

on rocky slopes. Map 2.

Vouchers: Merxmiiller & Giess 3244 (BR; M; PRE;

WIND); Schlechter 11444 (B; BM; BOL; G; GRA; K;

PRE).

The species cannot be confused with any other Rubia-

ceae occurring in the area. Extensively browsed plants are

often low, more or less cushion-like and have spine-tipped

branches (“Crocyllis intricatissima" !). flowers in a male

state (protandry!) should not be mistaken for cf flowers.

Anthospermeae

1,2:7

Tribe ANTHOSPERMEAE

by C. Puff*

Anthospermeae Cham. & Schlechtd. ex DC., Prodr. 4: 343, 578 (1830); Puff in Bot. J. Linn.

Soc. 84: 370(1982).

Anthospermeae (as ‘sectio’) Cham. & Schlechtd. in Linnaea 3: 309 ( 1828).

Shrubs, dwarf shrubs, short-lived subshrubs or perennial herbs. Flowers cf or sexes

variously distributed, plants quite often dioecious; corolla cylindrical to subcampanulate, in $ and

Cf always larger than in filaments inserted at base or at least below middle of corolla tube; disk

absent; ovary mostly 2- or 1 -locular, each locule with a single basally attached anatropous ovule;

style ± 0 or very short; stigmas long, hairy. Fruit dry or fleshy, dehiscing into 2 mericarps or into

exocarp-valves and endocarp plus seed, or indehiscent. Seed with membranous testa and copious

endosperm, remaining enclosed in endocarp. Chromosome numbers: x= 1 1 (2n=22, 44, 66 in the

Flora area).

A tribe centred in tropical, subtropical and temperate regions of the southern hemisphere.

All African and Madagascan taxa belong to subtribe Anthosperminae , which is distinguished by dry fruits and

unisexual and/or protandrous y flowers.

Wind-pollination (flowers small and inconspicuous, odourless, without disk, often unisexual, anthers dangling on

long, slender filiform filaments, stigmas long exserted, hairy all around, etc.) is characteristic for the entire tribe. See Puff

in Bot. J. Linn. Soc. 89: 357 ( 1982) for further details.

Key to genera

la Fruit separating into exocarp-valves and endocarp plus seed; flowers ^ or (less com-

monly) cf, mostly with only 1 fertile carpel and 1 stigma; corolla (4-)5-7-merous, lobes

distinctly hooded; calyx lobes mostly large, leaf-like; south-western Cape only

Carpacoce(p. 1 ,2: 52)

lb Fruit either separating into 2 indehiscent mericarps or indehiscent; flowers cf, cf, $,

sexes variously distributed, with 2 fertile carpels' and 2 stigmas; corolla 4-5-merous,

lobes not hooded; calyx lobes mostly small or subobsolete:

2a Perennial herbs with rhizomes or rootstocks; leaves strictly decussate, blades relatively

large and thin, distinctly petiolate; inflorescence terminal, paniculate to thyrso-pani-

culate; fruit without calyx lobes, separating into two mericarps, never supported by a

carpophore * 1 2; south-eastern Africa Galopina (p. 1 , 2: 48)

2b Large shrubs, dwarf shrubs, short-lived subshrubs or perennial herbs; leaves decussate

or in whorls of 3 or 4, blades mostly small and narrow, frequently ericoid, often

without distinct petioles; inflorescence mostly variously congested, frequently much

reduced and inconspicuous, in dioecious taxa often dimorphic (in 9 more conspicuous

than in cf ); fruit often with small calyx lobes, separating into two mericarps and sup-

ported by a carpophore 2 or indehiscent, without carpophore:

3a Growth form variable; leaves relatively large and broad to small and ericoid, decus-

sate or in whorls of 3 or 4; flowers cf, 9, sexes variously distributed; $, Cf

with long, cylindrical to short, broadly funnel-shaped to campanulate tubes; $

mostly with whitish to greyish (green) stigmas; fruit always dehiscing into two

merricarps always supported by a carpophore2; widely distributed in Southern

Africa Anthospermum (p. 1 , 2: 8)

3b Many-stemmed, often intricately branched dioecious dwarf shrubs; leaves frequently

ericoid, mostly decussate; cf flowers never with long, cylindrical tubes; mostly

with reddish-purplish stigmas; fruit dehiscent or indehiscent, sometimes distinctly

inflated, not always supported by a carpophore; centred in the south-western and

western Cape Nenax (p. 1 , 2: 37)

* University of Vienna, Austria.

1 Except for two Anthospermum species.

2 A ±U-shaped outgrowth on a pedicel which supports the two mericarps of a fruit (see Fig. 5).

Anthospermeae

1,2:8

8438 ANTHOSPERMUM

Anthospermum L., Sp. PI. 1058 (1753), Gen. PI., edn 5: 479 (1754); Hook. f. in Benth. & Hook,

f., Gen. Plant. 2: 140 (1873); Sond. in F. C. 3: 26 (1865); K. Schum, in Pflanzenfam. 4,4: 129

(1891); R. A. Dyer, Gen. 1: 622 (1975); Verde, in F.T.E.A. Rubiaceae 1: 324 (1976). Type

species: A. aethiopicum L.

Ambraria Heister [Syst. Plant. 11 (1748)] ex Fabr., Enum. Meth. Plant. (Hort. Med. Helmstad.), edn 2: 435 (1763)

[based on A. aethiopicum ]; non Ambraria Cruse [See Nenax).

Large shrubs, dwarf shrubs, short-lived subshrubs or perennial herbs. O', + $, or

occasionally c? + 9"+$>9"+9"orG' + $- Leaves decussate or occasionally in whorls of 3 O’

or cf (rarely 4), often seemingly in much larger numbers at nodes1, blades ± broad and large to ±

ericoid and small, mostly narrowed to base, acute to acuminate (seldom ± mucronate of ± obtuse)

at apex, shortly petiolate to sesile, with ± cup-shaped stipular sheaths bearing one to many seae2or

fimbriae on either side. Inflorescence frequently leafy and inconspicuous, made up of mostly

subsessile, many- to very few-flowered cymes, in dioecious taxa often sexually dimorphic (y

inflorescence contracted, ± cylinder-like; Fig. 3: 1,2). Flowers mostly subsessile, subtended by a

pair of leafy bracts, cf, $ or 9. 4-5-merous. Calyx: lobes large, conspicuous to small, indistinct

or ± lacking. 9", cf: Corolla : tube ± cylindrica, broadly funnel-shaped to subcampanulate, lobes

recurved, ± lanceolate; anthers yellowish to whitish, exserted, dangling on long slender filiform

filaments. $: corolla much smaller; tube cylindrical, lobes mostly erect, linear to ± lanceolate.

Ovary bicarpellate and biovulate, in 2 species one carpel reduced; style 0 or very short; stigmas 2,

only in A. ericifolium (no. 20) 1 , long exserted, hairy, greyish to greenish white, seldom purplich

red. Fruit crowned by persistent calyx lobes, supported by a ± U-shaped carpophore (Fig. 5),

dehiscing into two mericarps, each convex on dorsal side, plane to concave or sometimes hollowed

out and with a prominent to inconspicuous median longitudinal ridge on ventral side (Fig. 5:

6, S). Chromosome number : 2n=22, seldom 44, 66.

A genus of 39 species widely distributed in Africa south of the Sahara and in Madagascar. The highest concentration of

taxa is found in Southern Africa; 21 species (29 taxa) occur in the Flora area, of which 18 spp. (26 taxa) are endemic; of

these, 10 spp. (16 taxa) are endemic (or nearly so) to the south-western Cape Region. Map 3.

The inexperienced and those without field knowledge of the plants may often find it difficult to distinguish species

from each other. Some of the reasons for this are:

— Environment-induced morphological variability. Individuals exposed to fire and capable of resprouting from a

woody base often differ markedly from shrubby plants of the same taxon sheltered from fire. Browsing, especially in the

drier parts of the Flora area, frequently causes considerable morphological changes.

— More widely distributed taxa often contain a number of “Forms” (in the sense of ecotypes of geographical races)

which may be very distinct locally. They are, however, often connected with other “Forms” by a series of morphologically

intermediate populations so that it is not possible to clearly define and delimit them or to recognize them taxonomically . The

more prominent of these are pointed out in the text and included in the key [e.g. A. galioides subsp. galioides (‘Papillatum

Form’, etc.)].

— The occurrence of characters which are constant in some but variable in other taxa. Notably the arrangement of the

leaves-e.g. decussate or in whorls of 3 in A. bergianum (variable even within populations), or either strictly decussate or

strictly in whorls of 3 (e.g. A. spathulatum vs. A aethiopicum). For this reason (and also in view of the variable sex

distributions within a taxon; see below) taxa should be studied with care in the field and representative samples should be

collected.

— The occasional occurrence of flowers ‘transitional’ between 9 and ‘pure’ 9- Such ‘transitional’ flowers have

corollas intermediate in size between (jf and 9 and are characterized by the presence of small but clearly discernible, (±)

pollenless anther rudiments (Fig. 4d-f); ‘pure’ 9 with small corollas have no rudimentary anthers (fig. 4g). In some

essentially dioecious taxa, cf flowers do occasionally revert to in some it is only odd 9 flowers on a single cf plant of a

population, in others a considerable number of individuals may show this phenomenon. Also the reverse (transitions from 9

to cf ) is possible ( Note : the flowers are always protandrous; y flowers at anthesis-i.e. with stigmas which have not yet fully

elongated-should, therefore, not be confused with such transitional flowers). In both transitions cf — » 9" anc* V' — * & t^lere

are, however, no marked corolla size and shape differences.

1 Due to the presence of leafy much-contracted short shoots (Fig. 3: 3,4); this situation is referred to as ‘pseudoverticillate’

in the key and in the descriptions.

Single median setae are often flanked by a pair of minute gland-tipped setae. As the latter are often no longer discernible

on older leaves/stipular sheaths, no mention is made of them in the following descriptions.

Anthospermeae

1,2:9

— Sex dimorphism in dioecious taxa. Differences between the sexes may go beyond corolla size and shape differ-

ences; dimorphic inflorescences are rather common (contracted and often quite conspicuous in $; compare Fig. 3: 1 and 3:

2); occasionally cf and $ differ somewhat in leaf size and shape. It is recommended that both cf and $ are collected

whenever possible.

Because of the lack of other suitable and usable characters, it is often necessary to use calyx lobe, corolla or fruit sizes

or hair lengths as differential characters. It should be noted that measurements exact to tenths of millimeters are essential for

correct identifications.

In the following two regional keys taxa which may extend from one area to the other are included in both keys.

Especially in the eastern Cape a considerable mixing of south-western Cape species and species centred in the summer

rainfall area takes place, often making it extremely difficult to identify taxa with certainty.

KEY I

Key to taxa occurring in the Flora area excluding the Cape Floristic Region (the south-western Cape)

la Flowers in distinct terminal thyrsic or thyrso-paniculate inflorescences (from Transkei and eastern Cape to south-

western Cape 8 . A . paniculatum

lb Flowers in clusters of many to few at nodes; in 9 plants sometimes in ± condensed, cylindrical inflorescences:

2a Plants of coastal sand dunes; at least lower parts of stems usually buried in sand; mericarps densely covered

with spreading hairs to 0,8 mm long 3 .A. littoreum

2b Habitat and habit not as above; mericarps glabrous or variously hairy, but not as above:

3a Often scrambling, straggling or trailing perennial herbs; and cf flowers with cylindrical corolla tube to

3,7 mm long 10. A. herbaceum

3b Shrubs or subshrubs, erect, mat- or cushion-forming perennial herbs; Cf and cf flowers with corolla tube

much shorter and broader (funnel-shaped) or, if tube long and cylindrical, habit not as above:

4a Leaves densely covered with spreading hairs c. (0, 1— )0,3— 0,5 mm long;

5a Leaves 2,5— 5(— 6) x 0,8— 1 ,2(— 1 ,8 ) mm; mericarps 2-2,5 x 1-1,5 mm; dioecious, cushion- or

mat-forming dwarf shrubs of high altitudes (high Drakensberg areas of Cape, Natal, Lesotho and

Orange Free State) 7. A. basuticum

5b Leaves 5— 1 2(— 20) x 1— 3 ,5(— 5 ) mm; mericarps 1,7-2 x 0,7-1 mm; plants not dioecious, not occurring

above 2000(— 2300) m (from Transvaal and Swaziland to Transkei) 9. A. hispidulum

4b Leaves glabrous, or hairs on margins and/or midvein only:

6a Low, ± cushion-forming dwarf shrubs, rooting at nodes, occurring at high altitudes (above 2400 m;

Lesotho) 6. A. monticola

6b Growth form different, plants not rooting at nodes, confined to lower altitudes or, if at high

altitudes, perennial herbs:

7a Rounded dwarf shrubs, low subshrubs or perennial herbs; mericarps roundish in side view; plants

not dioecious:

8a Leaves linear to narrowly ovate-lanceolate, to 1 ,5(— 2) mm wide; mericarps glabrous or with ±

hooked or curled hairs:

9a Leaves mostly distinctly needle-like, to 0,8(— 1 ) mm wide; often mat- or cushion-forming, very

densely leafy dwarf shrubs (Natal South Coast only) 12. A. streyi

9b Leaves to 1 ,5(— 2) mm wide, sometimes linear but not distinctly needle-like; subshrubs with

numerous unbranched stems or ± robust dwarf shrubs:

10a Mostly subshrubs with numerous unbranched stems; leaves (4 — )6— 12(— 22) x

(0,5— )0,8— 1 ,5(— 2) mm, linear to narrowly lanceolate; 9" and cf flowers with corolla tube

to 1 ,4(— 1 ,7) mm long, lobes to 1 ,9(— 2,5) mm long (from Transvaal to eastern and north-

eastern Cape) 11 (a). A. pumilum subsp. pumilum

10b Fairly robust dwarf shrubs; leaves 3— 6(— 8 ) x 1-1,5 mm, ovate-lanceolate, often tough and

thickish; <jf and cf flowers larger, corolla tube to 1 ,5(— 1 ,9) mm long, lobes to 3(— 3,2) mm

long (Orange Free State and Cape excl. the south-western Cape)

11 (b). A. pumilum subsp. rigidum

8b Leaves broader, often ovate or ovate-lanceolate, to 6(— 10) mm wide; mericarps glabrous, ±

tuberculate or papillate, but never with curled or hooked hairs:

1 la Diffusely and divaricately branched robust dwarf shrubs with tough, thickish, ovate to oblong-

lanceolate leaves to 4(-4.5) mm wide; fruits without ± broad, conspicuous longitudinal

grooves between mericarps (Namaqualand and southern S.W. A. /Namibia)

15 (a). A. dregei subsp. dregei

1 lb Low cushion- or mat-forming perennial herbs or ± erect and tufted subshrubs; fruits mostly

tuberculate and with ± broad, conspicuous longitudinal grooves between mericarps (Fig. 5:5):

12a Leaves ± succulent in nature; stipular sheaths often with only 1 seta; near the sea (Natal

and Transkei) 10. A. herbaceum (‘Salt Spray Zone Form')

12b Leaves not succulent; stipular sheaths with several setae; growing at high altitudes or in

(burnt) grassland:

1,2: 10

Anthospermeae

13a Low, matted or cushion-forming perennial herbs; leaves small; partial inflorescences few-

to one-flowered 10. A. herbaceum (‘High Altitude’ or ‘Trampled Grassland Form’)

13b ± Erect, several- to many-stemmed subshrubs with ± massive, almost rosette-like ±

woody base 10. A. herbaceum ( ‘ Burnt Grassland Form')

7b Single- to several-stemmed, mostly erect shrubs to 3 m tall; mericarps elongated in side view;

plants dioecious:

14a Leaves in whorls of 3 (sometimes also decussate- 15b):

15a Leaves to 9(— 13) mm long, strictly in whorls of 3; mericarps glabrous or papillate; 9

plants mostly with conspicuous, condensed cylindrical inflorescences (Fig. 3.2) (south-western

Cape species extending into the eastern Cape) 4 .A. aethiopicum

15b Leaves to 20(— 27) mm long, sometimes also decussate; mericarps densely covered

with straight hairs c. 0, 1-0,2 mm long; 9 plants mostly not with very conspicuous, con-

densed intlorescences (Natal to eastern Cape) 2. A. galpinii

14b Leaves decussate:

16a Leaves (0,5— )0,8— 1 ,5(— 2) mm wide, narrowly oblanceolate to ± linear; mericarps densely

covered with straight hairs c. 0, 1-0,2 mm long 2. A. galpinii

16b Leaves wider or, if linear (-lanceolate), mericarps not shortly hairy:

17a Leaves (7—) 10— 35 x ( 1-)1 ,5-3,5 mm, oblanceolate to elliptic; stipular sheaths with 3— 5(— 7 or

8) setae; 9 plants mostly with conspicuous, condensed cylindrical inflores-

cences usually large, single-stemmed shrubs (Transvaal; and Cape: Mafikeng?)

l.A. welwitschii

17b Leaves (2— )3— 1 2 X 0,5-2 mm; stipular sheaths mostly with a single median seta; 9 plants

usually not with conspicuous cylindrical inflorescences; mostly small, several- to many-

stemmed shrubs:

18a Leaves (ob)ovate, oblong or (ob)ovate-lanceolate, (0,7—) 1—2 mm wide; fruits with con-

spicuous calyx lobes 0,3-0,6(-0,8) mm long (Natal, Lesotho, Orange Free State and

interior of Cape to Nieuweveld Mts) 6. A . monticola

18b Leaves lanceolate to linear-lanceolate, 0,5-1, 5 mm wide; fruits with indistinct or smaller

calyx lobes:

19a Mericarps narrowly obovate to oblong, not distinctly ribbed on dorsal side, often see-

mingly truncate above (eastern Cape; Humansdorp distr. to Great Kei River)

5(b). A. spathulatum subsp. uitenhagense

19b Mericarps broader, mostly with 3 ± distinct ribs on dorsal side and with calyx lobes to

0,4 mm long (south-western Cape taxon, extending into eastern Cape)

5(a). A. spathulatum subsp. spathulatum

KEY II

Key to taxa occurring in the Cape Floristic Region (the south-western Cape)1

la Leaves in whorls of 3:

2a Tall shrubs with small, ± needle-like glabrous leaves to 1 ,2(— 2) mm wide; corolla lobes 4 (widely distributed)

4. A. aethiopicum*

2b Subshrubs or dwarf shrubs; leaves mostly ± imbricate, to 3(— 3 ,5 ) mm wide, at least margins ciliate; corolla

lobes 5 (south-western Cape s.str.:) 19. A. bergiamim*

lb Leaves decussate:

3a Fruits with only I fertile, well developed carpel/mericarp (south-western Cape s. str . ):

4a Reduced carpel with 2 calyx lobes c. 1 ,4-2 mm long, fertile carpel with 3 much smaller ones; leaves needle-

like, to0,5 mm wide 21. A. bicorne*

4b Calyx lobes of fertile and reduced carpel subequal, c. 0,2-0, 6 mm long; leaves linear-lanceolate, to 1 ,2(— 1 ,4)

mm wide 20. A. ericifolium*

3b Fruits with 2 fertile carpels/mericarps:

1 The geographical delimitation of the Cape Floristic Region follows Bond & Goldblatt in J1 S Afr. Bot., suppl. 13 ( 1984).

i.e. roughly in the area from the Clanwilliam distr. south to the Cape Peninsula and to the Caledon and Bredasdorp distr.

Taxa endemic to the south-western Cape or nearly so (i.e. sometimes extending to the eastern Cape, into the drier interior

of the Cape, etc.).

Anthospermeae

1,2: 11

5a Flowers in distinct terminal thyrsic or thyrso-paniculate inflorescences (from George distr. to eastern Cape and

Transkei) 8. A. paniculatum

5b Flowers in clusters of many to few at nodes, or (seldom) solitary and terminal; in 9 plants sometimes in ±

condensed, cylindrical inflorescences:

6a Plants prostrate (but not mat-forming), with long, trailing stems radiating out from a common base, rooting

at nodes and bearing short, ascending to ± erect lateral branches, dioecious; only in coastal dune sand

(from West Coast to Port Elizabeth distr.) 14. A. prostratum*

6b Growth form different, if prostrate, stems not rooting at nodes and not with short ascending to erect lateral

branches, plants not dioecious:

7a Leaves, stipules and stems (or at least leaf margins near base) ± densely covered with whitish spreading

hairs c. 0,2— 1(— 1 ,5) mm long; corolla 5-merous:

8a Flowers solitary and terminal on branches, often overtopped by lateral shoots arising from below;

plants usually mat-forming; leaves 4— 6(-7,5) x ( 1 ,4 — )2— 3(-^4) mm, ovate to lanceolate (confined

to higher parts of Cape mountains) 17 (b). A. esterhuysenianum var. hirsutum*

8b Flowers in axillary clusters; plants not mat-forming; leaves larger, (4 — )5— 25(— 30) x (0,5-)0,8-3(-4,5)

mm, ovate-lanceolate to linear-lanceolate:

9a Plants mostly dioecious (cT plants occasionally with odd $ flowers), densely leafy; leaves mostly ±

imbricate, blades ( 4 — )5— 1 2(— 1 5 ) x (1 ,2—) 1 ,5— 3(— 3,5) mm; 9 plants with ± condensed cylindrical

inflorescences 19 . A. bergianum*

9b Plants not dioecious, not densely leafy; blades larger, (6— )8— 25 x (0,5-)0,8-3(-4,5) mm; 9 in-

florescences inconspicuous 18. A. hirtum*

7b Leaves and stems glabrous, very shortly hairy or papillate; if leaves with hairs to 0,4 mm long, hairs

confined to margins and corolla 4-merous:

10a Plants dioecious; 9 plants sometimes with ± conspicuous cylindrical inflorescences:

11a Tall single-stemmed shrubs or many-stemmed dwarf shrubs; c f corolla tube short, to 1,2 mm long,

(broadly) funnel-shaped to ± campanulate; mericarps narrow and elongated in side view, often

±triangular in section and 3-ribbed; (very variable and very widely distributed):

12a Mericarps ( 1 ,5— )1 ,7—2,3 x 0,6— 0,8(— 1 ) mm, ± oblong, glabrous or shortly hairy, not distinctly

ribbed; cf corolla lobes c. 1 ,2— 2(— 2,2) mm long; leaves (2,2— )2,5 — 4,5(— 5,2) x 0,5— 1(— 1 ,3)

mm (south-western Cape s.str.) 5 (e). A. spathulatum subsp. tulbaghense*

12b Mericarps larger, 2, 2^1, 2 x 1-2,2 mm, (broadly) obovate to ± oblong, often 3-ribbed, usually

glabrous; cf corolla lobes ( 1 ,5—) 1 ,9—3 ,4 mm long; leaves (3— )3,5— 18(— 25) x

0,5-2,5(-3) mm:

13a Leaves 8—1 8(— 25) X 0,5—1 ,8(— 2,2) mm, linear-(ob-)lanceolate to linear, fine and ± needle-

like; mostly single-stemmed (± narrowly) cylindrical, densely leafy shrubs c. 1-2 m tall,

in habit and appearance resembling A. aethiopicum; 9 plants with rather conspicuous,

condensed inflorescence zones 5 (d). A. spathulatum subsp. ecklonianum*

13b Leaves smaller and relatively broader, (3— )3 ,5— 1 2 x 0,5-2,5(-3) mm; growth form not as

above, plants often smaller, single- to many-stemmed shrubs or dwarf shrubs; 9 plants

with less conspicuous condensed inflorescence zones:

14a Several- to many-stemmed low cushion- or ± mat-forming dwarf shrubs with rather tough,

closely spaced leaves; partial inflorescences 3-1-flowered; mericarps mostly with con-

spicuous calyx lobes to 1 ,5(— 1 ,9) mm long (confined to highest parts of Cape moun-

tains) 5 (c). A. spathulatum subsp. saxatile*

14b Rather tall, single-stemmed shrubs or small, often several-stemmed dwarf shrubs; partial

inflorescences ± many- to few-flowered; mericarps with calyx lobes to 0,6(-0,9)

mm long; (variable and widely distributed) 5 (a). A. spathulatum subsp. spathulatum*

lib Subshrubs or low dwarf shrubs; cf corolla tube to 1 ,7(— 2) mm long, ± cylindrical to narrowly

funnel-shaped; mericarps broad and roundish in side view, ± semi-terete in section, never

ribbed:

15a Leaf blades distinctly recurved (at least near tip); cf corolla lobes 1 ,4-2,4(-2,9) mm long; meri-

carps c. 2-2,4 mm long; (± widely distributed, especially in mountains)

13 (b). A. galioides subsp. reflexifolium*

15b Leaf blades straight (but margins revolute); cf corolla lobes 2, 7-3, 2 mm long; mericarps

2,7-3 mm long (only in the interior; often in dry, karroid valleys) 16. A. comptonii*

10b Plants not strictly dioecious; sex distributions variable (plants + 9- 9- etc.); 9 plants never

with conspicuous ± condensed cylindrical inflorescences:

16a Plants distinctly woody and dwarf shrubby, erect and ± cylindrical to ± rounded:

17a Leaf blades distinctly recurved (at least near tip); (± widely distributed, especially in moun-

tains) 13 (b). A. galioides subsp. reflexifolium*

17b Leaf blades not distinctly recurved:

1,2: 12

Anthospermeae

18a Fruits papillate (or occasionally shortly hairy); blades narrowly lanceolate to linear (due to

strongly revolute margins), c. 5-10 x (0,5-)0,8-l,5 mm, margins glabrous (south-

western Cape s.str.) 13 (a). A. galioides subsp. galioides* (‘Papillatum Form’)

18b Fruits mostly glabrous; blades broader, oblong or obovate-lanceolate (less commonly linear-

lanceolate), 3— 9(— 12) X 1— 2,5(— 3) mm, margins often very shortly hairy:

19a Several-stemmed, ± erect dwarf shrubs; mericarps 2— 2,6(— 2,8) mm long; cf and $ corolla

glabrous, lobes ( 1 ,7-)2,2-3(-3,2) mm long; (only in the Gouritz River valley)

11 (b). A. pumilum subsp. rigidum

19b Often rounded, low dwarf shrubs; mericarps ( 1 ,5— )1 ,8— 2,2 mm long; cf and 9 corolla

mostly a little hairy outside, lobes 1 ,5— 2,5(— 3) mm long; (only in western part of south-

western Cape s.str. , confined to TMS areas) 15 (b). A. dregei subsp. ecklonis*

16b (Short-lived) perennial herbs or subshrubs; if ± dwarf shrubs, plants prostrate and mat-forming;

20a Straggling, trailing or ± ascending, sometimes rather short-lived perennial herbs; leaves

mostly ± lanceolate, to 30 mm long; stipular sheaths with 3-5 setae; cf and corolla tube

cylindrical, to 3,7 mm long; fruits often ± tuberculate and with ± broad, conspicuous

longitudinal grooves between mericarps (Fig. 5:5); (± afromontane species; from Heidelberg

distr. eastwards) 10. A. herbaceum

20b Growth form different; leaves smaller, to c. 10 mm long; stipular sheaths mostly with 1 seta;

Cf and 9" corolla tube to 1 ,8(— 2) mm long; fruits not as above:

21a Plants often ± mat-forming, stems prostrate to ± ascending; leaves relatively broad, to

j(-4; mm wide, ovate, oolong to ± lanceolate:

22a Flowers solitary and terminal on branches, often overtopped by lateral shoots arising

from below; corolla 5-merous; (confined to higher parts of Cape mountains)

17 (a). A. esterhuysenianum var. esterhuysenianum*

22b Flowers in axillary clusters; corolla 4-merous; (not occurring in Cape mountains)

13 (a). A. galioides subsp. galioides* (‘Prostrate Form’)

21b Plants not mat-forming, stems erect to ± ascending; leaves narrower, to 2(— 3) mm wide,

linear-lanceolate to ovate-lanceolate:

23a Leaf blades distinctly recurved (at least near tip) 13 (b). A. galioides subsp. reflexifolium*

23b Leaf blades straight, spreading to ascending (but margins occasionally revolute)

13 (a). A. galioides subsp. galioides*

1. Anthospermum welwitschii Hiern,

Cat. Afr. PI. Welw. 2: 500 (1898); Brenan in

Mem. N.Y. bot. Gdn 8: 455 (1954); Verde, in

F.T.E.A. Rubiaceae 1: 330 (1976). Type: An-

gola, Huila, Panda forests, near Erne, Wel-

witsch 5335 (BM; G; K, iso. !). '

1 The holotype, according to Verdcourt (op. cit.) in LISU,

was not amongst LISU material sent to me on loan.

A. cliffortioides K. Schum. in Bot. Jb. 30: 416 (1901).

Type: Tanzania, Usafua, Beya-Berg (=Mbeya Mt.),

Goetze 1082 (B, holo.t; BM; BR; E; G; K, iso.!).

A. ammannioides sensu auett. Afr. austr., non S. Moore.

Dioecious shrub, single- or (seldom) seve-

ral-stemmed, ± erect, c. 1-3 m tall. Stems

mostly much branched above, branches usually

± regular, paired, often ascending. Leaves de-

cussate, mostly pseudo verticillate; blades

(7-)10-35 x (1— )1 ,5—3,5 mm, (narrowly) ob-

lanceolate, elliptic to ± linear-lanceolate,

mostly glabrous; petioles ± 0-1 mm long; sti-

pular sheath with 3-5 (-7,8) setae, the longest

c. (0,5-)0,7— 4,5(-5,6) mm. Flowers subsessile

to shortly pedicellate (pedicels to 0,7 mm in

Cf), in clusters of many (very many: $) at

nodes, inflorescences dimorphic, in $ often

quite contracted, dense, ± cylindrical inflores-

cence zones; corolla 4-merous, greenish yellow

to pale yellow, occasionally purplish tinged

outside, mostly glabrous. cf: tube

(0,5-)0,7-l ,2 mm long, funnel-shaped, lobes

( 1 ,2—) 1 ,5— 2,2(— 2,7) x (0,4-)0,6-0,9(-l ,1)

mm; anthers 1-2 mm long; small rudimentary

ovary with 4 minute calyx lobes. 9: tu^e

0,2-0, 5 mm long, lobes 0,2-0, 7 x 0, 1-0,2

mm; style ± 0-1 mm long, stigmas 2,

Anthospermeae

1,2: 13

3— 7,5(— 10) mm long; ovary c. 0,6-1 x 0.3-0, 6

mm, with 4 sometimes unequal calyx lobes.

Fruit mostly reddish brown, shiny; mericarps

1,5-2, 7 x 0,7-1 ,2 mm, oblong, elliptic to ±

obovate, glabrous or (seldom) ± sparsely

shortly hairy, with 2 ± triangular calyx lobes c.

(0,3-)0,5-l(-l,2) x 0,2-0, 5 mm, one occasio-

nally longer than the other. Chromosome

number: 2n=22.

Known from Transvaal; extending to tropical East

Africa and Angola. Typically growing at the edge of afro-

montane forest or scrub, sometimes in disturbed sites. Map

4.

Vouchers; Acocks 23578 (BR; K; M; PRE); Edwards

4055 (K; PRE; SRGH); Scheepers 851 (B; BM; K; M; PRE;

SRGH).

A record from Mafikeng (Cape, 2525-DC: Kassner

1541, E!) appears highly doubtful (not mapped). In the

Flora area A. welwitschii is rather uniform in its characters.

A. ammannioides, with which it has consistently been con-

fused, occurs in the eastern highlands of Zimbabwe and on

Gorongosa Mt. and is distinguished by its leaves in whorls

of three.

2. Anthospermum galpinii Schltr. in J.

Bot., Lond. 35: 342 (1897). Type: Transkei,

West Gate above Port St. Johns, Galpin 3472

(BOL, holo. !; GRA; K; PRE; iso. !).

Dioecious shrub, mostly single-stemmed,

erect and ± cylindrical to roundish, c.

0,6-2(-2,5) m tall [several-stemmed, often not

more than 0,25 m tall if burnt]. Stems much-

branched above, branches mostly ± erect to

ascending, often in threes. Leaves in whorls of

3 or (seldom) decussate, pseudo- verticillate;

blades (6-) 10-20(— 27) x (0,5— )0,8— 1 ,5(— 2,5)

mm, ± narrowly oblanceolate, linear-lanceo-

late to ± linear, glabrous or sometimes lower

half of midrib below shortly hairy; petioles sub-

obsolete; stipular sheath with a small seta.

Flowers subsessile or cf occasionally with pe-

duncles to 2 mm long, in clusters of many to

few at nodes, 9 inflorescences sometimes ±

contracted and conspicuous; corolla 4-merous,

yellow to greenish yellow, shortly hairy or gla-

brous (especially 9)- Cf: tube 0,4—1 mm long,

(broadly) funnel-shaped to subcampanulate,

lobes 1,4— 2,2 x 0,7-1 mm; anthers 1,3-2 mm

long; minute rudimentary ovary present. 9:

tube 0,5-0, 8 mm long, lobes 0,2-0, 5 x

0, 1-0,2 mm; style 0 or c. 0, 2-0,4 mm long;

stigmas 2, (2,5— )3— 5(— 7) mm long; ovary c. 1

X 0,8 mm, with 4 small calyx lobes. Fruit red-

dish brown to greyish; mericarps c. 1,8-3 x

0,7-1, 2 mm, elliptic, oblong to ± obovate,

densely covered with whitish hairs or papillae

c. 0, 1-0,2 mm long, with 2 rounded to ±

triangular calyx lobes c. 0, 1-0,3 mm long or

calyx lobes ± lacking. Chromosome number:

2n=22.

Occurring from northern Natal to eastern Cape. Mostly

growing on rocky outcrops or at the edge of gorges or

kloofs, often in grassland-forest/scrub borders; mainly on

(confined to ?) TMS. Map 6.

MAPS 5-6: — 5. Anthospermum littoreum

6. Anthospermum galpinii

1,2: 14

Anthospermeae

Vouchers: Edwards 2263 (NU; PRE); Rudatis 1113

(BM; G; K; PRE; S; W); Strey 8682 (K; NH; PRE; SRGH).

In the eastern Cape, A. galpinii (especially cf plants)

may be difficult to separate from the closely allied A. ae-

thiopicum (no. 4); also some forms of A. spathulatum

subsp. spathulatum (no. 5) occurring in that area are some-

times very similar. Some collections with decussately ar-

ranged leaves from the Natal Midlands and northern Natal

approach A. monticolum (no. 6) in leaf size and shape and

habit; again, (f specimens may be problematic.

3. Anthospermum littoreum L. Bol. in

Ann. Bolus Herb. 2: 96 (1917). Syntypes:

Cape, East London, Sim 1483 (BOL!; PRE!),

Rattray 38 (BOL, lecto.!; GRA!; US!); Kentani

distr. , nr. Black Rock Cove, Pegler 2139

(BOL!; PRE!; US!; W!); Transkei, Port St.

Johns, Galpin 2850 (BOL!; GRA!; K!; PRE!).

A. ambiguum Greves in J. Bot., Lond. 63: 203 (1925).

Type: Transkei, Port St. Johns, Moss 2456 (BM, holo.!; J,

iso.!).

Shrub, dioecious or ± dioecious (cf plants

occasionally with odd flowers), ± procum-

bent, somewhat straggling or ± erect, at least

older parts often deeply buried in sand. Stems c.

0,5-3 m long, mostly much-branched, branches

spreading to ± ascending, buried parts often

rooting at nodes. Leaves decussate, pseudo- ver-

ticillate; blades (6-)8— 15(— 20) X (2-)2,5-5(-7)

mm, obovate, oblanceolate, oblong to ± ellip-

tic, ± thick and fleshy in nature, glabrous or

lowermost part of midrib below hairy, often ±

discolourous; petioles ± 0-1,5 mm long; stipu-

lar sheath with a small seta. Flowers mostly

subsessile, in clusters of several (to many: $) at

nodes, inflorescences often ± conspicuous in

9; corolla 4-merous, creamy yellow, yellow to

greenish yellow, hairy on outside at least near

base (CO or glabrous ($). cf(9): tube

(0,3-)0,4-0,6(-0,8) mm long, broadly funnel-

shaped to subcampanulate, lobes 2-2,4 x

0,6-0,8(-l) mm; anthers 1,7-2 mm long; min-

ute, densely hairy rudimentary ovary present.

tube c. 0,3-0,7(-l)mm long, lobes

0,2-0,5(-0,7) x 0,2 mm; style ± 0-1 mm

long; stigmas 2, (3— )4 — 6, 7(— 10) mm long;

ovary c. 0,8—1 ,2 mm long, densely hairy, with

4 minute calyx lobes. Fruit greyish brown or

greyish; mericarps (1 ,7— )2— 3(— 3 ,5) x (0,7-)

1—1 ,4(— 1 ,8) mm, oblong to ± elliptic, densely

covered with whitish spreading hairs c. (0,2-)

0,4— 0,8 mm long, with 2 ± triangular calyx

lobes c. 0, 1-0,4 x 0, 1-0,2 mm, usually hidden

amongst hairs. Chromosome number. 2n=22.

Fig. 5:9.

Occurring along the Indian Ocean coast from Natal

(Zululand) to eastern Cape (Alexandria distr ). Growing

mostly in dune sand, on the seaward- facing side of the first

dunes, in stabilized dunes amongst scrub or in grassy

patches, or at the edge of dune forest. Map 5.

Vouchers: Galpin 5664 (GRA; K; PRE); Strey 11204

(E; K; MO; NH; NU; PRE; SRGH; WAG); Venter 1836

(NH; PRE).

In areas where A. littoreum and the allied A. galpinii

(above) grow in close proximity, the occasional presence of

hybrids was observed.

4. Anthospermum aethiopicum L., Sp.

PI. 1058 (1753), edn 2: 1511 (1754); Cruse in

Linnaea 7: 132 (1832). Type: Cape, no collec-

tor given (LINN 1233.1 , holo. !).

[Linnaeus, Hortus Cliffortianus: 455: t. 27 (1737) (BM!)]

[Frutex africanus ambram spirans, Plukenet, Almagesti

Mantissa Botanici 159, t. 183, fig. 1 (1700)].

‘A. aromaticum’ , Salisb., Prodr. 59 (1796).

‘Ambraria heisteri’ Willd., Sp. PI. 4: 732 (1806).

A. aethiopicum L. var. ternifolium Cruse, Rub. Cap. 11

(1825), in Linnaea 6: 9 (1831); Sond. in F.C. 3: 28 (1865).

Type: none given.1

Dioecious shrub, single- stemmed, erect,

c. (0,5— )0,75— 2 m tall. Stems sparingly to much

branched above, branches often regular and ter-

nate, curving upwards to ± erect, in cf fre-

quently more numerous than in 9- Leaves

in whorls of 3, jsseudo-verticillate; blades

3— 9(— 1 3) x (0,3-)0,5-l ,2(— 2) mm (in cf of-

ten smaller than in 9)> narrowly lanceolate or

oblanceolate to linear, glabrous or occasionally

very shortly hairy near base; stipular sheath

with a small triangular to subulate seta. Flowers

subsessile, in clusters of very many (to many:

Cf) at nodes, inflorescences dimorphic, in 9

very conspicuous, much contracted, ± cylindri-

cal inflorescence zones c. (35— )50— 170 mm

long; corolla 4-merous, creamy yellow, green-

ish yellow or pale yellow, glabrous or ± papil-

late. Cf: tube (0,2-)0,4-0,9(-l ,2) mm,

(broadly) funnel-shaped, lobes ( 1 ,2— )1 ,6—

2,5(— 2,9) X 0,6—1 ,2(— 1 ,4) mm; anthers 1—1 ,8

mm long; small rudimentary ovary present. 9:

tube 0,l-0,4(-0,6) mm long, lobes 0, 1-0,5

(-0,7) x 0, 1— 0,2(— 0,4) mm; style ± 0-0,7 mm

long; stigmas 2, 4—6,3 mm long; ovary c.

0,6-0, 9 x 0,5-0, 7 mm. Fruit reddish brown to

dark grey brown; mericarps ( 1 ,4 — ) 1 ,7—2,5

1 Presumably any of Ecklon’s A. aethiopicum collections

from Table Mt. (e.g. Ecklon 24; GOET!; PRE!; S!). See

Cruse in Linnaea 7: 132(1832).

Anthospermeae

1,2: 15

FIG. 3. — 1-3, Anthospermum aethiopicum: 1, part of cf plant, x 0,5; 2, part of 9 plant, note congested inflores-

cence regions, X 0,5; 3, node with long and short shoot leaves (‘pseudoverticillate’ leaf arrangement), long shoot leaves

(shaded) in whorl of 3, x 5 ( Puff 790911-2/1). 4, A. spathulatum subsp. spathulatum, long shoot leaf pair with short

shoot leaves, X 5 (Davies s.n. sub PRE 41520).

1,2: 16

Anthospermeae

(-2,7) x 0,8— 1 (—1 ,2) mm, oblong, elliptic to

narrowly obovate, glabrous and often with

large, conspicuous epidermis cells or, less com-

monly, papillate, with 2 indistinct rounded to ±

triangular calyx lobes c. 0, 1-0,2 mm long or

calyx lobes ± lacking. Chromosome number:

2n=22. Fig. 3: 1-3.

Very widely distributed in the Cape Floristic Region

and extending to the eastern Cape (Albany, Bathurst and

Peddie distr.) and, in the north, to the Calvinia distr. (Nieu-

woudtville area). Found in all principal vegetation types of

the Cape Flora (except for the Strandveld) and perhaps most

abundant in areas where a moderately high rainfall is expe-

rienced and in relatively fertile (clayey) soils; in drier areas

often in moist to ± wet sites Map 7.

MAPS 7-10: — 7. Anthospermum aethiopicum

8. Anthospermum spathulatum subsp.

spathulatun

9. Anthospermum spathulatum subsp.

ecklonianum

10. Anthospermum spathulatum subsp.

uitenhagense

Vouchers: Bayliss 957; Boucher 3289 (PRE; STE);

Galpin 4 102 (GRA; PRE); Muir 84 1 ; 1955 .Thompson 1854

(PRE; STE).

A. aethiopicum is distinguished from A. spathulatum

(below), the other (large) shrubby South-western Cape

species, by its strictly ternately arranged leaves. It should

not be confused with A. spathulatum subsp. ecklonianum

(below) which is often very similar in habit and appearance.

On the eastern fringes of its distribution range the distinc-

tion of A. aethiopicum from A. galpinii (no. 2) can become

problematic .

5. Anthospermum spathulatum Spreng.,

Syst. Veg. 1: 399 (1825); Cruse, Rub. Cap. 13

(1825), in Linnaea 7: 133 (1832). Type: Cape,

no collector given.1

In the past, A. spathulatum has consistently been con-

fused with A. aethiopicum (above) and has, as ‘opposite-

leaved’ varieties (var. oppositifolium, var. ecklonianum,

var. montanum), been included in that species.

A. spathulatum is perhaps the most complex and most

variable of the Anthospermum species centred in the south-

western Cape. It is also the most widely distributed Cape

species and extends beyond the limits of the Cape Floristic

Region (1) to the north and east, and (2) into the karroid and

drier interior of the Cape. It occurs from sea level to the

highest mountain tops and over a wide array of substrates. It

is the only Anthospermum species known, to date, to com-

prise di-, tetra- and hexaploids.

5 subspecies are recognized:

5 (a), subsp. spathulatum.

A. aethiopicum L. var. oppositifolium Cruse, Rub. Cap.

11 (1825), in Linnaea 6: 10 (1831); Sond. in F.C. 3: 28

(1865); Salter in J1 S. Afr. Bot. 3: 109 (1937), in Adamson

& Salter, FI. Cape Penins. 732 (1950). Type: none given.2

A. aethiopicum L. var. montanum Sond. in F.C. 3: 28

(1865). Type: none given.3

A. tricostatum Sond. in F.C. 3: 28 (1865). Syntypes:

Cape, Rietvallei, Ecklon & Zeyher s.n. (S!); between Drie-

koppen and Bloodrivier, Drege 9550 (S, lecto.!).

Dioecious shrub or dwarf shrub, single- to

several-stemmed, ± erect, cylindrical to

rounded, c. 0,3-l,5(-2) m tall. Stems much to

sparsely branched above or from the base up-

Cruse [1825, op. cit. . and in Linnaea 6: 10 (1831)] claims having

seen and studied “Sprengel’s” specimens of A. spathulatum and

stated (1831) that specimens determined by Sprengel are in herbar-

ium B(t). A specimen in G, labelled 'A. spoilt. Sprg.’ (no collec-

tor given) and clearly marked as being from the Berlin herbarium,

is most likely an isotype of A. spathulatum.

2 In his 1831 publication [but not in Rub. Cap.) Cruse noted that

several Ecklon specimens are 'var. oppositifolium' . None of these

are in B(t?) nor is there an Ecklon collection in any other herbar-

ium which bears a handwritten identification by Cruse. It, ne-

vertheless, appears to be safe to select Ecklon 5 (Table Mountain;

S!) as a lecto-(neo-?) type.

3 Sonder considered var. montanum to be identical to A. aethiopi-

cum L. t, alpinum Eckl. & Zeyh. [Enum.: 366 (1836) (nomen non

valide publ.; since descr.)]. Ecklon & Zeyher 2307 t, [‘in vertice

montis "Kasteelsberg" (altit. V), in montibus prope "Simons-

town" (Cap), et in montibus “Hottentottshollandsberge'' supra

"Palmietrivier" (Stellenbosch).] can, therefore, be considered to

be the type (SAM!, holo.!; FI!. G!; LY!; M!; MO!; P!; PRE!,

iso.!).

Anthospermeae

1,2: 17

wards, branches often regular, paired, ascen-

ding to ± divaricate or ± irregular and intricate

(if browsed). Leaves strictly decussate, pseudo-

verticillate; blades (3-)4-12 x 0,5— 2,5(— 3)

mm, obovate, oblanceolate, lanceolate to li-

near-lanceolate, glabrous, upper surface often

± shiny, with conspicuous, small epidermis

cells; stipular sheath with a small seta. Flowers

subsessile or fruiting $ sometimes with pedi-

cels c. 0,2— 1(— 1 ,4) mm long, in clusters of

many to 2 at nodes, $ sometimes in ± con-

densed inflorescence zones; corolla 4-merous,

yellow to greenish yellow or pale whitish yel-

low, occasionally purplish tinged outside, gla-

brous. cT: tube 0,5-1 ,2 mm long, (broadly)

funnel-shaped, lobes 2-3,4 x 0,7-l,2(-l,5)

mm; anthers 1,2-2, 5 mm long; minute rudi-

mentary ovary with 4 small calyx lobes. $:

tube 0, 1-0,4 mm long, lobes 0,3-0, 8 x

0, 1-0,3 mm; style ± 0-0,5 mm long; stigmas

2, (1 ,5— )2— 7,6 mm long; ovary c. 0,9-1 ,5 x

0,5-0, 8 mm, with 4 calyx lobes. Fruit shiny

reddish brown to dull greyish brown; mericarps

2, 4-4, 2 x 1,2-2, 2 mm, (broadly) obovate to ±

elliptic, distinctly to rather obscurely 3-ribbed

(hence mericarps often ± triangular in section),

glabrous or, very rarely, shortly hairy, with 2

distinct to obscure ± triangular calyx lobes

0,3-0,6(-0,9) x 0,5-0, 7 mm or, less often,

calyx lobes ± lacking. Chromosome numbers:

2n=22, 44, 66. Fig. 3:4.

Very widely distributed in the south-western Cape Re-

gion and extending north to the Namaqualand distr. and east

to the King William’s Town and Keiskammahoek distr.;

also extending slightly into the drier interior parts of the

Cape. Growing on rocky slopes and ridges, on outcrops, in

gravelly areas or on sandy flats or sand dunes; usually in

relatively dry habitats; occurring mostly overTMS, granite,

Witteberg quartzite or limestone. Map 8.

Vouchers: Acocks 24076 (K; LISU; PRE; US); 14893

(K; PRE); Bolus 12701 (BOL; NH; PRE); Bos 510 (K; M;

PRE; STE; WAG); Esterhuysen 5840 (BOL; PRE); Laidler

148 (PRE; STE).

Subp. spathulatum is extremely variable, especially in

growth form (single-stemmed, rather tall plants to ±

rounded, several-stemmed dwarf shrubs; very irregularly

branched browsed forms are not uncommon in drier areas),

in the extent of the 9 inflorescences, in the fruits (dorsal

ribs; width of mericarps; size of calyx lobes), in leaf size

and shape and in chromosome number (polyploids morpho-

logically inseparable from diploids).

In spite of this variability, some ‘Forms’ (morpho- or

ecotypes) can be distinguished:

— ‘Typical’ subsp. spathulatum: Single- to few-

stemmed, often rather tall shrubs with typically regular,

paired, ± ascending branches; leaves (narrowly) obovate to

± lanceolate; 9 inflorescences somewhat condensed and

conspicuous, many- to several-flowered; fruits distinctly

ribbed and mostly sessile. Widely distributed and ecologi-

cally eurytopic.

— ‘Latifolium Form’: rather robust shrubs with broader

leaves than in the ‘typical’ form; 9 inflorescences inconspi-

cuous, partial inflorescences few- to 1-flowered; fruits

usually larger than in the ‘typical’ form, relatively broad,

conspicuously ribbed and mostly distinctly pedicellate.

Seems to be concentrated in the interior of the Cape (from

southern Calvinia distr. to Karroopoort, Witteberg and

Seven Weeks Poort); entirely absent in the south-western

Cape s.str.

— ‘Dune Sand Form’: Rather small, bushy, several- to

many-stemmed (dwarf) shrubs with rather fine, ± linear

leaves [small branches, especially of Cf plants, should not

be confused with subsp. ecklonianum (d), below, which

may look similar but tends to have longer leaves; specimens

without information on growth form and habitat may be

difficult to place]; 9 inflorescences rather inconspicuous,

partial inflorescences usually few-flowered; the subsessile

fruits tend to be smaller and less distinctly ribbed than in the

other Forms. Observed from the Cape Peninsula east to the

Port Elizabeth distr.

Some collections from the easternmost localities such

as Amatole Range, Hogsback Mt., Pirie (Rattray 231,

BOL, PRE; Sim 1328, PRE, STE; Story 3660) are some-

what atypical.

Subsp. spathulatum (especially d" specimens) should

not be confused with Nenax (notably N. acerosa subsp.

macrocarpa and N. divaricata), which may be similar in

appearance. The latter are distinguished vegetative ly by

having (at least a few) short hairs on the lower part of the

leaf margins and blades which are not narrowed to the base

as in subsp. spathulatum.

5(b). subsp. uitenhagense Puff, subsp.

nov., mericarpiis anguste obovatis vel ob-

longis, sine costis distinctis dorsalibus, saepe

apice ut videtur truncatis a subsp. spathulato

differt.

[A. aethiopicum L. var. uitenhagense Eckl. & Zeyh.,

Enum. 365 (1836), nomen non valide publ. (since descr.)].

Type: Cape, ‘Zwartkopsrivier inque planitie inter Kraka-

kamma et montes Van Stadensriviersberge altit. I, II (Uiten-

hage)’, Ecklon & Zeyher 2307(3 (SAM, holo.!; BOL;

GOET; LY; M; MO; S, iso.!).

Dioecious dwarf shrub, several- to many-

stemmed, mostly ± erect and cylindrical or

somewhat reclining, c. 0,2-0, 6(4), 9) m tall,

with thick, woody root; often with massive, ±

rosette-like woody base if burnt regularly.

Stems with several to many, ± erect to ascend-

ing branches (unbranched if burnt), quite

densely leafy above. Leaves : blades

(3-)3,5-8(-10) x 0,5-1, 2 mm, narrowly lan-

ceolate or oblanceolate to ± linear, mostly

tough and thickish; stipular sheath (broadly)

cup-shaped and conspicuous, with a minute

seta. Flowers subsessile, in clusters of several

to few at nodes, $ plants often with more flow-

ers than cf ; corolla 4-merous, glabrous or occa-

sionally papillate near tip. cf : tube 0,4-0, 9 mm

1,2: 18

Anthospermeae

long, lobes ( 1 ,5—) 1 ,9—2,5 x 0,5-1 mm; anthers

0,8-1 ,5 mm long. tube 0,2-0, 5 mm long,

lobes 0,3-0, 7 x 0, 1-0,2 mm; style ± 0-0,7

mm long; stigmas 2, ( 1 ,7— )2— 5 , 1 mm long;

ovary 0,6-0, 7 x 0,4 mm, with 4 minute, often

± indistinct calyx lobes. Fruit shiny reddish

brown to greyish green-brown; mericarps 2,2-3

x 1-1,3 mm, (narrowly) obovate, elliptic to

oblong, sometimes ± obscurely 3-ribbed, gla-

brous, often seemingly truncate above due to 2

sometimes ± joined (broadly) triangular calyx

lobes c. 0,1-0,5(-0,7) x 0,4— 0,5 mm. Chro-

mosome number: 2n=22.

Endemic to the south-eastern and eastern Cape and

occurring from the Humansdorp and Hankey distr. north-

east to the Cathcart distr. and the Great Kei River mouth.

Frequently associated with rocky outcrops in grassland, oc-

casionally in grassland or on dry, rocky slopes; often over

Witteberg quartzite. Map 10.

Vouchers: Flanagan 612 (GRA; PRE; SAM); Killick

780 (K; PRE); Sidey 3126 (PRE; S).

Very distinct in the eastern part of its distribution range

but sometimes difficult to separate from coastal 'Dune Sand

Forms' of subsp. spathulatum (see above) from the Port

Elizabeth and Uitenhage distr. westwards; especially cf

specimens may be problematic.

5(c). subsp. saxatile Puff, subsp. nov., ha-

bitu decumbenti vel pulvinato, internodiis bre-

vibus, foliis tenacibus et lobis calycis conspic-

uis longisque a subsp. spathulato distinguitur.

Type: Cape, Hex River Mts, Milner Peak,

5500-6000 ft., Jan. 1959, Esterhuysen 28089

(BOL, holo.!;WU, iso.!).

Dioecious dwarf shrub, several- to many-

stemmed, often decumbent, cushion- or ± mat-

forming. Stems to c. 250 mm long, occasionally

rooting at lowermost nodes, mostly much

branched, branches ± ascending to erect,

densely leafy. Leaves : blades (3 ,5— )5— 8(— 1 2) x

1-2 mm, lanceolate to oblanceolate or narrowly

obovate, tough and thickish; stipular sheath

with a small seta. Flowers subsessile, in clus-

ters of several to 2 at nodes; corolla 4-merous,

glabrous, cf: tube 0,8-1, 4 mm long, lobes

2, 2-2, 5 X 0,8-1 mm; anthers 1, 5-2,1 mm

long; minute rudimentary ovary with distinct

calyx lobes. 9: tube 0,2-0, 5 mm long, lobes

0,5—1 (—1 ,4) x 0, 2-0,4 mm; style ± 0-1 mm

long; stigmas 2, 2, 7-9, 2 mm long; ovary

1,4-1, 7 x 0,5-0, 8 mm, with 4 conspicuous

calyx lobes. Fruit shiny reddish brown; meri-

carps 2,4-3(-3,4) x 1,2-1 ,6 mm, obovate to

elliptic, ± indistinctly 3-ribbed (± triangular in

section), glabrous, with 2 ± triangular calyx

lobes 0,3—1 ,5(— 1 ,9) x 0,5-0, 7 mm. Chromo-

some number: 2n=44.

Confined to the south-western Cape mountains and

occurring from c. 1 500- 1800(- 1900) m. Growing on rocky

slopes, ledges or cliffs or in steep rocky gulleys; over TMS

and in damp to moist, ± shady, sheltered places. Map 1 1 .

MAPS 11-12.— 11. Anthospermum spathulatum subsp.

saxatile

12. Anthospermum spathulatum subsp.

tulbaghense

Vouchers: Esterhuysen 6407 (BOL; K; NBG; PRE);

1 1018 (BOL; K; NBG; PRE; SAM); 27943 (BOL; WU).

Very distinct in the field (cushion-forming or ± matted

habit; habitat!); perhaps closest to subsp. spathulatum.

5(d). subsp. ecklonianum (Cruse) Puff,

comb, et stat. nov. Syntypes: Cape: ‘Bavians-

berg bei Genadenthal’, Ecklon s.n. (S, lecto.!);

‘Schwarze Berg bei Caledon-Bad’, Ecklon s.n.

(S!; SAM!).1

A. aethiopicum L. var. ecklonianum Cruse in Linnaea 6:

10 (1831); Sond. in F.C. 3: 28 (1865).

A. spathulatum Spring, var. ecklonianum (Cruse) Cruse

in Linnaea 7: 133(1832).

Dioecious shrub, single-stemmed, ± erect

and typically (narrowly) cylindrical, c. (0,5-)

1-2 m tall. Stems ± much branched above,

branches mostly erect to ± ascending, usually

regular, paired, quite densely leafy. Leaves:

blades 8— 1 8(— 25) x 0,5-1, 8(2, 2) mm, linear-

(ob-) lanceolate to ± linear, often fine and ±

needle-like; stipular sheath with a seta to

1 ,2(— 1 ,9) mm long. Flowers subsessile or

1 The label of the lectotype specimen was written by Cruse

and signed ‘W. Cruse 1829’. Although no reference is

made on the label to Ecklon, there is no doubt that this is

one of the specimens referred to by Cruse (1831, op. cit. ).

The second-mentioned collection may have later been dis-

tributed as Ecklon & Zeyher 2307 e [‘inter frutices (altit.

II, III) montis “Zwarteberg” hand longe a thermis ( =

Caledon baths), nec non in collibus prope "Baby Ions

Toorensberg” (Caledon)’: Ecklon & Zeyher 1836: 366]

(BOL!; FI!; G!; GOET!; M!; MO!; P! ; S ! ; SAM!; W!).

Anthospermeae

1,2: 19

sometimes shortly pedicellate (cf), in clusters

of many to several at nodes, 9 sometimes in

somewhat condensed, ± cylindrical inflores-

cence zones; corolla 4-merous, glabrous, cf:

tube (0,3— )0,5— 1 ,4 mm long, lobes 1,9-2, 9

(-3,4) X 0,7— 1 (— 1,3) mm; anthers 1,3-2 mm

long. £: tube 0, 1-0,3 mm long, lobes

0,2-0,6(-0,8) x 0, 1-0,3 mm; style ± 0-0,3

mm long; stigmas 2, 2-4,4 mm long; ovary c.

0,5-0, 9 x 0,4-0, 6 mm, with 4 minute calyx

lobes. Fruit shiny reddish brown; mericarps

2,5-3, 1 x 1-1,5 mm, ± obovate to oblong,

rather obscurely 3-ribbed, glabrous or, very

rarely, shortly hairy, with 2 ± indistinct calyx

lobes c. 0,5 x 0,5-0, 7 mm, or calyx lobes ±

lacking. Chromosome number. 2n=44.

Endemic to the south-western Cape and rather widely

distributed. Mostly growing on rocky, well drained moun-

tain slopes, at the base of cliffs or along streams; very often

in granite-derived clayey soils. Map 9.

Vouchers: Kruger M26 (STE); Van Rensburg 425 (K;

PRE; STE); Williams 3070, 3071 (NBG; PRE).

Subsp. ecklonianum in habit and appearance resembles

A. aethiopicum (no. 4., above) and morphologically ap-

pears somewhat intermediate between A. spathulatum and

A. aethiopicum. Despite its tetraploidy there is no concrete

evidence for a hybrid origin of subsp. ecklonianum ; its

strictly decussate leaves (ternate in A. aethiopicum) are a

strong argument against such a speculation.

Some collections from the eastern part of its range

(George to Humansdorp distr.) appear to be ± atypical and

often do not fully match material from the south-west; spe-

cimens from that area may approach the ‘Dune Sand Form'

of subsp. spathulatum (see above).

5(e). subsp. tulbaghense Puff, subsp.

nov., mericarpiis parvis, plus minusve

oblongis, sine costis distinctis dorsalibus, flor-

ibus masculis minoribus et foliis parvis a subsp.

spathulato differt.

[A. aethiopicum L. var. tulbaghense Eckl. & Zeyh.,

Enum. 365 (1836), nomen non valide publ. (sine descr.)].

Type: Cape, '. . . non procula catarractis vallis "Tulbagh"

(Worcester)', Ecklon & Zevher 2307 (SAM, holo.!; FI;

LY, iso.!; S?).

Dioecious dwarf shrub, several- to many-

stemmed, erect to ± reclining and spreading.

Stems (100-)200-600(-750) mm long, mostly

much branched, branches ± irregular and rather

diffuse to ± regular, paired (9 plants occasio-

nally less branched than cf). Leaves : blades

(2,2— )2,5 — 4,5(— 5 ,2) x 0,5— 1(-1 ,3) mm, (nar-

rowly) obovate, oblanceolate, lanceolate to ±

linear-lanceolate; stipular sheath with a minute

seta. Flowers subsessile, in clusters of several

to ± many at nodes, 9 plants often more-flow-

ered than cf; corolla 4-merous, glabrous, cf:

tube 0,5-0,8(-l) mm long, lobes 1 ,2— 2(— 2,2)

x 0, 5-0,9 mm; anthers 0,9-1, 4 mm long. 9:

tube 0,2-0, 3 mm long, lobes 0,2-0, 5 x

0, 1-0,2 mm; style ± 0; stigmas 2, 1,4—3 mm

long; ovary 0,5-0, 8 x 0,4-0, 7 mm, with 4 ±

indistinct calyx lobes. Fruit shiny, light reddish

brown to dark purplish brown; mericarps

( 1 ,5—) 1 ,7—2,3 x 0,6 — 0,8(— 1 ) mm, ± oblong to

(narrowly) obovate or elliptic, glabrous, papil-

late or, less commonly, very shortly hairy, with

2 indistinct ± triangular calyx lobes c.

0,l-0,3(-0,4) x 0,3 mm or calyx lobes ± lack-

ing. Chromosome number. 2n = 22.

Endemic to the south-western Cape and occurring from

the Piketberg distr. south and south-east to the Cape, Wor-

cester and Caledon distr. Often found in coastal renoster-

veld, in degraded fynbos or disturbed sites; mostly growing

in clayey soils. Map 12.

Vouchers: Acocks 24512 (MO; PRE); Boucher 3386

(PRE; STE); Kruger KR557 (K; PRE).

Distinguished from the other subspecies primarily by

its smaller mericarps, which, furthermore, are distinctly

narrower (over two times longer than broad) and usually

lack distinct ribs. 9 plants, especially when in fruit, differ

quite markedly in habit (low dwarf shrubs, branches

densely beset with fruits) from the other subspecies. Leaves

and cf flowers are also rather smaller but are too variable in

size to allow an absolute separation from the other subspe-

cies, namely subsp. spathulatum. Subsp. tulbaghense may

deserve specific rank.

6. Anthospermum monticola Puff, sp.

nov., tubo corollae floris masculi breviore,

fructibus obovatis lobis calycis conspicuis coro-

natis et saepe foliis brevioribus ab A. spathu-

laato, et foliis fructibusque glabris ab A. basu-

tico differt.

Type: Cape, Witteberge, Lundean’s Nek-

Belmore rd., c. 8,5 km beyond (SSE of) New

England turnoff, towards Belmore, Puff

790113-3/1 (WU, holo.!; NBG; NU; PRE,

iso. !).

Dioecious shrub, few- to several-stem-

med, ± erect and cylindrical or roundish, low,

± cushion-forming if browsed excessively, c.

(0,2— )0,3— 1 ,5(— 2) m tall, often with thick,

woody tap root. Stems mostly much branched,

branches ± spreading or curving upwards,

usually paired. Leaves decussate, often pseudo-

verticillate; blades (2— )3— 1 0(— 1 3) x (0,7—) 1—2

mm, (ob)ovate, oblong, (ob)ovate-lanceolate to

1,2: 20

Anthospermeae

± lanceolate, glabrous; petioles subobsolete;

stipular sheath with a small seta. Flowers sub-

sessile, in clusters of (2— )6 (rarely more) at

nodes; corolla 4-merous, yellowish or greenish

yellow, occasionally reddish purplish tinged

outside, glabrous, d": tube 0,3-0, 5 mm long,

broadly funnel-shaped to (sub)campanulate,

lobes 2,2-2,7(-3) x (0,5— )0,8— 1 ,2(-l ,5) mm;

anthers 1-2 mm long. $: tube 0,4— 0,5 mm

long, lobes 0,3-0, 4 x 0,2-0, 4 mm; style 0;

stigmas 2, c. 3-5 mm long; ovary c.

( 1— ) 1 ,3—1 ,8 x 0,8-1 mm, with 4 conspicuous

calyx lobes. Fruit reddish brown to greyish

brown; mericarps 2-3 x 1,5-2 mm, (broadly)

ovate or obovate, oblong or ± elliptic, some-

times (faintly) 2- or 3-ribbed, glabrous, with 2

± triangular, erect to ± spreading calyx lobes

0,3-0,6(-0,8) mm long. Chromosome number.

2n—22.

Known from mountainous areas of Orange Free State.

Natal, Lesotho and Cape; in the Cape mainly on the higher

mountains of the interior (from Drakensberg and Witteberg

west to the Nieuweveld Mts). Growing on rocky slopes, on

outcrops, around rock sheets or sometimes in scrub along

streams; found over sandstone, dolerite or (less often)

basalt. Map 13.

18° 20° 22° 24° 28° ZB" 30° 32°

MAP 13. — Anthospermum monticola (scale bar for leaf

silhouettes is 10 mm)

Vouchers: Acoeks 15844 (K; PRE); Hilliard <6 Bunt

13552 (E; NU; PRE; WU); Killick 4446; Scheepers 1833

(K; LMU; MO; PRE; SRGH; WAG).

Very variable in leaf size and shape. Collections from

Natal and from the Natal-Lesotho-Orange Free State border

area tend to have larger and relatively narrower leaves (see

Map 13) and sometimes also slightly narrower fruits which

often lack distinct dorsal ribs; also their habit is somewhat

different (see also A. galpinii, no. 2).

Browsed individuals and plants growing at very high

altitudes (above 2400 m; Lesotho, Maluti Mts) tend to form

cushions; their stems may root at the nodes. They should

not be confused with Nenax microphylla which can be simi-

lar vegetatively.

A. monticola is allied to both A. spathulatum (subsp.

spathulatum ; = A. tricostatum sensu auct.), for which it has

often been mistaken, and A. basuticum (below). It is known

to form hybrids with the latter in areas where the species

occur together.

7. Anthospermum basuticum Puff, sp.

nov., A. hispidulo simile sed habitu dioecio,

floribus masculis minoribus, tubo corollae bre-

viore et fructibus longioribus latioribusque dif-

fert; A. monticolo affine sed folds fructibusque

pilosis praeclare distinguitur.

Type: Lesotho, Sehlabathebe National

Park, across stream from Motsea, 2550 m,

5.11 .1976, Hoener 1607 (PRE, holo. !).

Dioecious dwarf shrub, many-stemmed,

cushion- or mat-forming or ± erect and cylin-

drical. Stems (40-)10(k-300(-500) mm long,

erect to ± prostrate, mostly much branched,

branches often ascending, densely hairy, ±

densely leafy. Leaves decussate, often pseudo-

verticillate; blades often curved upwards or

ascending, 2,5— 5(— 6) x 0,8—1 ,2(— 1 ,8) mm,

narrowly lanceolate, often seemingly terete due

to strongly revolute margins, upper surface

densely covered with white spreading hairs

0,l-0,3(-0,6) mm long, lower surface often

glabrous except for midrib; petioles subobso-

lete; stipular sheath with a hairy seta. Flowers

subsessile, usually paired at nodes; corolla 4-

merous, yellowish to yellowish green, usually

hairy outside, cf: tube 0,7-1 mm long, broadly

funnel-shaped to subcampanulate, lobes (1,4—)

1,7-2, 3 x 0, 6-1,1 mm; anthers 1-1,6 mm

long. tube ± 0-0,3 mm long, lobes 0,5-1 x

0,2-0, 5 mm; style 0; stigmas 2, (2— )3— 6(— 7 ,5)

mm long; ovary 1-1,2 x 0,8 mm, mostly

densely hairy, with 4 calyx lobes. Fruit greyish

or greyish brown; mericarps 2-2,5 x 1-1,5

mm, ± obovate, mostly densely covered with

white spreading hairs 0, 1— 0,3(-A),5) mm long,

with 2 somewhat hairy, ± triangular erect to

spreading calyx lobes 0,5-0, 8 x 0,3-0,5(-0,7)

mm. Chromosome number: 2n = 22. Fig. 5: 4.

Anthospermeae

1,2:21

Known from Orange Free State, Lesotho, Natal and

eastern Cape; largely confined to the Drakensberg escarp-

ment and mostly occurring above c. 2300 m in basalt. Map

19.

Vouchers: Galpin 6649 (BOL; GRA; K; PRE); Hil-

liard & Burn 7121 (E; K; MO; NU; PRE; S); Killick &

Marais 2196 (K; NH; NU; PRE).

In the past, often confused with A. hispidulum (no. 9),

a non-dioecious ‘hairy’ species with larger cf and 9 flow-

ers and (narrowly) funnel-shaped, longer corolla tubes,

smaller, narrower fruits and an entirely different distribu-

tion range, which is probably only superficially similar. See

also A. monticola (above).

8. Anthospermum paniculatum Cruse,

Rub. Cap. 15 (1825), in Linnaea 6: 15 (1831);

Sond. in F.C. 3: 31 (1865). Type: Cape, ‘in

terra Houtniquas ad Promontorium bonae spei

mense Ianuario 1820', Mund(t) & Maire s.n.

(B, holo. t; G, iso. !).

A. paniculatum Cruse var. confertum Eckl. & Zeyh.,

Enum. 367 (1836). Type: Cape, ... ad “Zwartehoogdens”

non longe ab urbe “Grahamstown” (Albany), in monte

“Winterberg” prope fluvium “Katrivier” (Ceded Terri-

tory), Ecklon & Zevher 2314(3 (FI!; G!; LY!; M!; MO!; P!;

SAM!; W!).1

A. paniculatum Cruse var. elongation Eckl. & Zeyh.,

Enum. 367 (1836). Type: Cape. ... prope sylvas "Kraka-

kamma” et in montibus "Van Stadensriviersberge” (Uiten-

hage), Ecklon & Zevher 2314y (MO!; SAM!).1 [?also Eck-

lon or Ecklon & Zevher 349 (Vanstadensrivier Mts.) (BOL!;

BM!; K!; M!; PRE!; SAM!), 349 bis (BM!)].

'A. confertum Cruse’, Walp.. Repert. 2: 462 (1843).

Probably an error [A. paniculatum Cruse var. confertum

Eckl. & Zeyh.].

Dwarf shrub or subshrub, ± dioecious (cf

plants occasionally with some, rarely with

numerous $ flowers), few to several-stemmed;

with numerous erect, mostly unbranched flow-

ering stems from a woody base, if burnt regu-

larly. Stems ( 1 00—) 1 50 — 400(— 700) mm long,

erect to ascending, less commonly decumbent

and sometimes basal portions rooting at nodes,

branches erect to ± spreading. Leaves decus-

sate, pseudo-verticillate; blades 5—1 2(— 1 7) x

(0,3-)0, 5-2, 5( — 4) mm, linear, lanceolate to

1 On numerous sheets, cut-out parts from Ecklon & Zey-

her’s “Enumeratio” are used as labels. These often com-

prise the entire bottom part of page 367 (“2314. Antho-

spermum paniculatum Cruse”, (3. confertum. y. clonga-

tum"), so that it is not always quite certain to what variety

a particular specimen is meant to belong.

narrowly obovate, seldom (in fire-exposed indi-

viduals) ± broadly oblanceolate or obovate,

glabrous or sometimes ± papillate on upper

surface and/or midrib below; petioles ± 0-0,5

mm long; stipular sheath with a small 3-angular

to subulate seta. Flowers in terminal thyrsic or

thyrso-paniculate, ± narrowly cylindrical inflo-

rescence, peduncles (0,8—) 1 ,2—3 mm long,

pedicels to 0,7 mm long, slightly elongating in

fruit; corolla 4-merous, creamy yellow to yel-

low or greenish, glabrous. cf(9): tube

0,8-1 ,2(-l ,7) mm long, funnel-shaped, lobes

2, 2-2, 7 x 0, 5-1,1 mm; anthers 1,5-2, 2 mm

long, cf: ± prominent rudimentary ovary with

4 ± massive calyx lobes, often also rudimen-

tary stigmas present; $ : gynoecium as in 9- 9:

tube ± 0-0,3 mm long, lobes 0,5-0, 6 x

0, 1-0,2 mm, corolla sometimes absent al-

together; style 0-0,6 mm long; stigmas 2,

(3-)4-8 mm long, reddish to reddish purplish;

ovary 1,5-2, 4 x 0,9-1 ,6 mm, with conspi-

cuous calyx lobes. Fruit brown; mericarps

2, 5-3, 5(^1) x 1,7-2 mm, oblong to ± rectan-

gular or ± obovate, often with two faint ribs

below 2 ± triangular, often massive calyx lobes

0,5-1 ,2(-2,2) x 0,4-1 mm. Chromosome

number. 2n = 22.

Occurring from Transkei to south-western Cape

(George distr. ); centred in the eastern Cape. Mostly grow-

ing in grassveld, bush-clump veld or on rocky ridges; occa-

sionally in disturbed sites (roadsides, fire-breaks, etc.).

Map 25.

Vouchers: Britten 1205; Flanagan 215 (BOL; NH;

PRE; SAM); 2284 (BOL; K; P; PRE; SAM); Keel 1073

(GRA; PRE; STE).

The terminal, ‘typical’ thyrsic to thyrso-paniculate in-

florescence of A. paniculatum is unique in the genus. This

feature as well as the unusual, relatively large fruits (ana-

tomically resembling those of Nena.x ) and the reddish stig-

mas put the species into a somewhat isolated position.

(Regular) exposure or non-exposure to veld fires

appears to be the main factor responsible for the sometimes

striking growth form differences.

9. Anthospermum hispidulum E. Mey.

[in Drege in Flora 26, Bes. Beigabe 164 (1843),

nom. nud.] ex Sond. in F.C. 3: 29 (1865). Syn-

types: Transkei, Omsamwabo to Omsamcabo,

Drege s.n. (S, lecto.!; G!; K!; LY!; MO!; W!);

Natal, Zululand, Gerrard [& “ M'K ”

(=McKen)] 1361 (BM!;K!;W!).

A. burkei Sond. in F.C. 3: 29 ( 1865); Compton. FI. Swa-

zild 587 (1976). Types: Transvaal, Magaliesberg, ["Burke

& Zeyher" = ] Burke 86 (BM; K; PRE; SAM. iso.!). Zevher

770 (S, holo. !; BM; G; LY; K; P. iso.!).

1,2: 22

Anthospermeae

A. arenicolum Greves in J. Bot., Lond. 63: 203 (1925).

Type: Transvaal, Witpoortjie Kloof nr. Johannesburg, Moss

9806(BM,holo.!;PRE, iso.!).

A. rubricaule K. Schum., nom. nud.

Dwarf shrub, $ , $ + $, $, (I? + Cf or

(rarely) cf, usually several- to many-stemmed,

often with thick, woody (tap-)root. Stems c.

(100-)150-400(-600) mm long, ascending to

erect, sparingly to much branched, branches

ascending to ± erect, hairy. Leaves decussate,

usually pseudo-verticillate; blades often ascend-

ing, 5—1 2(— 20) x 1— 3 ,5(— 5) mm, oblanceolate,

ovate-lanceolate to narrowly lanceolate, less

commonly ± obovate, usually both surfaces

densely covered with whitish spreading hairs c.

(0,1-)0,3-0,5 mm long, sometimes less hairy

below except for midrib; margins mostly revo-

lute; petioles subobsolete; stipular sheath hairy,

with a seta (0,5-)0,8-l ,4 mm long. Flowers

subsessile, in clusters of c. (9— )6— 2 at nodes;

corolla 4-merous (very rarely also 5-merous in

9"), greenish yellow to yellowish, at least lobes

hairy on outside. $?, Cf: tube 1,4-1 ,6 mm (cf)

or -2(-2,4) mm (<J?) long, (narrowly) funnel-

shaped, lobes (1 ,7— )2— 2,7 x 0,6-0, 9 mm;

antners 1,4— 1,9 mm long; stigmas 2, c.

2,9-4, 1 mm long, ovary c. 0,7-1 ,2 x 0,5-0, 9

mm, hairy, with 4 calyx lobes; cf: small rudi-

mentary ovary, sometimes also rudimentary

stigmas present. 9: tube 0,3-0, 7 mm long,

lobes 0,2-0, 7 x 0, 1-0,4 mm; style ± 0-0,8

mm long; stigmas 2, 3-6,5 mm long; ovary as

in Cf. Fruit reddish brown; mericarps c. 1,7-2

x 0,7—1 mm, ± oblong, ± densely (less com-

monly ± sparsely) covered with whitish spread-

ing hairs c. 0,2-0, 5 mm long, with 2 often ha-

iry, ± triangular calyx lobes c. (0,2-)

0,4— 0,7(-0,9) x 0,l-0,3(-0,4) mm. Chromo-

some number: 2n=22.

Known from Transvaal, Swaziland, Natal and

Transkei; also on Orange Free State side of Vaal River (near

Parys; one record). Growing on rocky ridges, outcrops,

koppies, krantzes or in cracks of rock sheets, less com-

monly in rocky grassland; often occurring over sandstone

(Natal, Transkei), quartzite or old granite (Transvaal). Map

14.

Vouchers: Codd 7673 (K; NH; NU; PRE); Galpin

13079 (K; PRE; US; W); Strey 9015 (K; NH; PRE); Van der

Schijff 5996 (K; PRE; W).

Rather variable. Flowering shoots of recently burnt

individuals ('A. arenicolum' ) may look very different from

typical material. Hybridogenous contact with A. pumilum

subsp. pumilum (no. 11) is suspected in parts of the

Transvaal where both taxa may grow in immediate vicinity.

Allied to the similar A. whyteanum Britten (Zimbabwe

to southern Tanzania) which is distinguished by strictly

temately arranged leaves.

MAP 14. — Anthospermum hispidulum

10. Anthospermum herbaceum L. /.,

Suppl. 440 (1781); Murr., Syst. Veg. 919

(1784); Brenan in Mem. N.Y. bot. Gdn 8: 455

(1954); Agnew, Upl. Kenya Wild Flow. 407 &

fig., p. 406 (1974); Verdcourt in F.T.E.A. Ru-

biaceae 1: 325 & fig. 46 (1976). Type: Cape,

Thunberg s.n. (LINN 1233.5, holo.!).

A. lanceolatum Thunb., Prodr. 1: 32 (1794), FI. Cap.,

edn Schultes 157 (1823); Cruse, Rub. Cap. 12 (1825), in

Linnaea6: 12 (1831); Sond. in F.C. 3: 30 ( 1865); De Wild.,

PI. Bequaert. 2: 301 (1923); Robyns, FI. Parc. 2: 371

(1947); Compton, FI. Swazild 587 (1976). Type: Cape,

Thunberg (sheet 23314, A. lanceolatum ‘a’, UPS, holo.!;

G; S, iso.!).

A. muriculatum Hochst. ex A. Riel;., Tent. FI. Abyss. 1:

345 (1847); Hiem in F.T.A. 3: 229 (1877). Syntypes:

Ethiopia, Prov. Ouedgerate-, [Quartin-Dillon &] Petit s.n.

(P, lecto. ! ); ‘Koubi-, Schimper 732 ( BR ! ; G!; K!; S!; Pi-

mixed with A . pachyrrhizum) .

A. ferrugineum Eckl. & Zeyh., Enum. 366 ( 1836). Type:

Cape, Van Stadensriviersberge (Uitenhage), prope Gra-

hamstown (Albany) et ad flumen Makasanirivier prope

montem Chumiberg (Adelaide), Ecklon & Zeyher 2309

(SAM. holo.!; BOL; FI; G; GOET; M; MO; S; W, iso. !).

A. hedyotideum Sond. in F.C. 3: 30 (1865). Syntypes:

Cape, ‘Caffraria. Kreili's Country', Bowker (595] (S,

lecto.!; K!; PRE-photo.!); Keiskamma, Drege s.n. ('Herb.

Drege, Sender'; not located).

Anthospermeae

1,2: 23

A. lanceolatum Thunb. var. hedyotideum (Sond.)

Kuntze, Rev. Gen. 3: 1 17 (1898).

A. lanceolatum Thunb. var. latifolium Sond. in F.C. 3:

30 (1865). [A. latifolium E. Mey. in Drege in Flora 26, Bes.

Beigabe 164 (1843), nom. nud.] Lectotype: Port Natal,

Gueinzius ‘468’ (S ! ; as s.n.: G; SAM sub 16062, iso-

lecto.!).

A. nodosum E. Mey. in Drfege in Flora 26, Bes. Beigabe

164 (1843), nom nud. [‘Port Natal’ Drege s.n. (E!; G!; K!;

LY!; S !)] .

A. mildbraedii K. Krause in Wiss. Ergebn. Dt. Zentral-

Afr. Exped. 2: 341 (1914). Type: Rwanda, west of Lake

Mohasi, Mildbraed 497 (B, holo.t; BR; iso.!).

Perennial herb or ± subshrub, $f, $ + 9>

9, $ + Cf, Cf + $ + 9’ less commonly cf or

Cf + 9» several- to many-stemmed, somewhat

woody near base and with often ± thick, woody

root; sometimes rather short-lived. Habit varia-

ble: plants scrambling, straggling or trailing and

with stems to 2,5(-3) m long; sometimes ±

erect, cylindrical to rounded, to c. 0,3(-0,5) m

tall and with ± thick, almost rosette-like base;

occasionally low and ± mat- or cushion-form-

ing with shoots c. (20-)75-150(-200) mm

long. Stems unbranched to much branched;

branches often ± regular, arising in pairs, fre-

quently with short branches of a higher order,

often with short shoots bearing rather small

leaves. Leaves decussate; blades 5-55 x

( 1— )2— 25 mm, ± ovate, ovate-lanceolate, lan-

ceolate to ± linear-lanceolate, cuneate to

rounded at base, glabrous, ± densely papillate

or shortly hairy, often distinctly discolourous;

petioles 0,7-6, 5 mm long; stipular sheath typi-

cally with (3— )5(— 7) filiform setae*, seldom

with only 1 ± triangular seta, the longest

(0,3-)0,7-6,l mm. Flowers mostly on short

lateral branches, in ± sessile to somewhat

elongated clusters of many to 6 (less commonly

only 2) at nodes; corolla 4-merous, greenish to

yellow or yellowish, sometimes reddish pur-

plish tinged, mostly papillate or shortly hairy at

least near tip. S' : tube ( 1 ,5— )2— 3,7 mm long

(in 9" often longer than in cf), cylindrical,

lobes (1 ,5-)2-2,7(-3,4) x 0,3-0, 7 mm; an-

thers (0,9—) 1 ,2— 2(— 2,5) mm long; cf: small ru-

dimentary ovary and stigmas present, the latter

hidden in corolla tube; gynoecium as in 9

but stigmas only c. (2-)2,4-5(-6,4) mm long.

* Setae should not be confused with small, sometimes ±

linear short shoot leaves crowded in axils of long shoot

leaves!

9: tube 0,3-0,7(-l,2) mm long, lobes

0,3-0,7(-l) x 0, 1-0,3 mm; style ± 0-0,8 mm

long; stigmas 2, (2-)4— 10,2 mm long; ovary

0,5-1 ,2 x 0,4-0, 7 mm, sometimes with 4 of-

ten indistinct calyx lobes. Fruit yellowish

brown or reddish brown, mostly with ± broad,

conspicuous longitudinal grooves between me-

ricarps; mericarps ( 1 ,5—) 1 ,7— 2,5(— 2,8) x

0,9-1, 6 mm, elliptic, oblong to ±obovate, ±

densely covered with ± tuberculate structures,

shortly hairy, papillate or subglabrous, occasio-

nally with 2 obscure, ± triangular to rounded

calyx lobes c. 0,2-0, 3 x 0,2-0, 3 mm. Chro-

mosome number : 2n=22. Fig. 4; 5: 5 & 6.

Occurring from Transvaal to the south-western Cape;

extending north to Ethiopia and the south-western Arabian

Peninsula. Growing in forest edge vegetation, scrub, rive-

rine thicket, at the edge of marshes, in (fire-prone) grass-

land, on rocky slopes or occasionally in littoral scrub and

near the sea in the salt spray zone. Map 15.

MAP 15. — Anthospermum herbaceum

Vouchers: Thompson 1814 (PRE; STE); Van der

Schijff 5596 (K; PRE); Ward 4316 (K; NH; NU; PRE).

Highly variable in most of its characters. A. herba-

ceum contains a number of ‘Forms’ or ecotypes which in

places are quite distinct and well discemable. Some of the

more conspicuous are:

1,2:24

Anthospermeae

FIG 4. — Anthospermum herbaceum, flowers from a single plant: a, $ , X 17,5; b. cf . x 17,5; c, , rudimentary

gynoecium, corolla tube opened to show rudimentary stigmas, x 25; d & f, functionally 9 with rudimentary anthers,

x 17,5; e. detail of rudimentary anther, x 25; g. ‘pure' 9 without anther rudiments, x 17,5 (Puff 780409-1/1) (SEM-

graphs).

Anthospermeae

1 ,2: 25

— The (‘typical’) ‘Afromontane Forest (Edge) Form’:

Rather robust plants with the longest (scrambling, strag-

gling) stems and the largest leaves; mostly growing in

shady, moist to wet places.

— The ‘(Burnt) Grassland ('A. hedyotideum' ) Form’:

± erect, several- to many-stemmed plants, hardly more

than 300 mm tall; stems mostly unbranched, arising from a

± massive woody base; partial inflorescences normally

very few- to one-flowered.

— The 'High Altitude (Grassland) Form’; Often

found at the base of boulders or small rocks in high altitude

grassland. Low prostrate plants which often form mats or

cushions; stems usually very short (c. 100 mm), leaves

small and relatively broad, inflorescences much reduced,

flowers often only paired at nodes. Plants occurring in tram-

pled areas (at lower altitudes) are similar in appearance but

do not usually form dense cushions.

— The ‘Salt Spray Zone Form': Mostly growing in

cracks of rocks near the sea. Glabrous, ± prostrate plants

with rather broad leaves which tend to be quite succulent;

the stipular sheaths often only bear a single seta; the partial

inflorescences are frequently only 1 -flowered Not very

common, from the Natal South Coast to the Transkei (Wild

Coast).

In the south-westernmost part of its distribution range,

subglabrous and rather short-lived (?) forms with narrow,

lanceolate leaves and rather reduced inflorescences are pre-

dominant in places. Similar forms, but with more slender

stems and longer intemodes, occur on the Natal South

Coast.

In the south-western Cape, A. herbaceum should not

be confused with Carpacoce spermacocea (vegetatively

rather similar but characterized by a strong foetid odour, by

flowers with only one stigma and one fertile carpel and by

fruits with leaf-like calyx lobes which do not separate into

mericarps) and ecotypes of A. galioides (no. 13) with red-

dish stems and relatively broad, discolourous leaves (distin-

guished from A. herbaceum by smaller flowers, a differ-

ing fruit morphology and a different habit).

11. Anthospermum pumilum Sond. in

F.C. 3: 31 (1865). Type: Orange Free State,

Caledon River, Zeyher s.n. (S, holo.!; LY;

SAM sub no. 16064, iso.!).

A. rigidum sensu auctt. Afr. austr., non Eckl. & Zeyh

(1836).

A. humile N.E.Br. inKewBull. 1895: 145 ( 1895). Type:

Natal, Ulundi, Evans 370 (K, holo.!; NH, PRE-photo.,

iso.!).

A. ericoideum K. Krause in Bot. Jb. 39: 570 (1907);

Engler, Pflanzenwelt Afr. 1: 574, fig. 508 (1910); Launert

& Roessler in F.S.W.A. 115: 8 (1966). Type: South West

Africa/Namibia, Auasberge, Damara-Namaland, Dinter

291 (B, holo.t?).

A. pumilum Sond. var. pilosum Phill. in Ann. S. Afr.

Mus. 16: 112 (1917). Lectotype: Lesotho, Leribe Plateau,

A. Dieterlen 629 (SAM!; GRA; K; NH, PRE; STE. iso-

lecto.!).

A. spicatum Suesseng. in Trans. Rhod. Sci. Assoc. 43:

55 (1951). Type: Zimbabwe, Marandellas, Dehn 547 (M,

holo.!).

A. frutescens Dinter, nom. nud. [South West Africa/

Namibia, Auros, Otavi, Dinter 5659].

A species widely distributed in the Flora area (except

for the south-western Cape) and extending north to southern

Tanzania and to Angola. Belongs to the Anthospermum

galioides group, which is centred in the south-western and

western Cape (no. 13-16). The Natal South Coast

sandstone endemic, A. streyi (no. 12), also forms part of

this alliance.

Two subspecies are recognized:

11(a). subsp. pumilum.

Synonyms as above.

Subshrub, 9- 9 + 9’ 9’ occasionally d"

or 9 + O", rounded to cylindrical. Stems

numerous, unbranched, c. (50-)80-200(-300)

mm long, arising from a massive, often ±

rosette-like woody base (if exposed to fire) or

dwarf shrub, much branched, to 400 mm tall,

with thick, woody root; plant mostly ± densely

leafy. Leaves decussate, pseudoverticillate;

blades (4-)6-12(-22) x (0,5-)0,8-l ,5(-2)

mm, linear, linear-(ob-)lanceolate or narrowly

lanceolate, ± membranous; flat to somewhat

revolute margins mostly papillate, midrib be-

low often reddish brown and prominent; pe-

tioles subobsolete; stipular sheath with 1 (rarely

3) small seta(e). Flowers subsessile, in clusters

of 6-2 (occasionally more) at nodes; corolla 4-

merous, greenish to yellowish, often papillate

near tip. (f: tube (0,5-)0,7-l ,4(-l ,7) mm

long, narrowly funnel-shaped, lobes 1,2-1 ,9

(-2,5) x 0,3-4), 7 mm; anthers 1—1 ,8(— 2) mm

long; stigmas often shorter than in 9- 9:

tube 0,2-0, 5 mm long, lobes 0,2-0, 5 x 0,1 —

0,2 mm; style 0-0,5 mm long; stigmas 2, 2,4-

9,8 mm long; ovary c. 0, 5-0,9 x 0,3-0, 8 mm,

with 4 ± indistinct calyx lobes. Fruit reddish

brown, shiny; mericarps ( 1 ,5— )1 ,8—2.4 x

1-1,5 mm, elliptic to obovate, mostly ± gla-

brous, often with 2 ± broadly triangular to

rounded calyx lobes c. 0, 1-0,3 x 0, 3-0,4 mm.

Chromosome number: 2n=22. Fig. 5: 10.

Known from South West Africa/Namibia and south-

eastern Botswana and from Transvaal to the eastern and

north-eastern Cape. Usually growing in (rocky) grassland or

open woodland; mostly in fire-prone habitats. Map 16.

Vouchers: Flanagan 1217 (BOL; GRA; PRE; SAM);

Liebenberg 6949; McClean 659 (NH; PRE); Obermever

1 10, 2769.

1,2:26

Anthospermeae

clusters of 2-6 (rarely more) at nodes; corolla 4-

merous (very rarely 5-merous in $). cf, 9:

tube 0,5—1 ,5(— 1 ,9) mm long (in 9 often some-

what longer than in cf), narrowly funnel-

shaped to ± cylindrical, lobes ( 1 ,7— )2,2— 3

(-3,2) x 0,5-0, 8 mm, anthers 1 ,5— 2,2(— 2,5)

mm long; cf : small rudimentary ovary present,

often with rudimentary stigmas; 9 : stigmas of-

ten shorter than in $. 9: tube 0,2-0, 5 mm

long, lobes 0,3-1 X 0, 1-0,3 mm, style ± 0-1

mm long; stigmas 2, c. 3,4— 7,5 mm long; ovary

c. 0,6-0, 9 x 0,5-0, 7 mm. Fruit: mericarps

2— 2,6(— 2,8) x 1-1,5 mm, mostly ± sparsely

covered with whitish, curved, ± hook-like

hairs c. 0, 1-0,3 mm long. Chromosome

number. 2n=22.

MAP 16. — Anthospermum pumilum subsp. pumilum

Some populations from Transvaal and Natal are char-

acterized by an unusual ± cushion- or mat-forming habit,

slightly larger flowers than typical subsp. pumilum and a

trend towards unisexual flowers, i.e. dioecy ('A. humile').

They seem to be confined to ± wet seepage areas associated

with rock sheets, etc.

11(b). subsp. rigidum (Eckl. & Zeyh.)

Puff, comb, et stat. nov. Type: Cape, ‘in colli-

bus Karoo similibus (alt. II) prope fluvium

“Gauritzrivier” . . Ecklon & Zeyher 2315

(S, holo.!; LY; SAM, iso.!). [See comments

below],

A. rigidum Eckl. & Zeyh., Enum. 367 (1836).

Dwarf shrub, cf, 9 (occasionally 9 +

odd Cf or ^ + odd 9 ) or 9 , several- to many-

stemmed, ± erect and cylindrical to ±

rounded, with often thick, woody root; low and

± cushion-forming or with long, thin shoots

from a thick woody base if browsed. Stems

( 100-)200-400(-550) mm long, ± sparingly to

much branched; branching irregular if browsed;

intemodes mostly longer than leaves, plant not

densely leafy. Leaves decussate, sometimes ±

pseudo-verticillate (but short shoot leaves often

distinctly smaller than long shoot leaves);

blades 3-6(-8) x 1-1 ,5 mm (occasionally to 22

x 3 mm on new growth), narrowly ovate- or

obovate-lanceolate, lanceolate to linear-lanceo-

late, ± tough and thickish, glabrous or margins

papillate, upper epidermis cells often ±large

and conspicuous; stipular sheath with or with-

out a minute seta. Flowers subsessile (in fruit-

ing 9 pedicels occasionally to 0,5 mm long), in

Known from the Orange Free State and Cape, possibly

also in western Lesotho. Mostly growing in rocky places

(koppies, hillsides, etc); usually in relatively dry karroid

and ‘false karroid' vegetation types. Map 17.

MAPS 17-19: — 17. Anthospermum pumilum subsp rigi-

dum

18. Anthospermum streyi

19. Anthospermum basuticum

Anthospermeae

1 ,2: 27

FIG. 5. — 1-3, Anthospermum bicorne: 1, plant, x 0,56 (Bolus 341); 2, fruit in side view, x 10; 3, from back, sterile

carpel (arrow) with 2 enlarged calyx lobes, x 10 (Puff 800101-5/1). 4, A. basuticum, fruit in side view, x 10 (Puff

7901 1 3-5/3). 5 & 6, A. herbaceum: 5, fruit in side view, note broad longitudinal groove between mericarps, x 10; 6.

mericarp, ventral side, note longitudinal median ridge, x 10 (Puff 810916-3/1). 7, A. streyi, fruit in side view, x 10 (Puff

790426-3/1). 8, A. galioides subsp. galioides, mericarp, ventral side, with attached carpophore (arrow), x 10 ( Puff

7909 1 3-2/1 ). 9, A. littoreum, carpophore, x 10 (Puff 790415-5/1). 10, A. pumilum subsp. pumilum, longitudinal

section of ovary showing ovules, x \0(Puff 790112-2/1).

1,2: 28

Anthospermeae

Vouchers: Brynard 132 (K; PRE); Galpin 2013; Her-

man 348; Leistner & Joynt 2734 (K; KMG; PRE; SRGH).

The type locality of A. rigidum appears to be clearly

outside the distribution range of the taxon. The Gouritz

River valley both north and south of the Langeberge, how-

ever, is characterized by karroid vegetation rather than

south-western Cape fynbos. See Muir’s “Vegetation of the

Riversdale Area” [Mem. bot. Surv. S. Afr. 13 (1929)] for

details.

In a number of respects, subsp. rigidum is ± interme-

diate between A. dregei subsp. dregei (no. 15) and subsp.

pumilum. Some western collections of subp. rigidum (e g.

Acocks 17746) are rather similar to A dregei but in its

eastern and north-eastern range of distribution subsp. rigi-

dum shows much greater affinity to subsp. pumilum.

The separation of subsp. pumilum and rigidum is not

always clear-cut in the north-eastern Cape and in parts of

the Orange Free State where their distributions overlap.

Environmental factors in particular (fire-exposure and

browsing) often complicate the separation of the subspecies:

Shrubby plants of subsp. pumilum sheltered from fire may

somewhat resemble the more robust subsp. rigidum (occur-

ring in less fire-prone habitats); browsed plants of subsp.

rigidum may superficially look very similar to subsp. pumi-

lum. Subsp. rigidum differs in having larger (cf, 9") flow-

ers and in its clear trend to dioecy .

12. Anthospermum streyi Puff, sp. nov.,

habitu dissimili, foliorum laminis magis acero-

sis, saepe recurvatis et floribus hermaphroditis

majoribus ab A. pumilo distinguitur .

Type: Natal, Port Shepstone distr. , Beacon

Hill East, Strey 7248 (NH, holo.!; K; NU; PRE,

iso. !).

Dwarf shrub, 9" or $ (always?), procum-

bent and ± straggling, sometimes cushion-

forming. Stems c. ( 100-) 1 50-300(^)00) mm

long, ± sparsely to much branched, mostly

densely leafy. Leaves decussate, pseudoverti-

cillate; blades often curved, (5— )8— 1 2 x (0,3-)

0,5— 0,8(— 1 ) mm, linear to linear-lanceolate, of-

ten seemingly terete and needle-like when dried

due to strongly revolute margins, glabrous, mi-

drib below (and sometimes also upper surface)

reddish brown; petioles subobsolete; stipular

sheath mostly with a seta to 0,6 (-1) mm long.

Flowers subsessile, in clusters of 2-6 at nodes;

corolla 4-merous, yellowish, often with a few

short hairs near tip. 9": tube 1-1,3 mm long,

narrowly funnel-shaped, lobes (1 ,3-) 1 ,5-2,5

(-3) x 1 mm; anthers 1,3-2 mm long; style 0;

stigmas 2, 3-5 mm long; ovary c. 0,8-1, 2 x 1

mm, with 4 indistinct calyx lobes. 9: tube c.

0,3-0, 5 mm long, lobes 0,2-0, 5 x 0,2 mm;

gynoecium as in $ . Fruit reddish brown, ±

heart-shaped in side view; mericarps 2-2,5 x

( 1—) 1 ,2—1 ,5 mm, ovate toobovate, ± sparsely

covered with curled whitish hairs c. 0, 1-0,2

mm long, sometimes with 2 indistinct ± triang-

ular calyx lobes c. 0, 1-0,2 mm long. Chromo-

some number : 2n=22. Fig. 5: 7.

Narrowly endemic to the Natal South Coast (Port

Shepstone distr.). Growing on rocky outcrops in grassland,

amongst rocks, at the edge of krantzes. Map 18.

Vouchers: Huntley 752 (NH; NU); Nicholson 1 145.

A. streyi ‘replaces’ the closely allied and widely dis-

tributed A. pumilum subsp. pumilum (above) in the

sandstone areas of Natal’s southern border. It differs from

the latter in habit, in its more needle-like, often curved leaf-

blades, in having larger flowers and rounder (larger)

fruits.

13. Anthospermum galioides Reichb. f.

in Sprengel, Syst. Veg. 4: 338 (1827). Type:

‘C.B.S.% no collector given [C. W. Bergius

s.n. (B f ; G, neo.!)].1

A. ciliare L. , sensu auctt. 1

Subshrub or dwarf shrub with often thick,

woody root, $ + 9* ?’ *ess commonly d"

or 9" + Cf, several to many-stemmed, erect and

± cylindrical, rounded or prostrate and ± mat-

or cushion-forming. Stems c. (50-)80-300

(-500) mm long, unbranched to much

branched, not rooting at nodes in prostrate

forms. Leaves decussate, pseudoverticillate;

blades ascending to spreading, often recurved

at least near tip, (2,2— )2,7— 10(— 1 2) x (0,5-)

0,8-2,5(-3) mm, linear to lanceolate or ovate-

lanceolate [occasionally to 20 x 4 mm, (ovate-

lanceolate on young flowering plants or new

growth], ± membranous to ± tough and thick-

ish, sometimes distinctly discolourous, upper

surface often shiny and with large, conspicuous

epidermis cells; margins flat to strongly re vo-

lute with whitish spreading hairs c. 0, 1-0,4 mm

long, only minutely scabrid or entirely gla-

brous; petioles ±0-1 mm long; stipular sheath

with a (± broadly) triangular to subulate seta c.

(0,5 — )0,7— 1 ,5( — 2) mm long. Flowers subses-

sile, in clusters of 2-6 (rarely more) at nodes;

1 Cruse in Linnaea 6: 11 (1831) considered A. galioides to

be a synonym of ‘A. ciliare L.’, a conclusion reached

after studying ‘Reichenbach’s specimen'. Hence all spec-

imens (including the above Bergius collection) cited by

Cruse (op. cit. & Rub. Cap. 14. 1825) under ‘A ciliare

L.' can be considered to be identical to A. galioides.

Since Cruse's publications on Cape Rubiaceae (op. cit.)

the name A. ciliare has been consistently misapplied [the

type of A. ciliare L. (1763) corresponds to A. bergianum

Cruse (1825)]. It has been proposed to reject the name

Anthospermum ciliare L.. Sp. PI., edn 2: 1512 (1763);

see Puff in Taxon 31: 759 (1982) for details.

Anthospermeae

1 ,2: 29

corolla 4-merous (very rarely also odd 5-mer-

ous 0" and 3-merous $), greenish yellow to

yellowish, sometimes reddish purplish tinged

outside, mostly shortly hairy or papillate at least

near tip. cf, y: tube 0,7-1 ,7(-2) mm long, ±

cylindrical to (narrowly) funnel-shaped, lobes

1,4— 2,4(-2,9) x 0,3-0,8(-l) mm; anthers

(0,7— >1—1 ,8(— 2, 1 ) mm long; 9" : stigmas often

shorter than in tube 0,2-0, 5 mm long,

lobes 0,2-0, 8 x 0, 1-0,3 mm; style ± 0-0,6

mm long, stigmas 2, 2-8,1 mm long, whitish

grey, greyish or (seldom) purplish; ovary c.

0,3-0, 7 x 0,3-0, 5 mm, often with 4 ± indis-

tinct calyx lobes. Fruit reddish brown, shiny;

mericarps (1 ,5— )1 ,7—2,4 x 0,9-1 ,3(— 1 ,5) mm,

elliptic to obovate, glabrous, ± sparsely cov-

ered with very short whitish hairs or distinctly

papillate, occasionally with 2 indistinct calyx

lobes to 0,2 x 0,2 mm. Chromosome number.

2n=22.

One of the most common and widely distributed south-

western Cape species of Anthospermum, A. galiodes is eco-

logically eurytopic and highly variable in most of its charac-

ters. It is, nevertheless, possible to subdivide the species

into two subspecies which are (at least in part) ecologically

distinct and normally easily separated morphologically.

A. galioides and allied species (nos. 11 & 12, 14-16)

form a highly complex group. Some forms of A. galioides

may morphologically rather closely resemble A. pumilum

subsp. pumilum (no. 11), others may be similar to A. pros-

tratum (no. 14) or approach A. dregei subsp. ecklonis (no.

15) in certain characters. In terms of absolute mea-

surements, several characters often overlap considerably

between these taxa and, in addition, some characters are

variable within a given taxon. Differences sometimes only

become evident in the field and/or if their growth habit and

habitat are known.

Two subspecies are recognized:

13 (a), subsp. galioides.

A. ciliare L. var. angustifolium Eckl. & Zeyh.. Enum.

366 (1836). Type: Cape, ... ad radicem montis ‘Duyvels-

berg’, in monte prope "Tokay' (Cap), in montibus

‘Hottentottshollands- et Hauhoeksberge' (Stellenbosch),

prope ‘Caledon’, Ecklon & Zexher 2308 y (FI!; LY!; M!;

MO!; S!; SAM!; W!; as ‘51-8; Caledon': S!).

A. ciliare L. var. glabrifolium Sond. in F.C. 3: 29

(1865). Type: none given.

A. ciliare L. var. latifolium Eckl. & Zeyh., Enum. 366

(1836). Type: Cape, prope ‘Somerset’ in ‘Hottentottshol-

land’ (Stellenbosch), Ecklon & Zexher 2308P (FI1; GOET';

LY!; M!; MO!; S!; SAM!; W!).

A. ciliare L. var. papillatum Sond. in F.C. 3: 29 (1865);

Salter in J1 S. Afr. Bot. 3: 110 (1937). in Adamson &

Salter, FI. Cape Penins. 733 (1950). Type: Cape, Simon's

Bay and Rietvalley; no collector given.

A. ciliare L. var. scabrum Eckl. & Zeyh., Enum. 366

(1836), pro parte. Type: Cape, . . . inter ‘Coega- et

Zondagsrivier’ , in planitie prope ‘Krakakamma’ inque

alveo lapidoso fluminis ‘Zwartkopsrivier’ (Uitenhage),

Ecklon & Zeyher 2308 6 (B!; FI!; G!; GOET!; LY-one

sheet!; P-one sheet!; S!; SAM!; W!) [specimens mixed,

subsp. reflexifolium in other herbaria].

A. aethiopicum L. var. ciliare (L.) Kuntze, Rev. Gen. 3:

1 17 (1898). Type: as for A. ciliare.

Sex distributions and growth form as for

species as a whole. Stems : younger parts mostly

densely covered with short whitish hairs or

papillae. Leaves : blades linear to ovate-lanceo-

late, at least margins ciliate; if glabrous, blades

ascending or spreading and straight, not dis-

tinctly recurved. Fruit distinctly papillate or

shortly hairy; if glabrous, mericarps mostly not

longer than 2 mm. Fig. 5: 8.

MAPS 20-24: — 20. Anthospermum galioides subsp. gali-

oides

21. Anthospermum galioides subsp. re-

flexifolium

22. Anthospermum dregei subsp. dregei

23. Anthospermum dregei subsp ecklo-

nis

24. Anthospermum comptonii

1 ,2: 30

Anthospermeae

Widely distributed in the south-western Cape and, in

the east, slightly extending beyond the Cape Floristic Re-

gion to the Alexandria and Albany distr. Found in all princi-

pal vegetation types of the Cape Flora except for the West

Coast Strandveld and occurring on rocky slopes, in sandy

flats or gravelly areas and sometimes also in disturbed sites;

growing over a variety of different substrates (TMS, shale,

granite, Witteberg quartzite, limestone). Map 20.

Vouchers: Oliver 4368 (K; PRE; STE); Taylor 3805

(K; PRE; STE); Van Breda & Joubert 1927.

‘Typical’ subsp. galioides is characterized by having

ovate-lanceolate to linear-lanceolate (due to revolute mar-

gins) leaves with ciliate margins; plants are often many-

stemmed and ± densely tufted; fruits are either glabrous

and shiny or shortly hairy. This ‘typical’ form appears to be

centred in the Cape Peninsula.

Two more ‘forms’ (ecotypes) are occasionally quite

distinct:

— The 'Papillatum Form’: plants with ± few to seve-

ral, often ± erect stems; leaves (narrowly) lanceolate to

linear, glabrous; fruits mostly distinctly papillate. In habit it

may resemble A. pumilum subsp. pumilum (no. 1 la, which,

however, does not extend into the Cape Floristic Region).

This form may be ± confined to Mountain Fynbos and

TMS areas; recorded from the Cape Peninsula east to the

Caledon distr. and north(-east) to the Worcester, Ceres

(Cold Bokkeveld Mts), Piketberg and Clanwillian districts

(Cedarberg).

— The 'Prostratum Form': characterized by an often

entirely prostrate habit. Stems radiate out from a common

base and form ± dense mats; they do not root at the nodes.

Leaves are small but relatively broad, margins are glabrous

or sometimes ciliate; fruits are glabrous or shortly hairy and

smaller than in the other forms (hardly 2 mm long). Occur-

ring from the Caledon distr. east to the Suurbergpas north of

Addo and north to the Ceres and Lainsburg distr.; possibly

also in the Cape Peninsula. Most frequently found in fertile

soils (often associated with Elytropappus rhinocerotis),

occasionally in limestone areas or, inland, over Witteberg

quartzites; apparently never in TMS areas. Should not be

confused with A . prostratum (no. 14).

13(b). subsp. reflexifolium (Kuntze) Puff,

comb, et stat. nov. Type: Cape, Swellendam,

Kuntze s.n. (US sub 554666, holo.!; K, iso.!).

A. aethiopicum L. var. reflexifolium Kuntze, Rev. Gen.

3: 117(1898).

A. ciliare L. var. scabrum Eckl. & Zeyh., Enum. 366

(1836), pro parte. Type: Ecklon & Zeyher 2308 5 (LY-one

sheet!; M!; MO!; P-one sheet!) [see also subsp. galioides].

Plants mostly cf, $ + 9 or densely

leafy, ± erect, cylindrical and quite woody to

less densely leafy, ± rounded, weaker and ±

subshrubby. Stems: younger parts often only

with two rows of short hairs. Leaves: blades

linear-lanceolate to lanceolate, recurved (at

least near tip) to strongly recurved, mostly firm,

thickish and almost leathery, glabrous (save

for, sometimes, minute prickles on margins).

Partial inflorescences mostly only 1-flowered.

Fruit shiny, glabrous, epidermis cells often

large and conspicuous, but mericaps never dis-

tinctly papillate or shortly hairy; mericarps rela-

tively large, c. 2-2,4 mm long.

Occurring from the Porterville Mts (Schurweberge)

south to the Caledon distr.; eastwards following the moun-

tain ranges to the Groot Winterhoekberge, the Kouga Mts

and the Humansdorp distr. Growing in dry places in Moun-

tain Fynbos, sometimes also at the edge of riparian commu-

nities, in wooded kloofs and well drained slopes; perhaps

confined to TMS areas. Map 2 1 .

Vouchers: Kruger 1423 (PRE; STE); Schlechter 9149

(BM; BOL; COI; E; G; K; MO; P; PRE; S; US; W); Taylor

9664 (PRE; STE).

In the eastern part of the south-western Cape Region,

plants often appear more robust and shrubbier than else-

where, are densely leafy and bear small, quite firm and

thickish leaves to c. 5 mm long. Some of these forms (e g.

Acocks 23082) are so different from ‘typical’ subsp. reflexi-

folium in habit and appearance that they should perhaps be

recognized taxonomically (as a closely allied new species?).

In the Caledon and Worcester distr., on the other hand,

specimens may be so close to subsp. galioides in certain

characters, that the separation of the subspecies sometimes

becomes difficult.

Within subsp. reflexifolium there appear to be two ±

well defined ecotypes:

— ‘Typical’ subsp. reflexifolium, which seems to be

confined to moderately dry areas in Fynbos vegetation (the

above mentioned eastern forms belong here).

— Plants of moister habitats which differ in habit:

dense, rounded bushes to 0,5 m in diam.; stems often much-

branched; intemodes long, plants thus appear less densely

leafy.

14. Anthospermum prostratum Sond. in

F.C. 3: 28 (1865); Salter in J1 S. Afr. Bot. 3:

110 (1937), in Adamson & Salter, FI. Cape

Penins. 733 (1950). Type: only given for

varieties cited below.

A. prostratum Sond. var. glabrum Sond. in F.C. 3: 28

(1865). Type: Cape Flats, Ecklon s.n. (S, holo.!; LY, iso.!)

[selected as type of A. prostratum].

A. prostratum Sond. var. velutinum Sond. in F.C. 3: 28

(1865). Type: Cape Flats, C. Wright [491] (S, holo.!).

Dwarf shrub or subshrub with basally

woody stems, dioecious (very seldom C? with

odd 9"), prostrate, often with ± thick, woody

tap root. Stems trailing, rooting at nodes, to c. 1

m long, with ascending to ± erect short lateral

branches, c. 10-50(-100) mm long. Leaves de-

cussate, pseudoverticillate; blades (5— )7,5— 1 2

x 0,7-2(-2,3) mm, lanceolate, oblanceolate to

± linear-lanceolate, glabrous or (lower half of)

margins papillate; petioles subobsolete; stipular

sheath mostly with a small ± broadly triangular

Anthospermeae

1,2: 31

seta. Flowers subsessile, on short lateral

branches, single or paired at nodes (seldom in

paired 3-flowered cymes); corolla 4-merous

(very seldom also 5-merous in cf ), yellowish to

greenish yellow, frequently tinged reddish

purplish outside, glabrous, cf (odd $): tube

1.2- 2, 3 mm long, narrowly funnel-shaped,

lobes (2-)2,4— 3(-3,7) x 0,7-1 mm; anthers

1,5-2, 2 mm long; cf: small rudimentary ovary

present. 9: tube (0,5— )0,7— 0,9(— 1 ,1) mm long,

lobes 0,5-1 x 0, 1-0,2 mm; style 0-1,3 mm

long; stigmas 2, 4,8-10 mm long (shorter in

odd <J?); ovary 0,7-1 ,4 x 0,5-1 mm, not

crowned by conspicuous calyx lobes. Fruit red-

dish (brown), shiny; mericarps (2— )2,3— 2,9 x

1.2- 1 ,7 mm, (ob)ovate, glabrous or with very

short whitish hairs, not crowned by distinct ca-

lyx lobes. Chromosome number : 2n = 22.

Endemic to the south-western Cape and occurring in

coastal areas from the West Coast (Malmesbury distr.) and

Cape Peninsula east to the Port Elizabeth distr. Confined to

dunes and sandy flats, c. 0-50(-75) m. Map 26.

Vouchers: Acocks 20675 (K; M; PRE; SRGH): Bohnen

7605 (PRE; STE); Tolken 35 (PRE; STE).

Well distinguished by its growth form and habitat.

Relatively uniform in its characters except for the fruits

(glabrous or shortly hairy).

15. Anthospermum dregei Sond. in F.C.

3: 29 (1865); Launert & Roessler in F.S.W.A.

115: 7-8 (1966). Type: Cape, between Koussie

and Zilverfontein, Drege 3016 (S, holo. ! 1 ; E;

G; K; LY; MO; P; S'; SAM; W, iso.!).

Dwarf shrub, cf, <?, 9" + 9 or 9’ many-

stemmed, diffusely to ± regularly branched,

often ± rounded. Stems c. 150-400 mm long,

branches ± irregular (especially if browsed),

often divaricate. Leaves decussate, sometimes

± pseudoverticillate; blades 4-14(-20) x

1— 4( — 4,5) mm, ± ovate, oblong-lanceolate to

± lanceolate, ± membranaceous to thickish,

glabrous or with a few papillae on margins,

upper and/or lower surface; upper surface

sometimes distinctly purplish brown, lower sur-

face (light) green; margins ± revolute, con-

voluted rolled or ± flat; petioles ± 0-1 mm

1 There are two sheets in S. Only one of them, the sheet

containing two specimens (‘3016 cf’ and ‘3016 $’),

bears the addition Dregei Sond.’ and ‘8’ ( = species

number in Flora Capensis) in Sonder's handwriting; this

must be considered the holotype.

long; stipular sheath with or without a minute

seta. Flowers subsessile (only in 9 pedicels

occasionally to 1 mm long), in clusters of 2-6 at

nodes; corolla 4-merous (in 9 occasionally also

5-merous), greenish yellow to creamy yellow or

reddish purple, at least outside, sometimes

papillate, cf, 9" : buds 2,5-4 mm long; tube

0,7-1 ,4(— 1 ,8) mm long, (narrowly) funnel-

shaped to ± cylindrical, lobes 1,5-3, 2 x

0,5-1 ,2 mm; anthers 1 ,3-2,8 mm long; 9 : stig-

mas 2, c. 1-3 mm long, greyish white, ovary c.

0,7-1 ,5 x 0,5-1 mm, often with 4 indistinct

calyx lobes; cf: small rudimentary ovary pre-

sent. 9: tube ± 0 or 0, 1-0,7 mm long, lobes

0,2-l,3(-2) x 0, 1-0,5 mm; style 0-0,7 mm

long, stigmas 2, (2,5-)3-8,5 mm long, greyish

white or (seldom) reddish purplish; ovary as in

9" or somewhat larger. Fruit reddish brown,

shiny; mericarps (1 ,5-) 1 ,8-2,7 x 0,8-1, 5 mm,

± oblong, glabrous or with a few papillae,

mostly with 2 indistinct ± triangular calyx

lobes to c. 0,2-0, 3 mm long. Chromosome

number. 2n = 22.

Two subspecies are recognized:

15 (a), subsp. dregei.

A. thymifolium Dinter & Krause in Feddes Reprium 15:

91 (1917), nom. non valide publ. (sine descr.).

Robust dwarf shrub, thick-stemmed, to 400

mm tall, often irregularly (diffusely) and divari-

cately branched (especially if browsed);

browsed plants sometimes ± cushion-forming

and low; intemodes normally much longer than

leaves. Leaves : blades often broad and large, to

14(-20) x 4(-4,5) mm, ovate, ovate-lanceolate

or oblong-lanceolate, thickish and tough,

mostly papillate on lower and/or upper surface

and margins, distinctly discolourous, upper sur-

face often very dark; petioles usually c. 0,5-1

mm long. Plants mostly cf, 9’ 9 fairly uncom-

mon. cf (9*): buds c. 3,5-4 mm long; corolla

lobes 2, 3-3, 2 x 0,7-1, 2 mm, glabrous. Fruit

shiny, glabrous, carpophore c. 0,6-0, 7 mm

long; mericarps (1 ,8— )2, 2-2,' 7 x 1-1,5 mm.

Known from the Namaqualand distr. and adjacent

parts of South West Africa/Namibia (Liideritz-Sud, Warm-

bad and Keetmanshoop distr.). Growing in semi-desert and

desert areas; most commonly found in crevices, amongst

rocks or sheltered under boulders on granite hills and out-

crops, sometimes over mica-slate, seldom over sandstone.

Map 22.

1,2:32

Anthospermeae

Vouchers: Goldblatt 2397 (BR: MO; PRE; WAG);

Marloth 12225 (PRE; STE); Merxmuller & Giess 3426

(BR;M; PRE; WIND).

Easily distinguished from subsp. ecklonis (below) by

its more robust habit, its tough, thickish leaves and its clear

trend to dioecy. These characters, however, are often only

obvious in the field. Sheets only consisting of new shoots of

browsed plants are difficult to identify (similar in both

subspecies). Subsp. dregei occasionally closely approaches

A. pumilum subsp. rigidum (no. lib) in some morphologi-

cal characters but the distribution ranges of the two taxa do

not overlap.

15 (b). subsp. ecklonis (Sond.) Puff, comb,

et stat. nov. Type: Cape, on the Olifantsriver

and near Villa Brakfontein, Ecklon s.n. (S,

holo. !; LY, iso. !).

A. ecklonis Sond. in F.C. 3: 32 (1865).

Rather weak dwarf shrub, thin-stemmed,

often rounded, to c. 200 mm tall (less com-

monly more erect and to 400 mm tall), ± regu-

larly branched (unless browsed), branches often

numerous, ascending to erect; intemodes often

not much longer than leaves (except on new

growth). Leaves', blades often narrow and short,

4 — 9(— 1 2) x 1— 2,5(— 3) mm, (narrowly) oblong-

lanceolate to lanceolate, ± membranaceous,

glabrous or papillate (mainly on margins), ±

discolourous; margins often revolute; petioles

usually subobsolete. Plants 9” , 9 + $ or $,

rarely cf. $(Cf): buds c. 2,5-3,2(-3,8) mm

long; corolla lobes 1 ,5— 2,5(— 3) x 0,5-0, 8 mm,

mostly a little hairy or papillate outside. Fruit

shiny, glabrous or covered with some papillae,

carpophore c. 0,3-0, 5 mm long; mericarps

( 1 ,5—) 1 ,8—2,2 x 0, 8-1,1 mm.

Endemic to the western part of the Cape Floristic Re-

gion; occurring from the south-western Calvinia distr. and

Clanwilliam distr. south to the Tulbagh distr. where its

distribution range closely follows the distribution of TMS.

Growing in crevices, in gravelly soil or sandy places be-

tween rock(sheets) or in depressions of large open rock

surfaces in areas of very dry Fynbos. Map 23.

Vouchers: Acocks 15194 (K; PRE); 17479 (K; PRE);

Oliver 3836 (K; PRE; STE).

Should not be confused with A. comptonii (below) and

with the closely allied and very variable A. galioides subsp.

galioides (no. 13a), some forms of which may rather clo-

sely resemble A. dregei subsp. ecklonis in leaf size and

shape and fruit morphology.

16. Anthospermum comptonii Puff, sp.

nov., ab A. dregei subsp. ecklone mericarpiis

longioribus et floribus masculis majoribus dif-

fer!.

Type: Cape, south side of Witteberg, farm

Fisantekraal, c. 1200-1300 m. Puff 7909 14-3/2

(WU, holo. !; BOL; NBG; PRE; STE, iso. !).

Dioecious dwarf shrub, many-stemmed, ±

regularly to diffusely branched, rounded to ±

cylindrical or, if browsed, low cushion-like;

with thick woody, sometimes twisted (tap-)root

to c. 10(— 20) mm in diam. Stems to c.

(200-)300 mm long, branches few to numer-

ous, ascending to ± erect, often irregular and ±

spine-tipped if browsed. Leaves decussate;

blades 5— 8(— 10) x 0,8-1 ,2 mm, narrowly

ovate-lanceolate to linear-lanceolate (broader,

to 2 mm, on new growth of browsed plants),

thickish and ± tough, glabrous except for odd

papillae on margins; upper surface shiny, with

large conspicuous epidermis cells, midrib

below often ± prominent and broad; margins

usually revolute; petioles subobsolete; stipular

sheath with an often indistinct minute seta.

Flowers subsessile (in fruiting 9 pedicels occa-

sionally c. 0,2-1 mm long), single or paired at

nodes; corolla 4-merous, (greenish) yellow or

(dark) purplish at least outside, glabrous or

occasionally papillate near tip. cf: buds

3, 5-4, 2 mm long; tube 1-1,7 mm long, (nar-

rowly) funnel-shaped, lobes 2, 7-3, 2 x 0,7-1

mm; anthers 1,9-2, 4 mm long; small rudimen-

tary ovary present, often with broad ± rounded

calyx lobes to 0,3 mm long. 9 : tube 0, 1-0,2

mm long or ± 0, lobes 0,7-0,8(-l) x 0,2-0, 3

mm; style ± 0-0,7 mm long; stigmas 2,

2, 7-5, 6 mm long; ovary 1-1,4 x 0,5-0, 9 mm,

with ± indistinct calyx lobes. Fruit reddish

brown, shiny; mericarps 2,7-3 x 1,5-1, 7 mm,

± obovate, glabrous or (rarely) with odd papil-

lae, with 2 ± indistinct, rounded to ± triangu-

lar calyx lobes c. 0, 1-0,3 x 0,3 mm. Chromo-

some number. 2n—22.

Occurring inland of the major south-western Cape

folded ranges from the southern Schurweberge and the

Swartruggens to the Witteberg and the Touwsberge (Ceres,

Lainsburg and Ladismith districts); also further east in the

western Uniondale distr. Growing in rock fissures and crev-

ices, among boulders or at the base of rocks in Arid Fynbos

but also intruding into (open succulent) Karoo vegetation;

often in Witteberg quartzite areas. Map 24.

Vouchers: Compton 3662 (BOL); Levxns 7482 (BOL);

Taylor 5917 (PRE; STE).

Closely allied to A. dregei susbp. ecklonis (above) but

distinguished by its larger mericarps and larger cf flowers.

A. comptonii, furthermore, is strictly dioecious, occurs in

more arid areas than the former and does not grow on TMS-

derived substrates. Browsing appears to account for most of

the variability in the species. Continued browsing results in

low cushion-like growth forms.

Anthospermeae

1,2: 33

C f specimens should not be confused with Nenax el-

sieae which is similar in habit and, in part, occurs in the

same general area. The latter differs in having needle-like

leaves which are ± triangular to semiterete in section.

17. Anthospermum esterhuysenianum

Puff, sp. nov., combinatione characterum: ha-

bitu implexo vel caulibus serpentibus, floribus

solitariis terminalibus ramis brevibus saepe su-

peratis, corollis 5-meris facile distinguitur .

Type: Cape, Ceres-Tulbagh distr. , Wit-

zenbergen, Swartgat Peak, Esterhuysen 27926

(BOL, holo.!).

Subshrub, ± woody near base and with

woody tap root or ± dwarf shrub, mat-forming

or trailing, ^ $ or occasionally (j.

Stems c. (60-) 120-300 mm long, prostrate,

mostly much branched, branches often single at

nodes or, if paired, frequently of unequal

length. Leaves decussate, sometimes ± pseudo-

verticillate, occasionally slightly anisophyllous;

blades 4— 6(— 7 ,5) x ( 1 ,4 — )2— 3C— 4) mm, oblong,

elliptic, ovate, ovate-lanceolate to lanceolate,

glabrous or margins and often also upper sur-

face with white spreading hairs c. 0, 2-0,4 mm

long; petioles ± 0-0,5 mm long; stipular sheath

with a small seta. Flowers solitary, terminal on

shoots, often overtopped by lateral branches

arising below, shortly pedicellate, pedicels c.

0,2-0,3( — 1) mm long, often slightly elongated

and curved in fruit; corolla 5-merous (very rare-

ly 4-merous), yellowish to greenish yellow, of-

ten dark purplish tinged outside, glabrous or

hairy. $ , cf: tube 1—1 ,8(— 2) mm long, ± cylin-

drical or narrowly funnel-shaped, lobes

(2— )2,2— 3(— 3,5) x 0,5— 1 (—1 ,3) mm; anthers

1,4-1, 9 mm long; cf: rudimentary ovary

minute; $ : gynoecium as in 9- tube ±

0-0,3 mm long, lobes 0,6-0, 9 x 0,4 mm; style

0; stigmas 2, (4,5— )5 ,5— 7,5(— 9) mm long; o-

vary c. 0,8-1 ,2 x 0,7—1 mm, with 5 minute

calyx lobes. Fruit brownish; mericarps 1 ,9-2,5

x 1 ,3-1 ,5 mm, ovate, ± obovate or oblong, ±

densely papillate, mostly with 2 or 3 indistinct

calyx lobes (0,2-)0,3-0,5 x 0,2 mm. Chromo-

some number: 2n—22.

Occurring in the higher parts of the south-western

Cape mountains in rocky places; apparently confined to

shale bands and shale-derived soils.

A very distinct and unmistakable Cape mountain

endemic. It should not be confused with low-growing eco-

types of A. galioides subsp. galioides (no. 13a) which do

not occur in the mountains and differ in branching pattern,

inflorescence and their 4-merous flowers. Resemblances to

A. galioides are superficial and do not point to a close

relationship; the species occupies a rather isolated position

in the genus.

Two varieties are recognized:

17 (a), var. esterhuysenianum.

Leaves and corollas glabrous; leaf blades

oblong, elliptic or ovate, apices ± obtuse or

acute.

Confined to the high mountains in the triangle Clanwil-

liam-Stellenbosch-Worcester. Map 27.

Vouchers: Esterhuysen 22599 (BOL; K; PRE); 22599a

(BOL; K; NH; PRE); 34159a (BOL; PRE).

MAPS 25-32: — 25. Anthospermum paniculatum

26. Anthospermum prostratum

27. Anthospermum esterhuysenianum

var. esterhuysenianum

28. Anthospermum esterhuysenianum

var.hirsutum

29 Anthospermum bergianum

30. Anthospermum ericifolium

31. Anthospermum bicorne

32. Anthospermum hirtum

17(b). var. hirsutum Puff, var. nov., a

var. typica corolla extra et foliis margine sal-

tern patule a I bo- pi I os is differt.

Type: Cape, Ceres distr., northern Cold

Bokkeveld, Schurweberg Peak, Esterhuysen

29458 (BOL, holo.!).

1,2: 34

Anthospermeae

Outside of corolla, leaf margins and often

also upper surface covered with white spreading

hairs c. 0,2-0, 4 mm long; leaf blades lanceolate

to ovate-lanceolate, apices mostly ± distinctly

mucronate.

Only known from the Central Cedarberg and the

Schurweberg (northern Cold Bokkeveld). Map 28.

Vouchers: Esterhuysen 34826 (BOL; K; WU); 29431

(BOL; PRE).

18. Anthospermum hirtum Cruse, Rub.

Cap. 11 (1825), in Linnaea 6: 13 (1831); Sond.

in F.C. 3: 30 (1865); Salter in J1 S. Afr. Bot. 3;

110 (1937), in Adamson & Salter, FI. Cape

Penins. 733 (1950). Type: Cape, ad promonto-

rium bonae spei in monte Diaboli Orientem ver-

sus, mense Augusto 1816, C. W. Bergius s.n.

(B, holo. t; G, iso.!).

A. rubiaceum Reichenb. f. in Spreng., Syst. Veg. 4: 338

(1827). Type: ‘C.B.S.’, no collector given.*

'A. hirsutum' DC., Prodr. 4: 580 (1830).

Subshrub or distinct dwarf shrub, cf,

9" + 9,$or^ + cr, few- to several-

stemmed, ± cylindrical, rounded or low, ±

matted and weak, often with ± thick, woody

(tap-)root. Stems c. 0,2—1 (—1 ,5) m long, often

± much branched; ± densely covered with

whitish spreading hairs c. (0,2— )0,5— 1 ,4 mm

long, or hairs sometimes only in two rows, ra-

rely only around stipules. Leaves decussate, oc-

casionally ± pseudoverticillate; blades (6-)

8— 25(— 30) x (0,5-)0,8-3(-4,5) mm, narrowly

oblanceolate to ± linear-lanceolate, often dis-

tinctly discolourous, upper surface shiny, epi-

dermis cells large, conspicuous, lower epider-

mis distinctly papillate; margins, at least near

base, and often also upper surface ± densely

covered with white spreading hairs c.

0,4— 1 (— 1 ,5) mm long; petioles ± 0-1 mm long;

stipular sheath with a subulate hairy seta c.

(0,5—) 1— 2,5(— 3) mm long. Flowers subsessile

to shortly pedicellate (in fruiting 9 pedicels

occasionally to c. 0,7 mm long), in clusters of

6-2 at nodes; corolla 5-merous (occasionally

also 4-merous in 9). greenish yellow or (dark)

* Cruse in Linnaea 6: 13 (1831) claims having seen ‘the

author’s’ specimen when putting A. rubiaceum into syno-

nymy with his A. hirtum, but he did not cite the collec-

tion. Sieber FI . Cap. No. 413 (G ! ; MO!; W!) is likely to

be the type specimen. The sheet in W bears the label

'Anthospermum rubiaceum, Cap, Reichenb.’ (likely to be

in Reichenbach's handwriting) next to Sieber's distinct

printed ‘FI. Cap’ label which is directly attached to the

plant.

purplish red, with whitish spreading hairs at

least near tip. cf , tube 0,5-1 ,5 (-2) mm long,

(narrowly) funnel-shaped, lobes (1,7-)

2—3 ,4(— 3 ,7) x 0,5-0,8(-1) mm; anthers

( 1— ) 1 ,4 — 2(— 2,2) mm long; cf: small rudimen-

tary ovary and sometimes also rudimentary stig-

mas present; $ : style 0-1 mm long; stigmas 2, c.

2— 3(— 5 ,5) mm long, greenish white, greyish or

(seldom) reddish purple; ovary c. (0,5-)

0,9-1 ,2 x 0,5-1 mm, not crowned by distinct

calyx lobes. 9: tube 0,3— 0,8(— 1 ,2) mm long,

lobes (0, 1— )0,3— 1 x 0, 1-0,3 mm; gynoecium

as in 9 but stigmas 3, 7-7, 5 mm long. Fruit

reddish brown; mericarps 1,9-2, 7 x

( 1— )1 ,2—1 ,7 mm, ± oblong, with large conspic-

uous epidermis cells or (rare-ly) with odd hairs

or papillae, not crowned by distinct calyx lobes.

Chromosome number. 2n = 22.

Endemic to the south-western Cape and occurring from

the Cape Peninsula north to the Clanwilliam and south-east

to the Bredasdorp distr. Typically growing in ± moist to

damp sites (along streams, well drained slopes, base of

cliffs etc.) in partial shade, occasionally in drier places

(amongst rocks or boulders in sandy to gravelly soil); occur-

ring mainly on TMS-derived substrates, sometimes in Gran-

ite (clay) soils. Map 32.

Vouchers: Salter 6221 (BOL; K); Schlechter 10722

(BM; BOL; COI; E; G; GRA; K; MO; PRE; S; US; W);

Taylor 6883 (PRE; STE).

A record from the Riversdale distr. ( Muir sub Marloth

5655; PRE) appears very doubtful (dot with question mark

in Map 32); Muir, in ‘Vegetation of the Riversdale area,

Cape Prov.' [Men bot. Surv. S. Afr. 13 ( 1929)] does not list

A. hirtum as occurring in that area.

A variable species. ‘Dwarf’ or stunted forms with rela-

tively small leaves appear to occur in drier habitats and, in

general, under less favourable environmental conditions

(particularly obvious in the dwarf Fynbos of the Elim

flats!).

Some collections from the Piketberg area (3218-BC)

differ from ‘typical’ A. hirtum in being ± weak and thin-

stemmed, in having hairs around the stipules and base of the

leaf blades only and in their 1 -flowered partial inflores-

cences [e.g. Drege 7667 (E!;G!; K!; LY!; MO!; P!; S!; W!;

as ‘64.9’: FI!); Bolus 8505 (BOL!)]. ‘Typical’ A. hirtum

occurs in the same area and some collections are ± interme-

diate. Further field observations would be required to deter-

mine whether this form deserves formal taxonomic recogni-

tion.

Anthospermeae

1,2: 35

19. Anthospermum bergianum Cruse',

Rub. Cap. 9 (1825), in Linnaea 6: 7 (1831);

Sond. in F.C. 3: 29 (1865); Salter in J 1 S. Afr.

Bot. 3: 110 (1937), in Adamson & Salter, FI.

Cape Penins. 733 (1950). Type: Cape, in plani-

tie capensi versus Tygerberg, Aug. 1816, C. W.

Bergius s.n. (B, t); Cape Flats, Aug. 1818,

Mund(t) 91 (S, neo.!)2.

? Cliffortia spicata Reichb. f. [in sched.] ex Sprengel,

Syst. Veg. 4, Cur. Post. 209 (1827). Type: ‘C B S ', no

collector given.* 1

Subshrub or ± short-lived dwarf shrub,

mostly dioecious (seldom also cf + odd $ or

$), single- to few-stemmed, usually erect, with

woody tap root to 220 mm long. Stems (50—)

80-150 mm long in younger, occasionally to

450(-750) mm long in older flowering speci-

mens, sparingly to much branched, branches

ascending to erect, densely leafy above. Leaves

in whorls of 3, occasionally in whorls of 4 or

decussate, often pseudoverticillate; blades

mostly erect or ascending, ± imbricate,

(4 — )5— 1 2(— 1 5) x (1 ,2— )1 ,5— 3(— 3,5) mm, lin-

ear-lanceolate to ovate-lanceolate, margins, so-

metimes also mid-rib below or entire lower sur-

face, covered with white spreading hairs

(0,3-)0,5-l mm long, often reddish brown

above; petioles 0; stipular sheath with or wi-

thout a minute seta. Flowers subsessile, in clus-

ters of 3-9 at nodes, in 9 sometimes more; 9

inflorescences dense and ± cylindrical, c.

(20— )30— 1 20(— 1 70) x 10-20 mm; corolla 5-

merous (in 9 sometimes also 4-merous), yel-

lowish, sometimes reddish tinged outside, with

white hairs c. 0,3-0, 5 mm long at least near tip

or glabrous (especially 9)- Cf (odd 9"); tube

0,7-1 mm long, funnel-shaped, lobes 1 ,9-3 x

0,6-1, 1 mm, anthers 1-1,6 mm long; rudimen-

1 IMPORTANT NOTE: The name A. bergianum Cruse is

used on the assumption that the ‘Proposal to reject the

name A. ciliare L. (1763) (Rubiaceae)’ (proposal 685) is

accepted. Otherwise the name A. ciliare will have to re-

place A. bergianum. See also A. galioides (no. 13) and

Puff in Taxon 31: 759 ( 1982) for details.

: Specimen seen by Cruse and bearing the inscription 'A.

bergianum mihi. Cruse’. He may, in error, have attributed

the specimen to Ecklon, because in Linnaea 6: 7 ( 1831 ) he

wrote ‘in planitie capensis legerunt d. Bergius nec non

C. F. Ecklon’ but he did not cite Mund(t).

1 Weimarck [Monograph of the genus Cliffortia 160 ( 1934)]

states that ‘the Reichenbach type [of C. spicata] corre-

sponds to Anthospermum bergianum Cruse’ . As I

was unable to trace the type specimen I could not check

the correctness of this statement.

tary ovary (cf) to c. 0,2-0, 5 mm long. 9: tube

(0, 1— )0,3— 0,6 mm long, lobes 0,5-1, 2 x

0,2-0, 4 mm; style 1-2,8 mm long; stigmas 2,

(6— )8— 1 3 ,5 mm long; ovary 0,8-1, 5 X 0,6-1

mm, not crowned by distinct calyx lobes. Fruit

brown or reddish brown, often ± shiny; meri-

carps 1,8-2, 5 x 0, 9-1,1 mm, ± obovate to

oblong, ± densely papillate or some-

times with white spreading hairs to c. 0,2 mm

long, not crowned by distinct calyx lobes.

Chromosome number: 2n = 22.

Endemic to the south-western Cape and occurring from

the Clanwilliam distr. (Pakhuis Pass) south to the Caledon

distr. and the Cape Peninsula. Mostly growing on (dry)

rocky slopes in gravelly to fine sandy soil; usually asso-

ciated with TMS; often in recently burnt areas. Map 29.

Vouchers: Esterhuysen 13059 (BOL; PRE); Goldblatt

2652 (MO; PRE; WAG); Thompson 1556 (PRE; STE).

A. bergianum is killed by fire and regenerates from

seed. Young flowering plants, often only a few cm tall and

(much) less than one year old, are most common in recently

burnt areas. The plants, however, are not annuals; if not

exposed to fires they become ± shrubby.

Although variable in the fruit indumentum, in leaf

arrangement and in the degree of the hairiness of the leaves,

A. bergianum is one of the most easily recognizable Cape

species of Anthospermum.

20. Anthospermum ericifolium (Licht.

ex Roem. & Schult.) Kuntze, Rev. Gen. 3: 1 17

(1898) as ericaefolium. Type: Cape, nr. Rivier

Zonderend (=Riviersonderend), Lichtenstein

s.n. (?B,t).

Spermacoce ericaefolia Licht. [Spicileg. FI. Cap., MS.]

ex Roem. & Schult., Syst. Veg. 3: 281 ( 1818).

Anthospermum lichtensteinii Cruse, Rub. Cap. 15

(1825), in Linnaea 6: 16 ( 1831 ); Sond. in F.C. 3: 32 (1865).

Type: as above.

Dwarf shrub, ^ , 9 ’ seldom ^ + Cf or cf ,

few- to many-stemmed, ± erect. Stems

( 1 50-)250-400(-600) mm long, sparsely to

much branched, branches ± erect or ascending,

densely leafy above. Leaves decussate, pseudo-

verticillate; blades ascending to erect, 4—7 x

(0,6— )0,8— 1 ,2(— 1 ,4) mm, linear-lanceolate,

often broadly triangular and shallowly concave

above in section; margins, at least lower half,

with white spreading hairs c. 0,3-0, 4 mm long,

upper surface and midrib below often reddish

brown; petioles 0; stipular sheath with or with-

out a minute seta. Flowers subsessile, in

clusters of (2— )6— 10(— 1 4) at nodes; corolla 5-

merous, yellowish!?), with white spreading

hairs c. 0,2-0, 3 mm long at least near tip. <j

tube 0,7-1 mm long, (narrowly) funnel-shaped

to ± cylindrical, lobes, 1,2-1 ,7 x

1,2:36

Anthospermeae

0,5-0, 8(-l) mm; anthers 1-1,2 mm long: cf:

rudimentary ovary minute, hardly discernible; :

gynoecium as in 9. 9 : tube c. 0,2 mm long,

lobes 0,4-0, 6 x 0, 1-0,2 mm; style 0; stigma

one, 3-7 mm long; ovary with only one fertile

carpel, c. 1-1,3 x 0,5-0, 8 mm, densely hairy;

reduced carpel with 2, fertile carpel with 3

calyx lobes. Fruit greyish brown to greyish;

fertile mericarp (2,5-)2,8-3,5(^l) x (1-)

1,3-1, 8 mm, ± ovate to oblong, often curved

lengthwise, densely covered with straight white

hairs 0,4-0, 6 mm long; reduced carpel ± strap-

like, 2, 5-3, 5 x 0,5-0, 7 mm; fertile carpel with

3 triangular-lanceolate calyx lobes 0,3— 0,7(— 1 )

x0, 2-0,4 mm, reduced carpel with 2 some-

times marginally larger ones.

Endemic to the south-western Cape and occurring from

the Cape Flats to the western Worcester and the Caledon

distr. Growing in sandy to gravelly soil; mostly in sandveld

Fynbos. Map 30.

Vouchers: Esterhuysen 35553 (BOL; PRE; WU);

Schlechter 10242 (BM; BOL; COI; E; G; GRA; K; MO; P;

PRE; S; US; W).

A rather rare but unmistakable species (gynoecium

with only one fertile carpel and one stigma). Allied to A.

bicorne (below) but easily distinguished by ovaries and

fruits with 5 subequal calyx lobes.

21. Anthospermum bicorne Puff, sp.

nov., carpello sterili lobis calycis duobus multo

amplificatis coronato ab A. ericifolio praeclare

distinguitur .

Type: Cape, Caledon distr., Houw Hoek,

Gillen 829 (BOL, holo. !; STE, iso. !).

Dwarf shrub or subshrub, (f + £ or 9 ,

many-stemmed, usually ± erect. Stems

( 100-)200-550 mm long, sparsely to ± much

branched, branches ascending to erect, not

densely leafy. Leaves decussate, pseudoverti-

cillate; blades erect or ascending, (6,5-)

8— 16(— 19) x 0,4-0, 5 mm, linear, needle-like,

mostly ± triangular to ± semiterete in section,

glabrous or margins (often lower third only)

very shortly hairy; petioles 0; stipular sheath

with a minute seta. Flowers subsessile, in clus-

ters of 2-6 at nodes; corolla 5-merous, yellow-

ish or purplish outside, papillate or shortly hairy

mostly near tip. 9" : tube 0,8-1 ,3 mm long, fun-

nel-shaped, lobes 1,7-2, 5 x 0,4-0, 7 mm;

anthers 1,6-2, 4 mm long; style c. 0,7-0, 8 mm

long; stigmas 2, c. 2,5-4 mm long; ovary with

only one fertile carpel, 0,8-1 ,5 x 0,5-0, 7 mm,

densely hairy; reduced carpel with 2 large, fer-

tile lobes with 3 minute, indistinct calyx lobes.

9: tube (0,l-)0,3-0,4 mm long, lobes 0,3-0, 6

x 0, 1-0,2 mm; gynoecium as in ^ but stigmas

to 6,5 mm long. Fruit reddish brown to greyish;

fertile mericarp 2, 5-3, 5 x 1-1,7 mm, ± ovate,

sometimes distinctly curved lengthwise, den-

sely covered with white spreading hairs 0,3-0, 5

mm long, sometimes with 3 minute calyx lobes

c. 0, 1-0,4 mm long; reduced carpel ± strap-

like, c. 2,5— 3,5 x 0,6-0, 8 mm, less hairy than

fertile one or glabrous, with 2 large, ± divari-

cate calyx lobes 1,4—2 x 0,2-0 ,4 mm. Chro-

mosome number: 2n=22. Fig. 5: 1-3.

Endemic to the south-western Cape and occurring from

the Clanwilliam distr. (Cedarberg Mts) south to the Cape

Rats and to the Caledon and the Bredasdorp distr. Growing

mostly in dry (coarse) sandy soil, usually over TMS. Map

31.

Vouchers: Bolus 341 (BM; G; K; PRE; SAM; US; W;

as ‘5377’: BOL); Esterhuysen 7428 (BOL; NBG; PRE;

SAM); Hugo 907 (K; PRE; STE).

Its habit and needle-like leaves are reminiscent of

Nenax species but A. bicorne is distinguished from both

Nenax and A. ericifolium (above) by the two greatly en-

larged calyx lobes borne on the modified sterile carpel of

the fruit.

Excluded species

Anthospermum calycophyllum Sond. in F.C. 3: 31 (1865) is

Otiophora calycophylla (Sond.) Schltr. <4 K. Schum. in

Bot. Jb. 30: 416 (1901); see Puff in J 1 S. Afr. Bot. 47:

311 (1981).

Anthospermeae

1,2: 37

8439 NENAX

Nenax Gaertn., Fruct. 1: 165, t. 32, f. 7 (1788); Hook. f. in Benth. & Hook, f., Gen. Plant. 2: 140

(1873); K. Schum. in Pflanzenfam. 4,4: 129 (1891); Salter in J1 S. Afr. Bot. 3: 111 (1937), in

Adamson & Salter, FI. Cape Penins. 734 (1950); R. A. Dyer, Gen. 1: 622 (1975). Type species:

N. acerosa Gaertn.

Ambraria Cruse [non Heisterex Fabr.], Rub. Cap. 16 (1825), in Linnaea 6: 18 (1831); Sond. in F.C. 3: 33 (1865). Type

species: A. glabra Cruse.

Dwarf shrubs, dioecious, mostly many-stemmed and often much and intricately branched,

with thick woody roots. Leaves decussate or (rarely) in whorls of 3 but often seemingly in much

larger numbers at nodes*, small and often ± ericoid, ± acute, acuminate to mucronate at apex,

(sub)sessile, with small, ± cup-shaped stipular sheaths with or without l(-3) minute setae on

either side. Inflorescence frequently leafy and inconspicuous, made up of subsessile, mostly 3-1-

flowered cymes, arranged in pairs or single at nodes or flowers solitary and terminal on shoots.

Flowers subsessile, subtended by a pair of leafy bracts, 4-5(-6)-merous. Calyx : lobes small or

subobsolete. cf: Corolla : tube short, ± cylindrical to broadly funnel-shaped, lobes recurved, ±

lanceolate; anthers yellowish to whitish, exserted, dangling on long slender filiform filaments;

minute rudimentary ovary usually present. corolla much smaller; tube cylindrical, lobes linear

to ± lanceolate, erect to ± spreading. Ovary biovulate; style 0; stigmas 2, long exserted, hairy,

often purplish red. Fruit crowned by persistent calyx lobes, sometimes supported by a small ± U-

shaped carpophore, dehiscent, occasionally ± inflated, or hard, indehiscent, without carpophore.

Chromosome numbers : 2n=22, 44.

A genus of 11 species centred in the south-western and western Cape. One species extends into South West Africa/

Namibia, another is widely distributed from the central Cape to the Orange Free State and western Lesotho. Map 44.

Nenax is very closely allied to Anthospermum and, for the unexperienced, the distinction of the two genera may be

rather troublesome. Most characters, if compared one by one, overlap to some extent, but there is a conspicuous trend

towards more derived character states in Nenax (e.g. fruits dehiscent— >indehiscent; di — >tetraploidy; dioecy in the entire

genus; trend to very reduced, few-flowered inflorescences). Nenax, without exception, consists of distinctly woody, long-

lived dwarf shrubs, whereas Anthospermum is more variable in growth form (tall shrubs— >perennial herbs). The combina-

tion of the characters (1) distinctly woody, dwarf shrubby habit, (2) (±) needle-like leaves, (3) reduced, few-flowered

inflorescence and (4) dioecy is unique to Nenax.

Most taxa of Nenax are difficult to distinguish and some experience, including field knowledge, is often required to

ensure a definite identification, cf specimens of different taxa can be quite similar to each other and do, in general, not

provide easy-to-use key characters (o of some species should not be confused with Anthospermum'. ). The following key is,

therefore, based on (9) fruiting material; only occasionally are cf flowers referred to. If no mature fruits are present on a 9

individual, older stems should be searched for carpophores (cf. Fig. 8g): the presence of carpophores indicates the presence

of dehiscent (not inflated) fruits (species 3-6); the absence of carpophores indicates the presence of (a) inflated, dehiscent

fruits (species 1-2) or of (b) indehiscent fruits (species 7-11). It is recommended that always both cf and 9 specimens of a

given taxon are collected.

Nenax is presumably a (relatively) ‘young’ genus and probably in the process of actively and rapidly producing new

species. This may explain the rather frequent occurrence of ‘Forms’ (in the sense of geographically isolated and morpholo-

gically anomalous populations or distinct geographical or ecological races) which in some respects do not fully match

typical material of a given taxon. They may be extremely difficult to place; some are discussed with the respective taxa.

la Fruit dehiscent:

2a Fruit inflated, ± round in outline, not supported by an obvious carpophore:

3a Fruit c. 1,7-3, 2 mm in diam., subglobose to ± ellipsoidal; leaves glabrous or margins ± papillate

1 . N. microphylla

3b Fruit large, c. (5— )6— 8 mm in diam., laterally compressed, often ± round or broadly obovate in outline;

leaves silvery grey to greyish green due to dense cover of very short, often curled whitish hairs or

papillae 2. N. cinerea

2b Fruit not inflated, mericarps much longer than wide, supported by a U-shaped carpophore:

4a Mericarps covered with ± curled whitish hairs, c. 0,2-0, 5 mm long; leaves needle-like, small, 3-5 X 0,5-0, 8

(-1) mm; in drier parts of Worcester and Ceres distr 6. N. elsieae

4b Mericarps glabrous or papillate; leaves not distinctly needle-like or, if (±) needle-like, to 9(-l 1) mm long:

Due to the presence of leafy much-contracted short shoots; this situation is referred to as ‘pseudoverticillate’ in the key

and the descriptions.

1.2:38

Anthospermeae

5a Leaves not distinctly needle-like, 2,5^L5(-6) x 0, 9-1,4 mm; mericarps small, c. 2, 1-2,6 mm long;

corollas 5-merous, tube (cf) relatively long and ± cylindrical; mainly in Namaqualand 3. N. namaquensis

5b Leaves needle-like, (3— )3 ,5— 9(— 1 1) x 0,4—1 mm; mericarps (2,4—) 2,7-5 mm long:

6a Mericarps large, 3-5 x 1 ,9— 2,5(— 3) mm, crowned by relatively large calyx lobes c. 0,6-1, 2 mm long; cf

corolla 5-merous; dwarf shrubs with often pseudodichotomous and ± irregular branching; narrowly

confined to southern Van Rhynsdorp distr. and adjacent Clanwilliam distr 4 . N. coronata

6b Mericarps smaller, c. (2,4 — )2,7— 3,4 x 1,2-1 ,7 mm, crowned by small calyx lobes c. 0,3-0, 4 mm long;

Cf corolla 4-merous; dwarf shrubs with mostly ascending regular opposite branches; ± widely distri-

buted from Calvinia distr. south to Sir Lowry's Pass 5. N. divaricata

lb Fruit indehiscent:

7a Fruit ± soft, easily squashed between two fingers 10. Netiax sp. A

7b Fruit too hard to be squashed between two fingers:

8a Fruit small, c. 2-2.8 mm long:

9a Fruit reddish, shiny, glabrous, obscurely ribbed; leaves decussate, needle-like 1 1. Nenax sp. B

9b Fruit greyish, densely covered with whitish spreading hairs c. 0, 1-0,2 mm long; leaves in whorls of 3

7 (a). N. hirta subsp. hirta

8b Fruit larger, 3-8 mm long:

10a Fruit 5-8 mm long, entirely glabrous; partial inflorescences 1-flowered, flowers widely spaced, paired

or single at nodes; West Coast Strandveld 9. N. arenicola

10b Fruit 3-5 mm long, shortly hairy or ± glabrous:

11a Leaves to 3,5 mm long, decussate or occasionally in whorls of 3, widely spaced; fruits c. 3-3,7 mm

long, shortly hairy; intricately branched, robust dwarf shrubs; only on the West Coast (Langebaan

Peninsula and immediately south) over limestone 7 (b). N. hirta subsp. calciphila

lib Leaves to 1 2(— 15) mm long, strictly decussate, often pseudoverticillate; fruits 3-5,5 mm long, often

± crowded on shoots:

12a Stems reddish (at least when young), densely leafy, leaves usually pseudoverticillate, needle-like;

plants often only distinctly woody below; fruiting inflorescence ± conspicuous and spike-like,

flowers in clusters of 6(— 2) at nodes; fruits reddish, glabrous or ± papillate

8 (a). N. acerosa subsp. acerosa

12b Stems grey(ish), mostly with dimorphic leaves (long, needle-like and smaller, broader); distinctly

woody dwarf shrubs; inflorescence less conspicuous, flowers mostly paired at nodes; fruits

greyish (brown), mostly ± densely covered with short whitish spreading hairs

8 (b). N. acerosa subsp. macrocarpa

1 . Nenax microphylla (Sond.) Salter in J1

S. Afr. Bot. 3: 113 (1937); Hobson & Jessop,

Veld PI. S. Afr. 220 & pi. 21 (1975). Types:

Orange Free State, Sandrivier, Burke 506

(BM!; K!; PRE!; SAM!) Zevher 769 or s.n.

(SAM, lecto.!; BM!; G!: K!; PRE!).

Ambraria microphylla Sond. in F.C. 3: 34 (1865).

Dwarf shrub, rounded to ± cylindrical,

intricately branched, c. 0, 1— 0,3(— 0,4) m tall

and c. 0, 1-0,7 m in diam.; lower, cushion- to

± mat-forming if browsed. Stems ascending or

erect, often ± spine-tipped if browsed; some-

times arching downward or ± prostrate, occa-

sionally rooting at nodes and producing new

plantlets. Leaves decussate, often pseudo-verti-

cillate; blades (1 ,5-)2^4(-5,4) x 0,7-1 .2 mm,

(broadly) ovate to ± elliptic (often longer, to 7

mm, and ± linear-lanceolate or lanceolate on

new growth), frequently recurved, glabrous or

sometimes margins ± papillate. Flowers in

clusters of 2 (seldom 4—6) at nodes, inflores-

cences often ± conspicuous in fruiting co-

rolla 4-merous, pale yellow to greenish yellow,

sometimes reddish (purplish) tinged outside.

Cf : tube 0,5-0, 7 mm long, broadly funnel-

shaped, lobes 1,5-2, 2 x 0,6-0, 8 mm; anthers

1.2- 1 ,9 mm long. 9: tube 0.2-0, 4 mm long,

lobes 0,2-0, 5 x 0, 1-0.2 mm, corolla some-

times absent altogether; stigmas 2,8— 4,4 mm

long, whitish grey (also purplish red?); ovary

1.3- 1 ,7 x 0,6-0. 9 mm, with 4 minute calyx

lobes. Fruit inflated, (sub)globose or ± ellip-

soidal, dehiscent, not supported by a distinct

carpophore, c. 1,7-3, 2 mm in diam. (bright)

reddish brown or reddish, glabrescent or papil-

late, with 4 ± triangular calyx lobes c. 0,3-0, 4

x 0.2-0, 3 mm. Chromosome number : 2n = 22.

Fig. 6; 1; 8a & b.

Anthospermeae

1,2:39

rhJw4

FIG. 6. — 1, Nenax microphylla, part of fruiting browsed plant, x 0,5 (Saunders 10). 2 & 3, N. cinerea: 2, branch

with cf buds and flowers, X 0,5; 3, leaf pair x 2,5 (Acocks 16947). 4 & 5, N. divaricata: 4, part of fruiting plant, XDA

5, fruit in side view, x 5 (Acocks 17445). 6 & 7, N. acerosa subsp. acerosa: 6, fruiting plant, x 0,5; 7, fruit, x 5 (bmit i

5146).

1,2:40

Anthospermeae

Known from the central, northern and eastern Cape.

Orange Free State and lower-lying parts of Lesotho. Grow-

ing mostly in Karoo or karroid vegetation with few or no

trees or tall shrubs; often in rocky or gravelly grassy areas,

on stony hills or in crevices of rock sheets. Map 33.

MAPS 33-39: — 33. Nenax microphylla

34. Nenax coronata

35. Nenax elsieae

36. Nenax acerosa subsp. acerosa

37. Nenax acerosa subsp. macrocarpa

38. Nenax hirta subsp. hirta

39. Nenax hirta subsp. calciphila

Vouchers: Forward A. M. 1590; Leistner 962; Smith

4129.

An excellent fodder bush primarily for sheep; accord-

ing to Hobson & Jessop (op. cit.) regrowth is excellent

when rains are reasonable and its drought resistance is

outstanding Browsed plants remain low and become very

irregularly branched and deformed.

Newly produced cf flowering shoots of V. micro-

phylla with relatively longer and narrower leaves should not

be confused with Anthospermum pumihtm which may grow

in the same area. The two taxa apparently can form viable

hybrids but these seem to be very rare.

2. Nenax cinerea (Thunb.) Puff, comb,

nov. Type: Cape, Thunberg (sheet 23686, UPS,

holo.!; S, iso.!).

Cliffortia cinerea Thunb., Prodr. 2: 93 (1800).

N. dregei L. Bol. in Ann. S. Afr. Mus. 9: 215 & pi. VI, A

(1917); Launert & Roessler in F.S.W.A. 115: 19 (1966).

Types: Cape, Bot River bed, between Calvinia and Holle

River, Pearson 3966 (BOL, lecto.!; K!; NBG!); c. 24 km N

of Alewyn’s Fontein ( = Aalwynsfontein); Pearson 3930

(BOL!); between Klipplaat and Bitterfontein, Pearson 3295

(BOL!; STE!); between Anenous and Chubiessis Outspan,

Pearson 5979 (BOL!; K!).

N. hantamensis Schltr. , nom. nud.

Rigid dwarf shrub or shrub, intricately

branched, c. 0,5-1 m tall; much lower, c. 0,1-

0,3(-0,4) m, occasionally cushion-forming if

browsed. Stems erect, ascending or sometimes

± prostrate, much branched; branches often

single at nodes or, if paired, branches unequal,

often spine-tipped if browsed. Leaves decus-

sate; blades (2— )3— 7(— 10) x (1,2-) 1,5-2 mm,

ovate- or linear-lanceolate, shallowly concave

above, convex below, appearing greyish, sil-

very grey or greenish grey due to dense cover of

very short, often curled whitish hairs or papil-

lae. Flowers single or (less commonly) paired

at nodes, rarely in clusters of 4-6; corolla 5-

merous, yellowish to greenish yellow, some-

times reddish tinged outside, usually densely

papillate, cf : tube 0,8—1 ,3(— 1 .7) mm long,

broadly funnel-shaped, lobes 2, 5-3, 2 (-3,5) x

(0,6-)0,8-l mm; anthers ( 1 ,5—) 1 ,8—2,5 mm

long. 9: tube (0,l-)0,3-0,6 mm long, lobes

(0,l-)0,3-0,7(-0,9) x 0,l-0,2(-0,4) mm, co-

rolla sometimes absent altogether; stigmas

( 1 ,9— )2,5 — 7,5 mm long, dark purple, seldom

greyish; ovary 1,4-2 x 0,8—1 ,2(— 1 ,7) mm,

with 5 small calyx lobes. Fruit inflated, late-

rally compressed, ± round, broadly obovate or

± heart-shaped in outline, dehiscent, not sup-

ported by a distinct carpophore, c. (5— )6— 8 mm

in diam., light reddish brown, papillate to gla-

brescent, with 5 rounded to ± triangular calyx

lobes c. 0, 2-0,4 x 0,3 mm. Chromosome

number. 2n = 22. Fig. 6: 2 & 3.

Occurs in the interior of the western Cape (Laingsburg

distr. to Namaqualand) and in southern SWA/Namibia (as

far north as Aus). Growing mostly along ephemeral water-

courses. in dried up river beds, in crevices or at the edge of

rock sheets. Map 40.

Vouchers: Acocks 16947; Dinter 4132; Van der West-

huizen 328. 329.

Vegetatively distinguished from all other Nenax

species by its greyish, densely papillate or very shortly

hairy leaves.

Anthospermeae

1,2:41

3. Nenax namaquensis Puff, sp. nov.,

laminis foliorum glabris et fructibus minoribus

haud inflatisque a N. cinerea praeclare dis-

tinguitur.

Type: Cape, Namaqualand, a little north of

Middelkraal, Pearson 5615 (BOL, holo.!; K,

iso.!).

Dwarf shrub, intricately branched, ±

rounded to spreading, c. 0,15-0,6 m tall; often

± cushion-forming if browsed. Stems often

much branched; branching irregular, branches

mostly single at nodes, spine-tipped if browsed.

Leaves decussate; blades ascending to ±

spreading, often slightly curved, 2,5-4,5(-6) x

0,9-1 ,4 mm, narrowly ovate-lanceolate to

linear-lanceolate, shallowly concave above and

convex below to ± triangular in section, gla-

brous except for papillate margins, fresh green.

Flowers single (rarely paired) at nodes, some-

times also terminal and overtopped by lateral

shoots arising below; corolla 5-merous (in cf

very rarely, in $ occasionally also 4-merous),

yellowish, reddish tinged outside, cf: tube

1-2,1 mm long, funnel-shaped or occasionally

± cylindrical, lobes 2,4-3 ,4 x 0,7-1 mm;

anthers 1,5-2, 2 mm long; rudimentary ovary

often with conspicuous calyx lobes. 9: tube

0,2-0,4(-0,7) mm long, lobes 0,4-0, 7(-l ,2) x

0, 1— 0,2(— 0,3) mm; stigmas 1, 4-2,4 mm long;

ovary 1-1,2 x 0,8-1 mm, with 5 small calyx

lobes. Fruit dehiscent, supported by a some-

times large U-shaped carpophore or (seldom)

carpophore ± absent, dark reddish brown;

mericarps 2, 1-2,6 x 1-1,4 mm, ± obovate,

glabrous or sparsely to densely papillate, with 3

or 2 ± triangular calyx lobes c. 0,4-0, 7 X

0, 4-0,7 mm. Chromosome number: 2n=22.

Endemic to the western Cape; only known from the

Namaqualand distr. and the northern Clanwiiliam distr.

Growing in sandy to gravelly soil amongst rocks or in

crevices. Map 41 .

Vouchers: Hutchinson 836 (BOL; GRA; K; PRE): Van

der Merwe 190 (PRE; STE).

Vegetatively rather similar to N. cinerea (above) but

lacking the characteristic leaf indumentum of that species.

Its dehiscent fruits, moreover, are small and not inflated.

Perhaps most closely allied to A. coronata (below).

The cf collection HafstrAm & Acocks 1441 (PRE; S)

from near Sutherland (3220-BC) appears to be allied to N.

namaquensis but differs in having ± glabrous, flat leaves

(± oblong in section) and short, broadly funnel-shaped

corolla tube; The absence of fruiting ( $ ) material makes it

impossible to place it with certainty; it may be a new spe-

cies.

4. Nenax coronata Puff, sp. nov., meri-

carpiis majoribus, lobis calycis longioribus,

corollis 5-meris et habitu dissimili a N. divari-

cata facile distinguitur.

Type: Cape, west side of Pakhuis Pass, a

little west of Leipoldt grave, Puff 800902-6/4

(WU, holo. !; BOL; NBG; PRE: STE, iso. !).

Dwarf shrub, ± erect, often with thick,

woody root. Stems c. 0,25-0,4 m long, much-

branched, branches usually single at nodes,

branching often pseudodichotomous* and ±

irregular. Leaves decussate; blades (3— )5— 9

(-11) x 0,7-1 mm, linear, needle-like and ±

rigid, semiterete to ± triangular in section, gla-

brous or margins ± papillate, upper surface and

midrib sometimes (dark) brownish red. Flowers

mostly single at nodes, sometimes also terminal

and overtopped by short lateral shoots arising

below; corolla 5(rarely 4)-merous, yellowish

(?). cf: tube c. 0,8 mm long, funnel-shaped,

lobes 2, 5-3, 4 (-4) x 0,6-1 mm; anthers

1,4-2, 4 mm long; minute rudimentary ovary

sometimes with well discernible calyx lobes.

9: tube 0, 1-0,2 mm long, lobes 0,3-0, 8 x

0, 1-0,2 mm; stigmas ( 1 ,5— )2— 4 mm long;

ovary 1 ,2-2 x 0,7-1 mm, with 5 conspicuous

calyx lobes. Fruit dehiscent, supported by a

small carpophore or carpophore sometimes ±

absent, reddish brown; mericarps 3-5 x

1 ,9— 2,5(— 3) mm, oblong to obovate, glabrous

or papillate, with 3 or 2 triangular calyx lobes c.

0,6—1 ,2 x 0,3-0, 5 mm. Chromosome number:

2n=22.

Endemic to the western Cape; confined to the southern

Vanrhynsdorp distr. (Giftberg) and adjacent Clanwilliam

distr. Growing in sandy areas between rocks or in cracks of

rocks in dry Mountain Fynbos. Map 34.

Vouchers: Esterhuysen 22044 (BOL); Schlechter

8652, 10805 (BOL; GRA; PRE).

Allied to the remaining Nenax species with non-in-

flated, dehiscent fruits (species 3, 5 and 6) but perhaps

closest to N. divaricata, which differs primarily in having

4-merous cf flowers and smaller fruits with rather minute

calyx lobes. In the field, N. coronata and N. divaricata can

be easily distinguished by their growth form. In N. coronata

branches usually arise singly at nodes, branching may be-

come seemingly dichotomous and, subsequently, shoots

often show a distinct zig-zag pattern. In N. divaricata

branching is more regular, the divaricate branches mostly

arise in pairs at nodes.

* Vigorous lateral branches push main axes to the side; a Y-

like appearance of the branchings results.

1,2:42

Anthospermeae

FIG. 7. — Nenax divaricata: I, cf flower (ro: rudimentary ovary), X 20; 2, flower, X 20 {Puff 790712-2/1) (SEM-

graphs).

Anthospermeae

1,2:43

5. Nenax divaricata Salter in J1 S. Afr.

Bot. 3: 113 (1937). Type: Cape, Worcester

distr. , near Tulbaghskloof, Ecklon & Zeyher

2319 (or ‘1.9’) (SAM, holo. !; E!; FI!; GOET!;

LY!; M!; MO!; NBG!; PRE!; S!; US!; W!;

WU!).

Ambraria acerosa Sond. in F.C. 3: 34 (1865), pro parte

(C ?)'.

N. acerosa sensu Eckl. & Zeyh., Enum. 368 (1836), non

Gaertn.

Dwarf shrub, usually divaricately branch-

ed, rounded to ± cylindrical, c. (0,15-)

0,3-0,6(-l) m tall. Stems erect to ascending,

much branched; principal branches usually op-

posite, held at c. 45°. Leaves decussate; blades

(3-)3,5-8(-9,5) x 0,4— 0,5 mm, linear, needle-

like, semiterete to ± triangular in section, gla-

brous except for ± papillate margins. Flowers

mostly single at nodes, sometimes also terminal

and overtopped by short lateral shoots arising

below; corolla 4-merous, yellowish green, of-

ten purplish or purplish brown tinged or

streaked outside, cf: tube (0,8— )1— 1 ,5 mm

long, funnel-shaped, lobes 2, 7-3, 2 x 0,7-1

mm; anthers 1 ,4—1 ,7 mm long. $: tube 0,2-0, 3

mm long, lobes 0, 3-0,5 x 0, 1-0,2 mm; stig-

mas 3, 4-4, 6 mm long, purplish red to dark

purple; ovary 0,7-1, 2 x 0,4— 0,8 mm, with 4

small calyx lobes. Fruit tardily dehiscent, sup-

ported by a small carpophore, reddish brown,

greyish to dark grey; mericarps (2,4—)2,7-3,4

x 1,2-1 ,7 mm, ± obovate, glabrescent to

densely papillate, with 2 rounded to ± triangu-

lar calyx lobes c. 0,3-0, 4 mm long. Chromo-

some number: 2n=22. Fig. 6: 4 & 5; 7; 8g.

Endemic to the (south-)westem Cape; occurring from

Sir Lowry’s Pass north to the southern Vanrhynsdorp and

Calvinia distr. Growing in sandy to gravelly soil, primarily

in dry Mountain Fynbos. Map 42.

Vouchers: Acocks 17445, 17458 (K; PRE); Esterhuy-

sen 8 145 (BOL); Thompson 2090 (K; PRE; STE).

N. divaricata should not be confused with dioecious

Anthospermum species (A. spathulatum in particular,

‘Forms’ of which may resemble N. divaricata in having

shoots with similarly regular, opposite branches). Vegeta-

tively N. divaricata is distinguished from any of these by its

distinctly needle-like leaves (semiterete to ± triangular in

section).

An investigation of the specimens studied by Sonder (now in her-

barium S) revealed that the description of Ambraria acerosa Sond.

is based on cf N. divaricata (bei Brakfontein, Ecklon s.n., S!) and

9 N. acerosa Gaertn. (Tulbaghskloof, Ecklon & Zeyher '11-9' ,

SI). Salter (op cit.) had apparently only seen Ecklon & Zeyher

23 19 or ' 1 .9' , Cf collections quoted by Sonder, when pointing out

that Ambraria acerosa Sond. is a very different species from N.

acerosa Gaertn.

MAPS 40-44; — 40. Nenax cinerea

41 . Nenax namaquensis

42. Nenax divaricata

43. Nenax arenicola

44. Nenax, all taxa

Some collections from the southernmost locality. Sir

Lowry’s Pass (e.g. Acocks s.n., S !), tend to be much more

densely leafy and more branched than ‘typical’ N. divari-

cata from further north. More field observations and addi-

tional herbarium material would be needed to determine

whether these forms deserve taxonomic recognition.

6. Nenax elsieae Puff, sp. nov . , fructibus

lanatis et lobis calycis relative magnis a specie-

bus Nenacis fructibus dehiscentibus hand infla-

tisque (N. namaquense, N. coronata, N. divari-

cata) praeclare distinguitur .

Type: Cape, Worcester distr., Bonteberg,

Eikenbosch Hoek, 1070-1220 m, Esterhuvsen

3656 (BOL, holo.!).

Low dwarf shrub, rounded to ± cylindri-

cal, with thick woody root. Stems c. 80-200

(-300) mm long, ascending to erect, much

branched; branches usually single at nodes, ±

erect. Leaves decussate, sometimes ± pseudo-

1,2:44

Anthospermeae

verticillate; blades 3-5 x 0,5— 0,8(— 1 ) mm, lin-

ear(-lanceolate), needle-like, semiterete to ±

triangular in section, margins often papillate,

midrib sometimes reddish brown below. Flow-

ers usually single at nodes; corolla 4-merous,

yellowish green, cf: tube c. 0,8-1 mm long,

(broadly) funnel-shaped, lobes ( 1 ,7— )2— 3 x

0,8-1 mm; anthers 1,7-2 mm long; rudimen-

tary ovary usually crowned by 4 conspicuous

calyx lobes. 9: tube 0,2-0, 5 mm long, lobes

0,3-0, 8 x 0,2-0, 3 mm; stigmas 3,4-4, 4 mm

long, (dark) purplish red; ovary 1-1,4 x

0,7-0, 8 mm, hairy, with 4 conspicuous calyx

lobes. Fruit dehiscent, supported by a minute

carpophore, reddish brown; mericarps 2, 7-3, 5

x 1 ,8-2,2 mm, obovate, densely covered with

(±) curled whitish hairs c. 0,243,5 mm long,

with 2 glabrous, broad, rounded to ± trapezi-

form calyx lobes c. 0,5-0, 7 x 0,543,9 mm.

Fig. 8 e & f.

Endemic to the interior of the south-western Cape;

only known from the Worcester and Ceres distr. Growing

on rocky slopes amongst rocks in very dry Fynbos. Map 35.

Vouchers; Pi# 7909 13-6/1 (WU); Taylor 5874 (PRE;

STE).

This apparently quite rare species is well distinguished

by its dehiscent fruits with longish ± curled hairs and rela-

tively large calyx lobes, cf N. elsieae should not be con-

fused with Anthospermum comptonii ; the leaves of N. el-

sieae are ± linear and needle-like (semiterete to ± triangu-

lar in section), those of A. comptonii are much broader

(although the margins are often revolute so that the blades

may appear ± terete).

7. Nenax hirta (Cruse) Salter in J1 S. Afr.

Bot. 3: 113 (1937), in Adamson & Salter, FI.

Cape Penins. 735 (1950). Types: Cape,

Mund(t) & Maire s.n. (B, syn. t), at base of

Lion’s Mt. towards Drieanckerbay, C. W. Ber-

gius s.n. (B, syn. t); slopes of Signal Hill

above Three Anchor Bay, Salter 6407 (BOL,

neo. & topo.!).

Ambraria hirta Cruse, Rub. Cap. 17 (1825), in Linnaea

6: 19 (1831); Sond. in F.C. 3: 34 ( 1865).

Two subspecies are recognized:

7 (a), subsp. hirta.

Dwarf shrub, rounded to ± cushion-form-

ing, diffusely branched, to c. 0,3 m tall and

0,3-0, 5 m in diam. Stems ascending, prostrate

or ± erect, much branched; branches usually

single at nodes, shortly hairy. Leaves in whorls

of 3, pseudoverticillate; blades ( 1 ,2—) 1 ,5—3 ,5

x 0,443,9 mm, linear(-lanceolate) (often

longer, to 6,4 mm, on new growth), margins

and sometimes also midrib below with whitish

± straight hairs c. 0,143,2(43,4) mm long.

Flowers usually in clusters of 3 at nodes, inflo-

rescence sometimes ± conspicuous in fruiting

9; corolla 4-merous, yellowish (green), some-

times a little hairy outside, cf: tube 0,7-1, 2

mm long, broadly funnel-shaped, lobes (1,5-)

1, 7-2,1 X 0, 6-0,9 mm; anthers 1-1,7 mm

long. 9: lube 0,343,4 mm long, lobes (0,3-)

0,5-1 x 0, 1-0,3 mm; stigmas (2,2-)2,5-4,8

mm long, purplish red; ovary 1-1,2 x 0,6-1

mm, hairy, with 4 conspicuous calyx lobes.

Fruit indehiscent, hard, not supported by a car-

pophore, 2-2,8 x (1 ,5-) 1 ,7-2,2 mm, obovoi-

dal to spheroidal, greyish, covered with whitish

spreading hairs c. 0,143,2 mm long, with 4

glabrous rounded calyx lobes c. 0,343,4 x

0,4—0,5 mm. Chromosome number: 2n—22.

Fig. 8d.

Endemic to the south-western Cape; occurring from

the north end of the Cape Peninsula east and north-east to

the Worcester and the Piketberg distr. Growing mostly in

gravelly ground or in red, clayey soils in Coastal Renoster-

veld or in white sand in Coastal Fynbos. Map 38.

Vouchers: Salter (BM; BOL; K); Schlechter 10724

(BM; BOL; BR; E; G; K; MO; PRE; S; US; W); Thompson

2560 (PRE; STE).

Easily recognized by its small indehiscent hairy fruits,

its small (cf) flowers and its small temately arranged

leaves.

7(b). subsp. calciphila Puff, subsp. nov.,

fructibus majoribus, foliis minus congestis et

habitu robustiore a subsp. typica differt.

Type: Cape, Langebaan Peninsula, Oude

Post Private Nature Reserve, Boucher 2964

(STE,holo.!;PRE, iso.!).

Much more robust dwarf shrub than subsp.

hirta, c. 0,3-1 m tall, intricately branched;

branches often spine-tipped (browsed). Leaves

much less crowded than in subsp. hirta, ±

widely spaced, mostly decussate (but some

leaves occasionally in whorls of 3), less dis-

tinctly pseudoverticillate (few leafy much con-

tracted short shoots); blades to 3,5 mm long.

Flowers unknown. Fruit with somewhat shorter

hairs, larger, c. 3-3,7 x 2,7 mm, not as

crowded and numerous as in subsp. hirta.

Endemic to the western Cape; only known from the

Langebaan Peninsula south to Yzerfontein. Apparently con-

fined to exposed limestone ridges and sandy limestone soils;

characteristic for Nenax-Ma\tenus-Z\goph\Uum Limestone

Evergreen Shrubland. Map 39.

Vouchers: Acocks 14514 : Puff 791226-2/1 (WU).

Anthospermeae

1 ,2: 45

FIG. 8. — a & b, Nenax microphylla: a, fruit in side view showing the two mericarps, x 17,5; b, mericarp, ventral

side, x 17,5 (Puff 790413-1/1). c, N. arenicola, indehiscent fruit from above, x 17,5 (Puff 790714-2/1). d, N. hirta

subsp. hirta, indehiscent fruits with persistent calyx lobes, x 17,5 (Bolus 12764). e & f, N. elsieae; e, fruit (two mericarps)

in side view, x 17,5; f, upper part of mericarp with persistent calyx lobes, X 17,5 (Eslerhuxsen 3656). g, N. divaricata,

carpophore, x 17,5 (Puff 790713-1/1) (SEM-graphs).

1,2:46

Anthospermeae

Langebaan specimens originate from areas with heav-

ily trampled and grazed vegetation, but trampling or grazing

alone is not likely to be responsible for the characteristic

growth form of subsp. calciphila (obviously browsed speci-

mens of subsp. hirta differ markedly in habit).

Acocks 14523 from c. 12 km NE of Yzerfontein is ±

intermediate between the two subspecies.

8. Nenax acerosa Gaertn., Fruct. 1: 165,

t. 32, f. 7 (1788); Salter in J1S. Afr. Bot. 3: 112

(1937), in Adamson & Salter, FI. Cape Penins.

734 (1950). Type: Cape, Masson s.n. in herb.

Banks (BM, holo.!).

Cliffortia acerosa MS.

Ambraria glabra Cruse, Rub. Cap. 17 (1825), inLinnaea

6: 18 (1831); Sond. in F.C. 3: 33 (1865). Nenax glabra

(Cruse) Kuntze, Rev. Gen. 3: 121 (1898). Type: Cape,

Bergius s.n. (B.holo. t).

Ambraria glabra Cruse var. papillata Sond. in F.C. 3: 34

(1865). Type: none cited, but almost certainly ‘Capflache’,

1841, Ecklon s.n. (S, holo.!) [label bearing Sonder’s

handwriting].

Ambraria glabra Cruse var. tulbaghica Sond. in F.C. 3:

34 (1865). Type: Cape, waterfall, Tulbagh, no collector

cited but without doubt Pappe s.n. (S, holo.!; K, iso.!)

[label bearing Sonder’s handwriting].

Ambraria acerosa Sond. in F.C. 3: 34 (1865), pro parte

(9; see also A. divaricata, footnote).

Two subspecies are recognized:

8 (a), subsp. acerosa.

Synonyms as above.

Dwarf shrub, erect to ± diffuse or, if burnt

regularly, subshrubby and often with massive,

woody base and numerous, thin aerial stems.

Stems c. (0,l-)0,15-0,3(-0,4) m long, un-

branched to ± much branched, branches as-

cending to erect, glabrous to ± densely papil-

late, reddish (brown) and shiny above. Leaves

decussate, pseudoverticillate; blades (5— )8— 1 2

(-15) x 0,5-1 mm, linear, ± needle-like and ±

triangular in section or (on long shoots) some-

times linear-lanceolate and shallowly concave

above, glabrous or margins with whitish hairs

c. 0, 1—0,2 (-0,3) mm long, upper surface and

midrib often dark brown. Flowers in clusters of

6-2 at nodes, inflorescence spike-like, ± con-

spicuous, especially in fruiting 9i corolla 5-

merous (in cf very rarely, in 9 occasionally

also 4-merous), greenish yellow to creamy yel-

low, often reddisn pumle tinged, occasionally a

little hairy near tip. <j: tube 0,7-1, 5 mm long,

broadly funnel-shaped, lobes 2,4— 3,7 x 0,8-1

mm; anthers 1,4-2 mm long. 9: tube 0,3-0, 7

mm long, lobes 0,5-1 x 0,2-0, 3 mm; stigmas

c. 3,4-6,3(-7) mm long, purplish red or, less

commonly, greyish or greyish green; ovary c.

1,4-2 x 0,9-1, 5 mm, sometimes obscurely

ribbed below 5 small calyx lobes. Fruit indehis-

cent, hard, not supported by a carpophore, c.

3, 2-5, 5 x 2-3,4 mm, obovoidal to ellipsoidal,

reddish brown, glabrous or ± papillate, with 5

trapeziform, ± obovate to triangular calyx

lobes c. 0, 5-0,9 x 0,7-0, 8 mm. Chromosome

number. 2n=44. Fig. 6: 6 & 7.

Endemic to the south-western Cape; occurring from

the Cape Peninsula north-east to the Worcester and Ceres

distr. , and east to the Bredasdorp distr. Growing in sandy to

gravelly flats, sometimes in sand over clay, mostly in ± dry

habitats. Map 36.

Vouchers: Esterhuysen 26583 (BOL; WU); Schlechter

9747 (BM; BOL; BR; E; G; K; MO; PRE; S; US; W); Van

Breda 169 (BOL; PRE).

Two collections from around Faure (3418— BB; Ester-

huysen 11932, BOL; Compton 15998, NBG) and a collec-

tion from near Ratel River mouth (3419-DA?; Bolus 21875

ex parte, BOL) are very condensed, distinctly woody, much

branched low dwarf shrubs with unusually small (less than 3

x 2 mm) but seemingly mature shortly hairy fruits. They

may represent an odd state of subsp. acerosa (or a new

taxon?).

8(b). subsp. macrocarpa (Eckl. & Zeyh.)

Puff, comb, et stat. nov. Type: Cape, on the

Breederivier, Swellendam, Mund(t) s.n. sub

Ecklon & Zeyher 23180 (SAM, holo.!; S,

iso.!).

Ambraria hirta Cruse var. macrocarpa Eckl. & Zeyh.,

Enum. 368 (1836).

Dwarf shrub, often ± rounded or low, ±

cushion-forming, c. 0, 1-0,5 m tall and to c. 0,5

m in diam. , often with thick, woody root. Stems

c. 0,2-0, 5 m long, ascending to ± prostrate or

erect, mostly much branched, ± densely papil-

late, mostly greyish or greyish brown above.

Leaves often distinctly dimorphic; blades 2-6 x

0,4-1 mm, narrowly ovate-lanceolate and ob-

long to ± elliptic in section on old long shoots,

linear-lanceolate to linear, needle-like, relati-

vely longer and ± round in section on short

shoots and/or new growth, glabrous except for

± papillate margins. Flowers paired (seldom in

clusters of 4—6) at nodes, inflorescence some-

times ± conspicuous in 9; corolla 4-5-merous

(in 9 more commonly 4-merous), glabrous. 0”:

tube 0,7-1, 2 mm long, lobes 2,1-3 x 0,8-1

mm; anthers 1, 7-2,1 mm long. 9: tube ± 0 or

0,2-0, 3 mm long, lobes 0,5—1 x 0,2-0, 3 mm;

stigmas 3-5 mm long; ovary 1,2-1, 6 x 1-1,4

Anthospermeae

1,2:47

mm, shortly hairy, with 5 small calyx lobes.

Fruit c. 3-5 x 2, 8-3, 5 mm, greyish or greyish

brown, densely covered with short whitish

spreading hairs (if ± glabrescent, fruits ±

subglobose), with 5 triangular to ± rounded

calyx lobes c. 0,5-1 x 0,4— 0,7 mm. Chromo-

some number. 2n=(c.)44.

Endemic to the south-western Cape; from the Caledon

distr. east to the Riversdale distr. and north to the southern

Clanwilliam distr. Growing on hillsides, in sandy flats or in

Coastal Fynbos over sand; also in limestone areas. Map 37.

Vouchers: Acocks 23998; Compton 1 1898 (NBG); Puff

800920-2/2 (WU).

The relatively small but broad long shoot leaves are

similar in shape to those of N. namaquensis (no. 3) or N.

cinerea (no. 2). Specimens without mature fruits in which

none of the longer, ± needle-like short shoot leaves are

developed may be difficult to identify.

From the eastern Bredasdorp distr. to the Riversdale

distr. the separation of the two subspecies can become pro-

blematic (e.g. Van der Menve 1007 from Windhoek Plateau

is ± intermediate).

Collections from near The Fisheries (=Gouritsmond,

3421-BD; Acocks 21559; Puff 790910-4/1 , 800927-2/1,

WU) are difficult to place; their fruits do not fully match

either of the subspecies (± subglobose, relatively small and

less distinctly hairy than ‘typical’ subsp. macrocarpa ). The

habit of these collections, moreover, is highly variable;

low, intricately branched, ± cushion-shaped, with small

leaves in abandoned fields and on hard-packed, tram-

pled!?), sandy soil; rounded bushes with widely spaced

longer and narrower leaves in undisturbed Coastal Fynbos

on calcareous sand.

9. Nenax arenicola Puff, sp. nov., fructi-

bus majoribus solitariis vet paribus oppositis et

late dispersis, et habitu dissimili a N. acerosa

differt.

Type: Cape, c. 3 km south-east of Graaf-

water-Lambert s Bay rd., on road to Leipoldt-

ville, Puff 800915-2/1 (WU, holo.!; BOL;

NBG; PRE, iso.!).

Dwarf shrub, intricately branched, often

rounded, c. 0,3-0, 5 m in diam. and c.

0,2— 0,75(— 1 ) m tall, frequently with thick,

woody root. Stems ascending to erect, ± much

branched; branches typically paired at nodes,

spreading to ascending, branching very irregu-

lar if browsed. Leaves decussate, sometimes

pseudoverticillate; blades (5— )7— 1 3(— 1 6) x

0,5—1 mm, linear, often shallowly concave

above and convex below or ± semiterete in

section, glabrous or margins ± papillate. Flow-

ers paired or single at nodes, widely spaced;

corolla 4(-5)-merous, greenish yellow, some-

times purplish tinged, cf: tube 0,4-0, 8 mm

long, broadly funnel-shaped, lobes 2-2,9 x

0,6-0, 8 mm; anthers 1 ,6—1 ,9 mm long. 9: tube

± 0 or 0, 1-0,3 mm long, lobes 0,5— 1(— 1 ,3) x

0, 1-0,3 mm; stigmas 3-5,4 mm long, purplish

red or whitish grey tinged purplish; ovary c.

2, 2-2, 9 x 1-1,4 mm, with 4-5 small calyx

lobes. Fruit indehiscent, hard, not supported by

a carpophore, 5-8 x 2-3,5 mm, ellipsoidal,

reddish brown to dark grey, glabrous, with 4-5

often ± indistinct rounded calyx lobes to c.

0,4— 0,5 mm long. Chromosome number.

2n=44. Fig. 8c.

Endemic to the western Cape; occurring from the

south-western Namaqualand distr. south to the western

Clanwilliam distr. Growing in sandy coastal plains and oc-

casionally in heavier, stony soils of the adjacent foothills.

Map 43.

Vouchers: Acocks 14936 (K; PRE); 19635 (K; M;

PRE; SRGH); Van der Westhuizen 1 14.

Allied to N. acerosa (no. 8) but easily distinguished by

its less leafy appearance, a different habit, more reduced

inflorescences and a more north-western distribution range.

10. Nenax sp. A

Cape, Laingsburg distr., (farm) Cabidu

[north of Konstabel, 3320-AB], Compton

22209 (NBG).

Dwarf shrub with greyish stems, ± much

branched; branches mostly paired at nodes, ±

ascending. Leaves decussate, widely spaced;

blades of long shoot leaves ovate-lanceolate to

linear-lanceolate, c. 2-3 x 1 mm, those of

short shoot leaves narrower and to 6(-8) mm

long. Flowers unknown. Fruit indehiscent, ±

soft, easily squashed between two fingers, not

supported by a carpophore, c. 3, 5-4,5 mm in

diam., subglobose, greyish to greyish brown,

shortly hairy, crowned by minute to subobso-

lete calyx looes.

Allied to the other indehiscent-fruited Nenax species

(no. 7-9 and 11) and veeetatively similar to N. acerosa

subsp. macrocarpa (no. 8b). Probably a good new species

but so far only known from the collection cited above.

1 1 . Nenax sp. B

Cape, Zonder Einde Mts ( = Riviersonder-

end Mts), Boesmanskloof [3419-BA], Levyns

9200 (BOL).

Dwarf shrub with reddish stems, ± much

branched; branches mostly paired at nodes, ±

spreading. Leaves decussate, pseudoverticil-

late; blades c. 4—6 x 0,5 mm, ± needle-like,

glabrous, shiny. Flowers unknown. Fruit inde-

hiscent, hard, not supported by a carpophore, c.

2-2,5 mm in diam., subglobose, shiny red or

reddish brown, obscurely ribbed, crowned by

minute calyx lobes.

Probably allied to N. acerosa (no. 8). Most likely a

good new species but so far only known from the collection

cited above.

1,2:48

8435

GALOPINA

Anthospermeae

Galopina Thunb., Nov. Gen. PI. 1: 3 (1781); Cruse, Rub. Cap. 18 (1825); Sond. in F.C. 3: 26

(1865); Hook. f. in Benth. & Hook, f., Gen. PI. 2: 139 (1873); K. Schum. in Pflanzenfam. 4,4:

128 (1891); R.A. Dyer, Gen. 1: 622 (1975); Compton, FI. Swazild 586 (1976). Type species: G.

circaeoides Thunb.

Oxyspermum Eckl. & Zeyh., Enum. 365 (1836). Type species: O. asperum Eckl. & Zeyh.

Phyllis sensu Cruse in Linnaea 6: 19 (1831).

Perennial herbs with branched, often ± woody rhizomes or rootstocks, 9" + $, 9> or

occasionally Cf,£'+C?,d' + 9' + 9,Cf + 9- Leaves decussate, broadly ovate to lanceolate,

distinctly petiolate, with stipular sheaths bearing 3-5(-7) setae on either side. Inflorescence termi-

nal, paniculate to thyrso-paniculate, bracteate. Flowers <9 , 9 or (9 , 4(-5)-merous. Calyx obsolete.

<9, Cf: Corolla : tube (very) short, broadly funnel-shaped to campanulate, lobes recurved, ±

lanceolate; anthers yellowish to whitish, exserted, dangling on long slender filiform filaments. 9:

corolla much smaller, tube cylindrical, sometimes ± 0, lobes erect to spreading, ± linear. Ovary

biovulate; style 0; stigmas 2, long exserted, in <9 often shorter and thinner than in 9 , hairy, greyish

white, yellowish grey or greenish. Fruit dehiscent, not supported by a carpophore; mericarps ±

glabrous, tuberculate or covered with long hairs, dorsal side convex, ventral side plane to concave.

Chromosome number : 2n — 22.

A small south-east African genus. One of the four species is more widely distributed and extends to both the south-

western Cape and to Zimbabwe. Mozambique and southern Malawi.

la Ovary and fruit villous; stems typically without short shoots 4. G. crocyllioides

lb Ovary and fruit smooth, wrinkled or tuberculate, without hairs: stems often with leafy short shoots:

2a Inflorescence ± narrowly cylindrical; peduncles and pedicels not conspicuously divergent in fruit; fruiting

pedicels (0,8— )1— 2(— 3) mm long; short shoot leaves conspicuous, often almost as long as long shoot leaves;

petioles to 2(— 3) mm long 3. G. aspera

2b Inflorescence ellipsoidal to ± spheroidal; peduncles and pedicels divergent in fruit; fruiting pedicels(2-)3-26

(-37) mm long; short shoot leaves much smaller than long shoot leaves; petioles to 14 mm long:

3a Peduncles and pedicels (2— )3— 6(— 7) mm long in fruit, ± stiff; inflorescence pyramidal to ellipsoidal; leaves

typically ± densely tomentose, blades (27— )35— 50(— 62) x (15-)20-35mm, broadly ovate-lanceolate or

ovate 2.G. tomentosa

3b Peduncles and pedicels (7—) 1 2—26 (-37) mm long in fruit, filiform and slender; inflorescence lax, broadly

pyramidal to ± spheroidal; leaves glabrous or sparsely hairy, blades ( 28— )40— 80(— 1 05 )

x (8-)12-30(-37)mm, ± lanceolate to ovate-lanceolate 1. G. circaeoides

1. Galopina circaeoides Thunb., Nov.

Gen. PI. 1: 3 (1781); Cruse, Rub. Cap. 18

(1825); Sond. in F.C. 3: 26 (1865); Brenan in

Mem. N.Y. bot. Gdn 8: 454 (1954); Compton,

FI. Swazild 586 (1976). Type: Cape, in sylvis

Hautniquas, Groot Vaders-Bosch, aliisque.

Thunberg in herb. Thunberg (sheet 23313,

UPS, holo. !), in herb. Montin (S!) and in herb.

Gasstrdm (S!).

G. circaeoides Thunb. var. glabra Kuntze, Rev. Gen.

3,2: 120 (1898). Types: Cape. Swellendam, Kuntze s.n. (K,

lecto. ! ), Perie-Wald, Kuntze s.n. ( K ! ).

G. circaeoides Thunb. var. pubescens Kuntze, Rev.

Gen 3,2: 120 (1898). Type: Cape, Cathcart, Kuntze s.n.

(K, holo.!; US sub 554842!).

Anthospermum galopina Thunb., Prod. 1: 32 (1794).

Phyllis galopina (Thunb.) Cruse in Linnaea 6: 20 (1831).

Type: as for G. circaeoides.

Perennial herb, several- to many-stemmed, <9>

9" + 9, 9 or occasionally <9 + Cf or cf + 9"

+ 9- Stems (0,3-)0,4— 1 ,2(-l ,5) m long, as-

cending to erect or occasionally decumbent and

rooting at nodes, glabrous or occasionally spar-

sely hairy; often with much-contracted short

shoots bearing small leaves. Leaves : blades

(28-)40-80(-105) x (8-) 12-30(-37) mm, ±

lanceolate to ovate-lanceolate, narrowed to

base, acute at apex, glabrous or with some short

whitish spreading hairs along midrib and princi-

pal veins and/or scattered on both surfaces; pe-

tioles (3— )4 — 14 mm long; stipular sheath with 3

or 5 (-7) setae, free portion of the longest

(3,5-)4— 8(— 10,8) mm. Inflorescence broadly

pyramidal to ± spheroidal, lax, c.

( 1 90-) 1 50-300 x (80-) 130-280 mm; pedun-

cles and pedicels filiform, slender, glabrous,

strongly divergent in fruit, elongating to

Anthospermeae

1 ,2: 49

ft

FIG. 9. — Galopina circaeoides: 1, part of plant with fruiting inflorescence, x 0,6; 2, part of stipular sheath

with setae, x 1 1; 3, ybud, X 1 1 ; 4, ^ flower, x 1 1 ; 5, 9 flower, X 1 1 ; 6, fruit (two mericarps) in side view, x 1 1 (1 & 2:

Jacobsz 1598; 3-5: Puff 78 1203- 1/1; 6: Puff 780326-2/1).

1,2:50

Anthospermeae

(7— )12— 26(— 37) mm. Flowers : corolla whitish,

creamy white, yellowish or greenish, occasio-

nally reddish purplish tinged outside, glabrous

or papillate. C?: tube 0,3-0,5(-0,6) mm

long, lobes ( 1 , 1— )1 ,3—1 ,7(— 1 ,9) x (0,3-)

0,4-33,6 mm; anthers 1 , 1-1 ,6 mm long. $ : tube

±0-0,2(-0,3) mm long, lobes 0 ,6— 0,8(— 1 ) X

0,l-0,2(-0,3) mm; stigmas (2,3— )3— 5(— 6,5)

mm long; ovary c. 0,5-0,8(-l) x

(0,5-)0,8-l,3 mm, ± densely warty. Fruit

blackish to black; each mericarp c. 1,5-2 x

(0,7— )1— 1 ,4 mm, oblong to ± obovate, ±

densely warty. Fig. 9.

Occurs from Transvaal to the south-western Cape; also

in eastern Zimbabwe and adjacent parts of Mozambique and

in southern Malawi. Growing mostly in kloof or gallery

forests, in scrub along streams or in bushclumps. Map 45.

MAP 45. — Galopina circaeoides

Vouchers; Bayer & McClean 62 (BOL; GRA; PRE);

Galpin 8124; Scheepers 548 (M; MO; PRE; SRGH).

Variable in stem and leaf indumentum and leaf size

and shape, but easily distinguished from its nearest ally, G.

tomentosa (below), by its lax, broad and extensive inflores-

cence and its less dense and shorter indumentum.

2. Galopina tomentosa Hochst. in Flora

27: 555 (1844). Type: Natal, nr. Umlaas R.,

Krauss 52 (BM!; G!; K!; M!; MO!; W!).

G. hirsuta E. Mey. in Drfege in Flora 26, Bes. Beigabe

186 (1843), nom. nud.

Perennial herb, few- to many-stemmed,

9 + $, $ or occasionally cf, + C? or c? +

9 +9- Stems (0,3-)0,4-0,8(-l) m long, as-

cending to ± erect or ± prostrate and rooting at

nodes, mostly densely covered with longish

whitish or yellowish white hairs; often with

short shoots bearing small leaves. Leaves :

blades (27-)35-50(-62) x (15— )20-35 mm,

broadly ovate-lanceolate or ovate, cuneate,

truncate or ± subcordate at base, subacute to ±

obtuse at apex, both surfaces (particularly mid-

rib and prominent lateral veins) ± densely cov-

ered with longish yellowish to whitish spread-

ing hairs, very rarely glabrescent; petioles

(3— )4 — 6(— 9) mm long; stipular sheath with

3(-5) setae, the longest (l,5-)2^1(-5) mm. In-

florescence pyramidal to ellipsoidal, (60-)

10Q-200(-250) x (40-)70-150(-200) mm;

peduncles and pedicels thin but ± stiff, hairy to

subglabrous, divergent in fruit, elongating to

(2— )3— 6(— 7) mm. Flowers: corolla greenish,

greenish yellow or creamy yellow, sometimes

reddish brown tinged outside, glabrous or papil-

MAPS 46-47: — 46. Galopina tomentosa

47. Galopina crocyllioides

Anthospermeae

1,2:51

late near tip. cf: tube ± 0-0,4(-0,5) mm

long, lobes 1,1— 1,5(— 1 ,7) x 0,4-0,6(-0,9)

mm; anthers 0,9-1 ,2 mm long. $: tube

0-0,2(-0,3) mm long, lobes 0,6-0, 9 x

0,1-0,2(-0,3) mm; stigmas (2,2-)3-4,5(-5)

mm long; ovary c. 0,6-0, 9 x 0,8-1 ,3 mm, ±

wrinkled to ± warty. Fruit greenish brown,

brown to brownish black; each mericarp

(1— )1 ,2—1 ,8(— 2) x 0,8—1 ,4(— 1 ,6) mm, obovate

to elliptic, ± smooth, wrinkled or ± warty.

Occurs from Natal (Tongaland) to Transkei (Wild

Coast); growing mostly at the edge of (coastal or dune)

scrub or forest, occasionally also in open grassland. Map

46.

Vouchers: Rudatis 332 (BM; E; G; K; PRE; S; STE;

W); Strey 9561 (BR; NH; PRE; S); Ward 5488 (NH; NU;

PRE).

Some collections from coastal areas of Tonga- and

Zululand are atypical (less densely hairy or even glabres-

cent) and approach G. circaeoides (above) somewhat.

3. Galopina aspera (Eckl. & Zeyh.)

Walp., Repert. 2: 462 (1843), pro ‘Galopina

? aspera’ . Type: Cape, Katriviersberg, above

Philipstown (Ceded Territory), and on Chumi-

berg (Kafferland), Ecklon & Zeyher 2305

(SAM, holo.!; FI; G; GOET; M; MO; S; W,

iso.!).

Oxyspermum asperum Eckl. & Zeyh., Enum. 365

(1836). G. aspera (Eckl. & Zeyh.) Sond. in F. C. 3: 26

(1865).

G. oxyspermum Steud.,Nom. Bot. edn 2,1: 662 (1841),

nom. illeg.

Perennial herb, single- to several-

stemmed, cf, <jf, $ + $ or occasionally cf

+ $ + 9 or (very rarely) cf + $• Stems

(0,3-)0,5-l ,2(— 1 ,4) m long, ± erect to ascend-

ing, mostly unbranched, usually shortly hairy;

often with contracted short shoots bearing

relatively large leaves. Leaves: blades (25-)

30-45(-50) x (8-)12-25(-30) mm, ovate, el-

liptic or ovate-lanceolate, ± cuneate or ± trun-

cate at base, subacute to obtuse at apex, both

surfaces (particularly midrib and prominent la-

teral veins) ± densely covered with very short

whitish to yellowish spreading hairs or seldom

± papillate or subglabrous; petioles 0,5— 2(— 3)

mm long; stipular sheath with 5 or 3(-4) setae,

the longest c. 3 — 4( — 4,5) mm. Inflorescence ±

narrowly cylindrical, c. (80-)150-300(-380) x

(20-)40-80(-100) mm; peduncles and pedicels

thin but stiff, hairy to subglabrous, not conspic-

uously divergent in fruit, peduncles elongating

to c. ( 1 ,5— )2,5— 5(— 6) mm, pedicels to (0,8-)

1— 2(— 3) mm. Flowers: corolla greenish, green-

ish yellow, purplish or brownish red tinged,

often densely papillate outside, cf, (f: tube

(0,l-)0,2-0,5 mm long, lobes 1—1 ,5(— 1 ,6) x

(0,4— )0,5-0,6(-0,7) mm; anthers (0,5-)

0,7—1 , 1(— 1 ,2) mm long. 9: tube 0-0.2 mm

long, lobes (0,5— )0,6— 1 x (0,l-)0,2-0,4 mm;

stigmas (2-)2,5-4(-4,5) mm long; ovary c.

1-1,5 X 0,8-1 mm, densely tuberculate. Fruit

brownish; each mericarp 1,5-2 x (0,7-)

1—1 ,2(— 1 ,5) mm, conspicuously and densely

warty-tuberculate.

Known from Transvaal (along Drakensberg escarp-

ment), Swaziland, Natal, Transkei and eastern Cape; grow-

ing mostly at the edge of forest or scrub in ± sunny places.

Map 48.

MAP 48. — Galopina aspera

Vouchers: Acocks 9431; Schlechter 4549 (BM; BOL;

G; K; PRE; W; WU); Strey 6472 (NH; PRE).

Characterized vegetatively by its conspicuous leafy

short shoots (short shoot leaves often almost as long as long

shoot leaves).

1,2:52

4. Galopina crocyllioides Bar ex Schinz in

Vjschr. naturf. Ges. Zurich 68: 437 (1923); Hil-

liard & Burtt in Notes R. bot. Gdn Edinb. 32:

387 (1973). Types: Natal, Howick, Jutiod 238

(Z, lecto. !); Station Dumisa, Umgaye Flats,

Rudatis 867 (BM!; E!; K!; S!; W!).

Perennial herb, few- to single-stemmed,

0\ 9 + 9 or occasionally 9 + cf. Stems

(0,5-)0,6-l(-l,2) m long, erect, unbranched,

glabrous or younger parts ± papillate; usually

without short shoots. Leaves: blades

(35-)40-60(-70) x 12-25(-30) mm, ovate,

ovate-lanceolate to ± lanceolate, narrowed at

base, ± obtuse to subacute at apex, glabrous or

± papillate; petioles (0,5—) 1— 2(— 2,5) mm long;

stipular sheath with (2-)3^4(-5) setae, the long-

est 2,5-5 mm. Inflorescence ± narrowly cylin-

drical, c. (70-) 100-300 x 15-50(-70) mm; pe-

duncles and pedicels thin but stiff, not divergent

in fruit, peduncles elongating to c. ( 1— )2— 4(— 5)

Anthospermeae

mm, pedicels to 0,7-1, 5 (-2,5) mm. Flowers:

corolla greenish yellow or tinged brownish,

usually hairy outside. cf, 9: tube 0-0,5 mm

long, lobes 1-1,5 x 0,4— 0,7 mm; anthers

0,5-0,8(-l) mm long. 9: tiabe ± 0, lobes c.

0,5 x 0, 1-0,2 mm; stigmas 2-3(-3,5) mm

long; ovary c. 0,8-1, 2 X 0,7-1 mm, densely

hairy. Fruit reddish brown to blackish; each

mericarp l,3-2,2(-2,6) x 0,8-1, 2 mm,

densely covered with whitish spreading hairs c.

(0,3-)0,5-0,8 mm long.

Centred in the Natal Midlands and Coastal Regions;

also in Swaziland and in the Transkei. Growing mostly

among tall grass in grassland or in forest edge scrub. Map

47.

Vouchers: Compton 25527 (NBG; PRE); Hilliard &

Burtt 8072 (E; K; MO; NBG; NU; PRE; S); Flanagan 2848.

Well distinguished by its villous ovaries and fruits and

by the absence of leafy short shoots.

8443 CARPACOCE

Carpacoce Sond. in F.C. 3: 32 (1865); Hook. f. in Benth. & Hook, f., Gen. Plant. 2: 141 (1873);

K. Schum. in Pflanzenfam. 4,4: 130 (1891); Salter in J1 S. Afr. Bot. 3: 113 (1937), in Adamson &

Salter, FI. Cape Penins. 733 (1950); R. A. Dyer, Gen. 1: 623 (1975). Type species: Anthosper-

mum scabrum Thunb., i.e. Carpacoce scabra (Thunb.) Sond.

Lagotis E. Mey. in Drege in Flora 26, Bes. Beigabe 197 (1843), nom. nud.

Dwarf shrubs, rarely perennial herbs, 9 or occasionally d\ 9 + 9> 9 + Cf. Leaves

decussate or (rarely) in whorls of 3, often ericoid, ± pungent, acuminate or acute at apex, (sub)ses-

sile, with cup- or funnel-shaped stipular sheaths bearing minute setae or (less commonly) longer

bristles on either side. Inflorescence frequently very inconspicuous, made up of flowers paired or

solitary at nodes, or of mostly few-flowered cymes. Flowers 9> $ or occasionally Cf- Calyx

3-5(-6)-merous, lobes leaf-like, often unequal in size. Corolla 4-7-merous, tube cylindrical to ±

narrowly funnel-shaped, lobes linear to ± lanceolate, hooded (Fig. 1 la-d, f), spreading to spread-

ing-recurved (9 , cf) or ± erect (9); 9* Cf: corolla much larger than in 9- Stamens 4-7, anthers

purplish red to dark purplish brown, exserted, dangling on long slender filiform filaments. Ovary

with 1(2) fertile ovule(s); style 0; stigmas 1(2), long exserted, hairy, purplish red, greenish grey,

greyish or whitish. Fruit l(-2)-seeded, crowned by persistent calyx lobes; exocarp dehiscing into

valves, releasing endocarp with enclosed seed*. Chromosome number: 2n=22.

A small genus of 7 species endemic to the south-western Cape; only one species extends east to the Bathurst and

Albany distr. Map 58.

Note: In several Carpacoce species, flowers ‘transitional’ between 9 and ‘pure’ 9 may be found occasionally. Such

'transitional' flowers have corollas intermediate in size between and 9 and are characterized by the presence of small but

clearly discernible, (±) pollenless anther rudiments ; ’pure’ 9 with small corollas have no rudimentary anthers.

la Leaves (35— )50— 70(— 80) mm long; fruits crowned by 3 calyx lobes 2. C. gigantea

lb Leaves smaller, 3— 30(— 50) mm long; fruits crowned by 4-6 calyx lobes:

2a Leaves lanceolate or ovate-lanceolate, to 5(— 8) mm wide, not distinctly ericoid:

Endocarp plus enclosed seed (see Fig. 1 2 f-k) are referred to as 'diaspore’ (dispersal unit) in the following descriptions.

Anthospermeae

1,2: 53

3a Stipular sheath with a single minute seta, or seta absent; leaf blades rigid, conspicuously recurved,

(•4— )6— 1 0(— 12) mm long; corolla and calyx always 4-merous 3. C. curvifolia

3b Stipular sheath with numerous bristles, 1— 2,5(— 3) mm long; leaf blades ± membranaceous, not conspicuously

recurved (except for tips), 10— 30(— 35) mm long; corolla and calyx mostly 5-merous:

4a Plants slender, lax; partial inflorescences few- to 1-flowered; fruiting pedicels 3— 12(— 20) mm long; widely

distributed, east to the Heidelberg distr 1(a). C. spermacocea subsp. spermacocea

4b Plants more robust, distinctly woody at least near base; partial inflorescences several- to ± many-flowered;

fruiting pedicels 1-3 mm long; from the Mossel Bay distr. eastwards

1(b). C. spermacocea subsp. orientalis

2b Leaves ericoid, often ± needle-like, linear, linear-lanceolate or narrowly ovate-lanceolate, not wider than 2

mm;

5a Stipular sheath broadly cup-shaped, (4 — )5— 7 mm wide, 3-5 mm long; fruits crowned by 4 calyx lobes; corolla

tube (C^) 3-5,5 mm long 7. C. heteromorpha

5b Stipular sheath funnel-shaped or cup-shaped, small; fruits crowned by 5-6 calyx lobes; corolla tube

(<^, Cf ) 1-2,5 mm long:

6a Stipular sheath funnel-shaped, ( 1 ,5-)2-3(^t) mm long; fruits (4— )5— 7,5 X 2-3 mm 6. C. vaginellata

6b Stipular sheath cup-shaped, not longer than 0,5-1 ,2 mm; fruits 2-4 X (2,5-)3-3,5 mm:

7a Flowers with 1 stigma; fruits 1-seeded; leaves small, 3— 7(— 9) X 0,7— 1(— 2) mm 5. C. burchellii

7b Flowers with 2 stigmas; fruits 2-seeded; leaves (3— )5— 15(— 18) X (0,6-)0,8-2 mm:

8a More or less erect dwarf shrubs, intemodes (except near shoot tips) frequently longer than leaves;

fruiting pedicels c. 1-7 mm long 4(a). C. scabra subsp. scabra

8b Decumbent, ± much branched dwarf shrubs, intemodes mostly shorter than leaves; fruiting pedicels

to c. 1 mm long 4(b). C. scabra subsp. rupestris

1 . Carpacoce spermacocea (Reichb. f. )

Sond. in F.C. 3: 33 (1865); Salter in J1 S. Afr.

Bot. 3: 116 (1937), in Adamson & Salter, FI.

Cape Penins. 734 (1950). Type; none cited; a

specimen from ‘C.B.S.’. Neotype: 'Reich, [en-

bach] 63.9’ (E! ; LY!; WU!)'.

Anthospermum spermacoceum Reichb. f. in Sprengel,

Syst. Veg. 4: 338 (1827); Cruse in Linnaea 6: 17 (1831), in

Linnaea7: 134(1832).

Lagotis spermacocea (Reichb. f.) E. Mey. in Drege in

Flora 26, Bes. Beigabe: 197 (1843), nom. non valide publ.

Anthospermum foetidum Eckl. , nomen.

Two subspecies are recognized:

1(a). subsp. spermacocea.

Synonyms as above.

Perennial herb, 9 or occasionally 9" +

$, lax, straggling or scrambling, few- to seve-

ral-stemmed, with rhizomes or ± woody base,

(strongly) fetid. Stems c. 0,2-0,6(-0,9) m long,

sometimes rooting at lower nodes, branching

irregular, branches to 300 mm long, often with

leafy short shoots. Leaves decussate; blades

10—1 8(— 23) x ( 1 ,5— )2— 3(— 4) mm, narrowly

ovate-lanceolate to linear-lanceolate, ± spread-

1 These specimens are almost certainly duplicates of Ecklon

& Zeyher 2312 or ‘63.9’ (from ‘below Constantia, Tafel-

berg and in the Caledon and Swellendam Provinces’)

which must have been available to and studied by Rei-

chenbach.

tng, membranaceous, glabrous or margins

faintly scabrid; stipular sheath cup-shaped, with

3-5 (or sometimes more) bristles c. 1— 2(— 2,5)

mm long. Flowers in groups of few to 1 , lateral

and/or terminal, mostly on short lateral

branches, fruiting pedicels elongated,

3— 12(— 20) mm long; calyx (4— )5-6-merous,

lobes ( 1— )2,5— 5 mm long, linear-lanceolate, 1

or 2 often much larger than the others, erect to

spreading, or apices or upper thirds recurved;

corolla (4— )5(6-)-merous, greenish, greenish

yellow or dark purplish brown to blackish. 9:

tube 1-2 mm long, lobes 2-3,5 x 0,7-1 ,2 mm;

anthers 1,5-2, 2 mm long; stigma often shorter

than in 9- 9: tube 0,8-1 ,2 mm long, lobes

0,7-1 ,5 x 0,2-0, 4 mm; stigma 1, 5-13 mm

long; ovary c. 1,5-2 mm long. Fruit green,

turning dark brown, c. 2-3,5(-4) x 1-1,5 mm,

± obovate to cylindrical, often ± ribbed, 1-

seeded; diaspore c. 2— 3(^4-) x 1-1 ,5 mm, grey-

ish to black, ± cylindrical, hollowed out at base

(hollow sometimes filled with easily removable

‘plug’ of spongy tissue), ± rugose, occasio-

nally indistinctly ribbed. Chromosome number :

2n=22.

Occurring from the Cape Peninsula north-east to the

Tulbagh distr. and east to the Heidelberg distr. Growing in

wet or damp, often shady and sheltered localities (scrub

near streams, edge of kloof forests, base of cliffs, etc.).

Map 50.

1,2:54

Anthospermeae

FIG. 10. — 1 & 2, Carpacoce vaginellata: 1 , recently burnt plant, x 1;2, leaf pair with stipular sheath, x 4,5 ( Stokoe

s.n. sub SAM 64194). 3, C. scabra subsp. scabra, leaf pair with stipular sheath, x 4,5 (Story 4360). 4, C. curvifolia, leaf

pair, x 4,5 (Esterhuysen 27529). 5-7, C. spermacocea subsp. orientalist 5, stipular sheath, X 4,5 (Esterhuysen 27492);

6, longitudinal section of ovary showing ovule (ov) in endocarp case and spongy tissue at base (hatched), x 12; 7, diaspore,

note hollowed out base and 'plug’ of spongy tissue (hatched), x \2(Puff 790909-5/2).

Anthospermeae

1,2: 55

FIG. 11. a-e, Carpacoce spermacocea subsp. orientalis: a, Cf (or cT?) bud, note apical appendages of petals,

17,5, b, O flower (upper part of stigma broken off), x 17,5; c, 5 flower, x 17,5; d, apical appendage of petal (Cf),

* wfo^maticLh?irS (note pollen Srains>’ x 67 (Puff 790909-5/2). f, C. vaginellata, 9 flower, x 17,5

(Puff 790912-1/1) (SEM-graphs).

1,2: 56

Anthospermeae

Vouchers: Boucher 1 144 (PRE; STE); Schlechter 9225

(BM; BOL; BR; E; G; K; MO; PRE; S; US; W); Taylor

3550 (PRE; STE).

1(b). subsp. orientalis Puff, subsp. nov.,

habitu robustiore, floribus pluribus majoribus-

que et saepe fructibus longioribus a subsp. ty-

pica dijfert.

Type: Cape, Montagu Pass, Puff 790909-

5/2 (WU, holo.!; BOL; GRA; NBG; PRE; STE,

iso.!).

Perennial herb with woody base or distinct

dwarf shrub, $, occasionally $ + $ or

(seldom) 9" + Cf, often quite robust, strag-

gling, scrambling or sometimes ± erect, few-to

several-stemmed, (strongly) fetid. Stems c.

(0,l-)0,2-l,5 m long, branching often quite

regular, branches usually not more than 200

mm long, ascending to ± spreading, often ±

densely leafy. Leaves : blades ( 1 5— )20— 30(— 35)

x (2-)3-5(-8) mm, narrowly ovate-lanceolate

to linear-lanceolate. Flowers in groups of ± 8

or less, ± sessile, fruiting pedicels not longer

than c. 1-3 mm; calyx 5-6-merous, lobes 2-5

mm long; (cf ): corolla tube 0,8-1 ,5(-2) mm

long, lobes 2-3, 5(^1) x 0,8-1, 2 mm; anthers

1 ,7-2,5 mm long; $ : stigma shorter than in 9;

Cf: small rudimentary ovary present. 9: tube

0,6-1 mm long, lobes 0,6-l(-l,5) x 0,2-0, 4

mm; stigma 1 , 5-12 mm long; ovary c. 1 ,5-2,5

mm long. Fruit c. (3— )3 .5—5 x 1 ,5—2,5 mm;

diaspore c. 3—3 ,5( — 4) x 1,2-1, 7 mm. Chromo-

some number. 2n=22. Fig. 10: 5-7; 1 la-e; 12 a

& e.

Occurring in the eastern part of the Cape Floristic Re-

gion from the Mossel Bay distr. to the Uitenhage and Port

Elizabeth distr. Habitat as for subsp. spermacocea. Map 5 1 .

Vouchers: Acocks 21177 (K; M; PRE); Esterhuysen

27493 (BOL; PRE); Thode A 856 (K; MO; NH; PRE). '

The two subspecies cannot always be satisfactorily

separated morphologically; their distribution ranges, how-

ever, do not overlap. Overall, subsp. orientalis tends to be

more robust, is generally larger ana has more flowers; it is,

furthermore, often distinctly woody. Odd luxurious forms

of subsp. spermacocea (e.g. Williams 2604; K; NBG; PRE)

approach subsp. orientalis in leaf size and in having more

flowers per partial inflorescence than usual.

Within subsp. orientalis two ± distinct ‘Forms’ (eco-

types) can be distinguished: (1) ‘typical’ orientalis: luxu-

riant, straggling or scrambling plants with stems to 1,5 m

long. (2) Erect, ± stunted, distinctly dwarf shrubby and

rather densely leafy plants to 0,5 m tall; they appear to be

confined to the mountains (750 m and above) where they

occur in wet, rocky areas (base of cliffs, etc.).

C. spermacocea should not be confused with Antho-

spermum herbaceum. C. spermacocea is distinguished by

its nauseating odour, flowers with hooded corolla lobes

(Fig. 1 1 a-d) and 1 stigma and fruits with 1 or 2 enlarged

calyx lobes.

2. Carpacoce gigantea Puff, sp. nov.,fo-

liis multo longioribus et fructibus lobis calycis

persistentibus foliaceis tribus coronatis a cete-

ris speciebus Carpacocis praeclare distinguitur .

Type: Cape, ‘in the ascent of the Craggy

Peak in the Great Range at Swellendam’

[Langeberg north of Swellendam], Burchell

7320 (K, holo.!).

Shrub, $ or 9> — erect. Stems to 0,9 m

long, ± much branched, densely leafy. Leaves

decussate; blades (35-)50-70(-80) x (2-)3

(-7) mm, linear-lanceolate, glabrous, margins

MAPS 49-58: — 49. Carpacoce vaginellata

50. Carpacoce spermacocea subsp. sper-

macocea

51. Carpacoce spermacocea subsp.

orientalis

52. Carpacoce scabra subsp. scabra

53. Carpacoce scabra subsp. rupestris

54. Carpacoce curvifolia

55. Carpacoce burchellii

56. Carpacoce gigantea

57. Carpacoce heteromorpha

58. Carpacoce, all taxa

Anthospermeae

1,2: 57

flat to slightly revolute; stipular sheath cup-

shaped. Flowers solitary on short lateral

branches, mostly hidden amongst foliage, pedi-

cels to 4 mm long; calyx 3-merous, lobes

(6— )8— 12 mm long, ± erect, sometimes ± un-

equal in size; corolla 5(-6)-merous. tube

4-5,5 mm long, lobes 4—4,5 x 0,7-1 mm;

anthers 3-3,5 mm long. 9: tube 2,5 mm long,

lobes 0,7 X 0, 1-0,2 mm; stigma 1, c.

10-15mm long; ovary 2-2,5 mm long. Fruit

grey, 2,5-5 x 1,2-2 mm, ± ellipsoidal to cy-

lindrical, 1-seeded; diaspore (2— )2,5— 3 x 1,5

mm, greyish to black, elliptic to ± rectangular

in outline, rugose.

Only known from the mountains above Swellendam;

growing on moist, well drained mountain slopes. Map 56.

Voucher: Taylor 424\ (PRE;STE).

A very distinct species (large leaves; fruits crowned by

3 calyx lobes) but possibly extinct. Extensive searches for

the plants in recent years were unsuccessful.

3. Carpacoce curvifolia Puff, sp. nov.,

foliis rigidis nitidis conspicue recurvisque a

ceteris speciebus Carpacocis facile distinguitur .

Type: Cape, Uniondale distr., Tsitsikam-

ma Mts near Joubertina, Esterhuysen 27334

(BOL,holo.!;WU, iso.!).

Perennial herb with woody base or dwarf

shrub, 9 or 9’ many-stemmed, decumbent,

seldom ± erect. Stems c. (0,l-)0, 15-0,5

(-0,6) m long, ± sparsely to much branched.

Leaves decussate or (rarely) in whorls of 3;

blades (4 — )6— 1 0(— 1 2) x (l-)2^4 mm, lanceo-

late to ovate-lanceolate, usually strongly re-

03curved, rigid, shiny, glabrous or margins

faintly scabrid; stipular sheath cup-shaped.

Flowers single or paired at nodes, ± sessile,

largely hidden in stipular sheaths or, if termi-

nal, with pedicels up to 4 mm long; calyx 4-

merous, lobes (0,8-)l-2,4 mm long, lanceolate

to triangular, ± spreading, 2 often up to twice

as long as the others; corolla 4-merous. tube

( 1— )1 ,5— 2(— 3) mm long, lobes c. 2-3 x 0,8-1

(-1 ,5) mm; anthers 0,8-1 ,5(— 1 ,7) mm long. 9 '■

tube 0,5-1 mm long, lobes 0,5-1 X 0,3 mm;

stigma 1, 4-6 mm long; ovary 1-1,2 mm long.

Fruit greyish brown, c. 2-2,5 x 1-1,5 mm, ±

obovate, lower half often laterally compressed

and hidden in stipular sheath, 1-seeded; dias-

pore c. 1,2-2 x 0,8-1, 2 mm, black, ellipsoi-

dal, base often with an appendage of (easily

removable) spongy tissue, ± rugose. Fig. 10:

4.

Occurring on the mountains in the eastern part of the

Cape Floristic Region, from the Uitenhage to the Ladismith

district (Anysberg). Growing on steep rocky slopes, on

ledges and in crevices; appears to favour moist, sheltered

situations. Map 54.

Vouchers; Esterhuysen 27093; 27529; 28043 (BOL;

PRE).

Easily distinguished by its rigid, shiny, recurved

leaves and small fruits crowned by relatively small broad

calyx lobes.

Esterhuysen 25973 (BOL; WU) from the Anysberg,

the westernmost, ± isolated locality, appears to be a rather

atypical, depauperate form.

4. Carpacoce scabra (Thunb.) Sond. in

F.C. 3: 33 (1865); Salter in J1 S. Afr. Bot. 3:

115 (1937). Syntypes: Cape, Thunberg (sheets

23317, 23318, UPS!).

Anthospermum scabrum Thunb., Prodr. 1: 32 (1794), FI.

Cap. 1: 573 (1813), FI. Cap., edn Schultes 158 (1823);

Cruse, Rub. Cap. 14 (1825), in Linnaea 6: 14(1831).

4 (a), subsp. scabra.

Dwarf shrub, $ , 9 or occasionally +

9, 9 + Cf or cf, many-stemmed, often

rounded to ± cylindrical, sometimes slightly

fetid. Stems (80-) 150-400(^450) mm long,

sparsely to ± much branched, branches ascend-

ing to ± spreading, not densely leafy. Leaves

decussate; blades (3— )5— 15(— 1 8) x (0,6-)0,8-2

(-2,5) mm, linear(-lanceolate), ascending to

erect, ericoid, often shallowly concave above or

semiterete in section, scabrid at least on mar-

gins; stipular sheath cup-shaped. Flowers soli-

tary or paired, ± sessile to shortly pedicellate,

fruiting pedicels sometimes elongating to c. 7

mm; calyx 5(-6)-merous, lobes (2,5-)4— 6 mm

long, linear-lanceolate, subequal, spreading to

± erect; corolla 5(-6)-merous, yellowish green,

often with reddish brown or brownish purple

streaks. 9> Cf: tube 1,5-2, 5 mm long, lobes

2,5 — 4 x 1-2 mm; anthers 2— 2,5(— 3) mm long;

9 : gynoecium as in 9 ; Cf : small rudimentary

ovary and stigmas present. 9: tube 0,3-0, 6 mm

long, lobes 0,5-1, 2 x 0,4— 0,5 mm; stigmas 2,

(4— )5-9 mm long; ovary c. 1,5-2 mm long.

Fruit green, turning grey-brown, c. 2-3 x

(2,5-)3-3,5 mm, ± turbinate to subglobose, 2-

seeded; diaspore c. 1,5-2, 5 x 1-1 ,5 mm, grey-

ish white to dark grey, ± elliptical to obovate in

dorsal view, dorsal side convex, ventral side

plane to slightly concave, ± rugose and ribbed.

Chromosome number. 2n=22. Fig. 10: 3; 12:

b, f, j & k.

1,2:58

Anthospermeae

FIG 12 — Fruits (a-e) and diaspores (f— k) of Carpacoce species: a & e, C. spermacocea subsp. orientalis: a, fruit,

x 9; e, fruit from above, x 9 (Puff 790909-5/2). b, f, j & k, C. scabra subsp. scabra: b, fruit, x 9; f, diaspore, ventral

side, and two exocarp valves, x 17,5; j, diaspore in side view, x 17,5; k, diaspore, dorsal side, x 17,5 (Esterhuysen

10961). c, h & i, C. heteromorpha: c, fruit, x 9; h, diaspore in side view, x 17,5; i, diaspore, ventral side, x 17,5 (Puff

800917-4/1). d & g, C. vaginellata: d, fruit, x 9; g, diaspore, ventral side, x \1 ,5 (Puff 790912-1/1) (SEM-graphs).

Anthospermeae

1,2: 59

In drier inland areas, from the Calvinia distr. south-

east and south to the Laingsburg and Worcester distr. Grow-

ing mostly in sandy areas on mountain slopes or on ridges

and amongst rocks; confined to arid Fynbos types. Map 52.

Vouchers: Acocks 17377 (K; PRE); 23705; Bolus 9054

(BOL; K; NBG; PRE).

4(b). subsp. rupestris Puff, subsp. nov.,

habitu decumbenti, internodiis plerumque foliis

brevioribus et pedicellis in fructu usque ad 1

mm longis a subsp. typica differt.

Type: Cape, Ceres-Tulbagh distr., Swart-

gat Peak, Witzenbergen, Esterhuysen 27930

(BOL, holo. !; WU, iso. !).

Dwarf shrub, 9 or occasionally $ +

5 or $ 4- cf, several-stemmed, decumbent or

sometimes cushion-forming. Stems 100-200

(-250) mm long, ± much branched, densely

leafy. Leaves : blades (6-)8-12(-15) X

0,8-1 ,5(-2) mm, linear-lanceolate to narrowly

oblanceolate, ± erect, ± triangular in section

or ± flat and with a prominent midrib below.

Flowers solitary, ± sessile, fruiting pedicels

elongating to c. 1 mm; calyx lobes 2,5—4 mm

long; Cf: tube 1— 2(— 2,5) mm long, lobes

2,5— 4(— 5) x 0,8-1 ,2 mm; anthers 1,5-2, 5 mm

long. $: tube 0,3-0,5(-l) mm long; lobes

0,5-0, 8 x 0,2-0,3(-0,5) mm, often hidden be-

tween calyx lobes; stigmas 2 (rarely 1), 5-9 mm

long; ovary c. 1—1 ,5(— 2) mm long. Fruit 2-3,5

x (1 ,5— )2— 3 ,5 mm, ± obovate, 2- (rarely 1-)

seeded; diaspore c. 1,5-2, 5 x 1,2-1 ,5 mm,

black.

In the mountains of the Worcester distr. and on the

Witsenberg range. Growing on rocky slopes amongst rocks,

on ledges, etc.; only found on the upper parts or tops of the

highest mountains, c. 1 200-2 000 m. Map 53.

Vouchers: Esterhuysen 26491 (BOL); 31146 (BOL;

WU); Taylor 6567 (PRE; STE).

5. Carpacoce burchellii Puff, sp. nov.,

habitu C. scabrae similis sed ovariis/fructibus

ovulo/semine fertili solitario differt.

Type: Cape, Caledon distr., south slopes

of Riviersonderend Mts near Greyton, Ester-

huysen 20782 (BOL, holo. !).

Dwarf shrub $ or many-stemmed, ±

erect and cylindrical. Stems c. (50-)80-300 mm

long, unbranched to sparsely branched. Leaves

decussate; blades 3— 7(— 9) x 0,7-l(-2) mm,

linear, ericoid, glabrous or margins slightly sca-

brid; base of blades ± sac-like, stipular sheath

cup-shaped. Flowers in groups of 1-4 at nodes,

± sessile, or sometimes terminal, shortly pedi-

cellate; calyx 5-merous, lobes ( 1 ,5— )2— 3 ,5 mm

long, 1 often distinctly larger than the others;

corolla 5-merous. $ tube 1-2 mm long, lobes

(2-)3-4 x 0,5-1 mm; anthers 1,5-2 mm long;

stigma often shorter than in $?• tube 0,3-0, 6

mm long, lobes 0,4-1 x 0,2-0, 3 mm; stigma

1, (4—)5,5-8 mm long; ovary (1— )1 ,5—2,2 x

1-1,5 mm, with 1 fertile ovule. Mature fruit not

seen.

Only known from a few mountains in the Paarl and

Caledon distr. Growing on moist, well drained mountain

slopes, occasionally in marshy situations and in shady

kloofs. Map 55.

Vouchers: Burchell 7736 (K); Esterhuysen 20092

(BOL; K; PRE).

In habit, C. burchellii can be very similar to C. scabra

(above); it is, however, easily distinguished by its ovaries/

fruits which have only one fertile ovule/seed.

Plants resprouting after fire produce rather long un-

branched flowering shoots which bear relatively larger,

wider and less distinctly ericoid leaves (e.g. Esterhuysen

13707, BOL; K; PRE). Stokoe sub Marloth 1 1009 (PRE) is

an unusually dwarfed form with flowering shoots only a few

centimetres tall.

6. Carpacoce vaginellata Salter in J1 S.

Afr. Bot. 3: 113 (1937), in Adamson & Salter,

FI. Cape Penins. 734 (1950). Type: Cape Pen-

insula, Muizenberg Mt., Salter 6271 (BOL,

holo.!; K; NBG, iso.!).

Dwarf shrub, or occasionally $ +

0\ $ + cf, or cf, many-stemmed, cylindri-

cal to ± rounded, occasionally decumbent,

sometimes faintly fetid. Stems (0,1-)

0,15-0,4(-0,7) m long, ± much branched (un-

branched or sparsely branched in plants res-

prouting after fire), branches ascending to ±

erect. Leaves decussate; blades (7— )10— 25(— 30)

x 0,7-1 ,5(-2) mm, linear, ± erect, ericoid,

semiterete to ± triangular in section, margins

scabrid or with stiff whitish hairs to 0,2 mm

long; stipular sheath long and funnel-shaped.

Flowers mostly solitary, lateral and/or terminal

on branches, ± sessile or shortly pedicellate;

calyx 5(-6)-merous, lobes 3-8 mm long, lin-

eal-lanceolate), subequal, ± erect; corolla

5(-6, rarely -7)-merous, greenish yellow, red-

dish brown to dark purplish brown. , cf : tube

(1— )1 ,5—2,5 mm long, lobes (3— )3 ,5— 5(— 5 ,5) x

(0,7—) 1-2 mm; anthers (1 ,5-)2-3(— 4) mm long;

stigma always shorter, ovary sometimes

smaller than in $ ; cf : small rudimentary stigma

and ovary present, tube 0,5-1, 5 mm long,

1,2:60

Anthospermeae

lobes 0,5-1, 5 x 0, 1-0,5 mm, usually hidden

between calyx lobes; stigma 1 (very rarely 2),

(6— )8— 12(— 17) mm long; ovary c. 2-4 mm

long. Fruit green, turning dark grey-brown,

(4 — )5— 7,5 x 2-3 mm, ± obovate, l-(very

rarely 2)-seeded; diaspore (3— )3,5— 5 x 1,5-2

mm, grey to blackish, often laterally ± com-

pressed, ± elliptic to obovate in dorsal view,

rugose to muricate. Chromosome number :

2n=22. Fig. 10: 1,2; Ilf; 12d&g.

Widely distributed and extending east to the Albany

and Bathurst distr. Mostly growing in Coastal and Mountain

Fynbos, often on sandy to rocky slopes, plateaux or flats; in

the easternmost localities also in grassveld. Map 49.

Vouchers: Acocks 22841 (K; PRE); Esterhuysen 17290

(BOL; NBG; PRE); Taylor 7518 (K; PRE).

Specimens with 2-seeded fruits or with both 1- and 2-

seeded fruits are (very) uncommon. Plants rather frequently

bear ‘pseudo-fruits’ without well developed seeds; they are

(much) smaller than fertile fruits and are often reddish at

first and then brownish grey.

Superficially, C. scabra (no. 4) often closely resem-

bles C. vaginellata but the latter is distinguished vegetati-

vely by its long, funnel-shaped stipular sheaths

[( 1 ,5— )2— 3(— 4) mm vs. 0,5-1 mm long; compare Fig. 10: 2

and 10: 3].

7. Carpacoce heteromorpha (Buek) L.

Bol. in J. Bot., Lond. 34: 25 (1896). Type:

Cape, Caledon distr., on Babylonstorensberg,

Zwart & Marais sub Ecklon & Zeyher 2421

(SAM, holo. !).

Merciera heteromorpha Buek in Eckl. & Zeyh., Enum.

387(1837).

Dwarf shrub, single- to several-

stemmed, ± cylindrical and erect to rounded.

Stems c. 0,15-0,45 m long, ± sparsely to much

branched, branches often long, ascending to ±

erect, very densely leafy. Leaves decussate;

blades 10-25(-30) x 0,8-1, 5 mm, linear,

spreading-recurved to strongly recurved, ± te-

rete, semiterete to ± triangular in section, mar-

gins (lower half) with fine white hairs c.

0,2-0, 4 mm long; stipular sheath broadly cup-

shaped, hairy. Flowers solitary, lateral and/or

terminal on short branches, ± sessile, mostly

hidden amongst foliage; calyx 4-merous, lobes

(5— )6— 8 mm long, linear, subequal, ± erect,

hairy on margins; corolla (5-)6-7-merous,

greenish yellow to yellow, tube 3-5,5 mm

long, lobes (3— )3,5— 4,5(— 5) x 0,5-1 mm,

sometimes with a few stiff, erect bristles near

tip; anthers 2-2,7 mm long; stigma 1 (very ra-

rely 2), 8-13 mm long; ovary c. 1,5 mm long.

Fruit ash-grey, c. 1,5-3 mm in diam., subglo-

bose to ± turbinate, l-(very rarely 2)-seeded;

diaspore c. 1 ,2-2,5 x 1 ,2-2 mm, black or dark

grey, subglobose to ± pyriform, with a distinct

vertical groove, muricate or ± rugose. Chro-

mosome number. 2n=22. Fig. 12c, h & i.

Occurring from the Worcester and Somerset West

distr. south-east to the western Bredasdorp distr. Growing

mostly in dry areas, in gravelly to sandy soil amongst rocks;

in Coastal or Mountain Fynbos, less commonly in Coastal

Renosterveld. Map 57.

Vouchers: Boucher 782 (PRE; STE); 1623; Schlechter

7648 (BM;E; K; PRE; US).

Forms with less distinctly recurved leaf blades should

not be confused with the allied C. vaginellata (above). C.

heteromorpha is easily distinguished by the rather long

whitish hairs on the stipular sheaths and lower half of the

leaf blades, by having flowers with unusually long corolla

tubes and by its ± round fruits crowned by 4 long calyx

lobes (Fig. 12c); both flowers and fruits are usually hidden

amongst the dense foliage and plants give the impression of

being permanently in a vegetative state.

Rubieae

1,2:61

Tribe RUBIEAE

by C. Puff*

Herbs. Leaves and leaf-like stipules forming (pseudo)whorls. Flowers 9 . occasionally also

Cf, 9; corolla rotate to campanulate, tube mostly very short; disk present; ovary 2-locular, each

locule with a single ovule affixed to septum. Fruit dry and dehiscing into 2 mericarps, or fleshy;

one carpel occasionally aborted. Seed with membranous testa adhering to fruit wall, dorsal side

convex, ventral side plane to concave. Chromosome numbers'. x= 1 1 , rarely x= 10 (2n=22, 44; 40

in the Flora area).

Note : In the following descriptions the morphologically more precise but awkward term “leaves and leaf-like stipules

in whorls of is abbreviated to “leaves in whorls of”. Note also that leaf descriptions and measurements strictly refer to

leaves from the middle region of flowering shoots; in leaf width measurements the reflexed margins are not taken into

account.

Only two genera of this ± cosmopolitan tribe occur in Southern Africa:

Leaves petiolate; corolla mostly 5-merous; fruit fleshy, glabrous Rubia (below)

Leaves (sub)sessile; corolla mostly 4-merous; fruits dry, glabrous or variously hairy

Galium (p. 1,2: 65)

8489 RUBIA

Rubia L., Sp. PI. 109 (1753), Gen. PI. edn 5: 47 (1754); Sond. in F.C. 3: 34 (1865); R. A. Dyer,

Gen. 1: 626 (1975); Verde, in F.T.E.A. Rubiaceae 1: 380 (1976); Puff in J1 S. Afr. Bot. 50: 348

(1984). Type species: R. tinctorum L.

Scrambling, creeping or climbing perennial herbs, somewhat woody near the base; roots quite

woody, reddish (yielding a red dye). Stems branched, 4-angled, ± brittle, mostly beset with

recurved prickles. Leaves in whorls of 4-8(-12), petiolatet, blades cordate or broadly ovate,

ovate-lanceolate to linear, rounded to ± cordate at base or narrowed to base, with 1— 5(— 7) promi-

nent veins. Flowers in axillary and/or terminal cymes, 5(rarely 4 or 6)-merous. Calyx : lobes

obsolete. Corolla rotate to subcampanulate, greenish, yellowish green to yellow; lobes ± triangu-

lar, (long-)acuminate. Anthers exserted. Ovary 2-locular, one ovule in each chamber, crowned by

a minute disk; style branches 2, joined below; stigmas capitate. Fruit fleshy, glabrous, consisting

of 2 round mericarps, each with a single seed, round on dorsal side, plane to convex on ventral

side; one mericarp often aborted. Chromosome number. 2n=22 (in the Flora area).

A widely distributed genus (Europe, Asia and Africa) of c. 60(?) species; only few (3, or 4?) species occur in Africa south

of the Sahara.

In Southern Africa, Rubia is easily distinguished from Galium (fleshy vs. dry fruits, petiolate vs. epetiolate leaves),

but elsewhere the distinction between the two genera may be highly problematic [see comments in Verdcourt in F.T.E.A.,

Rubiaceae 1: 381 (1976)].

la Leaves in whorls of 4 (very rarely 3 or 5), blades with 5 (7; very rarely 3) prominent veins

1 . R . cordifolia subsp. conotricha

lb Leaves in whorls of 6— 8(— 1 2), blades with 3 prominent veins or only midvein distinct;

2a Leaf blade length to width ratio less than 3:1; blades with 3 prominent veins, to 25(-30) mm long and 5-15 mm

wide, cordate or ovate-lanceolate to broadly ovate, rounded to cordate at base 2. R. petiolaris

2b Leaf blade length to width ratio more than 3:1; blades with only midvein prominent, to 50(— 70) mm long and

1-8(10) mm wide, lanceolate to linear, narrowed the base; blades occasionally difficult to distinguish from

petiole , 3. R. horrida

* University of Vienna, Austria.

t The European species R. peregrina L. (with sessile leaves) is included in Flora Capensis (Sonder, op.cit.). A Thunberg

collection of a plant supposedly introduced to the Cape is cited. As I have not been able to trace this or any other

collection of R. peregrina from South Africa, this species is not included here.

1,2: 62

Rubieae

1. Rubia cordifolia L., Syst. Nat., edn

12,3 (App.) 229 (1768), Mant. Alt. 197 (1771);

DC., Prodr. 4: 588 (1830). Type: Majorca, L.

Gerard (ubi?).

subsp. conotricha (Gand.) Verde, in Kew

Bull. 30: 323 (1975), in F.T.E.A. Rubiaceae 1:

381, fig. 57 (1976); Puff in J1 S. Afr. Bot. 50:

349 (1984). Type: [East Griqualand], near Um-

zimhlawa R., Schlechter 6550 (LY, holo.;

BM!;K!;S!).

R. conotricha Gand. in Bull. Soc. bot. Fr. 65: 35 (1918).

R. longipetiolata Bullock in Kew Bull. 1932: 497 (1932).

Type: Kenya, Mt Elgon, Lugard 204 (K, holo. !; EA).

R. cordifolia sensu Sond. in F.C. 3: 35 (1865), non L.

Stems to c. 5 m long, distinctly 4-ribbed,

with recurved prickles on ribs, occasionally

with longish white hairs between ribs, at least

around nodes. Leaves in whorls of 4 (very

rarely 3 or 5); blades 27-55 x (3-)6-32 mm,

(narrowly) lanceolate to broadly ovate, rounded

to cordate at base, acute to acuminate at apex,

with 5 (7, very rarely 3) prominent veins; mar-

gins and veins below with recurved prickles,

upper surface ± glabrous to scabrid, lower sur-

face ± glabrous or sparsely to ± densely cov-

ered with longish white hairs, blades rarely gla-

brous; petioles (10-)15-60(-75) mm long,

mostly with recurved prickles and occasionally

also hairy. Inflorescence several- to many-flow-

ered, ± lax to rather dense; peduncles and pedi-

cels 1-8,5 mm long, glabrous, somewhat pu-

bescent or with small prickles; ultimate bracts

c. 1-2 mm long or sometimes absent. Flowers 5

(very rarely 6)-merous; corolla 3-5,7 mm in

diam.; filaments to 1 mm, anthers to 0,5 mm

long; ovary 0,4-0, 8 mm long. Fruit dark purple

to black, each mericarp 2,7-3 ,9 mm in diam.

Chromosome number. 2n=22.

Known from Transvaal (Soutpansberg distr. and along

Drakensberg escarpment), Swaziland, Natal, Transkei and

eastern Cape (to East London distr.); extending to tropical

East Africa and also in Angola. Grows at forest edges, in

clearings, scrub etc. (“typical” conotricha and “Soutpans-

berg-Drakensberg Escarpment Form”) or in coastal (sand

dune) scrub or dune forest (“Coastal Form”). For details on

ecotypes (“Forms”) see Puff in J1 S. Afr. Bot. 50: 350

(1984). Map 59.

Vouchers: Puff 790415-6/1 (BR; J; NU; WU); Schee-

pers 973 (K; PRE); Strey 8 103 (K; PRE).

2. Rubia petiolaris DC., Prodr. 4: 588

(1830); Puff in J1 S. Afr. Bot. 50: 353, fig. 2a

(1984). Type: Cape, around Krakakamma, Bur-

chell 4561 (K, iso.!).

R. petiolaris var. isophylla Sond. in F.C. 3: 35 (1865).

Syntypes: Cape, Zwartkopsrivier, Zeyher 2721 (S, lecto. !);

Algoa Bay, Port Elizabeth, Ecklon & Zeyher 2320 (S!;

SAM!: 2 sheets, one mixed with R. cordifolia subsp. cono-

tricha)-, Port Elizabeth, Drige s.n. (“/?. petiolaris DC.”;

K!; S!; SAM!); ‘CBS’, Thunberg rs.n. (“R. cordifolia L.”;

S!: several sheets).

Stems to c. 3 m long, distinctly 4-ribbed,

with recurved prickles on ribs, younger parts

often with hairs below nodes, occasionally with

some short whitish hairs between ribs. Leaves

in whorls of 6-8(-9); blades 8-25(-30) x 5-15

mm, cordate or ovate-lanceolate to broadly

ovate, rounded to cordate at base, acute at apex,

Rubieae

1,2: 63

FIG. 13. — Rubia petiolaris: 1, part of plant, X 1; 2, enlargement of stem, x 5; 3, leaf, x 3; 4, corolla with stamens,

x 10; 5, gynoecium, x 10; 6, fruit, x 3 (1-5: Giffen 1095; 6: PeglerAM).

1 ,2: 64

Rubieae

MAP 60. — • Rubia petiolaris

■ Rubia horrida

with 3 prominent veins; margins and at least

midvein below with recurved prickles, rarely

scabrid and/or with prickles above; petioles

1 0— 30(— 35) mm long, mostly with (2 rows of)

recurved prickles. Inflorescence few- to seve-

ral-flowered, rather lax; peduncles and pedicels

1-5 mm long, glabrous or (seldom) somewhat

hairy; ultimate bracts c. 2-4 mm long. Flowers

5 (very rarely 4)-merous; corolla 3-5 mm in

diam.; filaments to 0,9 mm long, anthers to 0,4

mm long; ovary c. 0, 5-0,9 mm long. Fruit

blackish, each mericaro c. 2,4-3 mm in diam.

Chromosome number. 2n=22. Fig. 13.

Endemic. Ranges from the Orange Free State, Leso-

tho, south-west Natal and eastern Cape to the Riversdale

distr. ; growing mostly in scrub. Map 60.

Vouchers: Dieterlen 331b (K; NBG: PRE; SAM);

Johnson 838; Puff 790112-12/1 (BR;J;W;WU).

Variable both morphologically (leaves: see Map 60!)

and ecologically (coastal fynbos to afromontane areas; c.

15-2200 m!). Closely allied to R. horrida (below); the two

species are occasionally difficult to separate in the area

between the Orange River and the Graaff-Reinet and Cra-

dock districts.

Rubieae

1 ,2: 65

3. Rubia horrida (Thunb.) Puffin Kew

Bull. 32: 432 (1978), in J1 S. Afr. Bot. 50: 357,

fig. 2b,c( 1984).

Galium horridum Thunb. in Hoffmanns Phytograph.

Blatter 1: 16 (1803) & FI. Cap. 1: 556 (1813); Sond. inF.C.

3: 37 (1865). Type: Masson, in herb. Thunberg (sheet

3323, UPS. holo.!).

R. petiolaris DC. var. heterophylla Sond. in F.C. 3: 35

(1865). Types: Orange Free State, on the Groot Vetrivier,

Burke & Zeyher 774 [ Burke s.n. (K!); Zeyher s.n. (S ! ; sub

SAM 1 6074!); Zeyher 774 ( BM ! ; K!)].

Stems to c. 2-3 m long, distinctly 4-

ribbed, with recurved prickles on ribs, younger

parts often with short hairs below nodes, occa-

sionally also with some short whitish hairs be-

tween ribs. Leaves in whorls of 6-8(-12),

blades (8-)12-50(-70) x 1— 8(— 1 0) mm, lan-

ceolate, linear-lanceolate to linear, mostly nar-

rowed to base, blades occasionally difficult to

distinguish from faintly winged petioles, acute

at apex, only midvein prominent; margins and

midvein below with recurved prickles, seldom

scabrid above; petioles (6-) 10-35(^15) mm

long, ± 3-angular in section or somewhat ca-

naliculate, mostly with (2 rows of) recurved

prickles. Inflorescence several- to many-flow-

ered, rather lax to ± dense; peduncles and pedi-

cels 1-6 mm long, glabrous or sometimes

slightly hairy; ultimate bracts c. 2-5 mm long,

linear. Flowers 5 (very rarely 4 or 6)-merous;

corolla 3-5,9 mm in diam.; filaments to 0,6

mm, anthers to 0,5 mm long; ovary c. 0,4-0, 8

mm long. Fruit blackish or dark purple, each

mericarp (2,2-)2,5-3,7 mm in diam. Chromo-

some number: 2n—22.

Known from South West Africa/Namibia (Windhoek

Bergland), south-east Botswana, Transvaal, Natal (Estcourt

and Weenen districts). Orange Free State and Cape; also in

Zimbabwe. Growing mostly in bush clumps, Acacia scrub,

open woodland or riverbank vegetation. Map 60.

Vouchers: Acocks 2115 (K; KMG; PRE); Puff

790516-1/1 (BR; J; NU; WU); Scheepers 1722 (K; PRE);

Wells 2409 (K; PRE).

Variable in leaf size and shape (see Map 60); leaf blade

length to width ratios help to distinguish R. horrida from

the closely allied R. petiolaris. See Puff in J1 S. Afr. Bot.

50: 360 (1984) for details.

8486 GALIUM

Galium L., Sp. PI. 105 (1753), Gen. PI. edn 5: 46 (1754); Sond. in F.C. 3: 35 (1865); R. A. Dyer,

Gen. 1; 625 (1975); Verde, in F.T.E.A. Rubiaceae 1: 383 (1976); Puffin J1 S. Afr. Bot. 44: 219

(1978). Type species: G. verum L.

Perennial herbs or annuals. Stems erect, creeping or climbing, often distinctly 4-angled,

glabrous, hairy or with recurved prickles. Leaves in whorls of 4-10, (sub)sessile, blades linear,

lanceolate to (ob)ovate, mostly with a prominent midvein (1-nerved). Flowers in axillary and/or

terminal many- to few-flowered cymes, (and protandrous), rarely and 4-merous. Calyx:

lobes obsolete. Corolla mostly rotate, greenish(yellow), creamy white to bright yellow, lobes

ovate to triangular, acute to acuminate. Anthers exserted. Ovary 2-locular, one ovule in each

chamber, crowned by an often 2-lobed disk; style branches 2, joined below; stigmas capitate. Fruit

dry, glabrous or variously hairy, dehiscing into 2 (sub)globose or reniform mericarps, each with a

single seed, round on dorsal side, ± excavated on ventral side; one mericarp occasionally aborted.

Chromosome number: 2n=22, 44, rarely 20 (in the Flora area).

A cosmopolitan genus of several hundred species, having its main range of distribution and highest species densities in

temperate regions. Most of the taxa occurring in the Flora area are endemic but show clear affinities to tropical African and

north-temperate species groups. Map 61.

la Leaves 3-nerved, strictly in whorls of 4 1 . G. thunbergianum

lb Leaves 1-nerved, in whorls of 4-10:

2a Inflorescences very reduced, flowers in groups of 3-1 (occasionally 4, rarely -6):

3a Leaves large, to 38(^45) mm long and 10 mm wide, with recurved prickles on margins:

4a Flowers mostly single on peduncles/pedicels ( 8— ) 1 2— 20(— 3 5 ) mm long 2. G. spurium subsp. africanum

4b Flowers in groups of 3, occasionally 4 or rarely 6:

5a Ovary and fruit with hooked hairs; corolla minute, to c. 2 mm in diam

‘Galium spurium-aparine complex’ (page 1 ,2: 67)

5b Ovary and fruit glabrous; corolla larger, to 4 mm in diam 3. G. undulatum

3b Leaves small, to 14 mm long and 2 mm wide, mostly without prickles on margins:

6a Leaves to 2,5 mm long

6b Leaves (much) longer:

8. G. bredasdorpense

1,2: 66

Rubieae

7a Flowers solitary, subtended by whorls of bracts 12. G. amatymbicum

7b Flowers in groups of 3 or 2, mostly without bracts:

8a Flowers in groups of 3 9. G. subvillosum

8b Flowers in groups of 2:

9a Leaves (8— >10—14 mm long, linear-lanceolate to lanceolate; fruiting pedicels to over 10 mm

long 10. G. mucroniferum

9b Leaves 5-7 mm long, oblanceolate to spathulate; fruiting pedicels to 6 mm long 1 1 . G. rourkei

2b Inflorescences ± extensive and many-flowered:

10a Flowers cf, 9, plants dioecious, peduncles and pedicels villous 4. G. tomentosum

10b Flowers 9", peduncles and pedicels glabrous, scabrous, or with a few ± straight white hairs, but never villous:

11a Leaves obovate to obovate-oblong, (6— >8— 1 0(— 11) mm wide, often glaucous, always in whorls of 6;

flowers with very short filaments, anthers almost sessile 3. G. undulatum

lib Leaves linear, lanceolate or (narrowly) oblanceolate, (0,6-)0,8^t(-5) mm wide, never glaucous, in

whorls of 6-8-10; flowers with ± long filaments, stamens c. half as long as corolla lobes:

12a Leaves with densely set recurved prickles on margins; pedicels with ± long straight white hairs or

glabrous 6. G. scabrelloides

12b Leaf margins glabrous, with ± long straight white hairs or only very few small recurved prickles;

pedicels glabrous or scabrous:

13a Leaf surfaces densely covered with ± long white spreading hairs, blades 0,6— 0,8(— 1) mm wide;

stems densely covered with short spreading hairs; corolla ( 1 ,5—) 1 ,7— 2(— 2 ,3) mm in diam.

7. G. monticolum

13b Leaf surfaces smooth or with small upward-directed prickles or short straight hairs,

blades (0,3-)0,5-2(-2,5) mm wide; stems glabrous, with minute prickles and/or very short curled

whitish hairs; corolla 2—3 ,5(— 4> mm in diam.:

14a Leaf surfaces with small, upward-directed prickles or short straight hairs; pedicels scabrous

(or subglabrous) 6 (b). G. capense subsp. namaquense

14b Leaves glabrous or with a few small, mostly recurved prickles on margins only; pedicels

mostly glabrous:

15a Leaves up to 1 2(— 15) mm long; mericarps glabrous 6 (a). G. capense subsp. capense

15b Leaves up to 20(-23) mm long; mericarps ± densely covered with short curled whitish

hairs, occasionally (sub)glabrous 6 (c). C. capense subsp. garipense

MAP 61 . — Galium, all taxa

1. Galium thunbergianum Eckl. &

Zeyh., Eum. 369 (1836); Verde, in F.T.E.A.

Rubiaceae 1: 387 (1976); Puff in J1 S. Afr. Bot.

44: 221 (1978). Type: Cape, Katriviersberg,

Ecklon & Zeyher 2321 (S, holo. !; SAM, iso. !).

Perennial with extensive rhizome. Stems

climbing, suberect or procumbent, (80-)100-

400(-600) mm long, glabrous or with short

spreading white hairs. Leaves in whorls of 4, 3-

nerved, (7—) 1 0— 1 8(— 22) x (3,5-)4-8(-10) mm,

ovate, elliptic or ± rhombic, ± acute or with a

short hyaline point at apex, narrowed to base;

glabrous or both surfaces and margins with

short spreading hairs. Inflorescence broadly py-

ramidal to ± cylindrical, cymes ± many-flow-

ered, ultimate branches with 1-2 minute bracts;

pedicels 1— 3(— 5) mm long, slightly elongating

after anthesis, glabrous or hairy, ± divaricate in

fruit. Flowers-, corolla (1— )1 ,5— 2,5(— 3) mm in

diam., often somewhat hairy outside, greenish,

greenish white to yellowish, lobes longer than

wide, acute; stamens c. half as long as lobes;

ovary c. 0,3-0, 5 mm long. Fruit densely cov-

ered with white tuberculate hooked hairs; meri-

carps subglobose, each 1-1,5 mm in diam.

Chromosome number: 2n=22.

Occurs from the Transvaal to the eastern Cape; also in

afromontane areas of tropical Africa. Grows mostly in for-

est margin vegetation, scrub, or rocky grassland. Map 62.

Rubieae

1,2: 67

MAP 62. — • Galium thunbergianum var. thunber-

gianum

★ Galium thunbergianum var. hirsutum

® both varieties

The following two varieties are ± equally well repre-

sented in Southern Africa; they have similar distribution

ranges and are often found side by side in the field. For

diagnostic characters see the descriptions.

1 (a), var. thunbergianum.

G. rotundifolium sensu Sond. in F.C. 3: 39 (1865), non

L.

G. natalense Rouy in FI. Fr. 8: 9 (1903), ad not. Type:

Natal, Drakensberge, Van Reenen Pass, Medley Wood 5562

LY; K: photo.!).

G. rotundifolium L. var. normale Kuntze, Rev. Gen. 3:

120 (1898). Type: Natal, Van Reenen Pass, Kuntze s.n. (K,

holo.!).

Stems, leaves, peduncles and pedicels gla-

brous or nearly so (occasionally a few hairs at

some upper nodes or along nerves of leaves).

Vouchers: Galpin 6650 (BOL; GRA; K; NH; PRE;

SAM); 11890; Af cense 10331 (PRE;S).

1(b). var. hirsutum (Sond.) Verde, in

Kew Bull. 30: 326 (1975). Type: Cape, Masson

in herb. Thunberg (sheet 3354, UPS, holo.!).

G. rotundifolium L. var. hirsutum Sond. in F.C. 3: 39

(1865).

G. rotundifolium sensu Thunb., FI. Cap. 1: 551 (1813),

non L.

G. dasycarpum Schweinf. , Beitr. FI. Aethiop. 135

(1867). Type: Ethiopia, Simien, Debra-Eski, Schimper s.n.

(Bt).

G. biafrae Hiem in F.T.A. 3: 245 (1877). Syntypes:

Fernando Po, Mann 605 Cameroun Mtn, Mann 1284 (K!).

Stems (mainly angles), leaves, peduncles and

pedicels ± densely covered with short spread-

ing white hairs.

Vouchers: Hilliard & Burn 8036 (E; K; MO; NU;

PRE; S); Jacot Guillarmod 799; Schlechter 4387 (BOL;

PRE; S; SAM; W; WU).

2. Galium spurium L., Sp. PI. 106

(1753). Type: plant grown in Uppsala Botanic

Garden (no specimen in LINN).

subsp. africanum Verde, in Kew Bull. 30:

324 (1975), in F.T.E.A. Rubiaceae 1: 390

(1976); Puff in J1 S. Afr. Bot. 44: 271 (1978).

Type: Kenya, Kiambu distr. , Muguga, Milne-

Redhead & Taylor 7147 (K, holo.!; NU, WU:

photos!).

G. horridum sensu Eckl. & Zeyh., Enum. 370 (1836),

non Thunb.

G. aparine sensu Sond. in F.C. 3: 38 (1865), non L.

Annual. Stems weak, prostrate to semi-

erect, (0,12-)0,3-c. 2 m long, sparingly

branched, with recurved prickles on angles.

Leaves in whorls of 6-8, 1-nerved,

( 8—) 1 5—3 8 ( — 45 ) x ( 1 ,5— )3— 6(— 7) mm, linear-

lanceolate to obovate, with a distinct ± filiform

acumen at apex, narrowed to base; upper sur-

face glabrous or with a few scattered ± straight

or curled hairs, midrib (beneath) and margins

with coarse recurved prickles. Inflorescence ex-

tremely reduced, flowers single or occasionally

paired on peduncles/pedicels (8-)12-20(-35)

mm long when in fruit, ± arcuate, with re-

curved prickles or (seldom) glabrous; pedicels

occasionally separated from peduncle by a

small ± linear bract. Flowers : corolla

( 1— )1 ,2—1 ,8(— 2) mm in diam., greenish, green-

ish white or creamy white, lobes longer than

wide, pointed; stamens very short; ovary c.

0,5-0, 8 mm long. Fruit covered with white

non-tuberculate hooked hairs; mericarps ±

globose, each (1 ,8— )2,5 — 4 mm in diam. Chro-

mosome number. 2n=40.

Known from Transvaal, Natal, Lesotho and Cape; ex-

tending to tropical Africa. Grows along riverbanks, at forest

edges and in cultivated and disturbed ground. Map 63.

Vouchers: Acocks 21429; Hoener 1571; Huntley 265

(NU; PRE).

In the Cape, collections differing in having 2^t (sel-

dom to 6)- flowered partial inflorescences occur next to typi-

cal material of the indigenous G. spurium subsp. afri-

canum. It is suspected that they may be introduced plants of

G. spurium subsp. spurium and/or G. aparinefl). For rea-

sons explained in Puff in J1 S. Afr. Bot. 44: 272 (1978),

such collections can, at present, not be identified with cer-

tainty and are provisionally treated as “Galium spurium-

aparine complex” (Map 63); vouchers: Acocks 22593 (K;

PRE); Olivier 152(a).

1,2:68

Rubieae

MAP 63. — • Galium spurium subsp. africanum

A Galium spurium-aparine complex

@ both

3. Galium undulatum Puff in J1 S. Afr.

Bot. 44: 262 (1978). Type: Cape, Lichtenstein

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

G. uncinatum Licht. in Bartl. & Wendl., Beitr. Bot. 2:

12 (1825), nom. illeg. Type: as forG. undulatum Puff.

G. glabrum sensu Sond. in F.C. 3: 38 (1865), non

Thunb.

Perennial with slightly woody rootstock.

Stems ascending to erect, c. 0,3-1 ,5 m long,

usually with few branches, glabrous or with a

few recurved prickles on angles, often ± glau-

cous. Leaves in whorls of 6, 1 -nerved,

(15-)20-30 x (6— )8— 10 mm, obovate to obo-

vate-oblong, with a short hyaline point at apex,

narrowed to base; glabrous except for ± coarse

recurved prickles on margins, often glaucous.

Inflorescence ± cylindrical, cymes 3- to ±

many-flowered, ultimate branches usually

ebracteate; peduncles (5-) 10-22 mm long, gla-

brous, pedicels (3— )5— 1 0(— 20) mm long,

slightly elongating after anthesis, glabrous, fil-

iform, divaricate in fruit. Flowers : corolla

(2,5-)3^4 mm in diam., whitish, lobes much

longer than wide, acute; stamens very short,

much less than half as long as lobes; ovary c.

0,5 mm long. Fruit glabrous, granulate; meri-

carps ± globose, each (2-) 2, 5-3, 5

mm in diam.

Endemic to the south-western Cape (more or less along

Langeberg range from Swellendam eastwards); growing at

forest edges, in riverbank vegetation or on damp, ± shady

mountain slopes. Map 64.

MAP 64. — Galium undulatum

Vouchers: Ecklon & Zeyher 2325 (PRE; S; SAM);

Esterhuysen 10464 (BOL; K).

4. Galium tomentosum Thunb., FI. Cap.

1: 551 (1813); Sond. in F.C. 3: 39 (1865); Puff

in J1 S. Afr. Bot. 44: 263, fig. 21 (1978). Syn-

types: Cape, Thunberg (sheets 3373, 3374,

UPS!; NU, WU: photos!).

G. asperum Thunb., Prodr. 1: 30 (1794); Sond. in F.C.

3: 38 (1865), nom. illeg. Type: Cape, Thunberg (sheet

3302, UPS, holo.!; NU, WU: photos!).

G. glabrum Thunb., FI. Cap. 1: 551 (1813). Type: Cape,

Thunberg (sheet 3313, UPS, holo.!; K, NU, WU: photos').

G. asperum var. villosum Eckl. & Zeyh., Enum. 369

(1836). Type: Cape, Clanwilliam distr. , near Brakfontein,

Ecklon & Zeyher 2323(3 (GOET; P; S, iso.!; WU: photo.!).

G. namaquense Schltr., nom. nud.

Rubia lanata Dinter, nom. nud.

Dioecious perennial with woody root-

stock. Stems climbing or scrambling, c.

(0,3-)0,5-2,5(-3) m long, much branched,

with recurved prickles on angles or (rarely)

subglabrous, upper parts of stems frequently

densely villous, often purplish. Leaves in

whorls of 6-8, 1 -nerved, (12-)15-25(-30) x

(2— )3— 8(— 10) mm, narrowly to broadly obo-

vate, broadly ovate or ± lanceolate, often acu-

minate or acute at apex, narrowed to base;

slightly reflexed margins and midrib beneath

with densely set recurved prickles. Inflores-

cence extensive, ± broadly pyramidal, cymes

many-flowered, of rather different appearance

in o and ^ plants. O': peduncles ( 1 — )2— 3 mm

long, ± thickish, subglabrous to villous, pedi-

cels 1-2 mm long, ± filiform, hairy, strongly

divaricate; peduncles ( 1 0—) 1 5 — 30( — 35 ) mm

long, somewhat elongating after anthesis, vil-

lous, pedicels (20-)25-60(-90) mm long after

Rubieae

1 ,2: 69

anthesis, ± thickish, villous, convergent.

Flowers : cf: corolla (2-)2,5-3,5(-4) mm in

diam., occasionally somewhat hairy outside,

greenish, greenish yellow to pale yellow, lobes

much longer than wide, acute; stamens very

short; rudimentary ovary present, disk small.

9: corolla 3-4 mm in diam., ovary c. 0,5-1

mm long, disk larger than in cf; rudimentary

stamens usually present. Fruit granulate, gla-

brous or sometimes with a few whitish hairs;

mericarps ± reniform, mostly with 3 grooves

on dorsal side, each (2-)2,5-3 x ( 1—) 1 ,5—2

mm. Chromosome number. 2n=44.

Occurs from the southern Namib to the south-western

and eastern Cape; also in the Little, Great and Upper Kar-

roo. Grows mostly in scrub on slopes, riverbanks or coastal

dunes. Map 65.

Vouchers: Acocks 14988 (cT); Dinter 8106 ($; B;

BOL; K; PRE; WIND); Olivier 152 (O'); Rourke 704 ( 9;

NBG; PRE; S).

Widely distributed and variable (leaf size and shape!).

Cf plants have less densely villous and less conspicuous

inflorescences than §; cf forms with less hairy inflores-

cence branches should not be confused with G. undulatum

(above).

5. Galium scabrelloides Puff in J1 S. Afr.

Bot. 44: 250 (1978). Type: Natal, Cathedral

Peak Forest Reserve, Puff 760314-1/2 (WU

holo.!).

Perennial with somewhat woody root-

stock. Stems climbing or decumbent, (0,3-)

0,5—1 ,0(— 1 ,2) m long, often much branched,

mostly with white spreading hairs at least on

angles. Leaves in whorls of (7— )8— 10, 1-

nerved, ( 1 2—) 1 5— 20(— 22) x (0,6-)0,8-4(-5)

mm, linear to linear-lanceolate or oblanceolate,

with a whitish mucro at apex, narrowed to base;

both surfaces, lower surface or at least midrib

below with white ± straight spreading hairs,

(strongly) reflexed margins with closely set

coarse recurved prickles. Inflorescence broadly

to narrowly pyramidal, cymes many-flowered,

ultimate branches with 3-1 tiny linear to lan-

ceolate bracts; peduncles 2-3 mm long, pedi-

cels 1— 2,5(— 3) mm long, ± filiform, hairy,

strongly divaricate in fruit. Flowers', corolla

1 ,8—3 ,5(— 4) mm in diam., usually somewhat

hairy outside, bright yellow, creamy yellow,

greenish yellow or greenish, lobes much longer

than wide, acute; stamens a little less than half

as long as lobes; ovary c. 0,4-0, 6 mm long.

Fruit (densely) covered with straight white

spreading hairs, rarely subglabrous; mericarps

subglobose to slightly reniform, each

(0, V— )0,8— 1 (— 1 ,2) mm in diam.; often only one

mericarp developed. Chromosome number:

2n=22.

Occurs from Transvaal and Swaziland to the eastern

Cape; growing mostly at forest edges or in scrub. Map 66.

MAP 66. — Galium scabrelloides

Vouchers: Flanagan 2858 (BOL; PRE); Schmitz 7481;

Trauseld 538 (NU; PRE).

Shows considerable variability in growth form, leaf

size and shape; narrow- leaved, weakish plants and atypical,

± glabrescent forms should not be confused with G. ca-

pense subsp. garipense (below). In the eastern Cape the two

taxa are occasionally difficult to distinguish.

6. Galium capense Thunb., Prodr. 1: 30

(1794); Sond. in F.C. 3: 36 (1865); Puff in J1 S.

Afr. Bot. 44: 228 (1978). Type: Cape, num-

bers (sheet 3308, UPS, holo.!; NU, WU: pho-

tos!).

1,2: 70

Rubieae

Perennial with sometimes slightly woody

rootstock. Stems scrambling, sprawling or

climbing, or ascending to erect, (80-) 150-900

mm long, sparingly to much branched, gla-

brous, with a few minute recurved prickles or

with short curled whitish hairs. Leaves in

whorls of 6— 8(— 10), 1-nerved, (3— )5— 20(— 23)

X (0,3-)0,5-2(-2,5) mm, linear to narrowly

lanceolate or oblanceolate, sometimes with a

brownish white mucro at apex; surface gla-

brous, with small upward-directed prickles or

with short straight hairs, margins often re-

flexed, glabrous or with a few recurved

prickles. Inflorescence broadly pyramidal to ±

cylindrical, cymes many- to several-flowered,

ultimate branches with (4— )3-l small linear

bracts or ebracteate; peduncles and pedicels

(1 — )1 ,5 — 4(— 5) mm long, ± filiform, glabrous

or scabrous, very rarely hairy, divaricate in

fruit. Flowers : corolla 2-4 mm in diam. , mostly

(bright) yellow, creamy yellow or greenish yel-

low, lobes (much) longer than wide, acute or

sometimes slightly acuminate; stamens c. 1/4 to

3/4 of lobe length; ovary c. 0,3-0, 6 mm long.

Fruit glabrous, ± tuberculate or granulate, or

covered with short straight or curled whitish

hairs; mericarps subglobose, each c. 1-2 mm in

diam. Chromosome number. 2n=22.

G. capense is subdivided into 3 subspecies; the separa-

tion of the subspecies may become difficult where their

ranges overlap; each of the subspecies is morphologically

very variable.

G. capense and G. monticolum (below) form a closely

allied pair.

6 (a), subsp. capense.

G. mucronatum Thunb., Prodr. 1: 30 (1794). Type:

Cape, Thunberg (sheet 3342, UPS, holo.!; NU, WU: pho-

tos!).

G. expansum Thunb., Prodr. 1: 30 (1794). Type: Cape,

Thunberg (sheet 3312, UPS, holo.!; NU, WU: photos!).

G. capense var. expansum (Thunb.) Sond. in F.C. 3: 36

(1865).

G. expansum var. elongatum Eckl. & Zeyh., Enum. 370

(1836). Type: Cape, between Hassaquaskloof and Breederi-

vier, Ecklon & Zeyher 2326(3 (BOL; SAM, iso.!; WU:

photo.!).

G. mucronatum var. densiflorum Eckl. & Zeyh., Enum.

370 (1836). Type: Cape, Vierentwintig Rivieren, Ecklon &

Zeyher 2327p (GOET; P; SAM; W, iso. !; WU: photo. !).

G. capense var. minus Sond. in F.C. 3: 36 (1865). Syn-

types: Cape, Thunberg (sheet 3342, UPS!; NU, WU: pho-

tos!); Wolvekop, Zeyher 773 (S!).

ered with short curled whitish hairs, often

purplish. Leaves in whorls of (6— )8(— 10),

(3— )5— 12(— 15) X 0,5-l,5(-2) mm, linear to

narrowly lanceolate or oblanceolate; surfaces

often shiny, glabrous, occasionally a few small

mostly recurved prickles on reflexed margins.

Peduncles and pedicels glabrous. Flowers : co-

rolla (2-)2,5^l mm in diam., bright to pale

yellow. Fruit glabrous, ± tuberculate or granu-

late; each mericarp c. 1 mm in diam. Fig. 14:

1-5.

Centred in the south-western Cape but also extending

into karroid areas and to the eastern Cape; growing mostly

in rocky, ± damp areas or near water courses. Map67.

MAP 67. — A Galium capense subsp. capense

■ Galium capense subsp. namaquense

• Galium capense subsp. garipense

B subsp. capense and namaquense

@ subsp. capense and garipense

Vouchers: Drege 7685 (E; K; P; W); Esterhuysen 9187

(BOL); Thompson 2179 (PRE; STE).

Growth form and habit are very variable (plants weak

to quite robust, inflorescences extensive and many-flowered

to rather few-flowered; this is probably to a considerable

extent environment-dependent). See also subsp. garipense

(subsp. c).

6(b). subsp. namaquense (Eckl. & Zeyh.)

Puff in J1 S. Afr. Bot. 44: 234 (1978). Type:

Cape, near Heerelogement, Ecklon & Zeyher

23^2 (S; SAM, iso. !; WU: photo. !).

G. namaquense Eckl. & Zeyh., Enum. 369(1836).

G. capense var. scabrum Sond. in F.C. 3: 37 (1865).

Syntypes: Cape, Modderfonteinsberg, Drege 7682 (E!; K! ;

P!; S ! ; WU: photo.!); between Pedroskloof and ‘Lilly Fon-

tein’ (Leliefontein), Drege 7683 (K!; P!; WU: photo.!).

Stems scrambling or climbing, (80-)150-

500(-700) mm long, sparingly to much

branched, at least younger parts densely cov-

Stems scrambling or sprawling, ( 1 50—)

300-900 mm long, usually much branched,

densely covered with very snort, mostly curled

RUBIEAE

1,2:71

PIG i4 Galium capense: 1-5, subsp. capense: 1, part of plant, x 1; 2, enlargement of stem, x 5; 3, leaf with

strongly revoiute margins, side view, X 7; 4, corolla with stamens, X 15; 5, gynoecium, x 15 (Thompson 2179). 6, subsp.

namaquense, leaf (from above), x 7 (Taylor 7485). 7 & 8, subsp. garipense: 7, leaf (from below), x 7; 8, fruit, x 10

(Hanekom 831).

1,2: 72

Rubieae

whitish hairs, often purplish. Leaves in whorls

of 6-8, (4— )6-8(-10) x (0,5— )1— 2(— 2,5) mm,

linear to narrowly lanceolate or oblanceolate;

upper and (sometimes) lower surface and mar-

gins (densely) covered with small, upward-di-

rected prickles or short straight hairs, margins

often reflexed. Peduncles and pedicels sca-

brous, more rarely subglabrous. Flowers : co-

rolla (2-)2,5-3,5 mm in diam., yellow or

greenish yellow. Fruit covered with short

straight whitish hairs or glabrous, ± tuberculate

or granulate; each mericarp 1-2 mm in diam.

Fig. 14: 6.

Endemic to the western Cape; growing mostly in river-

bank vegetation, in renosterveld, arid fynbos or scrub. Map

67.

Vouchers: Acocks 17374 (K; PRE); Taylor 7485 (PRE;

STE).

Distinguished from subsp. capense primarily by its

indumentum. The two subspecies are almost completely

allopatric.

6 (c). subsp. garipense (Sond.) Puffin J1

S. Afr. Bot. 44: 234 (1978). Syntypes: Cape,

on the Garip near Buffelvallei, Drege 767 4 (K!;

PRE!; S!; W!); Natal, Buffaloerivier, Gerr. &

M’K. (as ‘Port Natal’, Gerrard & McKen 1340

in S ! ; as Natal & Zululand, Gerrard 1340 in

K!).

G. garipense Sond. inF.C. 3: 37 (1865).

G. mucronatum var. subglabrum Eckl. & Zeyh., Enum.

370 (1836). Type: Cape, ‘Tambukiland’, on the right side

of the Keyrivier, Ecklon & Zeyher Tilly (GOET; P; PRE;

SAM;W, iso. !;WU: photo.!).

G. wittbergense Sond. in F.C. 3: 37 (1865). Type: Cape,

rocky wet places in the Wittbergen, Drege s.n. (in herba-

rium Sonder; specimen not located).

G. wittbergense var. glabrum Phill. in Ann. S. Afr. Mus.

16: 113 (1917). Syntypes: Lesotho, Leribe, Phillips 713

(K!; SAM!), Dieterlen 40 p.p. (SAM!).

G. capense subsp. garipense var. wittbergense (Sond.)

Puff in J1 S. Afr. Bot. 44: 237 (1978).

Stems usually ascending to erect,

(100-)200-500(-600) mm long, usually with ±

few lateral branches, glabrous, with a few min-

ute recurved prickles on angles and/or with ±

short whitish hairs. Leaves in whorls of 6-8,

(6-)10-20(-23) x (0,3— )0,5— 1 ( 1 ,5) mm, lin-

ear, glabrous, sometimes a few small, mostly

recurved prickles on often ± reflexed margins.

Peduncles and pedicels glabrous, very rarely

hairy. Flowers: corolla 2-3,5(^f) mm in diam.,

bright yellow, sometimes creamy yellow or

greenish yellow. Fruit ± densely covered with

short curled whitish hairs or occasionally

(sub)glabrous; each mericarp 1-1,8 mm in

diam. Fig. 14: 7 & 8.

Widely distributed from the Transvaal to the eastern

Cape; growing mostly along stream banks, in vleis, on ±

damp slopes or in open grassland. Map 67.

Vouchers: Devenish 440; 1238 (K; NH); Flanagan

1351 (BOL; NBG; NU; PRE; SAM); Hilliard & Burn 8279

(E; K; MO; NU; PRE); Jacobsz 1030; 2076 (NBG).

Var. wittbergense is no longer retained; subsp. gari-

pense is too variable to allow a certain distinction of varie-

ties. The separation of subsp. garipense and subsp. capense

can become rather troublesome in the eastern Cape, where

their distribution ranges overlap. Subsp. garipense tends to

be more robust and to have longer leaves.

Forms with ± needle-like leaves and a peculiar ‘dense’

habit (e.g. Moll 5157, NH; PRE), previously believed to be

confined to western Natal and thought to possibly represent

a separate taxon [see Puff in J1 S. Afr. Bot. 44: 238 (1978)],

also occasionally occur elsewhere and are perhaps best in-

cluded in subsp. garipense.

7. Galium monticolum Sond. in F.C. 3: 36

(1865); Puff in J1 S. Afr. Bot. 44: 243 (1978).

Type: Cape, mountains near Cape Town, Eck-

lon 84 (S, holo.!; WU: photo.!).

Perennial with ± woody rootstock. Stems

ascending to erect, c. 0,3-1 ,2 m long, usually

much branched, densely covered with short

spreading hairs, sometimes purplish. Leaves in

whorls of (6-)8-10, 1-nervea, 5— 8(— 1 2) x

0,6-0,8(-l) mm, linear, with a long whitish,

sometimes upturned mucro at apex, often appa-

rently terete due to strongly re flexed margins;

upper surface, midrib beneath and margins

densely covered with ± long white spreading

hairs. Inflorescence ± cylindrical, cymes seve-

ral- to ± few-flowered, ultimate branches with

(5-)3-l(-0) minute linear, hairy bracts; pedun-

cles ± thickish, hairy, pedicels filiform, gla-

brous, both 0,8-1 ,5(-2) mm long, divaricate

MAP 68. — • Galium monticolum

it Galium bredasdorpense

Rubieae

1,2: 73

after anthesis. Flowers : corolla ( 1 ,5— )1 ,7—2

(-2,3) mm in diam., colour unknown, lobes

longer than wide, acuminate; stamens ca. 1/4 of

lobe length; ovary c. 0,3-0, 5 mm long, gla-

brous, ± granulate. Mature fruit not seen.

Endemic to the (south- )westem Cape and apparently

very rare; growing on mountain slopes. Map 68.

Voucher: Pearson 5177 (BOL; K; MO; PRE).

Very closely allied to G. capense (no. 6) and perhaps

not specifically distinct; too few collections are known to

allow certainty about its status.

8. Galium bredasdorpense Puff in J1 S.

Afr. Bot. 44: 244 (1978). Type: Cape, Bredas-

dorp distr., 2 m S of Wydgelegen Post Office,

Acocks 23175 (PRE, holo.!; WU: photo.!).

Perennial. Stems decumbent or suberect,

100-250 mm long, much branched, at least

younger parts densely scabrous. Leaves in

whorls of 6, 1-nerved; 1,5-2, 5 x 0,7-1 ,2 mm,

± ovate-lanceolate, with a ± long whitish

mucro at apex, glabrous, shiny, margins ±

reflexed. Inflorescence very reduced, flowers in

groups of 3-1, subtended by 4(— 1) leaf-like

bracts; peduncles and pedicels 0,5-2 mm long,

filiform, glabrous, divaricate after anthesis.

Flowers', corolla 1,5-2 mm in diam., yellow-

ish, lobes much longer than wide, shortly acu-

minate; stamens c. half as long as lobes; ovary

c. 0, 2-0,4 mm long. Fruit glabrous, ±

wrinkled; mericarps subglobose, each c. 1 mm

in diam. Chromosome number: 2n = 22.

Endemic to the Bredasdorp district (south-western

Cape); confined to limestone areas. Map 68.

Vouchers: Burgers 1494; 1595.

Distinguished from all other Galium species occurring

in the south-western Cape by its closely spaced, small

leaves.

9. Galium subvillosum Sond. in F.C. 3: 38

(1865); Puff in J1 S. Afr. Bot. 44: 255 (1978).

Type: Cape, Du Toitskloof, Drege 7687 (S; W,

iso. !; NU, WU: photos!).

Perennial with ± woody rootstock. Stems

prostrate, ascending or scrambling, 150-500

(-600) mm long, ± much branched, covered

with short spreading white hairs, subglabrous or

glabrous, often purplish. Leaves in whorls of 6,

1-nerved, (5-)6-8(-ll) x (l,5-)2-3,5(-4,5)

mm, oblanceolate or ± lanceolate, with a dis-

tinct hyaline point at apex; both surfaces and

slightly reflexed margins with spreading white

hairs, subglabrous or glabrous. Inflorescence ±

cylindrical, very reduced, flowers in groups of

3, subtended by (0-)l-2 ± small lanceolate

bracts; peduncles (4 — )5— 10 mm long, thickish,

hairy or glabrous, pedicels (3-)4— 6(-8) mm

long, slightly enlongating after anthesis, thick-

ish to ± filiform, hairy or glabrous, divaricate

in fruit. Flowers : corolla 3-4 mm in diam.,

usually a little hairy outside, whitish yellow,

creamy yellow or purplish, lobes longer than

wide, acute; stamens nearly as long as lobes;

ovary c. 0,3-0, 5 mm long. Fruit wrinkled, cov-

ered with short straight hairs or glabrous; meri-

caips subglobose to ± reniform, each

2-2,5(-3) mm in diam., often only one meri-

carp developed. Chromosome number. 2n =

44.

Endemic to the mountains of the south-western Cape;

growing mostly in damp to moist, ± shady places, e.g. at

the base of cliffs, along watercourses, etc. Map 69.

MAP 69. — • Galium subvillosum var. subvillosum

★ Galium subvillosum var. subglabrum

® both varieties

Closely allied to G. mucroniferum (below) but easily

distinguished by its 3-flowered cymes.

Two varieties are recognized. For diagnostic charac-

ters see the descriptions.

9 (a), var. subvillosum.

Stems, foliage, peduncles, pedicels

(densely) covered with snort spreading white

hairs; occasionally also fruits hairy.

Vouchers: Esterhuysen 11331 (BOL; K; NBG; PRE);

Schlechter 9 1 84 (BOL; E: GRA; K; P: PRE; S; W).

More common and more widely distributed than var.

subglabrum (below).

9(b). var. subglabrum Puff in J1 S. Afr.

Bot. 44: 257 (1978). Type: Cape, Bains Kloof,

Compton 18632 (NBG, holo.!).

1,2:74

Rubieae

Stems, foliage, peduncles and pedicels

glabrous or nearly so.

Vouchers: Esterhuysen 1721 (BOL); 3512 (BOL).

10. Galium mucroniferum Sond. in F.C.

3: 37 (1865); Puff in J1 S. Afr. Bot. 44: 259

(1978). Syntypes: Cape, Du Toitskloof, Drege

7677 (E!; K!; WU: photo.!); near Genadenthal,

Drege 7686 (E!; K!; P!; PRE!; S!; NU, WU:

photos!); no localities given: Drege 7684, 7689

(S!); in ‘Caffraria’ , Ecklon & Zeyher s.n. (S!).

Perennial with ± woody rootstock. Stems

scrambling, ascending or erect, 100-550 mm

long, usually much branched, glabrous, with

few recurved prickles or with white straight

spreading hairs on angles. Leaves in whorls of

6(— 8), 1 -nerved, (8-)10-14 x (l-)l,3-2(-2,5)

mm, linear-lanceolate to lanceolate, with a long

hyaline point at apex, glabrous, slightly re-

flexed margins with a few recurved prickles or

upper surface, midrib below and margins with

white straight spreading hairs, often very shiny.

Inflorescene often narrowly cylindrical, very

reduced; flowers in groups of 2, subtended by

(3— )2— 1 (— 0) minute, ± linear bracts, peduncles

5— 1 2(— 1 5) mm long, filiform or ± thickish,

glabrous or hairy, pedicels (4 — )6— 1 0(— 13) mm

long, elongating after anthesis, ± filiform, gla-

brous or hairy, strongly divaricate in fruit.

Flowers: corolla (2,5-)3-4 mm in diam., some-

times a little hairy outside, greenish yellow,

creamy yellow or pale green, lobes much longer

than wide, acute; stamens nearly as long as

MAP 70. — • Galium mucroniferum var. mucronife-

rum

★ Galium mucroniferum var. dregeanum

(£) both varieties

lobes; ovary c. 0,5-0, 8 mm long. Fruit gla-

brous, ± granulate or wrinkled or covered with

short straight hairs; mericarps ± globose, each

(2-)2,5-3 mm in diam.; often only one meri-

carp developed. Chromosome number: 2n=44.

Endemic to the mountains of the south-western Cape;

growing mostly on rocky slopes or in ravines and other ±

cool, sheltered and shady places. Map 70.

Two varieties are recognized. For diagnostic charac-

ters see the descriptions.

10(a). var. mucroniferum.

Stems and leaves glabrous or with a few

recurved prickles on angles and margins (but

never with white spreading hairs); peduncles

and pedicles glabrous; fruits occasionally with a

few hairs.

Vouchers: Parker 4473 (BOL; K; NBG); Schlechter

9252 (BOL; E; K; P; PRE;S).

10 (b). var. dregeanum (Sond.) Puff in J1

S. Afr. Bot. 44: 260 (1978). Type: Cape, Du

Toitskloof, Drege 7688 (E; K; P; S, iso.!; NU,

WU: photos!).

G. dregeanum Sond. in F.C. 3: 38 (1865).

Stems (mainly angles), leaves (margins,

upper surface and midrib below), peduncles,

pedicels and fruits covered with white straight

spreading hairs.

Voucher: Duthie 577 (BOL).

Much rarer than var. mucroniferum and only known

from a few localities.

1 1 . Galium rourkei Puffin J1 S. Afr. Bot.

49: 185 (1983). Type: Cape, Kogelberg Forest

Reserve, Kuduberg, Rourke 1765 (NBG,

holo.!; PRE!; WU, iso.!).

Perennial. Stems delicate, ± prostrate, c.

100-200 mm long, ± much branched, at least

younger parts with curved or straight white

hairs on angles. Leaves in whorls of 4— 5(-6), 1-

nerved, 5-7 x (2-)2,5-5 mm, oblanceolate to

spathulate, with a distinct rigid point at apex, to

1 mm long; with ± long straight or curved hairs

on upper and lower surface and margins; mar-

gins straight. Inflorescence very reduced; flow-

ers in groups of 2; peduncles 1 ,5-5 mm long,

hairy; pedicels filiform, hairy, 1,5-3 mm long

at anthesis, to 6 mm long and divaricate in fruit.

Flowers: corolla (2,5-)3-3,5(^4) mm in diam.,

creamy white, lobes longer than wide, acute,

with a few white straight or curved hairs on

margins; stamens c. 1-1,5 mm long; ovary c.

Rubieae

1,2: 75

0,3-0, 4 mm long. Fruit covered with white

spreading hairs; mericarps ± globose, each c.

1,5-2 mm in diam., often only one mericarp

developed.

Endemic to the south-western Cape and so far only

known from the type locality; growing in permanent shade

under damp rock cliffs. Map 7 1 .

Allied to the two preceding species; easily distin-

guished by the shorter and broader leaves.

MAP 7 1 . — ▲ Galium rourkei

• Galium amatymbicum

12. Galium amatymbicum Eckl. &

Zeyh., Enum. 370 (1836); Puff in J1 S. Afr.

Bot. 44: 245 (1978). Type: Cape, ‘Tambuki-

land’, on the right side of the ‘Keyrivier’, Eck-

lon & Zeyher 2328 (GOET; S; SAM, iso.!;

WU: photo.!).

Perennial with ± extensive rhizomes.

Stems weak and thin, caespitose, (50-)70-200

(-300) mm long, with many, often short lateral

branches, covered with ± long white spreading

hairs or glabrous. Leaves in whorls of 4—6, 1-

nerved, 3-5(-6) x (0,7-)0,8-l mm, linear-lan-

ceolate to lanceolate, ± acuminate at apex,

with ± long white spreading hairs on upper and

lower surface or at least on margins, occa-

sionally subglabrous. Inflorescence extremely

reduced; flowers solitary, subtended by a whorl

of bracts, axillary or terminal on short lateral

branches; pedicels (0,5-)l(-2) mm long, ±

thickish, hairy or glabrous, ± arcuate in fruit.

Flowers : corolla c. 1-1,5 mm in diam., hairy

outside, whitish, lobes longer than wide, acute;

stamens very short. Fruit covered with long ±

straight hairs or subglabrous and wrinkled;

mericarps reniform, each c. 1-1 ,5 mm long.

Endemic to the eastern Cape and apparently quite rare;

growing in thorn veld. Map 7 1 .

Vouchers: Acocks 23930; Rattray 1306.

Easily recognizable by its solitary flowers and small,

reniform mericarps.

1,2:76

INDEX

1 ,2: 77

Ambraria Cruse 1 ,2: 37

Ambraria Heister ex Fabr 1,2:8

acerosa Sond 1,2: 43,46

glabra Cruse 1,2:46

var. papillata Sond 1 ,2: 46

var. tulbaghica Sond 1 ,2: 46

heisteri Willd 1,2:14

hirta Cruse 1 ,2: 44

var. macrocarpa Eckl. & Zeyh 1 ,2: 46

microphylla Sond 1 ,2: 38

ANTHOSPER.V1EAE 1,2:7

anthospermum L 1,2:8

aethiopicumL 1,2:14

var. alpinum Eckl. & Zeyh 1,2: 16

var. ciliare (L. ) Kuntze 1 ,2: 29

var. ecklonianum Cruse 1,2:18

var. montanum Sond 1,2:16

var. oppositifoliam Cruse 1,2:16

var. reflexifolium Kuntze 1 ,2: 30

var. ternifolium Cruse 1,2: 14

var. tulbaghense Eckl. & Zeyh 1,2: 19

var. uitenhagense Eckl. & Zeyh 1,2: 17

ambiguum Greves 1,2:14

ammannioides sensu auctt. Afr. austr., non S.

Moore 1,2: 12

arenicolum Greves 1 ,2: 22

aromaticum Salisb 1,2:14

basuticum Puff, sp. nov 1 ,2: 20

bergianum Cruse 1 ,2: 35

bicorne Puff, sp. nov 1,2: 36

burkei Sond 1,2:21

calycophyllum Sond 1,2:36

ciliare L 1,2:28

var. angustifolium Eckl. & Zeyh 1 ,2: 29

var. glabrifolium Sond 1,2: 29

var. latifolium Eckl. & Zeyh 1 ,2: 29

var. papillatum Sond 1 ,2: 29

var. scabrum Eckl. & Zeyh 1,2: 29,30

cliffortioides K. Schum 1,2:12

comptonii Puff, sp. nov 1 ,2: 32

confertum Cruse 1,2:21

crocyllis Sond 1,2:5

dregei Sond 1,2:31

subsp. dregei 1,2:31

subsp. ecklonis (Sond.) Puff, comb, et stat.

nov 1,2:32

ecklonis Sond 1,2:32

ericitolium ( Lichi . ex Roem. & Schult.) Kuntze 1,2: 35

ericoideum K. Krause 1,2: 25

esterhuysenianum Puff, sp. nov 1,2: 33

var. esterhuysenianum 1,2:33

var. hirsutum Puff, var. nov 1 ,2: 33

ferrugineum Eckl. & Zeyh 1 ,2: 22

foetidum Eckl 1 ,2: 53

frutescens Dinter 1,2:25

galioides Reichb.f. 1,2:28

subsp. galioides 1,2:29

subsp. reflexifolium (Kuntze) Puff, comb.

et stat. nov 1,2:30

galopina Thunb 1 ,2: 48

galpinii Schltr 1,2: 13

hedyotideum Sond 1,2:22

herbaceum L.f. 1 ,2: 22

hirsutum DC 1,2:34

hirtum Cruse 1 ,2: 34

hispidulum E. Mey. ex Sond 1,2:21

humile N.E. Br 1,2:25

lanceolatum Thunb 1 ,2: 22

var. hedyotideum (Sond.) Kuntze 1,2:23

var. latifolium Sond 1,2: 23

latifolium E. Mey 1,2: 23

lichtensteinii Cruse 1 ,2: 35

littoreum L. Bol 1,2: 14

mildbraedii K. Krause 1,2:23

monticolaPuff, sp. nov 1,2: 19

muriculatum Hochst. ex A. Rich 1 ,2: 22

nodosum E. Mey 1,2: 23

paniculatum Cruse 1,2:21

var. confertum Eckl. & Zeyh 1,2: 21

var. elongatum Eckl. & Zeyh 1 .2: 21

prostratum Sond 1 ,2: 30

var. glabrum Sond 1,2: 30

var. velutinum Sond 1,2: 30

pumilum Sond 1,2:25

subsp. pumilum 1,2:25

subsp. rigidum (Eckl. c£ Zeyh.) Puff, comb.

et stat. nov 1,2:26

var. pilosum Phill 1 ,2: 25

rigidum Eckl. & Zeyh 1 ,2: 26

rigidum sensu auctt. Afr. austr., non Eckl. &

Zeyh 1,2:25

rubiaceum Reichenb. f 1,2: 34

rubricaule K. Schum 1 ,2: 22

scabrum Thunb 1 ,2: 57

spathulatum Spreng 1,2:16

subsp. ecklonianum (Cruse) Puff, comb, et

stat. nov 1,2: 18

subsp. saxatile Puff, subsp. nov 1,2: 18

subsp. spathulatum 1,2:16

subsp. tulbaghense Puff, subsp. nov 1,2: 19

* An asterisk signifies exotic species which are not naturalized; synonyms are in italics.

1,2:78

subsp. uitenhagense Puff, subsp. nov 1,2: 17

var. ecklonianum (Cruse) Cruse 1,2:18

spermacoceum Reichb. f 1 ,2: 53

spicatum Suesseng 1,2: 25

streyiPuff, sp. nov 1,2:28

thymifolium Dinter & Krause 1 ,2: 3 1

tricostatum Sond 1,2: 16

welwitschii Hiern 1,2: 12

whyteanum Britten* 1,2:22

CARPACOCE Sond 1,2:52

burchellii Puff, sp. nov 1,2:59

curvifolia Puff, sp. nov 1,2: 57

gigantea Puff, sp. nov 1 ,2: 56

heteromorpha (Buek)L. Bol 1 ,2: 60

scabra (Thunb.) Sond 1,2: 57

subsp. scabra 1,2:57

subsp. rupestris Puff, subsp. nov 1,2: 59

spermacocea (Reichb. f.) Sond 1,2:53

subsp. orientalis Puff, subsp. nov 1 ,2: 56

subsp. spermacocea 1,2:53

vaginellata Salter 1,2:59

Cliffortia acerosa MS 1 ,2: 46

cinerea Thunb 1 ,2: 40

spicata Reichb. f 1 ,2: 35

CROCYLLIS E.Mey.exHook.f. 1,2:3

anthospermoides E. Mey. ex K. Schum 1 ,2: 5

intricatissima Dinter 1,2:5

Galium L 1,2:65

amatymbicum Eckl. & Zeyh 1 ,2: 75

aparine sensu Sond., non L 1,2:67

asperum Thunb 1 ,2: 68

var. villosum Eckl. & Zeyh 1 ,2: 68

biafrae Hiern 1,2:67

bredasdorpense Puff 1,2:73

capense Thunb 1 ,2: 69

subsp. capense 1,2:70

subsp. garipense (Sond.) Puff 1,2:72

var. wittbergense (Sond.) Puff 1,2: 72

subsp. namaquense (Eckl. & Zeyh.) Puff .... 1,2:70

var. expansum (Thunb.) Sond 1,2: 70

var. minus Sond 1,2: 70

var. scabrum Sond 1 ,2: 70

dasycarpum Schweinf 1 ,2: 67

dregeanum Sond 1 ,2: 74

expansum Thunb 1 ,2: 70

var. elongatum Eckl. & Zeyh 1 ,2: 70

garipense Sond 1 ,2: 72

glabrum Thunb 1 ,2: 68

glabrum sensu Sond. , non Thunb 1 ,2: 68

horridum Thunb 1 ,2: 65

horridum sensu Eckl. & Zeyh. , non Thunb. ... 1,2:67

monticolum Sond 1 ,2: 72

mucronatum Thunb 1,2:70

var. densiflorum Eckl. & Zeyh 1,2:70

var. subglabrum Eckl. & Zeyh 1 ,2: 72

mucroniferum Sond 1 ,2: 74

var. dregeanum (Sond.) Puff 1,2:74

var. mucroniferum 1,2:74

namaquense Eckl. & Zeyh 1 ,2: 70

namaquense Schltr 1 ,2: 68

natalense Rouy 1 ,2: 67

rotundifolium L. var. hirsutum Sond 1 ,2: 67

var. normale Kuntze 1 ,2: 67

rotundifolium sensu Sond. , non L 1 ,2: 67

rotundifolium sensu Thunb. , non L 1 ,2: 67

rourkei Puff 1 ,2: 74

scabrelloides Puff 1 ,2: 69

spurium-aparine complex 1,2:67

spuriumL 1,2:67

subsp. africanum Verde 1 ,2: 67

subvillosum Sond 1 ,2: 73

var. subglabrum Puff 1 ,2: 73

var. subvillosum 1,2:73

thunbergianum Eckl. & Zeyh 1 ,2: 66

var. hirsutum (Sond.) Verde 1 ,2: 67

var. thunbergianum 1,2:67

tomentosum Thunb 1 ,2: 68

uncinatum Licht 1 ,2: 68

undulatum Puff 1 ,2: 68

verumL.* 1,2:65

wittbergense Sond 1,2: 72

var. glabrum Phill 1,2:78

Galopina Thunb 1 ,2: 48

aspera (Eckl. & Zeyh.) Walp 1,2: 51

aspera (Eckl. & Zeyh.) Sond 1,2:51

circaeoides Thunb 1,2:48

var. glabra Kuntze 1 ,2: 48

var. pubescens Kuntze 1 ,2: 48

crocyllioides Bar ex Schinz 1,2:52

hirsuta E. Mey 1,2:50

oxyspermum Steud 1,2:51

tomentosa Hochst 1,2: 50

Hondbessen Adans 1,2:1

Lagotis E. Mey 1,2: 52

spermacocea (Reichb. f.) E. Mey 1,2: 53

Lecontea A. Rich 1,2: 1

bojerana A. Rich 1,2: 3

Lygodisodea Ruiz & Pav 1,2: 1

Merciera heteromorpha Buek 1,2:60

NENAX Gaertn 1,2:37

acerosa sensu Eckl. & Zeyh 1 ,2: 43

acerosa Gaertn 1 ,2: 46

subsp. acerosa 1,2:46

subsp. macrocarpa (Eckl. & Zeyh.) Puff,

comb, et stat nov 1 ,2: 46

arenicola Puff, sp. nov 1 ,2: 47

cinerea (Thunb.) Puff, comb, nov 1,2: 40

coronata Puff, sp. nov 1 ,2: 41

divaricata Salter 1 ,2: 43

dregei L. Bol 1,2:40

elsieae Puff, sp. nov 1 ,2: 43

glabra (Cruse) Kuntze 1 ,2: 46

hantamensis Schltr 1 ,2: 40

turn (Cruse) Salter 1,2:44

subsp. calciphila Puff, subsp. nov 1,2: 44

subsp. hirta 1,2:44

microphylla (Sond.) Salter 1,2:38

namaquensis Puff, sp. nov 1 ,2: 41

sp. A 1,2:47

sp. B 1,2:47

Otiophora calycophylla (Sond.) Schltr. & K.

Schum 1,2:36

Oxyspermum Eckl. & Zeyh 1 ,2: 48

asperum Eckl. & Zeyh 1 ,2: 51

PAEDERIAL 1,2: 1

bojeriana (A . Rich.) Drake as bojerana 1 ,2: 3

subsp. foetens (Hiern) Verde 1 ,2: 3

foetens (Hiern) K. Schum 1 ,2: 3

ligun Sweet 1,2:3

1,2: 79

PAEDERIEAE 1,2: 1

Phyllis galopina (Thunb.) Cruse 1 ,2: 48

Phyllis sensu Cruse 1 ,2: 48

RubiaL 1,2:61

conotricha Gand 1,2:62

cordifoliai, 1,2:62

subsp. conotricha (Gand.) Verde 1 ,2: 62

cordifolia sensu Sond. , non L 1 ,2: 62

horrida (Thunb.) Puff 1,2:65

lanata Dinter 1,2:68

longipetiolata Bullock 1 ,2: 62

peregrinaL.* 1,2:61

petiolaris DC 1,2:62

var. heterophylla Sond 1,2: 65

var. isophylla Sond 1,2:62

tinctorumL.* 1,2:61

RUBIEAE 1,2:61

Siphomeris Boj 1,2:1

foetens Hiern 1,2:3

ligun (Sweet) Boj 1 ,2: 3

Spermacoce ericaefolia Licht. ex Roem. &

Schult 1,2:35

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