FLORA OF SOUTHERN AFRICA
VOLUME 33 ASTERACEAE
Editor G. Germishuizen
Part 4: Anthemideae
Fascicle 1: Eriocephalus and Lasiospermum
by M.A.N. Muller, P.P.J. Herman and H.H. Kolberg
Digitized by the Internet Archive
in 2016
https://archive.org/details/floraofsoutherna334unse
FLORA OF SOUTHERN AFRICA
which deals with the territories of
SOUTH AFRICA, LESOTHO, SWAZILAND, NAMIBIA AND BOTSWANA
VOLUME 33: ASTERACEAE
PART 4: ANTHEMIDEAE
FASCICLE 1 : Eriocephalus and Lasiospermum
by
M.A.N. Muller, P.P.J. Herman and H.H. Kolberg
Scientific editor: G. Germishuizen
Technical editor: E. du Plessis
NATIONAL
Botanical
I N S T I T U
Pretoria
2001
Editorial Board
B.J. Huntley
R.B. Nordenstam
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National Botanical Institute, Cape Town, RSA
Swedish Museum of Natural History, Stockholm, Sweden
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ISBN 1-919795-59-6
CONTENTS
New taxa, new combinations and new statuses published in Volume 33, Part 4, Fascicle 1 . . iv
Introduction v
Preface vi
Eriocephalus 1
Lasiospermum 64
Index 75
Appendix:
Plan of Flora of southern Africa A- 1
FSA contributions in Bothalia A-3
Flora of southern Africa : alphabetical list of published taxa A-4
NEW TAXA, NEW COMBINATIONS AND NEW STATUSES
PUBLISHED IN VOLUME 33, PART 4, FASCICLE 1
Eriocephalus africanus L. var. paniculatus (Cass.) M.A.N. Miiller, P.P.J. Herman &
H.H.Kolberg, comb, et stat. nov.. p. 26
Eriocephalus ambiguus (DC.) M.A.N. Miiller, comb, et stat. nov., p. 42
Eriocephalus brevifolius (DC.) M.A.N. Miiller, comb, et stat. nov., p. 21
Eriocephalus ericoides (L.f.) Druce subsp. griquensis M.A.N. Miiller, subsp. nov., p. 49
Eriocephalus giessii M.A.N. Miiller, sp. nov., p. 34
Eriocephalus glandulosus M.A.N. Miiller, sp. nov., p. 49
Eriocephalus grandiflorus M.A.N. Miiller, sp. nov., p. 29
Eriocephalus karooicus M.A.N. Miiller, sp. nov., p. 31
Eriocephalus klinghardtensis M.A.N. Miiller, sp. nov., p. 19
Eriocephalus longifolius M.A.N. Miiller, sp. nov., p. 11
Eriocephalus merxmuelleri M.A.N. Miiller, sp. nov., p. 59
Eriocephalus microphyllus DC. var. carnosus M.A.N. Miiller, var. nov., p. 54
Eriocephalus microphyllus DC. var. pubescens (DC.) M.A.N. Miiller, comb, et stat. nov.,
Eriocephalus namaquensis M.A.N. Miiller, sp. nov., p. 57
Lasiospermum Lag. section Radiatum M.A.N. Miiller, sect, nov., p. 65
Date of publication: April 2001.
INTRODUCTION
This part was compiled in accordance with the Guide for contributors to the Flora of southern
Africa (compiled by Leistner, Ross & De Winter and available from the Editor, National Botanical
Institute, Private Bag X 1 0 1 , Pretoria, 0001 South Africa).
The maps show the distribution of the various taxa in the FSA region only.
The numbering of the genera is according to De Dalla Torre & Harms in their Genera
siphonogamarum ( 1900-1907), as adapted by Arnold & De Wet ( 1993, Plants of southern Africa:
names and distribution).
Background to this Fascicle
M.A.N. Muller completed a revision of the genera Eriocephalus L. and Lasiospermum Lag. for
which a Ph.D. degree was conferred upon him by the University of Stellenbosch in 1988. The
results of his thesis had not been published by the time he tragically died in a car accident in April
1997. Permission was granted by the University to publish the taxonomic part of his thesis in the
FSA format as it was considered to be a valuable contribution to plant taxonomy in southern Africa.
P.P.J. Herman extracted and adapted the relevant text from the thesis in co-operation with H.H.
Kolberg and translated it into English. However, not all specimens examined by Muller were seen
by Herman. The original line drawings by Blythe Loutit and Elna de Bruyn could not be traced.
The figures in this publication were scanned from a copy of the thesis.
v
PREFACE
The genera Eriocephalus L. and Lasiospermum Lag. belong to the tribe Anthemideae (Asteraceae),
which is characterised by an aromatic scent and dissected leaves; the pappus is often absent where-
as paleae are often present (Muller 1988, 'n Morfologiese en taksonomiese studie van die genusse
Lasiospermum Lag. en Eriocephalus L. ( Asteraceae ) in snidelike Afrika, unpublished Ph.D. thesis;
Bremer 1994, Asteraceae. cladistics and classification). The tribe is economically important as
many of its members are weeds ( Anthemis cotula L.), cultivated ( Dendranthema species) or used
medicinally ( Artemisia afra Jacq. ex Willd.).
The last revision of Eriocephalus and Lasiospermum in southern Africa was done by Harvey (1865,
in Flora capensis 3). The genus Eriocephalus is endemic to southern Africa and its distribution
covers the whole FSA region except Gauteng and KwaZulu-Natal, with the highest concentration
of taxa in the Western Cape. It is an important member of the karoo vegetation and often forms an
important fodder plant in these dry areas (Van Breda & Barnard 1991, 100 Veld plants of the win-
ter rainfall region. A guide to the use of veld plants for grazing-, Le Roux, Kotze, Nel & Glen 1994,
Bossieveld).
Except for Lasiospermum brachyglossum DC. var. sinaicum Asch. & O.Hoffm. (this should prob-
ably rather be treated as a subspecies), which occurs in the Sinai Desert, the genus Lasiospermum
is endemic to southern Africa and its distribution covers Namibia, North-West, Gauteng, the Free
State, Lesotho and the Northern, Western and Eastern Cape. It is economically important as some
members are reported to be poisonous to stock (Walsh 1909, South African poisonous plants',
Vahrmeijer 1981, Poisonous plants of southern Africa that cause stock losses).
Members of Eriocephalus display anomalous secondary growth, which leads to the splitting of
older plants resulting in independent daughter plants (Muller 1988). The branches can be divided
into long, ‘normal’ branches (dolichoblasts), with leaves either opposite or alternate, and dwarf
shoots (brachyblasts) containing leaf tufts. The most important characters used in the delimitation
of species are: paleae of marginal female florets free or connate, capitula radiate or disciform, the
indumentum of the leaves felty or sericeous and leaves opposite or alternate. Until now it was
believed that the involucre consisted of two rows of involucral bracts (De Candolle 1838,
Prodromus systematis naturalis regni vegetabilis 6; Harvey 1865; Bentham 1873, in Genera plan-
tarum 2; Phillips 1926, The genera of South African flowering plants’, Merxmiiller 1967, in
Prodromus einer Flora von Siidwestafrika 139; Bremer & Humphries 1993, in Bulletin of the
Natural History Museum, London (Botany series) 23,2; Bremer 1994). Muller ( 1988) conducted an
intensive ontogenetic and anatomical investigation of the capitula and came to the conclusion that
the involucre consists of only one row of involucral bracts and that the inner row represents the
paleae of the marginal female florets. These paleae can be either free or connate.
Herman (2001, in South African Journal of Botany 67: 66) described septate hairs on the paleae of
most of the Eriocephalus species and on the cypselas of two of the four Lasiospermum species and
it seems possible that these hairs could be a generic character of both Eriocephalus and
Lasiospermum.
VI
ASTERACEAE: Eriocephalus
9320000 1. ERIOCEPHALUS
by M.A.N. MULLERt, P.P.J. Herman* * & H.H. Kolberg**
(Literature references on p. 61)
Eriocephalus L., Species plantarum, edn 1: 1310 (1753); Murray: 795 (1784); Lam.: 387
(1786); Juss.: 186 (1789); Gaertn.: 428 (1791); Moench: 590 (1794); Thunb.: 168 (1800); Curtis:
t. 833 (1805); Pers.: 497 (1807); Thunb.: 724 (1823); Spreng.: 621 (1826); Cass.: 491 (1827);
Spreng.: 642 (1831); Less.: 268, 269 (1832); DC.: 145 (1838); Endl.: 441 (1838); Harv.: 185
(1838); Harv.: 199, 200 (1865); Benth.: 416 (1873); E.Phillips: 660, 661 (1926); Levyns: 261
(1929); Marloth: 261 (1932); Adamson & T.M. Salter: 800 (1950); Merxm.: 58, 59 (1967);
R. A. Dyer: 701, 702 (1975); M.A.N.Miiller: 155 (1988); K.Bremer & Humphries: 160 (1993);
K. Bremer: 473 (1994); P.P.J. Herman, Retief, M.Koekemoer & W.G.Welman: 136 (2000). Type
species: E. africanus L.
Many-stemmed, sparsely to much-branched, erect to spreading, sometimes spinescent shrubs,
rarely suffrutices, 0.25-2.0 m high and in diameter, often aromatic. Old stems mostly displaying
anomalous secondary growth; young branches sericeous to felty, often glabrescent: sometimes with
short-lived brachyblasts in leaf axils. Leaves mostly opposite, but sometimes alternate, densely
imbricate on brachyblasts, ericoid, linear or acicular, entire or pinnatisect with 1-7 linear lobes.
Capitula solitary on brachyblasts or in terminal umbellate racemes at ends of young shoots or
brachyblasts or in racemes or spikes; heterogamous radiate or disciform, with 2-60 florets: 1-8 ray
or marginal female and 1-60 functionally male disc florets. Involucre semiglobose, of 4-6 involu-
cral bracts in 1 row; bracts oblong, ovate, lanceolate to obovate, slightly keeled to flattened, often
with broad transparent membranous margin, pubescent or glabrous. Receptacle flattened, paleate.
Paleae as long as florets or shorter, those of marginal florets totally or partially connate or entire-
ly free from each other, hard and thick or membranous and transparent; free paleae ovate to lanceo-
late to linear, margins fringed, abaxially long-lanate, hairs septate. Marginal female florets with or
without strap-shaped lamina, shorter than, as long as, or longer than furcation of style, or distinct-
ly longer than style branches, strap-shaped to cuneate, to broadly cuneate, apex 2-4-dentate or
-lobed; corolla white, cream-coloured, pink or purple, rarely golden yellow. Style cylindrical, fur-
cate with 2 flattened, linear, acute branches. Ovary (and cypsela) oblong-linear to narrowly
obovoid, slightly flattened, sometimes slightly angular, lanate or pilose. Seed dark brown, smooth,
glabrous, oblong-ovoid, slightly flattened. Pappus absent. Paleae : those of disc florets ovate to
lanceolate to linear, flattened, membranous, margins fringed to long-lanate, abaxially long-lanate.
Disc florets functionally male with sterile ovary; corolla tubular, slightly widened, trumpet-shaped
to infundibuliform. 5-lobed; lobes triangular; corolla white, cream-coloured, yellow or purple.
Style cylindrical, unbranched, apex globose, with sweeping hairs, rarely 2-lobed. Stamens 5,
anthers laterally fused, each with lanceolate apical appendage, ecaudate and ecalcarate, endothe-
cial tissue polarised. Basic chromosome number : x = 9 (2 n = 18, 36, 54, 72).
la (lb: p. 3) Capitula radiate:
2a Rays golden yellow, pale yellow or cream-coloured:
3a Leaves pinnatisect, 3-7-lobed, alternate; peduncle 15 mm long or longer; disc florets
30-60 1 . E. pinnatus
t Late of the National Herbarium of Namibia, Windhoek. Namibia.
* National Botanical Institute, Private Bag X101, Pretoria, 0001 South Africa.
** National Plant Genetic Resources Centre. National Botanical Research Institute. Private Bag 13184. Windhoek. Namibia.
ASTERACEAE: Eriocephalus
3b Leaves mostly entire, rarely pinnatisect, 3-lobed. opposite but alternate on flowering
shoots; peduncle shorter than 10 mm; disc florets 8-12 2. E. macroglossus
2b Rays white, pale to dark red-purple or white with purple tinge:
4a Paleae of marginal florets free:
5a Plants not spinescent 18. E. dinteri
5b Plants spinescent:
6a Capitula sessile; Free State, Northern and Eastern Cape 17. E. karooicus
6b Capitula shortly pedunculate; peduncle 2. 5-4.0 mm long; endemic to Namibia . . .
19. E. giessii
4b Paleae of marginal florets partly or totally connate:
7a Capitula in terminal spike or spicate-racemose; peduncle very short, 0.3-0. 5 mm
long 3. E. capitellatus
7b Capitula solitary, racemose, umbellate-racemose or paniculate; peduncle absent or
up to 40 mm long:
8a Capitula sessile or subsessile:
9a Capitula large, 5-12 mm long; mainly solitary, rarely 2—4 in raceme, sometimes
spinescent 4.E. eximius
9b Capitula smaller, 3. 5-4.0 mm long, umbellate-racemose or paniculate
14b. E. africanus var. paniculatus
8b Capitula shortly to distinctly pedunculate:
10a ( 10b: p. 3) Leaves felty, glabrescent or felty-sericeous:
1 1 a Rays always pale to dark red-purple 6. E. purpureas
1 lb Rays white or sometimes pale red-purple or white with purple tinge:
12a Leaves alternate, 18-38 mm long; Northern Province 5. E. longifolius
12b Leaves decussate, opposite or opposite and alternate on flowering shoots:
13a Leaves decussate or opposite:
14a Leaves decussate, glabrous except for felty axillary buds . ... 8. E. aromaticus
14b Leaves opposite, felty, felty-sericeous, glabrescent:
15a Leaves sparsely felty, glabrescent; capitula 3-4 mm long; widespread
(Mpumalanga, the Free State, Lesotho, the Northern, Western and East-
ern Cape) 10. E. tenuifolius
15b Leaves felty to felty-sericeous, glabrescent, semisucculent; capitula 4-6
mm long; plants sometimes spinescent; endemic to Namibia
11 . E. klinghardtensis
13b Leaves opposite, alternate on flowering shoots:
16a Leaves succulent:
1 7a Peduncle 1 0-30(— 40) mm long; capitula large, 4-5 mm long; disc florets
40-45; leaves entire, rarely pinnatisect with 2 or 3 lobes; Namaqualand
l.E. pedicellaris
17b Peduncle (3.9— )6.0— 8.5(— 1 2.0) mm long; capitula smaller, 3.5— 4.0 mm
long; disc florets ( 1 2—) 1 6— 1 8(— 24); leaves palmatisect, 3-7-lobed or
pinnatisect, 3-lobed or entire; exclusively coastal
14a . E. africanus var. africanus
16b Leaves not succulent:
1 8a Leaves permanently felty-sericeous; capitula solitary or umbellate-race-
mose; Namaqualand (Northern Cape), Swartruggens-Roggeveld and
Swartberg Mountains (Western Cape) 12. E. brevifolius
1 8b Leaves felty, glabrescent:
ASTERACEAE: Eriocephalus
3
19a Peduncles mostly longer than subtending leaves; capitula mostly with
7-10 disc florets; leaves entire to pinnatisect with up to 3 lobes
9. E. punctulatus
19b Peduncles mostly shorter than or as long as subtending leaves; capitula
mostly with 13-22 disc florets; leaves entire 10. if. tenuifolius
10b ( 10a: p. 2) Leaves sericeous:
20a Plants spinescent 15. E. grandiflorus
20b Plants not spinescent:
21a Involucral bracts almost totally membranous with narrow green central strips
13. if. scariosus
21b Involucral bracts with membranous margins and broad green central strips:
22a Leaves sericeous, with a rough surface; restricted to the Langkloof
Mountains (eastern border of the Western Cape and the Eastern Cape) . . .
1 6. E. tenuipes
22b Leaves permanently sericeous or glabrescent but then without rough surface:
23a Capitula 4-5 mm long; solitary or in umbellate racemes, with light brown
long-pilose indumentum between involucral bracts and outer paleae after
anthesis; leaves entire 12. E. brevifolius
23b Capitula 3. 5-4.0 mm long, umbellate-racemose or paniculate, with white
long-pilose indumentum between involucral bracts and outer paleae after
anthesis; leaves entire or lobed:
24a Leaves distinctly succulent, 3-5-7-lobed or entire, (6.0— )8.2— 1 5.0(— 34.0)
x 0. 8-2.5 mm; spreading shrubs, up to 4 m in diameter; branches rigid;
exclusively coastal 14a. E. africanus var. africanus
24b Leaves not or weakly succulent, mostly entire, (5— )8 — 1 7( — 40) x 0.4-0. 8
mm; mostly erect shrubs, 0.3-0.6 m in diameter; branches flexible; inland
14b. E. africanus var. paniculatus
lb (la: p. 1 ) Capitula disciform:
25a Paleae of marginal florets connate:
26a Leaves permanently silver-sericeous, decussate, alternate on flowering shoots; capitula
usually solitary, rarely racemose; plants sometimes spinescent 21. E. decussatus
26b Leaves permanently grey-felty, alternate, rarely opposite; capitula racemose or pani-
culate:
27a Capitula sessile to very shortly pedunculate; peduncle up to 5 mm long; disc florets
(4— )7 — 9 20a. E. racemosus var. racemosus
27b Capitula distinctly pedunculate; peduncle (5-) 10- 15 mm long; disc florets 13-21
20b. E. racemosus var. affinis
25b Paleae of marginal florets free:
28a Capitula spicate, sessile 23. E. pauperrimus
28b Capitula solitary, racemose, umbellate-racemose, paniculate, pedunculate, if spicate,
not sessile:
29a Leaves alternate or rosulate on brachyblasts:
30a Plants mostly spinescent; capitula usually solitary, rarely in terminal racemes . . .
24. E. ambiguus
30b Plants not spinescent; capitula usually racemose or umbellate-racemose, some-
times solitary 25. E. luederitzianus
29b Leaves decussate, opposite or opposite and alternate on flowering shoots:
31a Leaves sericeous or felty-sericeous:
32a Plants spinescent:
4
ASTERACEAE: Eriocephalus
33a Capitula always solitary, sessile or peduncles up to 3.5(— 5.0) mm long; shrub
up to 1 m high, rigid; growing on sandy soils 30. E. spinescens
33b Capitula often racemose or solitary; peduncles 2.5-12.0 mm long; shrub up to
400 mm tall, flexible; growing on shist 31. E. namaquensis
32b Plants not spinescent:
34a Capitula relatively large, 4-8 mm in diameter; leaves opposite, decussate, per-
manently sericeous, semisucculent 22. E. kingesii
34b Capitula smaller; leaves opposite, rarely alternate on flowering shoots, felty-
sericeous or sericeous, glabrescent:
35a Leaves permanently felty-sericeous 28b. E. microphyllus var. pubescens
35b Leaves sericeous, glabrescent:
36a Leaves small, 1.8—4. 0 mm long; central Namaqualand (Northern Cape) . . .
28a. E. microphyllus var. microphyllus
36b Leaves 4 — 9(— 14) mm long; southern Namibia and Northern Cape
32. E. merxmuelleri
31b Leaves permanently felty or felty, glabrescent:
37a Capitula 4-6 mm long; leaves succulent; Worcester and Montagu Districts . . .
28c. E. microphyllus var. carnosus
37b Capitula shoiier than 3 mm; leaves not succulent:
38a Leaves permanently long-felty; between Orange River and Botswana border
(Northern Cape) 26b. E. ericoides subsp. griquensis
38b Leaves felty, glabrescent:
39a Disc florets 10-18 27. E. glandulosus
39b Disc florets fewer than 10:
40a Leaves 0.8-1 .6 mm long, decussate; capitula mostly solitary; much-branched
shrubs; side branches forming an angle of 70-90° with main axis, opposite
29. E. microcephalus
40b Leaves (0.75—) 1 .0— 3.0(— 5.0 ) mm long, opposite, decussate on brachyblasts,
rarely alternate on flowering shoots; capitula spicate racemose, racemose
or solitary; sparsely branched, conical or broom-like shrubs; branches
tending to be vertically orientated 26a. E. ericoides subsp. ericoides
Key to the species of Eriocephalus in different geographical areas of southern Africa
Namibia
la Capitula radiate:
2a Paleae of marginal florets connate:
3a Rays golden yellow; capitula 10-12 mm long; leaves pinnatisect 1. E. pinnatus
3b Rays white or pale red-purple; capitula up to 6 mm long; leaves entire, rarely dentate:
4a Leaves opposite or subopposite, felty to felty-sericeous, glabrescent; disc florets
12-15 11 . E. klinghardtensis
4b Leaves alternate, densely sericeous to glabrous; disc florets 4—9 13. E. scariosus
2b Paleae of marginal florets free:
5a Plants not spinescent; involucral bracts shortly appressed sericeous 18. E. dinteri
5b Plants spinescent; involucral bracts permanently long-sericeous to long-pilose
19. E. giessii
ASTERACEAE: Eriocephulus
5
lb Capitula disciform:
6a Capitula spicate, sessile 23. E. pauperrimus
6b Capitula solitary, racemose, umbellate-racemose or paniculate, shortly to distinctly
pedunculate:
7a Leaves alternate or rosulate on brachyblasts:
8a Capitula solitary, rarely in terminal racemes; plants spinescent 24. E. ambiguus
8b Capitula racemose or umbellate-racemose, rarely solitary; plants not spinescent . . .
25. E. luederitzianus
7b Leaves opposite, decussate, rarely alternate on flowering shoots:
9a Capitula large, 4-8 mm in diameter; leaves 6-12 mm long, permanently densely
sericeous, semisucculent 22. E. kingesii
9b Capitula smaller, up to 5 mm long; leaves up to 9(— 1 2) mm long, not succulent:
10a Leaves (0. 75-) 1.0-3. 0(— 5.0) mm long, initially densely felty, soon glabrescent,
shiny; sparsely branched, conical or broom-like shrubs; branches tending to be
vertically orientated 26a. E. ericoides subsp. ericoides
10b Leaves 4— 9(— 1 4) mm long, felty-sericeous, sericeous to glabrescent; much-
branched shrub; side branches opposite 32. E. merxmuelleri
Botswana
Plants mostly spinescent; capitula usually solitary, rarely in terminal racemes ... 24. E. ambiguus
Plants not spinescent; capitula usually racemose or umbellate-racemose, sometimes solitary
25. E. luederitzianus
Northern Province and Mpumalanga
la Capitula disciform; paleae of marginal florets free; leaves silver-grey sericeous
25. E. luederitzianus
lb Capitula radiate; paleae of marginal florets connate; leaves felty, glabrescent:
2a Peduncle 12-17 mm long; capitula 5-6 mm long; leaves alternate. 18-38 mm long;
Northern Province 5. E. longifolius
2b Peduncles (3-)4— 7(-10) mm long; capitula 3^4 mm long; leaves opposite, sometimes
alternate on flowering shoots, 4— 14(— 24) mm long; Mpumalanga 10. E. tenuifolius
Free State and Lesotho
la Capitula disciform; disc florets ( 1— )3— 5(— 7); sparsely branched, conical or broom-like
shrubs; side-branches tending to be vertically orientated . . 26a. E. ericoides subsp. ericoides
lb Capitula radiate; disc florets up to 35; much-branched shrubs:
2a Paleae of marginal florets free; disc florets 4-10; spinescent shrubs 17. E. karooicus
2b Paleae of marginal florets connate; disc florets more than 10; plants spinescent or not:
3a Capitula 5-12 mm long, mainly solitary, rarely 2-4 in a raceme, sessile or subsessile;
peduncle shorter than 0.5 mm; disc florets 26-35; leaves permanently silvery
sericeous 4. E. eximius
3b Capitula 3^1 mm long, in umbellate racemes; peduncle (3— )4— 7(— 10) mm long; disc
florets 13-22; leaves sparsely felty, glabrescent 10. E. tenuifolius
6
ASTERACEAE: Eriocephalus
North-West and Northern, Western and Eastern Cape
la Capitula radiate:
2a Rays cream-coloured to pale yellow 2. E. macroglossus
2b Rays white, pale to dark red-purple or white with purple tinge:
3a Paleae of marginal florets free 17. E. karooicus
3b Paleae of marginal florets partly or totally connate:
4a Capitula in terminal spike or spicate-racemose; peduncle very short, 0.3-0. 5 mm
long 3. E. capitellatus
4b Capitula solitary, racemose, umbellate-racemose or paniculate; peduncle absent or
up to 40 mm long:
5a Capitula sessile or subsessile:
6a Capitula large, 5-12 mm long, mainly solitary, rarely 2-4 in raceme; plants
sometimes spinescent 4. E. eximius
6b Capitula smaller, 3. 5^4.0 mm long, umbellate-racemose or paniculate
14b. E. africanus var. paniculatus
5b Capitula shortly to distinctly pedunculate:
7a Leaves felty, glabrescent or felty-sericeous:
8a Rays always pale to dark red-purple 6. E. purpureus
8b Rays white or sometimes pale red-purple or white with purple tinge:
9a Leaves decussate: plants glabrous except for felty axillary buds . . . 8. E. aromaticus
9b Leaves opposite, alternate on flowering shoots:
10a Leaves succulent:
1 la Peduncle 1 0— 30(— 40) mm long; capitula large, 4-5 mm long; disc florets
40-45; leaves entire, rarely pinnatisect with 2 or 3 lobes; Namaqualand
7 . E. pedicellaris
lib Peduncle (3.9— )6.0— 8.5(— 1 2.0) mm long; capitula smaller, 3. 5-4.0 mm
long; disc florets ( 1 2— ) 1 6— 1 8(— 24); leaves palmatisect, 3-7-lobed or pin-
natisect, 3-lobed or entire; exclusively coastal
14a. E. africanus var. africanus
10b Leaves not succulent:
12a Leaves permanently felty-sericeous; capitula solitary or umbellate-race-
mose; Namaqualand (Northern Cape), Swartruggens-Roggeveld and Swart-
berg Mountains (Western Cape) 12. E. brevifolius
12b Leaves felty, glabrescent:
13a Peduncles mostly longer than subtending leaves; capitula mostly with
7-10 disc florets; leaves entire to pinnatisect with up to 3 lobes
9. E. punctulatus
13b Peduncles mostly shorter than or as long as subtending leaves; capitula
mostly with 13-22 disc florets; leaves entire 10. if. tenuifolius
7b Leaves sericeous:
14a Plants spinescent 15. E. grandiflorus
14b Plants not spinescent:
1 5a Involucral bracts almost totally membranous with narrow green central strips
13. E. scariosus
15b Involucral bracts with membranous margins and broad central green strips:
16a Leaves sericeous, glabrescent, with a rough surface; restricted to the
Langkloof Mountains (eastern border of the Western Cape and the Eastern
Cape) 16. E. tenuipes
ASTERACEAE: Eriocephalus
7
1 6b Leaves permanently sericeous or glabrescent but then without rough surface:
17a Capitula 4-5 mm long, solitary or in umbellate racemes, with light brown
long-pilose indumentum between involucral bracts and outer paleae after
anthesis; leaves entire 12. E. brevifolius
17b Capitula 3. 5-4.0 mm long, umbellate-racemose or paniculate, with white
long-pilose indumentum between involucral bracts and outer paleae after
anthesis; leaves entire or lobed:
18a Leaves distinctly succulent, 3-5-7-lobed or entire, (6.0— )8.2— 1 5.0(— 34.0)
x 0.8-2. 5 mm; spreading shrubs, up to 4 m in diameter; branches rigid;
exclusively coastal 14a. E. africanus var. africanus
18b Leaves not or weakly succulent, mostly entire. (5— )8— 1 7( — 40) x 0.4-0.8
mm; mostly erect shrubs, 0.3-0. 6 m in diameter; branches flexible;
inland 14b. E. africanus var. paniculatus
lb Capitula disciform:
19a Paleae of marginal florets connate:
20a Leaves permanently silver-sericeous, decussate, alternate on flowering shoots; capi-
tula usually solitary, rarely racemose; plants sometimes spinescent . ... 21. is. decussatus
20b Leaves permanently grey-felty, alternate, rarely opposite; capitula racemose or pani-
culate:
21a Capitula sessile to very shortly pedunculate; peduncle up to 5 mm long; disc florets
(4— )7— 9 20a. E. racemosus var. racemosus
21b Capitula distinctly pedunculate; peduncle (5-) 10- 15 mm long; disc florets 13-21
20b. E. racemosus var. ajfinis
19b Paleae of marginal florets free:
22a Capitula spicate, sessile 23. E. pauperrimus
22b Capitula solitary, racemose, umbellate-racemose, paniculate; pedunculate, if spicate,
not sessile:
23a Leaves alternate or rosulate on brachyblasts 24. is. ambiguus
23b Leaves decussate, opposite or opposite and alternate on flowering shoots:
24a Leaves sericeous or felty-sericeous:
25a Plants spinescent:
26a Capitula always solitary, sessile or peduncles up to 3.5(— 5.0) mm long; shrub
up to 1 m high, rigid; growing on sandy soils 30. E. spinescens
26b Capitula often racemose or solitary; peduncles 2.5-12.0 mm long; shrub up to
400 mm tall, flexible; growing on shist 31. E. namaquensis
25b Plants not spinescent:
27a Leaves permanently felty-sericeous 28b. E. microphyllus var. pubescens
27b Leaves sericeous, glabrescent:
28a Leaves small, 1 .8 — 4.0 mm long 28a. E. microphyllus var. microphyllus
28b Leaves 4— 9(— 14) mm long 32 . E. merxmuelleri
24b Leaves permanently felty or felty, glabrescent:
29a Capitula 4—6 mm long, leaves succulent: Worcester and Montagu Districts . . .
28c. E. microphyllus var. carnosus
29b Capitula shorter than 3 mm; leaves not succulent:
30a Leaves permanently long-felty; between Orange River and Botswana border
(Northern Cape) 26b. E. ericoides subsp. griquensis
30b Leaves felty, glabrescent:
31a Disc florets 10-18
31b Disc florets fewer than 10:
27. E. glandulosus
ASTERACEAE: Eriocephalus
32a Leaves 0.8-1. 6 mm long, decussate; capitula mostly solitary; much-
branched shrubs; side branches forming an angle of 70-90° with main axis,
opposite 29. E. microcephalus
32b Leaves (0.75-) 1 .0— 3.0(— 5.0) mm long, opposite, decussate on brachyblasts,
rarely alternate on flowering shoots; capitula spicate racemose, racemose
or solitary; sparsely branched, conical or broom-like shrubs; branches
tending to be vertically orientated 26a. E. ericoides subsp. ericoides
1. Eriocephalus pinnatus O.Hoffm. in
Botanische Jahrbiicher 10: 277 (1889); Merxm.:
61 (1967). Type: Namibia, ‘Hereroland, Ubib.
in saxosis alt. 1000 m, Marloth 1440, Florif. m.
Jun. 1886’ (SAM, lecto.!, designated here;
BOL!, GRA!, PRE!).
Erect, many-stemmed, freely branched,
weakly woody shrubs or suffrutices, with annual
regrowth, 35CM-50 mm tall, 350 mm in diame-
ter. Old stem not displaying anomalous second-
ary growth, cylindrical, grey-yellow; young
shoots golden brown, densely long-pilose mixed
with dense felty indumentum, pubescence per-
manent; brachyblasts absent; dolichoblasts leafy,
without cushion-like thickenings on stems.
Leaves alternate, distinctly petiolate; lamina
20-75 mm long, pinnatisect, 3-7-lobed; lobes
linear, up to 48 x 1.0-1. 5 mm, adaxially basally
flattened to concave, abaxially convex; perma-
nently grey-golden hairy, indumentum sparsely
long-pilose mixed with felty mat; petiole slight-
ly broadened at base, temporarily retained after
leaf fall, abscising later. Capitula heterogamous
radiate, mostly racemose or in umbellate termi-
nal racemes, rarely solitary in leaf axils, rela-
tively large, 10-12 x 5-15 mm; peduncles
cylindrical, 15-46 mm long, densely felty.
Involucral bracts 5, 10-13 x 4-6 mm, with
abaxially hairy, central, green, herbaceous strip
and relatively broad, glabrous, membranous
margins. Paleae : those of marginal florets con-
nate into hard cylindrical sheath, 8-10 mm
long, margin densely lanate, hairs septate; those
of disc florets spathulate to linear, membranous,
6-8 mm long, abaxially densely lanate, indu-
mentum intertwined with that of neighbouring
paleae to form a mat, apex fringed. Ray florets
female, 4—8. 6-10 mm long, lamina broadly
cuncate. 6x5 mm, golden yellow, 3-lobed,
abaxially glandular, longer than style branches.
Style forked, branches flattened, linear. Ovary
(and cypsela) oblong, lanceolate, long-pilose/
lanate. Seed 5-6 mm long, flatfish trigonous.
Disc florets functionally male with sterile ovary,
30-60; corolla tube cylindrical to trumpet-
shaped, 5-lobed, golden yellow. Style un-
branched, cylindrical, apex globose, with
sweeping hairs. Stamens 5, exserted at maturity.
Receptacle after anthesis with long hairs
between involucral bracts and marginal connate
paleae. Chromosome number. In = 18. Flower-
ing time : March to May, sometimes continuing
into August; flowering is linked to rainfall,
which can be fairly sporadic in the distribution
range.
This Namibian endemic occurs in the north-
ern and central Narnib, on the escarpment and
mopane savanna (Giess 1971). These areas re-
ceive an average annual rainfall of less than 200
mm. Although the distribution of this species
extends over a large area, plants are extremely
scarce and occur fairly localised. They are usu-
ally found in groups of five to eight but some-
times of up to 20 plants. They grow mostly in
stony areas or on sandstone koppies. Map 1 .
E. pinnatus is unique in the genus in more
ways than one. It has distinctly pinnatisect
leaves, large golden yellow ray florets and is
herbaceous — all characters not found in the rest
of the genus. The whole plant is covered by an
indumentum of dense golden grey felt inter-
spersed with scattered long-pilose hairs. Some
other species of the genus also have a felty
indumentum, but it consists mostly of tempo-
rary, fine, soft, white hairs. Where a permanent
felty indumentum occurs, it is more felty
sericeous.
ASTERACEAE: Eriocephalus
9
Map i . — • Eriocephalus pinnatus; ▲ E. macroglossus;
■ E. capitellatus.
Pinnatisect leaves are not uncommon in the
genus Eriocephalus , but the degree of incision
reaches its peak in this species. E. pinnatus has
a distinct petiole and a blade with 3-7 lobes.
The stems are mostly herbaceous, woody at
base only. Anomalous secondary growth is
absent only in this species and in E. longifolius
(no. 5). The plant is browsed to the ground or
above-ground parts die back every year and
resprout after first good rains. It is the only
species with golden yellow ray florets. Yellow
disc florets are present inter alia in E. lueder-
itzianus (no. 25) and E. ambiguus (no. 24) and
pale yellow to cream-coloured rays in E.
macroglossus (no. 2). Although each capitulum
contains 4—8 female florets, very few cypselas
are formed. From 10 capitula with probably 60
female florets, only two mature seeds were
observed, thus a seed-set of only 3%. This pos-
sibly explains the scarcity of this species.
Common name: kapokbossie.
Vouchers: Boss A] 07 (PRE); Craven 1023
(WIND); De Winter & Hardy 8230 (PRE,
WIND); Giess 7925 (WIND); Hall 366 (BOL,
NBG).
2. Eriocephalus macroglossus B.Nord. in
Journal of South African Botany 30: 49-52
(1964). Type: Northern Cape, 10 km west of
Springbok, Maguire 374 (NBG, holo. !).
Vigorous, much-branched shrubs, 0.5- 1.2 m
high. Old stems and branches glabrous, display-
ing anomalous secondary growth, bark grey;
young shoots erect, straight, shortly sericeous.
Leaves opposite, alternate on flowering shoots,
clustered on brachyblasts, sessile on cushion-
like thickenings, linear, 3-20 x 1-5 mm, most-
ly entire, rarely pinnatisect, 3-lobed, adaxially
flattened, concave towards base, abaxially con-
vex, slightly keeled, permanently densely
appressed silvery sericeous, apex acute, base
slightly broadened. Capitula heterogamous
radiate, 4-10, umbellate-racemose, terminal,
5-7 mm long; peduncles 4—8 mm long, densely
sericeous. Involucral bracts 4, ovate, 4—5 x
2. 5^1.0 mm, central part herbaceous to slightly
coriaceous, apex obtuse, rarely acute, slightly
fringed, margins brownish or blackish; 2 bracts
narrow and slightly keeled, other 2 broader and
flattened, overlapped by margins of narrow
bracts. Paleae : those of marginal florets con-
nate at base, broadly ovate, keeled, 6-7 mm
long, coriaceous central part with membranous,
fringed margins, abaxially long-lanate, hairs
septate; those of outer disc florets ovate, of
inner ones narrowly oblong to lanceolate, 6-7 x
1- 3 mm, membranous, acute, margins and
abaxially long-lanate. Rays 2 or 3; corolla tube
2- 3 mm long, pale brown; lamina broadly
oblong or oblong-cuneate, 6-7 x 4-6 mm,
cream-coloured, obtuse, 3(or 4)-lobed. Style
branches flattened, linear, acuminate, 2. 5-4.0
mm long. Ovary (and cypsela) narrowly
oblong, densely long-lanate. Seed 2-3 mm long,
slightly flattened. Disc florets 8-12, functional-
ly male with sterile ovary; corolla light brown
to creamy, tubular, gradually widening distally,
3. 5^4.0 mm long, 5-lobed; lobes acute, 0.5 mm
long. Style unbranched with slight convex apex
surrounded by short sweeping hairs. Stamens 5,
1. 2-2.0 mm long, barely exserted at maturity.
Receptacle after anthesis with dense indumen-
tum between marginal paleae and involucral
bracts, white or tawny to brown. Chromosome
number. 2 n = 36. Flowering time: closely cor-
related with rainfall, June to August. The distri-
10
ASTERACEAE: Eriocephalus
bution area receives winter rain, 150-300 mm
annually.
E. macroglossus is currently known only
from northern Namaqualand. The record from
Botterkloof Pass is doubtful since the fragment
was mounted with material of E. purpureus
( Barker 9293, NBG). An attempt to locate it in
that area proved to be unsuccessful. The plants
are found ± 600 m above sea level on low
mountains in stony soil. Its distribution falls
into Acocks’s (1975) Namaqualand Broken
Veld. Map 1.
The species is distinguished by its leaves
with a silvery sericeous indumentum, umbellate
racemes, large capitula with distinct, large,
cream-coloured (pale yellow) rays and well-
developed long-lanate indumentum in the capi-
tula. It is closely related to E. grandiflorus (no.
15). The latter is a much-branched, slightly
spinescent shrub with pure white or pale to dark
purple rays, whereas E. macroglossus is not
spinescent and has pale yellow rays.
Common name: kapokbos.
Vouchers: Acocks 19572 (BOL, NBG. PRE);
Goldblatt 2353 (NBG, PRE); Marloth 12367b
(BOL, NBG. PRE); Muller 3553 (WIND);
Midler 4021 (WIND).
3. Eriocephalus capitellatus DC., Prodro-
mus: 146 (1838); Harv.: 201 (1865). Type:
Western Cape, ’Zwaanepoelspoortberg, auf
steinigen, trocknen Bergriicken, 2000-3000
Fuss, August’, Drege 2144 (G-DC, holo.; G!,
NBG!. P!, PRE. photo.!, SAM!).
Slender, erect, small, conical shrubs,
0.25-1 .2 m high. Old stems grey to grey-brown,
4 mm in diameter; dolichoblasts red-brown,
barely 0.5 mm in diameter, growing points
green-brown. Leaves mostly alternate or rarely
opposite, mostly palmatisect to pinnatisect, but
sometimes entire, 4.0-7. 5 mm long, 0.4-0.6
mm in diameter, basally adaxially slightly con-
cave, abaxially convex, blue-green to grey-
green, indumentum delicately sericeous with
underlying felted layer and extending down to
leaf base, those on dolichoblasts and brachy-
blasts of the same length; lobes linear, cylindri-
cal to clavate, apex obtuse to slightly acute.
Capitula heterogamous radiate, small, barely 2
mm long, in terminal spike or spicate-racemose;
peduncles 0.3-0. 5 mm long. Involucral bracts 4
or 5, oval to ovate, 1.7 x 1.0 mm, slightly keeled
to flattened, with central green part and broad
membranous margin, finely appressed seri-
ceous. Paleae : those of marginal florets connate
forming cylindrical sheath, membranous, up to
1.3 mm long, margins strongly fringed, abaxial-
ly long and densely lanate, nairs septate; those
of central florets small, barely 0.6 mm long,
transparent membranous, margins fringed,
abaxially long-lanate. Ray florets 1 or 2 (or 3),
up to 2.2 mm long with up to 1.2 mm long,
white, strap-shaped to cuneate, 3-lobed lamina.
Style branches strap-shaped, apex acute, up to
0.6 mm long. Ovary (and cypsela) oblong, flat-
fish. trigonous, long-lanate. Seed 1.5-2. 1 mm
long, ovoid, slightly flattened. Disc florets
1^4(— 1 3). functionally male with sterile ovary,
up to 1.6 mm long; corolla infundibuliform,
creamy white with red-purple tint, 5-lobed.
Stamens 5. St} >le undivided, apex globose.
Receptacle after anthesis with sparse white,
long-hairy indumentum between involucral
bracts and connate marginal paleae. Chromo-
some number. 2 n = 18. Flowering and fruiting
time: April to September (winter-rainfall area),
February to May (summer-rainfall area).
E. capitellatus occurs on the high mountains
of the Western and Eastern Cape. It grows at an
altitude of over 900 m in both winter- and sum-
mer-rainfall areas. The species never occurs in
dense stands and is sparsely distributed on
mountain slopes. Map 1 .
This shrub with its slender branches has
blue-green to grey-green leaves, which turn
darker upon drying. The pinnatisect to palmati-
sect leaves have extremely narrow lobes
(0.4-0. 6 mm). While other species of
Eriocephalus are very attractive during flower-
ing and/or fruiting, this species is not very con-
ASTERACEAE: Eriocephalus
spicuous because its small capitula lack a con-
spicuous long, dense indumentum at fruiting
and are hardly visible among other plants.
Common name: kapokbos.
Vouchers: Barker 4526 (NBG); Compton
5224 (BOL); Dahlstrand 2365 (NBG, PRE);
Esterhuysen 4517 (BOL, PRE); Marloth 9027
(NBG, PRE).
4. Eriocephalus eximius DC., Prodromus:
147 (1838); Harv.: 203 (1865). Type: Western
Cape, 'Auf steinigen, trocknen Bergriicken von
Sneeuberge, 400-500 Fuss, August’, Drege 2138
(G-DC, holo.; PRE, photo.!).
Much-branched, rigid shrubs, 0.3-0. 6 m
high. Old stems and branches bare, sometimes
spinescent, red-brown to grey-brown; young
branches initially shortly hairy, soon glabrous.
Leaves opposite, densely imbricate on brachy-
blasts, linear to triangular, 2-9 mm long, semi-
rounded, abaxially slightly keeled, adaxially
basally slightly flattened, entire, permanently
silvery sericeous, silvery white, apex acute,
base broadened, amplexicaul. Capitula heteroga-
mous radiate, mainly solitary, terminal on
brachyblasts, rarely 2—4 in terminal racemes,
5-12 x 3-8 mm, sessile or subsessile (pedun-
cles shorter than 0.5 mm). Involucral bracts 4.
broadly ovate, 4-6 x 3-5 mm, outer 2 slightly
flattened, inner 2 more keeled, margin purple,
abaxially sericeous, apex obtuse, rarely acute.
Paleae : those of marginal florets partly or
entirely connate forming a cylindrical tube,
glabrous except for apex; those of disc florets
narrowly oblong to lanceolate, 6-7 x ± 1 mm,
membranous, acute, long-pilose on margins, ab-
and adaxial surfaces glabrous. Rays 3 or 4; lam-
ina broadly oblong or oblong-cuneate, 5-8 x
3-6 mm, many-veined, glandular abaxially,
pale to dark red-purple or white, obtuse, 3-den-
tate, tubular part narrowly cylindrical, 2-3 mm
long. Style branches flattened, linear, 2. 0-3. 5
mm long. Ovary slightly lanceolate-flattened,
densely hairy. Seed ovoid, slightly flattened,
2-3 mm long. Disc florets 26-35, functionally
male with sterile ovary; corolla tubular, widen-
ing in upper third, 6-8 mm long; glandular
abaxially; corolla lobes acute, 0.5 mm long.
Stamens 5, up to 4 mm long. Style unbranched,
with slightly convex apex surrounded by short,
sweeping hairs. Receptacle after anthesis with
white, long-pilose indumentum between involu-
cral bracts and connate marginal paleae.
Chromosome number. 2 n = 18. Flowering time:
correlated with rainfall, January to April in
summer-rainfall areas, July to August in winter-
rainfall areas.
The distribution of E. eximius is restricted to
the high mountainous parts of the Free State,
Lesotho and the Northern, Western and Eastern
Cape. Its reported occurrence in the Prieska
area is doubtful. In the communities where the
species occurs, it is found singly or in small
groups, never as the dominant component of the
vegetation. Map 2.
Common name: grootbergkapok (Smith 1966).
Vouchers: Ferreira FI 91 (PRE); Galpin
6697 (BOL, GRA, PRE, SAM); Hoener 1885
(PRE); Marloth 5831 (NBG, PRE); Thompson
2328 (NBG, PRE).
5. Eriocephalus longifolius M. A. N. Midler,
sp. nov., folia 18-38 mm longa; caulis singu-
laris sine crassificatione secundaria characteris-
12
ASTERACEAE: Eriocephalus
Figure 1- — Eriocephalus longifolius: A, flowering shoot with inflorescences and dried up peduncle, x 1 ; B. branch with
leaves, x I ; C, capitulum, x 4; D, involucral bract, x 8; E. connate marginal paleae, x 4; F, central palea, x 4; Gl, ray floret, x
4; G2. branched style, x 16; HI, disc floret, x 4; H2, anthers, x 16; H3, style, x 16; 1. leaf surface, x 32 (Gerstner 6099, PRE),
ASTERACEAE: Eriocephalus
13
tica fissuram caulis efficienti ut in speciebus
ceteris Eriocephali.
Type: Northern Province, Soutpansberg, Farm
Llewellyn, Miiller 4032 (PRE, holo.; K, WIND).
Slender, erect, sparsely branched shrubs,
0.4-1 .5 m high. Old steins leafless, leaves only
at branch tips, brown-grey to dark grey, regular-
ly cylindrical not displaying anomalous sec-
ondary growth, growing points felty, glabres-
cent; young shoots brown. Leaves alternate, at
maturity adaxially glabrescent, abaxially basal-
ly with permanent felty strip, acicular, 18-38 x
0.4-0. 5 mm, mostly entire, sometimes pinnati-
sect with 3 lobes, semiconvex distally, abaxial-
ly flattened, main vein prominent in dried mate-
rial in proximal third to half of leaf, then shal-
lowly grooved to near apex, bright green, apex
acute, base adaxially flattened, not broadened;
young leaves felty/cobwebby, initially adhering
to each other, later free. Capitula heterogamous
radiate, umbellate-racemose, 5-6 x 6 mm;
peduncles felty, 12-17 mm long. Involucral
bracts 5, broadly ovate with narrow, green, cen-
tral, herbaceous part and broad, membranous
margin, 4. 5-5. 2 x 2. 2-3. 8 mm, felty to
glabrous, apex slightly fringed. Paleae: those of
marginal florets 4.2^16 mm long, totally con-
nate into cylindrical sheath with 3 or 4 lobes,
fringed, long-lanate abaxially, hairs septate;
those of outer disc florets lanceolate, keeled,
central ones oblong to linear, flattened, mem-
branous, 3. 5-6.0 x 1.2-0. 5 mm, apices long-
fringed, abaxially long-lanate. Ray florets 2 or
3, female, 6-7 mm long; corolla white with
broad cuneate to broad strap-shaped lamina,
distinct, 3-lobed or obtusely 3-dentate, 2.3 x 3.6
mm. Style branches linear, flattened, 0.7-1. 4
mm long. Ovary (and cypsela) oblong to
obovoid, slightly flattish. trigonous, long-
lanate. Seed lanceolate to narrowly ovoid, 3-4
mm long. Disc florets 10-18, functionally male
with sterile ovary; corolla white to creamy to
pale purple, trumpet-shaped, 3. 6-4. 8 mm long,
5-lobed. Stamens 5. Style undivided, apex trun-
cate with short, sweeping hairs. Receptacle after
anthesis with dense, white, long-hairy indumen-
tum between involucral bracts and connate.
marginal paleae. Chromosome number. 2 n = 18.
Flowering time', correlated with rainfall, De-
cember to March. Figure 1 .
To date, E. longifolius has been collected
only on the Soutpansberg and Waterberg in the
Northern Province. It grows only on mountain
tops, above 1 700 m, and forms part of Acocks’s
(1975) Sour Bushveld. It probably also occurs
on high mountains in Mpumalanga and North-
West. Although fairly rare, the species is not
endangered. Very few young plants were seen.
Most individuals in each community (± 15)
were already a few years old. The percentage
female florets producing fruit is low: less than
10% (86 capitula, with two or three female flo-
rets each, produced only 19 seeds). Map 2.
E. longifolius is distinguished from all other
species by the long, acicular, alternate leaves
and the sparsely branched, regularly thickened
stems. In nature, branching occurs only when an
inflorescence is formed. The umbellate racemes
develop terminally on stems and side branches
develop below this. If the upper part of the stem
is damaged, the stem resprouts from the base.
Common name: kapokbos.
Vouchers: Gerstner 6099 (PRE); Meeuse
10241 (LISC, PRE); Verdoorn 2232 (PRE).
6. Eriocephalus purpureus Burch.. Travels
in the interior of southern Africa: 232 (1822);
G.Don: 364 (1830). Type: Western Cape,
Sutherland Division: on the Wind Heuvel-
Koedoes Mountains, 22 July 1811, Burchell
1281 (Goudbloem Heights) (K. holo.!).
E. xerophilus Schltr.: 206 (1899). Type: Northern Cape.
‘In regione carrooidea: In collibus aridis, carroideis, prope
Matjiesrivier, in ditione Clanwilliam, alt. c. 2500 ped.. 4
Sept. 1896', Schlechter 8842 (B. holo.!; BOLL GRAI. PRE!.
SAM!, Z!).
Slender, erect, much-branched shrubs. 0.3-0.6
m high. Old stems dark grey, displaying ano-
malous secondary growth; young shoots yellow-
brown. sparsely felty, glabrescent: older branch-
es brown to brown-grey, striped; brachyblasts
14
ASTERACEAE: Eriocephalus
short-lived, up to 5 mm long. Leaves opposite,
but alternate on flowering shoots, decussate on
brachyblasts, densely imbricate, entire, linear,
2-6 mm long, adaxially slightly flattened, con-
cave towards base, abaxially convex, keeled dis-
tally, bright green, initially sparsely felty,
glabrescent, shiny because of glands in cavities
on leaves, apex acute, base semi-amplexicaul.
Capitula heterogamous radiate, 4-6 mm long, in
terminal umbellate-racemes as well as solitary
on brachyblasts; peduncles slender, sparsely
felty, 6-12 mm long. Involucral bracts 5, broad-
ly ovate to ovate-lanceolate, 4 x 2.5 mm, abaxi-
ally glabrous, with glands in cavities on surface,
central triangular part herbaceous, margin broad,
membranous, apex obtuse to acute. Paleae :
those of marginal florets connate with only apex
free, hard, coriaceous, margins fringed, base
abaxially densely lanate, hairs septate, glabrous
distally; those of disc florets lanceolate to nar-
rowly oblong, 3. 0-4.5 mm long, flattened, mar-
gins and abaxially long-lanate. Ray florets 2 or
3, female; corolla 6-8 mm long with strap-
shaped to cuneate, 3-lobed, 4 mm long lamina,
pale to dark purple, very conspicuous because of
size, abaxially glandular. Style branches flat-
tened, 1. 2-2.0 mm long. Ovary oblong to nar-
rowly obovoid, long-pilose. Seed oblanceolate,
2-4 mm long. Disc florets 5-20, functionally
male with sterile ovary; corolla trumpet-shaped,
5-7 mm long, 5-lobed, dark red-purple. Style
unbranched, apex truncate, globose, with sweep-
ing hairs. Stamens 5, exserted at maturity.
Receptacle after anthesis with long hairs
between involucral bracts and connate, marginal
paleae. Chromosome number : 2n = 36.
Flowering time : correlated with rainfall, June to
September with a peak from July to August.
The distribution of E. purpureus is restricted
to the winter-rainfall area and extends from
Loeriesfontein southwards to Matjiesfontein in
mountainous regions above 300 m. The eastern
boundary of its distribution overlaps with the
western distribution of E. ericoides (no. 26).
Map 3.
A.', purpureus has distinct, large, pale to dark
purple, strap-shaped ray florets, which easily
distinguish it from related species. However,
sterile and fruiting material presents problems
in identification as it closely resembles E. eri-
coides (no. 26). If no remains of the ray florets
are present, the two species can be distin-
guished by the connate, marginal paleae form-
ing a hard, cylindrical, coriaceous sheath in E.
purpureus as opposed to the short, yellow, mar-
ginal female florets and free, marginal paleae in
E. ericiodes.
Although E. purpureus had already been
described in 1822 by Burchell from material
collected in the Windheuvel-Koedoesberg, it
was later mentioned by Don (1830) only as a
plant known to British gardeners. De Candolle
(1838) and Harvey (1865) did not include it in
their studies on the genus Eriocephalus ,
although E. decussatus (no. 21) and E.
spinescens (no. 30), two species described by
Burchell (1822) in the same publication, were
indeed included. Only after the publication of E.
xerophilus by Schlechter (1899), was this
species noted and all herbarium material includ-
ed under this name.
Leipoldt 760 (BOL, SAM) mentioned that
this species was a very valuable fodder plant.
This can be confirmed by signs of heavy brows-
ing visible on some herbarium specimens.
Common name: kapokbos.
ASTERACEAE: Eriocephalus
15
Vouchers: Acocks 18863 (PRE); Compton
3236 (BOL, NBG); Goldblatt 2119 (NBG,
PRE); Middelmost 1604 (NBG); Salter 7346
(BOL).
7. Eriocephalus pedicellaris DC., Prodro-
mus: 146 (1838). Type: Western Cape, ‘Klein
Namaqualand, bei Mierenkasteel, karrooartige
Hohe, 1000-2000 Fuss, August’, Drege 6733
(G-DC, holo.; G!, NBG!, P!, PRE, photo.!,
SAM!).
E. pteronioides DC.: 146 (1838). Type: Western Cape,
(Olifants River) ‘Ebenezar, auf steinigen trocknen (kar-
rooartigen) Hiigeln, unter 500 Fuss, Nov.’, Drege 6035 (G-
DC, holo.; P!, PRE, photo.!).
E. punctulatus DC. var. pedicellaris (DC.) Harv.: 201
(1865). Type: as for E. pedicellaris DC.
Many-stemmed, slender, erect shrubs, 0.4-0. 9
m high. Stems brittle; old stems deeply grooved
basally, displaying anomalous secondary
growth; young shoots with conspicuous purple-
brown stems, glabrous; older branches with yel-
low-brown to grey to dark grey bark, smooth to
shallowly grooved; brachyblasts 5 mm long,
short-lived. Leaves mostly opposite to suboppo-
site, scattered on young shoots, alternate on
flowering shoots, sessile on cushion-like thick-
ening on stem, blue-green, oblong-lanceolate to
linear, 12-30 x 0. 5-2.0 mm, those on brachy-
blasts slightly shorter than those on young
shoots, entire, rarely pinnatisect with 2 or 3
lobes apically, succulent, distal part almost
terete, adaxially proximally flattened to con-
cave, abaxially convex, surface with cavities,
glandular, glands in cavities, apex obtuse to
slightly acute; young leaves sparsely felty,
glabrescent. Capitula heterogamous radiate,
relatively large, 4-5 x 5-8 mm, in terminal
racemes or umbels, never solitary on brachy-
blasts or axillary; peduncles 10—30(^40) mm
long, slender, red-brown to purple-brown,
sparsely appressed pilose to glabrous. Involu-
cral bracts 5, ovate, with triangular, central,
brown, herbaceous part with white to pale
brown membranous margin, apex fringed.
Paleae: those of marginal florets connate with
Map 4. — • Eriocephalus pedicellaris; ■ E. aromaticus.
distinct septa indicating the different paleae, 5
mm long, abaxially densely lanate, hairs sep-
tate, adaxially smooth, apex fringed; those of
disc florets 5 x 1.5 mm, oblong, membranous,
apex long-fringed. Ray florets 2 or 3, female;
corolla white; tube 4-5 mm long, strap-shaped
part 3x6 mm, broadly cuneate, 3-lobed, much
longer than style branches. Style branches
strap-shaped, flattened, apices acute. Ovary
oblong to obovoid, slightly flattened. Seed flat-
fish, trigonous, smooth, 2—4 mm long. Disc flo-
rets 30-45, functionally male with sterile ovary,
infundibuliform, 5 mm long, red-purple, 5-
lobed. Stamens 5, exserted at maturity. Style
unbranched, truncate, with sweeping hairs.
Receptacle after anthesis with long hairs
between involucral bracts and connate marginal
paleae. Chromosome number: 2n = 72. Flower-
ing time : closely corellated with rainfall, which
extends from June to October with a peak from
July to September.
This species grows mainly on sandy soils
and rocky slopes with good drainage. It is
restricted to the winter-rainfall area with an
average of less than 200 mm per anum. The dis-
tribution extends from the Richtersveld to
Nieuwoudtville along the west coast. Plants
occur mostly singly or sparsely distributed and
rarely form dense homogenous stands. Map 4.
16
ASTERACEAE: Eriocephalus
Few herbarium specimens could be traced
under the name E. pedicellaris. Most material
was found under E. punctulatus (no. 9) or
misidentified as E. africanus (no. 14). Although
closely related to E. punctulatus , E. pedicellaris
can be distinguished from that species by the
long, blue-green, succulent leaves which turn
noticeably darker upon drying, the relatively
large capitula, 4-5 x 5-8 mm, on long pedun-
cles, 10-30(^4-0) mm, and by the brittle stems.
E. pedicellaris grows mostly at 300-600 m alti-
tude, but often below 300 m, in contrast to E.
punctulatus which grows at and above 600 m.
E. pedicellaris is a very palatable shrub,
which is heavily and selectively browsed wher-
ever it occurs, possibly because of its succulent
leaves and soft shoots, in contrast to E. punctu-
latus, which is hardly browsed. Common name:
kapokbos.
Vouchers: Barker 7414 (BOL, NBG); Miiller
3576 (WIND); Oliver, Tolken & Venter 616
(PRE); Van Breda 4081 (PRE); Van Jaarsveld
6234 (NBG).
8. Eriocephalus aromaticus C.A.Sm. in
Kew Bulletin 1931: 100, 101 (1931). Type:
Western Cape, Laingsburg Division: slopes of
the Witteberg, 975 m, October, Compton 2681
(K, holo.!; BOL!).
Erect, much-branched shrubs up to 0.6 m
high. Old stems and branches dark brown, lon-
gitudinally grooved, displaying anomalous sec-
ondary growth, glabrous, rigid; young shoots
red-brown, thin, straight, internodes relatively
long, initially felty, glabrescent except for
dense, white felt in leaf axils. Leaves decussate,
oblong to linear-oblong, 2^4(-9) x 0.3-0. 6 mm,
entire, basally broadly amplexicaul, adaxially
basally concave, slightly flattened towards
apex, abaxially convex, semiterete, whole sur-
face with cavities, sometimes with glands in
cavities, shiny, glabrous except for felty, axil-
lary buds, apex mucronate; leaves of
dolichoblasts and brachyblasts of same size and
shape, those of brachyblasts imbricate. Capitula
heterogamous radiate, racemose or umbellate-
racemose, small, 4.0 x 3.5 mm; peduncles
3— 5(— 1 2) mm long, glabrous to sparsely shortly
pilose. Involucral bracts 4, 2 slightly keeled, 2
ovate, 2.5 x 2 mm, other 2 broadly ovate, 2.5 x
3.5 mm, with central part green to purple with
broad membranous margin, abaxially glandular,
apex slightly fringed. Paleae : those of ray flo-
rets connate to ± one third of their length, cori-
aceous, rigid, margins fringed, abaxially long-
lanate. hairs septate, adaxially glabrous, shiny;
those of disc florets oblong-linear, 2 x 0.5 mm,
membranous, transparent, weakly keeled to
flattened, margins fringed, abaxially long-
lanate, adaxially smooth. Ray florets 2 or 3,
female; corolla white, up to 6.5 mm long; lami-
na up to 3 mm long, broadly cuneate, 3-lobed,
with or without glands. Style branches at most 1
mm long, tapering (acuminate), only apices
exserted. Ovary (and cypsela) oblong-ovoid.
Seed slightly flattened, obovoid, up to 3 x 1
mm. Disc florets 13-18, functionally male with
sterile ovary, 3-4 mm long; corolla infundibuli-
form, distal widened part red-purple, 5-lobed,
lower part yellow. Stamens 5, ± as long as
corolla tube or exserted only up to 0.5 mm.
Style truncate. Receptacle after anthesis densely
long-sericeous between involucral bracts and
connate marginal paleae. Chromosome number:
2 n - 18. Flowering time: (May to) June to
October (to November), correlated with rainfall
which varies in this area from 350-600 mm per
annum.
E. aromaticus is restricted to the mountains
of the winter-rainfall area of the Western and
Eastern Cape, higher than 900 in above sea
level. It never occurs in dense stands, but is
rather sparsely distributed. The distribution of
E. aromaticus and E. punctulatus (no. 9), a
related species, overlaps only in the Witteberg
and Klein Roggeveld Mountains. Map 4.
In the past, E. aromaticus was often con-
fused with E. punctulatus, a closely related
species. It can be easily distinguished from that
species by the consistently opposite leaves,
even on flowering shoots. On brachyblasts the
leaves are distinctly decussate. The otherwise
ASTERACEAE: Eriocephalus
17
glabrous leaves have a felty floccose indumen-
tum in the axil. In his original description,
Smith (1931) mentioned that the paleae of the
marginal florets were free. Although they look
free superficially, a thorough study showed that
they are connate from the base for about one
third of their length.
Common name: kapokbos.
Vouchers: Acocks 18420 (PRE); Compton
18405 (BOL, NBG); Hafstrom & Acocks 1556
(PRE); Hutchinson 450 (BOL, GRA, PRE);
Marloth 14160 (PRE).
9. Eriocephalus punctulatus DC., Prodro-
mus: 146 (1838); Harv.: 201 (1865). Type:
Northern Cape, ‘Namaqualand: Vorberge der
Camisberge, bei Kasparskloof, Elleboogfontein
und Geelbekskraal, 300-400 Fuss, August’,
Dr'ege 2734 (G-DC, holo.; PRE & WIND,
photo.!).
Slender, erect, sometimes spreading shrubs,
0.5-1. 5 m high. Old stems dark grey, display-
ing anomalous secondary growth; young
shoots green to dark red to golden brown,
dense felty indumentum with glands in cavi-
ties on leaves; older branches brown to brown-
grey to dark grey; branches thin with relative-
ly long internodes; brachyblasts short-lived,
up to 20 mm long. Leaves mostly opposite, but
alternate on flowering shoots, linear to almost
clavate, (2-)4-7(-28) x 0.3-0. 5 mm, entire to
pinnatisect with up to 3 lobes, adaxially flat-
tened, but concave towards base, abaxially
convex, bright green, initially with felty indu-
mentum, glabrescent, apex obtuse to acute,
base semi-amplexicaul. Capitula heteroga-
mous radiate, umbellate-racemose, terminal on
young shoots and brachyblasts, 3-4 x 3 mm;
peduncles (3— )5— 8(— 16) mm long, felty to
glabrous, thin (0.2 mm in diameter), longer
than subtending leaves. Involucral bracts 4 or
5, 2.8 x 1.7 mm, outer slightly keeled, inner
more flattened, with green, herbaceous, central
part and broad membranous margin. Paleae:
those of marginal flore's connate into a cylin-
drical sheath, 3 mm long, thickly coriaceous,
margins fringed, abaxially densely lanate,
hairs septate, adaxially glabrous; those of disc
florets oblong to linear, 2.6 x 0.7-2. 6 x 0.3
mm, margins fringed, abaxially densely lanate,
adaxially glabrous, membranous. Ray florets
1-3, female, 3. 9-4. 2 mm long with cuneate
lamina 1 .5-1.9 mm long, mostly white or
occasionally pale red-purple. Style branches
flattened, apices acute. Ovary oblong, slightly
flattened, with dense, lanate indumentum.
Seed flattish, trigonous, obovoid, 1.3-2. 1 mm
long. Disc florets usually 7-10, functionally
male with sterile ovary; corolla red-purple,
2. 4-3. 3 mm long; corolla tube trumpet-shaped
to infundibuliform. Style not branched, apex
globose, with sweeping hairs. Stamens 5,
exserted at maturity. Receptacle after anthesis
with long hairs between involucral bracts and
marginal, connate paleae. Chromosome num-
ber. 2 n = 36. Flowering time : correlated with
rainfall, May to October with a peak from July
to September.
The distribution of E. punctulatus extends
from Springbok in the Northern Cape along
the western parts of the Western Cape (the
Roggeveld and Witteberg Mountains) where it
overlaps with the related E. aromaticus (no.
8). It is found mostly in high-lying mountain-
ous localities, above 300 m altitude. Although
the distribution is limited to the winter-rainfall
area, there are indications that the amount of
rain influences the growth form. In veld types
such as Namaqualand Broken Veld, Succulent
Karoo and Mountain Renosterbosveld (Acocks
1975) with an annual rainfall of 200 mm and
less, slender, rigid, erect shrubs with relative-
ly short leaves (2-7 mm long) and often shiny,
red stems are found. In veld types with an
annual rainfall of more than 200 mm, for
instance Fynbos and Coastal Renosterbosveld
(Acocks 1975), a strongly branched, slightly
spreading, open, bushy shrub with relatively
long leaves (4-28 mm long) and long, droop-
ing shoots is found. However, these two forms
cannot be separated from each other as there
is no clear-cut transition. Map 5.
18
ASTERACEAE: Eriocephalus
Map 5. — ■ Eriocephalus punctulatus; • E. tenuifolius.
In E. punctulatus the subtending leaves are
shorter than the peduncle and therefore the
umbellate racemes are very obvious. In con-
trast. in the closely related E. tenuifolius (no.
10) the subtending leaves are as long as or
longer than the peduncles, which are thus hid-
den among subtending leaves. E. punctulatus
has mostly 7-10 disc florets while E. tenuifolius
has mostly 13-22.
Common name: kapokbos.
Vouchers: Goldblatt 2404 (NBG); Leistner
336 (PRE); Muller 4072 (WIND); Oliver 4415
(NBG. PRE); Verdoom 1896 (BOL, PRE).
10. Eriocephalus tenuifolius DC., Pro-
dromus: 146 (1838). Type: Eastern Cape,
‘Sneeuwbergen, auf steinigen Hiigeln und an
trocknen Abhangen, 4000-5000 Fuss, Sep-
tember’, Drege 2139 (G-DC, holo.; G!, NBG!,
P!, PRE, photo.!).
E. punctulatus DC. var. tenuifolius (DC.) Harv.: 201
(1865). Type: as above.
Rigid, erect, many-stemmed shrubs, 0.3-1. 3
m high. Old stems displaying anomalous sec-
ondary growth, dark grey to brown-grey; young
shoots rigid, firm, chestnut-brown, sparsely
felty hairy, glabrescent, densely leafy. Leaves
opposite, but sometimes alternate on flowering
shoots, linear, 4—14(-24) x 0.4-0. 6 mm, entire,
adaxially flattened, concave towards base,
abaxially convex, keeled towards apex, pale
green to yellow-green, greenish shiny white,
glandular, glands in cavities on leaf surface,
smooth, apex acute, base semi-amplexicaul;
opposite leaves basally connate; young leaves
sparsely felty, glabrescent. Capitula heteroga-
mous radiate, in umbellate racemes, terminal on
dolichoblasts or on brachyblasts, 3-4 mm long;
peduncles as long as or shorter than subtending
leaves, rarely longer, (3-)4—7(-10) mm long,
felty to glabrous. Involucral bracts 5 (rarely 4),
3.2 x 2.2 mm, central triangular to spathulate
part green, herbaceous with broad, membranous
margin, keeled to slightly flattened, sparsely
felty to glabrous, central part containing cavi-
ties with glands. Paleae: those of marginal flo-
rets connate into cylindrical sheath, 4 mm long,
thickly coriaceous, abaxially lanate, hairs sep-
tate; those of disc florets lanceolate, 4 x 1.3-1 x
0.3 mm, membranous, margins fringed, abaxial-
ly densely lanate. Ray florets 2 or 3, female,
3. 5-5. 5 mm long; lamina cuneate, 3- or 4-lobed,
3. 5-5. 5 mm long, white, sometimes with red-
purple tinge. Style branched; branches flattened,
linear, up to 1.8 mm long, apex acute. Ovary
oblong, slightly flattened, long-lanate. Seed
oblong, 2-3 mm long. Disc florets usually
13-22, functionally male with sterile ovary,
3.CM-.5 mm long; corolla red-purple, trumpet-
shaped, 5-lobed. Style not branched, apex glo-
bose, with sweeping hairs. Stamens 5, slightly
exserted at maturity. Receptacle after anthesis
with dense, white, long-pilose indumentum
between involucral bracts and connate marginal
paleae. Chromosome number, unknown. Flower-
ing time : closely correlated with rainfall, occur-
ring from January to almost October, with
peaks in January to April in summer-rainfall
area and July to September in winter-rainfall
area.
This species is part of the vegetation of the
mountains and hills of southern Mpumalanga,
the Free State, Lesotho and the Northern,
Western and Eastern Cape. Most of the distribu-
tion area receives summer rain. Map 5.
ASTERACEAE: Eriocephalus
19
Although E. tenuifolius is closely related to
E. punctulatus (no. 9), the two species can be
separated by their leaf size, peduncle length, the
length of the subtending leaves of the peduncles
and their distribution. A further distinguishing
character is the fact that E. punctulatus is hard-
ly browsed in contrast to E. tenuifolius, which is
heavily browsed (Smith 1966).
In the past, the leaves were used as substitute
for buchu by the Griquas, hence the common
name boegoekapok (Smith 1966). Another com-
mon name is klein-bergkapokbossie.
Vouchers: Dieterlen 435 (GRA, PRE, SAM);
Muir 7764 (PRE); Muller 4081 (WIND); Smith
4478 (BOL, PRE); Thode 7943 (NBG).
1 1 . Eriocephalus klinghardtensis M.A.N.MUI-
ler, sp. nov., E. africani L. et E. scariosi DC.
affinis sed foliis semper oppositis vel subop-
positis et dense velutinis differt.
Type: Namibia, Klinghardt Mountains in
Diamond Area No. 1, Muller 695 (WIND, holo.;
M, PRE).
Many-stemmed, much-branched, bushy,
aromatic shrubs, 0.35-0.6 m high, 0.5 m in
diameter. Old stems grey-black to almost black,
displaying anomalous secondary growth;
young shoots yellow-brown to brown-purple,
densely felty; older shoots glabrescent, brown
to grey-brown to grey-black, sometimes
spinescent. Leaves opposite to subopposite,
even on flowering shoots, sessile on cushion-
like permanent thickening on stem, linear to
clavate, semisucculent, 5— 10(— 17) x 0.7-1. 2
mm, entire, by exception dentate with at most 2
teeth, silvery grey, adaxially flattened, slightly
concave towards base, abaxially convex or
semiterete, densely felty to felty sericeous,
apex obtuse, base hardly broader than rest of
leaf; older leaves glabrescent but never totally
glabrous; leaves on brachyblasts densely imbri-
cate without distinct decussate arrangement as
in rest of genus. Capitula heterogamous radi-
ate, mainly in terminal, umbellate racemes,
rarely racemose, on dolichoblasts, 4-6 mm
long; peduncles 7-10 mm long, permanently
felty. Involucral bracts 4, 2 strongly keeled and
2 slightly flattened, lanceolate to ovate to obo-
vate, 3. 2-3. 6 x 1. 5-3.0 mm, with central,
green, herbaceous strip and relatively broad,
purple to light brown to transparent, membra-
nous margin, abaxially permanently felty
sericeous, sometimes glabrescent. Paleae :
those of marginal florets connate into cylindri-
cal, coriaceous sheath, up to 4 mm long, mar-
gins and abaxially long-lanate, hairs septate;
those of central florets lanceolate to spathulate,
3. 0-3. 5 mm long, membranous, apices fringed,
abaxially long-lanate. Ray florets 2 or 3,
female, 3 mm long; corolla white, lamina
broadly cuneate, 2. 0-2. 5 x 3-4 mm, 3-lobed,
glandular below. Style branches up to 1.5 mm
long, flattened, linear, apex acute. Ovary (and
cypsela) oblong, long-lanate. Seed obovoid,
1.5-2. 2 mm long. Disc florets 12-15, function-
ally male with sterile ovary, 3. 6-4.0 mm long;
corolla tubular, widening distally, 5-lobed,
basal part creamy white, distal part red-purple,
abaxially glandular. Style cylindrical, apex glo-
bose, with sweeping hairs. Stamens 5, exserted
at maturity. Receptacle after anthesis with
dense, white, long-lanate indumentum between
involucral bracts and connate marginal paleae.
Flowering time : correlated with winter rainfall,
with a peak from June to August. Figure 2.
E. klinghardtensis is restricted to the
Klinghardt Mountains, an isolated mountain
range within the Desert and Succulent Steppe
(Giess 1971) of Namibia. This region receives
an average annual winter rainfall of less than
100 mm, supplemented by fog from the ocean
at night. Although restricted in distribution, this
sometimes weakly spinescent shrub is relative-
ly common and grows in association with E.
giessii (no. 19). Map 6.
E. klinghardtensis is closely related to E.
africanus (no. 14) and E. scariosus (no. 13)
from which it can be distinguished by the con-
sistently opposite leaves covered with a dense,
felty indumentum.
20
ASTERACEAE: Eriocephalus
Figure 2. — Eriocephalus klinghardtensis: A, flowering shoot with inflorescences, x 1; B, branch with leaves, x 2; C,
capitulum, X 6; Dl, D2 involucral bracts, x 10; E, connate marginal paleae, x 4; F, central palea, x 4; Gl, ray floret, x 8;
G2, branched style, x 16; HI, disc floret, x 8; H2, anthers, x 16; H3, style, x 16; I, indumentum, x 32 (Muller 695, WIND).
ASTERACEAE: Eriocephalus
21
Map 6. — • Eriocephalus klinghardtensis; ■ E. brevi-
folius.
Common name: kapokbos.
Vouchers: Dinter 3935 (BOL, SAM, Z);
Merxmiiller & Giess 32159 (M, WIND); Muller
3371 (WIND).
12. Eriocephalus brevifolius (DC.)
M. A. N. Muller, comb, et stat. nov.
E. punctulatus DC. var. brevifolius DC., Prodromus:
146 (1838). Type: Northern Cape: Namaqualand,
Modderfonteinsberg, Kamiesberge, Drege 6037 (G-DC.
holo.; PRE. photo.!, SAM!).
Erect conical shrubs up to 1 .2 m high. Old
stems grey-brown to grey-black, displaying
anomalous secondary growth, glabrous; young
shoots grey, felty to shortly sericeous. Leaves
opposite except flowering shoots where they are
sometimes alternate, entire, linear, clavate,
3.0^4.5(-15.0) x 0.8-1. 2 mm, with felty to
shortly sericeous, permanent grey-green indu-
mentum; long leaves on dolichoblasts slightly
falcate to the inside, apex obtuse; short leaves
on brachyblasts decussate, imbricate, ± navicu-
late. Capitula heterogamous radiate, solitary on
brachyblasts or in umbellate racemes, terminal-
ly on dolichoblasts and brachyblasts, 4-5 mm
long; peduncles 5— 10(— 20) mm long, perma-
nently felty. Involucral bracts 5, broadly lanceo-
late to ovate, flattened, 3. 0-3. 5 x 1. 3-2.0 mm,
central part triangular, green with broad, mem-
branous, straw-coloured to red-purple margin,
abaxially appressed, shortly sericeous. Paleae:
those of marginal florets partly connate, form-
ing cylindrical sheath with free lobes, sheath
abaxially long-lanate, hairs septate, adaxially
glabrous, 4-5 mm long; those of disc florets
narrowly lanceolate, flattened, 2. 2-5.0 x 0.1-
0.3 mm, transparent, membranous, apices
fringed, only abaxially long-lanate. Ray florets
2 or 3, female, 3. 0-3. 5 mm long, with conspicu-
ous 2. 6-3. 2 mm long, white lamina. Ovary (and
cypsela) oblong, flattish, trigonous, densely
long-lanate. Seed obovoid, slightly flattened.
1.3-2. 1 mm long. Disc florets (7— )10— 14(— 16),
functionally male with sterile ovary, 4. 0-4. 5
mm long; corolla red-purple, trumpet-shaped,
5-lobed, 4. 0-5. 5 mm long. Stamens 5, ± as long
as corolla tube, exserted 0.5 mm at most. Style
truncate, rarely with sweeping hairs. Receptacle
after anthesis with dense, brown, long-pilose
indumentum between involucral bracts and
connate marginal paleae. Chromosome number.
2 n = 54. Flowering time : correlated with the
rainy season, reaching a peak from July to
September.
E. brevifolius is one of the more poorly col-
lected species and initially all the known mate-
rial came from Namaqualand. Its occurrence in
the Swartruggens-Roggeveld and Swartberg
Mountains was subsequently established and it
is possible that it has an even wider distribution.
It occurs at altitudes above 900 m, mainly in the
winter-rainfall area. Map 6.
Although related to E. africanus var. panicu-
latus (no. 14b), it can be distinguished from that
taxon by the dense, felty, sericeous indumentum
of the leaves resulting in a grey-green appear-
ance, as opposed to the silvery white appear-
ance of E. africanus var. paniculatus. The capitu-
la have a light brown, long-pilose indumentum
in contrast to the white, long-pilose indumen-
tum of E. africanus var. paniculatus. E. afri-
canus var. paniculatus is hardly or not browsed,
while E. brevifolius is readily browsed where it
occurs.
22
ASTERACEAE: Eriocephalus
The permanent short-sericeous indumentum
of the peduncles together with the felty
sericeous indumentum giving the leaves a grey-
green colour, shows a close relationship with E.
africanus var. paniculatus , E. capitellatus (no.
3), E. scciriosits (no. 13) and E. klinghardtensis
(no. II). E. punctulatus (no. 9), on the other
hand, has a sparse felty indumentum, glabres-
cent with smooth, shiny leaves and numerous
multicellular glands in cavities on the leaf sur-
face. The distinct large capitula of E. brevifolius
(up to 3.5 mm long) with ( 7—) 1 0— 1 4(— 1 6) disc
florets show closer relationship with E.
africanus var. paniculatus Group IIA (see p. 27)
(up to 3.5 mm long) with ( 8—) 11—1 4(— 27 ) disc
florets than with E. punctulatus (capitula up to
2.8 mm long and with 7-10 disc florets). The
indumentum alone shows that E. brevifolius is
not related to E. punctulatus , but rather to E.
africanus. It is therefore incomprehensible why
De Candolle (1838) described E. brevifolius as
a variety of E. punctulatus.
Common name: kapokbos.
Vouchers: Bond 1702 (NBG); McDonald
669 (NBG); Muller 3563 (WIND); Rosch & Le
Roux 450 (PRE).
13. Eriocephalus scariosus DC., Prodro-
mus: 147 (1838); Harv.: 202 (1865). Type:
Northern Cape, ‘Namaqualand, zwischen Nat-
voet und Gariep’, Drege 2738 (G-DC, holo.
(fragment); G!, NBG!, P!, PRE, photo.!,
SAM!).
E. scariossisimus S. Moore: 1019 (1904); Merxm.: 62
(1967). Typo: Namibia: ‘Groot Namaland’, farm Inachab,
Dinier 33 (BM. holo.!; Z!).
E. rangei Muschl. in Dinter: 260 (1921); Range: 56
(1935). Type: Namibia. Garub, Range 512 (B. holo.f;
SAM!).
E. virgatus Dinler: 87 (1932). Type: Namibia:
'Kamcllager'. north of Aus. Dinler 3676 (B. holo.t: BOL!.
NBG!. PRE!, SAM!, WIND!. Z!).
Slender, erect, much-branched, almost ever-
green. strongly aromatic shrubs, 0.5-1. 5 m high,
1-2 m in diameter. Old stems dark brown,
deeply grooved; older branches yellow-brown,
fairly smooth to slightly grooved, thin, glabrous;
dolichoblasts green-yellow, cylindrical, smooth,
sparsely to densely sericeous with sessile
glands; brachyblasts short-lived, at most 2 mm
long. Leaves alternate, sparse on dolichoblasts,
dense on brachyblasts, densely sericeous to
glabrous, leaves of dolichoblasts and brachy-
blasts of the same size and length, linear-lanceo-
late, 4-12 x 0.5-1. 5 mm, entire, semisucculent,
silvery white to green-grey to bright green,
basally adaxially concave, abaxially convex,
with glands in cavities on leaves. Capitula het-
erogamous radiate, solitary on brachyblasts or
racemose, terminal on flowering shoots, 4—6 x
3-6 mm; peduncles slender, 6-12 mm long,
sericeous. Involucral bracts 4 or 5, oval, 4x3
mm, with a green central strip surrounded by a
broad membranous margin. Paleae: those of
marginal florets connate, 4 mm long, sometimes
only partially connate, fringed margins of con-
nate paleae intertwined, abaxially densely long-
lanate, hairs septate; those of disc florets nar-
rowly linear, 4.0 x 0.5 mm, membranous, mar-
gins fringed. Ray florets (1)2(3), 4 mm long,
female; corolla white, tubular, lamina strap-
shaped to narrowly cuneate, 3-lobed or 3-den-
tate, up to 6 x 2 mm, much longer than flattened
style branches. Ovary (and cypsela) oblong, flat-
tened. Seed obovoid, flattened, 2.5 mm long.
Disc florets 4-9, 4-5 mm long, functionally
male, with sterile ovary; corolla tubular, 5-
lobed, glandular abaxially. Sty’le cylindrical,
apex globose, with sweeping hairs. Stamens 5,
distinctly exserted at maturity. Receptacle after
anthesis with dense, white, long-pilose indu-
mentum between involucral bracts and connate
marginal paleae. Chromosome number. 2 n = 72.
Flowering time: correlated with rainfall, varying
from December to April and June to September.
The distribution of this species extends over
both summer- and winter-rainfall areas. It
grows on mountains and hills but never on open
plains. It forms part of the flora of the sandstone
hills and mountains extending from the Namib-
Naukluft Park southwards to the Orange River.
Map 7.
ASTERACEAE: Eriocephalus
23
Map 7. — ■ Eriocephalus scariosus; • E. africanus var.
africanus.
E. scariosus is probably the most aromatic
species of Eriocephalus. The leaves are semisuc-
culent and, like those of the related E. africanus
(no. 14), have a very variable indumentum. The
plants occurring furthest north in Namibia, thus
within the summer-rainfall zone, are relatively
sparsely sericeous to glabrescent. Plants growing
along the Orange River within the winter-rainfall
area, have a dense, sericeous indumentum, giv-
ing it a silvery white appearance. Intermediate
forms are found scattered irregularly throughout
the distribution area. As a result of the variation
in indumentum, the appearance of the plants
varies from bright green to silvery white. Similar
variation to that found in the indumentum occurs
in the shape and length of the lamina of the ray
floret. Moore (1904) separated E. scariosissimus
from E. scariosus inter alia on the grounds of the
variation in the lamina length of the ray florets.
They vary from narrowly to broadly oblong, nar-
rowly to broadly cuneate and 3-dentate to 3-
lobed. In fresh material the colour of the rays is
pure white, but it changes to bright yellow upon
drying, as in E. macroglossus (no. 2).
Despite the strong aroma of E. scariosus, it
is eagerly browsed by domestic and wild ani-
mals. Common name: kapokbossie.
Vouchers: Dinter 6616 (BOL, NBG, SAM,
Z); Galpin 14141 (BOL); Giess & Miiller
14347 (M, WIND); Hall 4574 (NBG, PRE);
Muller & Tilson 910 (WIND).
14. Eriocephalus africanus L., Species
plantarum, edn 1: 1310 (1753); Hill: 225 (1759);
L.: 18 (1759); Burm.f.: 25 (1768); Houtt.: 428
(1775); Giseke: 12 (1779); Reichard: 938
(1780); Murray: 795 (1784); Lam.: 387 (1786);
Aiton: 278 (1789); J.F.Gmel.: 1277 (1792);
Lam.: t. 717, fig. 1 (1797); Thunb.: 168 (1800);
Willd.: 2384 (1803); Curtis: t. 833 (1805); Pers.:
497 (1807); Thunb.: 724 (1823); Spreng.: 621
(1826); G.Don: 364 (1830); Loudon: 1074
(1838); Loudon: 742 (1855); Harv.: 200 (1865);
Adamson & T.M. Salter: 800 (1950). Iconotypes:
Dill., Hortus elthamensis 132, t. 110, fig. 134
(1732); Hill: fig. 79 (1759).
E. corymbosus Moench: 590 (1794). Iconotype: as for
E. africanus.
E. variifolius Salisb.: 211 (1796). Iconotype: as for E.
africanus.
E. frutescens R.Br.: 180 (1813). Iconotype: as for E.
africanus.
E. septifer Cass.: 494 (1827); DC.: 145 (1838). Type:
Cape Province, collector unknown (G-DC, holo.; WIND,
photo.!).
E. septulifer DC.: 145 (1838). Type: Western Cape,
'Kaapsche Vlakte', Drege 6040 (G-DC, holo.; PRE &
WIND, photos!).
Much-branched, spreading to erect, conical
shrubs, 0. 3-0.9 m high, up to 4 m in diameter.
Old steins displaying anomalous secondary
growth, grey-brown to grey-black; young
shoots red-brown to grey-green and densely
leafy, silvery grey to green-grey, permanently
hairy to glabrescent, internodes relatively long;
older branches and stems yellow-brown to grey-
brown to brown to shiny red-brown, rigid or
slender. Leaves mostly opposite, sometimes
even in whorls of 3, alternate on flowering
shoots, sparsely spaced on dolichoblasts,
(5— )8— 1 7(— 40) x 0.4-2. 5 mm, palmatisect with
3-7 lobes or pinnatisect with 3 lobes distally or
3 lobes proximally or entire, adaxially flattened
but basally almost triangular to concave, abaxi-
24
ASTERACEAE: Eriocephalus
I k.i ki 3 —Eriocephalus africanus var. africanus: A. flowering shoot with inflorescences, x 1; ( Miiller 3624a,
WIND). E. africanus var. paniculatus: B, flowering shoot with inflorescences, x 0.6: C, branch with leaves, x 4: D, capitu-
lum * f HI. E2. involucral bracts, x 8; F. connate marginal paleae, x 5: G. central palea. x 5: HI. ray floret, x 8; H2,
branched style, x 25: 1 1 . disc floret, x 8: 12. style, x 25: 13. anthers, x 16 ( Miiller 3628, WIND).
ASTERACEAE: Eriocephalus
25
ally convex, succulent or not, blue-green to
grey-green to silver-grey, appressed silver-
sericeous to densely felty sericeous, permanent-
ly hairy to glabrescent, apex obtuse to acute;
lobes linear to clavate, straight or slightly
curved inwards. Capitula heterogamous radiate,
in terminal or lateral umbellate racemes or pa-
niculate, 3. 5-4.0 mm long; peduncles almost
absent to 26 mm long, permanently sericeous to
glabrous. Involucral bracts 4-6, oblong to
ovate to obovate to lanceolate, 2. 0-3. 5 x
1.0-2. 5 mm, central part green, herbaceous with
light brown to red-purple membranous margin,
margin fringed to entire, permanently sericeous
to glabrous. Paleae : those of marginal florets
connate, forming cylindrical sheath with free
apices, adaxially smooth or sometimes with
septa, abaxially long-lanate, hairs septate; those
of disc florets lanceolate to oblong, slightly flat-
tened, membranous, margins and abaxially
long-lanate. Ray florets 3-5, female, 2-4 mm
long; corolla distinctly strap-shaped; lamina
2-5 mm long, white to pale red-purple, cuneate
to broadly cuneate. Style branches flattened,
apices acute. Ovary oblong to obovoid, slightly
flattish, trigonous, long-lanate. Seed obovoid,
flattened, 1.3-2. 5 mm long. Disc florets 2-27,
functionally male with sterile ovary, 2-5 mm
long; corolla tubular to trumpet-shaped, red-
purple or yellow in proximal part and red-pur-
ple in distal part. Style unbranched, cylindrical,
globose, with sweeping hairs. Stamens 5.
Receptacle after anthesis with dense, white,
long-pilose indumentum between involucral
bracts and connate marginal paleae. Chromo-
some number. 2 n = 18, 36.
E. africanus is very widely distributed, oc-
curring in a variety of vegetation types. This
complex species shows much variation in life
form, leaf shape, indumentum and flower com-
position in capitula, while hibridisation also
seems to occur. Two well-demarcated groups
can be distinguished. The one group occurs in
the dune areas of the Coastal Fynbos, from sea
level to about 100 m inland or on rocks arising
from the sea. The plants therefore grow in soil
with a high salt content. They have succulent
leaves and a spreading habit. The second group
occurs at a higher altitude, further inland. The
plants are more erect, with thin, slender stems
and nonsucculent or only slightly succulent
leaves.
Two varieties are distinguished:
Leaves distinctly succulent, 3-5-7-lobed
or entire, (6.0— )8.2— 1 5.0(— 34.0) x
0.8-2. 5 mm; shrubs spreading, up
to 4 m in diameter; branches rigid;
exclusively coastal
14a. var. africanus
Leaves not or weakly succulent, mostly
entire, (5-)8-17(-40) x 0.4-0.8
mm; shrubs mostly erect, 0.3-0.6 m
in diameter; branches flexible;
inland 14b. var . paniculatus
Note: in his thesis, Muller (1988) distinguished
these two taxa as subspecies. It was decided,
however, to change them to variety level as they
occur in the same geographical area.
14a. var. africanus.
Spreading shrubs up to 4 m in diameter;
branches rigid, up to 4 mm thick. Young shoots
red-brown to grey-green depending on indu-
mentum, densely leafy. Leaves mostly opposite,
sometimes even in whorls of 3, alternate on
flowering shoots, sessile on permanent cushion-
like thickening, (6.0-)8.2-15.0(-34.0) x 0.8-
2.5 mm, palmatisect, 3-7-lobed or pinnatisect,
3-lobed distally or entire, slightly widening dis-
tally, succulent, blue-green to grey-green.
Capitula terminal, in umbellate racemes, rarely
paniculate; peduncles (3.0— )6.0— 8.5(— 1 2.0) mm
long, permanently felty sericeous. Ray florets 3
or 4(5), 3. 2-4.0 mm long; lamina white, broad-
ly cuneate, 4-5 mm long. Disc florets (12-)
1 6— 1 8(— 24). Flowering time : correlated with
rainy season, from July to September, but flow-
ers can be found throughout the year as plants
receive moisture from sea mist. Figure 3.
This variety is mostly restricted to the coast
of the Cape Peninsula, but also occurs from
26
ASTERACEAE: Eriocephalus
Mossel Bay to Knysna. The habitat extends
from the high-water mark to about 100 m inland
and on rocks arising from the sea. Map 7.
The spreading habit and succulent, palmati-
sect, blue-green to green-grey leaves with 3-5-7
lobes on firm, relatively thick shoots (up to 4
mm thick), are diagnostic characters of this vari-
ety. The growing points are normally enveloped
by older leaves, giving it a quadrangular appear-
ance. Eastwards between Mossel Bay and
Knysna the plants are not as distinctly succulent
as on the Cape Peninsula. The leaves are slight-
ly smaller but have the same typical palmatisect
shape with 3-5-7 lobes. The distribution areas
of var. africanus and var. paniculatus overlap
and possible hybrids were observed, e.g. Tyson
3048 (NBG), Tyson 3051 (SAM) and Pearson
225 (NBG). In the case of Esterhuysen 32159a
(BOL), the variations are clearly visible.
Although this is the oldest described species
of the genus Eriocephalus, it is not widely
known by the public and has few common
names. It has the oldest known common name,
namely clustery leaved scentwort (Hill 1759).
Other common names are kapokbossie and
wilde roosmaryn (Smith 1966).
Vouchers: Bolus 364 (BOL, PRE, SAM);
Breyer sub PRE23892 (PRE); Muller 3624
(WiND); Pillans 3632 (BOL, PRE); Wilman
PRE43623 (PRE).
14b. var. paniculatus (Cass.) M.A.N. Miil-
ler, P.PJ. Herman & H.H.Kolberg, comb, et
stat. nov.
E. paniculatus Cass., Dictionnaire des sciences
naturelles 50: 493 (1827). Iconotype: Gaertn., De fructibus
et seminibus plantarum 2,3: 428, t. 168, fig. 7 (1791).
E. racemosus Gaertn.: 428. t. 168, fig. 7 (1791) non L.:
1311(1 753); Jacq.: 1 57, 1 58, t. 1 1 , fig. 2 ( 1 796); Lam.: 4, t.
717, fig. 2 (1797). Type: based on that of var. paniculatus.
Monochlaena racemosus Cass.: 496 (1827). Type: based
on that of var. paniculatus.
E. umbellulatus Cass.: 493 (1827); Sch.Bip.: 12 (1844);
Levyns: 261 (1929): Marloth: 261 (1932). Type: based on
that of var. paniculatus.
E. umbellulatus Cass. var. glabriusculus DC.: 147
(1838); Harv.: 202 (1865). Type: Western Cape, Paarl,
Drege 87 (G-DC. holo.; G!, K!, P!. PRE & WIND, photos!).
E. umbellulatus Cass. var. argenteus DC.: 147 (1838);
Sch.Bip.: 676 (1844); Harv.: 202 (1865). Type: Cape
Province, Little Namaqualand (precise locality unknown),
Drege 2737 (G-DC, holo.; G!, P!, PRE & WIND, photos!).
E. sericeus Gaudich. ex DC.: 145 (1838); Sch.Bip.: 676
(1844); Harv.: 201 (1865). Type: Cape Province, collector
unknown (G-DC, holo.; PRE & WIND, photos!).
Erect to slightly spreading shrubs, up to 0.6
m in diameter, heaves mostly opposite, alter-
nate on flowering shoots, linear, (5— )8— 17(— 40)
x 0.4-0. 8 mm, not or weakly succulent, mostly
entire, sometimes 1- or 2-dentate to pinnatisect
with 3 lobes; lobes acicular to clavate, straight
or slightly falcate inwards, tapering from base
to apex, cushion-like thickening absent, silver-
grey sericeous. Capitula in terminal or lateral
umbellate racemes or paniculate; peduncles
almost absent to 26 mm long, permanently
sericeous to glabrous. Ray florets 3 or 4, 2. 0-2. 5
mm long, lamina white to pale red-purple,
cuneate to broadly cuneate, 2. CM-. 5 mm long.
Disc florets 2-27 . Flowering time: peaking
from July to September, but January to March
in summer-rainfall areas. Figure 3.
The distribution extends mostly over the
winter-rainfall area. After E. ericoides (no. 26),
E. africanus var. paniculatus is the taxon with
the widest distribution. It must be regarded as
the taxon with most potential for hybridisation,
as it occurs together with so many other species.
The distribution extends over the Northern,
Western and Eastern Cape. The distribution area
covers various veld types, e.g. Succulent Karoo,
Fynbos, Coastal Renosterbosveld and Fynbos,
Succulent Mountain Scrub, Karroid Broken
Veld, False Fynbos, Knysna Forest, Alexandria
Forest, Valley Bushveld, Noorsveld, False
Karroid Broken Veld and False Upper Karoo
(Acocks 1975). Such a large number of differ-
ent habitats show the potential of the taxon to
adapt to different soil types and altitudes. It fur-
thermore grows in both winter- and summer-
rainfall areas. Map 8.
ASTERACEAE: Eriocephalus
27
Map 8. — Eriocephalus africanus var. paniculatus.
The phenotypic plasticity of var. paniculatus
is high. The variety shows much variation in
habit, leaf size and shape, and degree of hairi-
ness. Some varying characters can be ascribed
to environmental influences, but others are
genetically determined. Some characters are
correlated with geographic distribution, but at
this stage it seems best not to distinguish formal
infravarietal groups as the morphological varia-
tion is continuous and it has not been studied in
detail. The problem can be solved only by inten-
sive population studies. It will be necessary to
cultivate plants from the different areas under
similar conditions.
Based on herbarium and field studies, the
taxon can be divided into the following five
groups.
Group I
Plants of this group have glabrescent leaves.
The glabrescence varies, even on one plant. It
seems that leaves produced during dryer sea-
sons are more densely sericeous and tend to be
less glabrescent, while those produced during
the active growing season and flowering time
tend to be more glabrescent. The distribution of
this group is mainly along the western interior
in the Calvinia area, Vanrhynsdorp to Citrusdal,
with a few scattered localities like Paarl and
Scheepersrus in the Western (southern) Cape
and Somerset East and Tarkastad in the Eastern
Cape. Chromosome number. 2 n = 36.
Group II
Two variants can be distinguished in this
group:
IIA: Shrubs with early glabrescent rigid
stems, sometimes with a glossy red-brown to
chestnut-brown colour. The leaves are fairly
short, grey-white sericeous. Capitula are borne
on shortly pedunculate, sometimes almost ses-
sile, umbellate racemes, alternate, terminally on
shoots. The inflorescences develop mostly ter-
minally on brachyblasts. The colour of the ray
florets varies from pure white to pale red-purple
to almost dark red-purple. The distribution of
this group extends from the low-lying parts of
the Bredasdorp coastal areas to the high-lying
mountainous parts of the Hantam, Cedar,
Langeberg, Waboom and Swartberg Mountains.
Chromosome number. 2 n = 18.
IIB: In the Langeberg Mountains at Kog-
manskloof and Baden, very slender shrubs with
very small leaves and almost sessile to shortly
pedunculate (1. 5-3.0 mm) capitula are found.
This variant shows a close resemblance to E.
capitellatus (no. 3) from which it differs in
indumentum and leaf shape. It might be a
hybrid between E. africanus (no. 14) and E.
capitellatus. Chromosome number: 2 n =18.
Group III
This group is found in the low-lying areas
(mostly below 300 m) from Humansdoip to
Bushmans River mouth and inland to
Grahamstown. The leaves are fairly short and of
the same length, (3.5— )4.2— 7.4(— 1 0.5) mm on
the entire plant. Plants at the coast have a
strongly flattened habit. The leaves are distinct-
ly silver-white-sericeous. The ray florets are
white and the capitula shortly pedunculate,
1 .0— 3.0(— 5.5) mm, borne terminally in umbel-
late racemes on flowering shoots. With a few
28
ASTERACEAE: Eriocephalus
Figure 4. — Eriocephalus grandiflorus: A, flowering shoot with inflorescences, x 1; B, branch with leaves, x 4: C,
capitulum, x 4; Dl, D2, involucral bracts, x 5; E, connate marginal paleae, x 5; F, central palea, x 5; Gl, ray floret, x 5;
G2, branched style, x 20; HI, disc floret, x 5; H2, anthers, x 20; H3, truncate style, x 20 (Muller 4040, WIND).
ASTERACEAE: Eriocephalus
29
exceptions, most capitula have four involucral
bracts and two ray florets. Chromosome num-
ber: unknown.
Group IV
This group is restricted mainly to the moun-
tainous regions of the Cape Peninsula, namely
Table Mountain, Devil’s Peak, Signal Hill and
Muizenberg. The plants are compact, much-
branched, slightly spreading, small shrubs with
long, nonsucculent leaves, densely silver-white
appressed sericeous in dense groups on brachy-
blasts and on young shoots. Capitula are fairly
large, borne mainly in umbellate racemes or in
pseudopaniculate racemes. The distribution of
this group borders on that of E. africanus var.
africanus (no. 14a) and hybridisation between
the taxa does occur. Leaves of the hybrid indi-
viduals vary from succulent to nonsucculent.
However, they all show the slightly spreading
habit of E. africanus var. africanus. Chro-
mosome number. 2 n = 36.
Group V
This group is found from the Cape Flats fur-
ther inland. The small shrubs have slender, thin
branches with sparsely leafy dolichoblasts and
long internodes. The growing points are
exposed as they are not enveloped by older
leaves. Capitula, which are smaller than those
of Group IV and with fewer rays, are densely
grouped on flowering shoots on long-peduncu-
late, umbellate racemes or terminally on
branches or brachyblasts in pseudopanicles.
Chromosome number. 2 n = 36.
Common names: kapokbossie, renosten>ekl-
kapok, roosmaryn, rosemary.
Vouchers: Acocks 12154 (PRE); Dyer 21
(PRE); Goldblatt 2152 (NBG); Taylor 10566
(NBG); Van der Merwe 123 (NBG, PRE).
15. Eriocephalus grandiflorus M.A.N. Mid-
ler, sp. nov., E. africani L. et E. eximii DC.
valde affinis, sed a E. africani foliis capitu-
lisque majoribus, radiis distinctis, habituque
multi ramoso rigido differt; a E. eximio capitulis
pedunculatis differt.
Type: Western Cape, 1 1 km N of Matjies-
fontein, Muller 4074 (NBG, holo.; PRE, WIND).
Robust, rigid, spinescent, much-branched
shrubs, 200-450 mm tall. Old stems and branch-
es grey to grey-black; young shoots chestnut-
brown, initially densely silvery sericeous,
glabrescent, opposite branches forming an angle
of almost 180°. Leaves decussate, rarely alter-
nate on some flowering shoots or clustered on
brachyblasts, sessile on cushion-like thicken-
ings, 4. 5-9.0 x 1 .2-2.2 mm, entire, rarely with
single lobe, adaxially basally strongly concave
with glabrous, triangular, basal part where con-
secutive leaves press against each other, other-
wise densely appressed sericeous, distally weak-
ly concave to flattened, abaxially semiterete,
permanently silver-white sericeous, apex obtuse
to acute, leaf base slightly broadened. Capitula
heterogamous radiate, 4—7, terminally, umbel-
late or semi-umbellate, or solitary on brachy-
blasts, relatively large, 5-6 mm long; peduncles
4-10 mm long, densely appressed sericeous.
Involucral bracts 4 or 5, broadly ovate, 4.6 x 3.3
mm, triangular, herbaceous central part with
broad purple to red-purple membranous margin,
some slightly keeled, others flattened, appressed
sericeous, apex obtuse, rarely acute, slightly
fringed. Paleae: those of marginal florets con-
nate, 6.0-6. 5 mm long, 4-lobed, coriaceous,
hard, free apices fringed, abaxially long-lanate;
those of disc florets transparent, membranous,
lanceolate to narrowly linear, 5. 2-6.0 mm long,
outer slightly keeled, inner slightly flattened,
abaxially long-lanate, apex acute, fringed. Ray
florets 2-4, 4-6 mm long; lamina broadly
cuneate, 3- or 4-dentate or -lobed, 3.5 x 3. 5 — 4.2
mm, white or pale to dark purple. Style cylindri-
cal, forked, branches flattened, linear, acute,
2. 0-2. 6 mm long. Ovary (and cypsela) oblong to
oblanceolate, densely long-lanate. Seed slightly
flattened, obovoid, 3.0-3. 5 mm long. Disc flo-
rets 12-22, 5-6 mm long, functionally male
with sterile ovary; corolla trumpet-shaped to
infundibuliform, base creamy, widened distal
30
ASTERACEAE: Eriocephalus
Map 9. — • Eriocephalus grandittorus: ■ E. tenuipes;
▲ E. karooicus; ★ E. dinteri.
part red-purple, 5-lobed. Style unbranched, apex
slightly convex, with sweeping hairs. Stamens 5.
Receptacle after anthesis with dense, white to
light brown, long-pilose indumentum between
involucral bracts and connate marginal paleae.
Chromosome number. 2n - 54. Flowering time :
June to September. Figure 4.
This species is confined to the mountainous
area between the Roggeveld, Witteberg and
Swartberg Mountains. Map 9.
Closely related to E. africanus (no. 14) and
E. eximius (no. 4). Differs from E. africanus by
its larger leaves, larger capitula with large con-
spicuous ray florets and its much-branched rigid
habit. It differs from E. eximius by its peduncu-
late capitula.
It is one of the few species of Eriocephalus
that are highly palatable. E. africanus, in con-
trast, is not or hardly browsed. Common name:
kapokbos.
Vouchers: Acocks 14310 (PRE); Compton
11260 (NBG); Hanekom 464 (PRE); Levyns
11161 (BOL); Miiller 3610 (WIND).
16. Eriocephalus tenuipes C.A.Sm. in
Kew Bulletin 1931: 101, 102 (1931). Type:
Western Cape, Langkloof near Haarlem,
Fourcade 1334 (K, holo.!; BOL!).
Many-stemmed, slender, much-branched
shrubs, 0.4-1. 0 m high. Old stems displaying
anomalous secondary growth: young shoots
bright green to light brown; older branches
brown-black. Leaves mostly alternate, some-
times opposite near growing points and on
brachyblasts, sessile on cushion-like thicken-
ings, linear to linear-oblong to clavate, 3.5-10.5
x 0.6-0. 8 mm, entire, rarely pinnatisect with 1
or 2 lobes, adaxially flattened, slightly broad-
ened towards base and concave, abaxially con-
vex or semiterete, distally cylindrical, leaf sur-
face pitted and rough, glands in cavities, initial-
ly silvery sericeous, glabrescent and matt green,
turning olive-green to olive-green-brown to
almost black upon drying, apex obtuse to slight-
ly acute; leaves on dolichoblasts and on brachy-
blasts of equal length. Capitula heterogamous
radiate, 3. 5-5.0 mm long, in terminal and later-
al, umbellate racemes, each with 3-8 capitula;
peduncles slender, 5-7 mm long, longer than
subtending leaves on flowering shoots,
sericeous. Involucral bracts 4 or 5, elliptic-
oblong to broadly ovate, 1.8-2. 7 x 1.2-1. 7 mm,
with central olive-green herbaceous part and
broad membranous margin, abaxially sericeous
to glabrous, some bracts slightly keeled, others
flattened, margins fringed. Paleae : those of
marginal florets connate into cylindrical sheath,
2.5— 1.0 mm long, hard, coriaceous, apex and
abaxial surface long-lanate; those of disc florets
linear, 2.5 mm long, membranous, margins
fringed, abaxially long-lanate. Ray florets 2 or
3, female, up to 6.3 mm long; corolla with dis-
tinct white lamina, 3.4-4. 1 x 3.4-4.4 mm,
broadly cuneate, 3-lobed. Style branches linear,
flattened, 1. 4-2.0 mm long, apex acute. Ovary
oblong to obovoid, slightly trigonous. Seed flat-
tish, trigonous, 1 .7-2.0 mm long. Disc florets
8-12, 3. 0-6. 5 mm long, functionally male with
sterile ovary; corolla lube infundibuliform to
trumpet-shaped, yellow with red-purple tinge,
5-toothed. Style cylindrical, apex globose, with
sweeping hairs. Stamens 5, slightly exserted at
maturity. Receptacle after anthesis with long
hairs between the involucral bracts and connate
ASTERACEAE: Eriocephalus
31
marginal paleae. Chromosome number. 2 n = 36.
Flowering time: mainly June to September, but
January to March when it rains in summer.
The distribution area receives mainly winter
rain but often also summer rain. E. tenuipes is
restricted mainly to the high-lying Langkloof
Mountains where E. capitellatus (no. 3) also
occurs. It seems that these two species
hybridise, but it has to be confirmed. Map 9.
It is surprising that Smith (1931) considered
E. tenuipes to be closely related to E. punctula-
tus (no. 9). The sericeous indumentum of E.
tenuipes rather points to a close relationship
with E. africanus (no. 14), especially var. pa-
niculatus (no. 14b). Some specimens of E.
africanus var. paniculatus are also glabrescent
like E. tenuipes, but the leaf surfaces do not
have the characteristic rough appearance of E.
tenuipes. The leaves of E. africanus var. pa-
niculatus are highly variable in length and
appear bright green to silver-white (depending
on indumentum) upon drying as opposed to
those of E. tenuipes which are all of the same
length and change to olive-green or olive-green-
brown upon drying.
Common name: kapokbos.
Vouchers: Compton 4229 (BOL, NBG);
Fourcade 5001 (NBG, PRE); Muller 4086
(WIND); Rycroft 2494 (NBG); Thode A2444
(PRE).
17. Eriocephalus karooicus M.A.N. Mul-
ler, sp. nov., E. spinescentis Burch, affinis sed
capitulis sessilibus radiisque prominentibus dif-
fert.
Type: Free State, Fauresmith Botanical Re-
serve, Smith 4531 (BOL, holo.; PRE).
E. spinescens sensu DC.: 147 (1838), pro parte; sensu
Harv.: 203 (1865), pro parte.
Many-stemmed, much-branched, spinescent,
aromatic shrubs, 15CM-50 mm tall, spreading.
Old stems displaying anomalous secondary
growth, dark grey; young shoots yellow-brown,
shortly sericeous; older shoots brown-grey to
dark grey, terminally spinescent, 10-22 mm
long. Leaves \-4 mm long, always decussate,
sessile on cushion-like thickenings, entire, per-
manently silver- to green-grey sericeous, adaxi-
ally proximally concave, distally flattened,
abaxially semiterete, apex acute, base semi-
amplexicaul, bases of two opposite leaves con-
nate, leaves on brachyblasts obtuse-triangular to
lanceolate, densely imbricate, those on young
shoots linear to linear-lanceolate. Capitula het-
erogamous radiate, 2. 0^4. 5 mm long, sessile,
solitary, terminal on brachyblasts or spicate on
dolichoblasts, almost hidden among subtending
leaves. Involucral bracts 4, 3. 2-4.0 x 0.8-1. 4
mm, 2 slightly keeled with margins overlap-
ping, other 2 flattened; each with central thick-
ened green to purple part and transparent mar-
gin, abaxially appressed sericeous, adaxially
smooth. Paleae : those of marginal florets free,
oblong-ovate, 3.5 x 1.2 mm, hardened, keeled,
abaxially lanate, hairs septate, enveloping
female florets; those of disc florets narrowly
oblong to lanceolate, becoming narrower
towards centre, 3.8 x 1.0-3. 6 x 0.2 mm, with
membranous margins, abaxially lanate. Ray flo-
rets 2 or 3, 2. 5-2. 8 mm long; corolla white with
distinct 1.3-2. 2 mm long lamina. Style branch-
es linear, as long as or shorter than ray, rarely
longer. Ovary (and cypsela) oblong, slightly
flattened, densely lanate. Seed 2-3 mm long,
obovoid, slightly flattened. Disc florets 4-10,
2. 5-3.0 mm long, functionally male with sterile
ovary; corolla trumpet-shaped, 5-lobed, white
to pale yellow to yellow with red-purple lobes.
Stamens 5, exserted at maturity. Style un-
branched, apex truncate, with sweeping hairs.
Receptacle after anthesis white-lanate between
involucral bracts and marginal paleae. Chro-
mosome number: 2 n = 18. Flowering time: De-
cember to March in summer, July to September
in winter. Figure 5.
The distribution area receives both winter
and summer rain. E. karooicus is centred in the
Free State and bordering Northern and Eastern
Cape in the following veld types: False Karoo,
Cymbopogon-Themeda Veld, Transitional Cym-
32 ASTERACEAE: Eriocephalus
Figure 5. — Eriocephalus karooicus: A. Ilowering shoot with inflorescences, x 1.5; B, capitulum, x 3; C, involucral
bract, x 8; D, marginal palea, x 16; E, central palea, x 16; FI, ray floret, x 10; F2, branched style, x 15; Gl, disc floret, x
10; G2, anthers, x 12; G3, truncate style, x 16; H, indumentum, x 32 (Smith 4531 , PRE).
ASTERACEAE: Eriocephalus
33
bopogon-Themeda Veld, mainly on heavy soils,
and the Themeda Veld to Cymbopogon-Themeda
Veld Transition (Acocks 1975). Map 9.
This species is closely related to E.
spinescens (no. 30). When Burchell described
E. spinescens in 1822, he only mentioned the
spinescence and position of capitula, but not
the ray florets and whether the capitula were
sessile or not (‘branches spinescent terminally.
Capitula solitary, lateral’). A more complete
description of E. spinescens appeared in De
Candolle ( 1838). It was, however, described as
a plant with capitula solitary and sessile on ter-
minal brachyblasts and the corolla of the ray
florets strap-shaped and much longer than the
style. This description of De Candolle was
partly made from a second specimen men-
tioned by him, Drege 6041 (G-DC) from the
Sneeuw Mountains. This specimen is actually
aspecific and belongs to the current E.
karooicus (no. 17). It cannot be assumed that
De Candolle did not see Burchell’s specimen
as both are on one herbarium sheet housed in
the Herbarium, Conservatoire et Jardin
botaniques de la Ville de Geneve, under the
name E. spinescens.
Harvey (1865) made the same mistake as De
Candolle by, except for the type specimen of E.
spinescens (Burchell 1419) (G-DC) and Drege
6041 (G-DC) mentioned by De Candolle, quot-
ing Zeyher 279 (NBG) and Zeyher 858 (GRA,
NBG), which are all aspecific, under E.
spinescens. The species was classified under the
Phaenogyne (i.e. capitula with distinct ray flo-
rets, longer than the style and involucral bracts).
Again the description was partly based on rays
from the Drege specimen and partly on certain
other specimens actually representing the cur-
rent E. karooicus.
This misinterpretation by both De Candolle
and Harvey caused all the E. karooicus material
through the years to be misidentified as E.
spinescens (no. 30). Although these two species
look very similar superficially, the former can
easily be distinguished from the latter by the
sessile capitula and distinct ray florets.
Although the terminal spines are a conspicu-
ous character of E. karooicus, during years of
good rain, long shoots are produced without
these spines. Spines develop again later when
active growth slows down. As mentioned
before, spicate synflorescences are sometimes
produced terminally on young shoots.
These delicate, spinescent shrublets are very
similar to E. ericoides during the active grow-
ing period. They differ from that species by the
green-grey to silver-grey, small, imbricate
leaves looking remarkably like a feather. From
this stem the common names veerkapok,
veerkarookapok, veerkaroo. In areas where the
species occurs, plants form dense stands and are
eagerly browsed. They are known as excellent
fodder (according to Dr M.G.A. Henrici as
quoted by Mogg 13620, PRE). Smith 4333
(PRE) mentioned that specimens collected by
Amot in 1867 near Colesberg were locally
known as wilde dagga. Other common names
for this species in the Karoo and Fauresmith
area are: doringkapok(bossie), kleinkapok-
bossie, kleindoringkapokbos, silwerkapok-
bossie, veerkapok(bossie) and volstruiskapok
(Roux 1984; Smith 1966).
Vouchers: Badenhorst 78 (PRE); Brueckner
874 (BOL, PRE); Gilfillan in herb. Galpin 5539
(GRA, PRE); Muller 220 (NBG. PRE); Smith
4483 (PRE).
18. Eriocephalus dinteri S. Moore in
Bulletin de l'Herbier Boissier 2,4: 1018, 1019
(1904); Merxm.: 60 (1967). Type: Namibia,
Hereroland, Windhoek, Dinter 853 (BM. holo.!;
Z!).
E. pan’iflorus Dinter: 87, 88 (1932). Type: Namibia.
‘Gross-Namaland: Aus bei 1 400 m auf Granit in Bliite, 2
Juni 1922’, Dinter 3544 (B. holo.f: BOL!. PRE!. SAM!.
WIND!, Z!).
Slender, erect, many-stemmed, much-branched
shrubs, 0.3- 1.0 m high, 300-500 mm in diam-
eter. Old stems displaying anomalous secondary
growth, 10-20 mm in diameter, mostly grey-
brown, in some areas with grey-black bark;
34
ASTERACEAE: Eriocephalu:
dolichohlasts light yellow to yellow-brown,
slender, densely appressed sericeous, internodes
4—10 mm long. Leaves always decussate, green-
grey, linear to obtuse-triangular, scale-like,
entire, permanently appressed sericeous, adaxi-
ally basally concave, glabrous to middle, distal-
ly flattened, abaxially semiterete, distally
keeled, apex obtuse to slightly acute, base
0.25-0.50 mm wide, leaves on dolichoblasts
2.3— 5.0(— 1 3.0) mm long, those on brachyblasts
1 .2^4.4 mm long. Capitula heterogamous radi-
ate, terminal, racemose or umbellate-racemose
or solitary on brachyblasts, 3.6-4. 1 mm long;
peduncles 2. 3-8. 5 mm long, appressed silver-
sericeous. Involucral bracts 4, ovate, 1.7-3. 3 x
0.9-2. 4 mm, 2 slightly keeled, opposite,
enveloping other 2 flattened bracts, green, mar-
gin broad, membranous, shortly appressed
sericeous. Paleae: those of marginal florets
free, keeled, margins fringed, 1 .8-2.5 x 0.6-0. 8
mm, each enveloping a single floret (mostly
female), hard, coriaceous, abaxially densely
long-lanate, hairs septate; those of disc florets
linear, fringed, 2.0 x 0.3 mm, membranous,
abaxially lanate. Ray florets (1 or) 2, 1.7-3. 3
mm long; corolla white to red-purple; lamina
conspicuous, irregularly 3-lobed, (0.8—) 1 .2—2. 1
mm long, ± 2 mm broad, obovate. Style branch-
es linear, 0.6-1. 2 mm long, much shorter than
lamina of ray floret. Ovary (and cypsela)
oblong, slightly flattened, densely woolly. Seed
obovoid, slightly flattish, trigonous, shorter
than 2 mm. Disc florets 2-5, functionally male
with sterile ovary, 1.6-3. 2 mm long; corolla
trumpet-shaped to cylindrical, 5-lobed, white to
creamy at base with red-purple margins.
Receptacle after anthesis with dense, white,
long hairs between involucral bracts and mar-
ginal paleae. Chromosome number. 2n = 36.
Flowering time: January to March and some-
times to May in the northern summer-rainfall
area, July to September and/or January to April
in southern parts receiving winter and summer
rainfall respectively.
The distribution is mainly restricted to sum-
mer-rainfall areas, although the southern
extremes of its distribution near Aus fall in the
transitional zone between summer and winter
rainfall. E. dinteri is endemic to Namibia and
has a restricted distribution, occurring only on a
few high mountains, e.g. the Brandberg, Auas
Mountains and Aus Mountains, above 1 000 m
altitude. Population density at the different
localities varies from scattered to rare, the plants
occurring nowhere in dense stands. Map 9.
Only a single record of a hybrid between E.
dinteri and E. luederitzianus (no. 25), Midler 52
(WIND), could be positively identified.
Hybridisation may possibly occur between E.
dinteri x E. merxmuelleri (no. 32) and E. dinteri
x E. ambiguus (no. 24), but it is difficult to con-
firm this.
Common name: kapokbos.
Vouchers; Hardy 1997 (PRE, WIND); Mul-
ler 51; 945 (WIND); Range 1117 (SAM);
Rennie in herb. Levyns 1973 (BOL).
19. Eriocephalus giessii M.A.N. Midler, sp.
nov., E. dinteri S. Moore affinis sed habitu
ramosissimo spinescenti; bracteis involucral-
ibus indumento permanenti longe sericeo ad
longe piloso tectis; floribus radii prominentibus
anguste oblongis differt.
Type: Namibia, Liideritz District, Farm Aar
LUS 16. ‘Am Quartzitberghang’, Giess 13383
(WIND, holo.; M, PRE).
Many-stemmed, much-branched, spinescent
shrubs, 350^450 mm tall, up to 450 mm in diam-
eter. Old stems much distorted at base, displaying
anomalous secondary growth, yellow-grey;
dolichoblasts yellow-brown, appressed seri-
ceous, glandular, glabrescent; older branchlets
yellow-grey to grey, glabrous; brachyblasts
short, with restricted growth. Leaves opposite on
dolichoblasts, distinctly decussate on brachy-
blasts, linear to obtuse triangular or keeled,
2. 5-4.2 x 0.4— 0.6 mm, entire, silver-grey, perma-
nently silver-sericeous, adaxially flattened, con-
cave to base, abaxially convex (semiterete), apex
acute, base semi-amplexicaul, broadened. Capi-
tula heterogamous radiate, mostly terminal, race-
ASTERACEAE: Eriocephalus
35
mose or umbellate-racemose on dolichoblasts,
also solitary or umbellate-racemose on brachy-
blasts, 3. 2-4.0 mm long; peduncles short,
2.5 — 4.0 mm long, permanently sericeous to long-
pilose. Involucral bracts 4, 2 keeled, other 2
slightly laterally flattened, 3.0-3. 5 x 1.2-1. 5
mm, red-purple to red-brown, transparent mem-
branous margin absent, permanently long-
sericeous to long-pilose, with subsessile, multi-
cellular glands. Paleae : those of marginal florets
free, totally enveloping marginal florets (usually
female), keeled, lanceolate to ovate, rigid, mem-
branous, 3. 2-3.6 mm long, margins and abaxial-
ly long-lanate; those of disc florets broad to nar-
rowly lanceolate, up to 3.2 mm long, membra-
nous, margins and abaxially long-lanate. Ray flo-
rets female, 2, 2. 2-2.4 mm long; corolla white,
lamina up to 2.4 mm long, 3-lobed to 3-toothed,
narrowly oblong, glandular abaxially. Style
branches flattened, linear, apex acute. Ovaiy
oblong to oblanceolate, long-lanate. Seed lanceo-
late, slightly flattened, 1.6-2. 2 mm long. Disc
florets 3-5, functionally male with sterile ovary,
2.4—3.4 mm long; corolla infundibuliform, 5-
lobed, yellow with pale purple limb or entirely
red-purple, with subsessile multicellular glands
abaxially. Style cylindrical, apex globose, with
sweeping hairs. Stamens 5, slightly exserted at
maturity. Receptacle after anthesis with dense,
white, long-hairy indumentum between involu-
cral bracts and free marginal paleae. Chromo-
some number. 2n = 18. Flowering time', corre-
lated with rainfall, January to April and July to
September, depending on time of rainfall. Figure
6.
The distribution area receives both summer
and winter rain. E. giessii is restricted to the
Liideritz District of Namibia and occurs in and
adjacent to Diamond Area No. 1 . Plants grow in
mountainous parts some 1 000 m above sea
level and occur scattered. Map 10.
E. giessii is related to E. dinteri (no. 18) from
which it can be distinguished by the much-
branched, spinescent habit, permanent, long-
sericeous to long-pilose indumentum on involu-
cral bracts and the large, narrowly oblong ray
florets.
16- -
18 — y- -
-V
20
22
24
\ - - -
- .
iff
\
—
26
28 —
__
30
Map 10. — ■ Eriocephalus giessii; • E. racemosus var.
racemosus.
Common name: kapokbos.
Vouchers: Merxmiiller & Giess 32248 (M,
WIND); Merxmiiller & Giess 32250 (M,
WIND); Muller 825 (WIND); Muller 3345
(WIND).
20. Eriocephalus racemosus L., Species
plantarum, edn 1: 1311 (1753); L.: 26 (1760);
Burm.f.: 25 (1768); Murray: 795 (1784); Lam.:
387 (1786); J.F.Gmel.: 1277 (1792); Thunb.:
168 (1800); Willd.; 2385 (1803); Pers.: 497
(1807); W.T.Aiton: 180 (1813); Thunb.: 724
(1823); Spreng.: 621 (1826); G.Don: 364
(1830); DC.: 147 (1838); Loudon: 742 (1855);
Harv.: 203 (1865); Adamson & T.M. Salter: 801
(1950). Type: Cape Province, precise locality
unknown, collector unknown (LINN 1040.3,
holo., microfiche!).
E. simplicifolius Salisb.: 21 1 ( 1796). Type: based on that
of E. racemosus.
E. spicatus Burm. ex DC.: 147 (1838). Type: Western
Cape, between Knysnadrif and Gowkamma-station,
Burchell 5605 (G-DC, holo.; GRA!. WIND, photo.!).
Many-stemmed, slender, erect shrubs,
1. 2-2.0 m high. Old stems displaying anoma-
lous secondary growth, brown-grey; young
shoots grey, internodes either short, densely
36
ASTERACEAE: Eriocephalus
Figure 6. — Eriocephalus giessii: A, flowering shoot with inflorescences, x I; B, branch with leaves, x 10; C, capitu-
lum, x 5; Dl , D2, involucral bracts, x 8; E, marginal palea, x 5; F, central palea, x 10; G I, ray floret x 8; G2, branched style,
x 25; HI, disc floret, x 8; H2, anthers, x 25; H3, style, x 25; I, indumentum, x 32 ( Giess 13383, WIND).
ASTERACEAE: Eriocephalus
37
leafy or relatively long with leaves scattered;
brachyblasts short-lived. Leaves alternate,
rarely opposite, sessile on cushion-like thick-
enings on stem, linear to narrowly lanceolate
or obtuse-triangular, 3-30 x 0.5— 1 .5(— 2.0)
mm, entire, succulent, permanently grey-felty,
adaxially flattened, concave towards base,
abaxially convex, apex acute. Capitula het-
erogamous disciform, racemose or paniculate,
2. 5-4. 8 mm long, sessile to distinctly pedun-
culate; peduncles 0-15 mm long, felty. In-
volucral bracts 4, 3 x 2 mm, central part
herbaceous, green, red-purple towards mem-
branous margin, abaxially felty, 2 slightly
keeled, 2 laterally flattened, margins enve-
loped by the two keeled bracts. Paleae : those
of marginal florets connate into cylindrical
sheath, basally slightly globose, apices and
abaxially long-lanate, hairs septate; those of
disc florets lanceolate to linear, 1. 5-2.0 mm
long, apices fringed, abaxially lanate, some-
times absent in central florets. Marginal
female florets 1-3; corolla white to pink, ob-
ovate, constricted around style; narrowed part
very short. Style almost totally exposed, cylin-
drical, forked; style branches flattened, linear,
up to 1 mm long, apices acute. Ovary (and
cypsela) ovoid-oblong, abaxially densely
lanate, slightly flattened, 3x2 mm. Seed
ovoid, laterally compressed, 1.5-2. 3 mm long.
Disc florets 4-21. functionally male with ster-
ile ovary; corolla ± 3 mm long, widening dis-
tally, glandular abaxially, yellow-green to yel-
low with purple-red margins, rarely entirely
purple-red. Style cylindrical, unbranched, apex
broad, with sweeping hairs. Stamens 5, exsert-
ed at maturity. Receptacle after anthesis with
dense, white, long-hairy indumentum between
involucral bracts and connate marginal paleae.
Chromosome number. 2 n = 36.
After E. africanus, E. racemosus is the old-
est known species of Eriocephalus. Although it
is relatively easy to distinguish E. racemosus
from related taxa, a few herbarium specimens
have been incorrectly identified as E. africanus.
These misidentifications can be ascribed to a
description and associated illustration of E.
racemosus by Gaertner (1791), based on ma-
terial of E. africanus, which were accepted by
Lamarck (1796) and Jacquin (1796).
Obvious differences in capitulum structure
together with differences in leaf shape, have led
to the recognition of two varieties. This division
is supported by chromosome morphology.
Note: in his thesis, Muller (1988) distin-
guished these two taxa as subspecies. It was
decided, however, to lower them to variety level
as they occur in the same geographical area.
Capitula sessile to very shortly peduncu-
late; peduncle up to 5 mm long;
disc florets (4 — )7— 9
20a. var. racemosus
Capitula distinctly pedunculate; peduncle
(5-) 10- 15 mm long; disc florets
13-21 20b. var. affinis
20a. var. racemosus.
Leaves linear. Capitula 2. 5-3. 5 mm long,
sessile to very shortly pedunculate (at most 5
mm long). Paleae : those of marginal florets
connate, thin, membranous; those of disc florets
weakly developed and often absent in central
florets. Marginal female florets (1)2. Disc flo-
rets (4— )7— 9, 2.4-3. 2 mm long. Flowering time:
June to September, but depending on rain, con-
tinuing until November. Figure 7.
With a few exceptions, var. racemosus is
almost always found near the coast. It extends
as far east as Port Elizabeth and west to
Lambert’s Bay. Map 10.
This variety forms a dense, compact shrub,
which is densely leafy at branch tips. The syn-
florescence is a dense, drooping, spicate
raceme. Capitula are mostly sessile. In Henrici
3721 (PRE) and Bohnen 409/3 (NBG) the capit-
ula are shortly pedunculate (less than 5 mm)
and they could possibly be transitional forms to
var. affinis. However, in both these specimens
the capitula contain fewer than 10 disc florets, a
feature which assigns them to var. racemosus.
ASTERACEAE: Eriocephalus
Figure 7. — Eriocephalus racemosus var. racemosus: A, flowering shoot with inflorescences, x 1 ; B. branch with leaves
x 1.5; C, capilulum. x 4; Dl, D2, involucral bracts, x 8; E, connate marginal paleae, x 12; F. central palea, x 10; Gl, mar-
ginal female floret, x 12; G2, branched style, x 35; HI, disc floret, x 12; H2, anthers, x 25; H3, style, x 25; I, indumentum,
x 40 (Muller 3634, WIND). E. racemosus var. affinis: J. flowering shoot with inflorescences, x 1 (Muller 4003, WIND).
ASTERACEAE: Eriocephalus
39
Doubtful cases with shortly pedunculate capitu-
la can therefore be positively identified by the
number of disc florets.
E. racemosus var. racemosus which forms
part of the Coastal Fynbos (Acocks 1975), can
survive veld fires and has become an invader in
certain areas. In some areas it is well browsed
but in other areas it is not utilised at all. Ac-
cording to Smith ( 1966) the wool is used by the
Cape Penduline Tit ( kapokvoeltjie ), Antho-
scopus minutus, for making its nest, and by
rural people for stuffing pillows. Common
names: sandveldkapok, strandveldkapok , rivier-
kapok (Bredasdorp area) and kapkappie.
Vouchers: Bolus 6323 (BOL. PRE, Z);
Boucher 3322 (NBG); Brown in herb. Rogers
29217 (GRA, NBG, PRE); Parker 3590 (NBG,
PRE); Van Breda 1635 (PRE).
20b. var. affinis (DC.) Harv., Flora capensis
3: 203 (1865). Type: Western Cape, ‘Langevalei,
an und in der Valei. unter 1000 Fuss, Juli’, Drege
2736 (G-DC, holo.; PRE, photo.!).
E. affinis DC.: 147 (1838). Type: as above.
Leaves semisucculent to succulent, linear.
Capitula 3. 8^4.8 mm long; peduncle (5-) 10-15
mm long. Paleae: those of marginal florets con-
nate into a hard coriaceous, tubular sheath; cen-
tral florets each enveloped by well-developed
palea. Marginal female florets (2)3. Disc florets
13-21, 3. 6-4.5 mm long. Flowering time : June
to September, but plants in flower collected as
early as April and as late as November, depend-
ing on rain. Figure 7.
The distribution of var. affinis extends from
near the coast to 50 km inland and it has been
collected from Hondeklip Bay to Melkbos-
strand. Most material has been collected away
from the coast, on sandy soil. It forms dense
stands, often dominating plant communities.
Map 11.
Common name: kapokbos.
Map 1 1 . — • Eriocephalus racemosus var. affinis: ▲ E.
decussatus: ■ E. kingesii.
Vouchers: Acocks 14527 (PRE); Hugo 2874
(NBG); Le Roux 2588 (NBG); Levyns 11690
(BOL); Muller 4003 (NBG, PRE. WIND).
21 . Eriocephalus decussatus Burch.. Travels
in the interior of southern Africa: 272 (1822);
G.Don: 364 (1830). Type: Northern Cape,
between Karree River and Klein Quaggas Fontein.
near Frazerburg, 24-26/8/1 811, Burcliell 1407 (K.
holo.!; PRE, photo.!).
E. aspalathoides DC.: 148 (1838): Harv.: 203 (1865);
non E. aspalathoides sensu Merxm.: 60 (1967). Type:
Western Cape, 'Zwischen Zwarteberg und Aasvogelberge,
Namaqualand'. Drege 2142 (G-DC, holo.; P!, PRE.
photo.!).
Shrubs much-branched from base, some-
times spinescent, 0.6- 1.5 m high and in diam-
eter; branches conspicuously opposite. Old stems
grey-black, displaying anomalous secondary
growth; dolichoblasts yellow-brown, silver-
sericeous; older branches brown-grey. Leaves
decussate, often alternate on flowering shoots,
imbricate, scale-like, triangular, all leaves short.
0.75—1 ,75(— 3.0) mm long, entire, permanently
densely appressed silver-sericeous, giving plant
silvery grey appearance, adaxially basally con-
cave, abaxially basally convex, apex acute, base
semilunate. Capitula heterogamous disciform,
solitary on brachyblasts, rarely in terminal
40
ASTERACEAE: Eriocephalus
racemes, 3. 5^4.0 mm long; peduncles (1.0-)
2.0-3.5(-6.0) mm, appressed sericeous. Invo-
lucral bracts 4, rarely 5, ovate, slightly acute,
3.5 x 2.4 mm, greenish purple to reddish purple
with silvery white sericeous indumentum, cen-
tral part thickened, margin narrow, membra-
nous. Paleae: those of marginal florets totally
connate, forming cylindrical sheath with 2-4
free apical lobes, coriaceous, free apices
fringed, abaxially long-lanate, hairs septate;
those of outer disc florets slightly keeled, hard,
coriaceous, those of central florets flattened,
membranous, 3.2 x 0.6 mm, free apices fringed,
abaxially long-lanate. Marginal female florets
2-4, creamy white, 3. 6-5.0 mm long, lamina
extremely short, 0.6- 1.2 mm, inconspicuous,
longer than point of furcation of style but short-
er than style branches, 3-lobed or -dentate. Style
branches strap-shaped. Ovary (and cypsela)
oblong, flattish, trigonous, long-lanate. Seed
1 .5-2.2 mm long, ovoid, flattened. Disc florets
(3— )5— 8(— 1 1 ), functionally male with sterile
ovary, 3. 7-4. 5 mm long; corolla tube trumpet-
shaped, basally cream-coloured, apex red-pur-
ple. Stamens 5, exserted at maturity. Style undi-
vided, cylindrical, apex truncate, with sweeping
hairs. Receptacle after anthesis with long hairs
between involucral bracts and marginal, con-
nate paleae. Chromosome number. 2 n = 18.
Flowering time : correlated with rainfall,
extending from January to April and from July
to September in the different rainfall regions.
The distribution of E. decussatus extends
over both summer- and winter-rainfall regions,
over the central Karoo and parts of Nama-
qualand, mostly on sandy soil. Although a large
number of capitula are produced, this species is
never found in dense stands but is rather scat-
tered. Map 1 1 .
E. decussatus is one of the species that,
because of incomplete descriptions coupled
with misinterpretations by later researchers,
kept taxonomists on the wrong track. Burchell
first collected it in 1811 and in 1822 a very
short, incomplete description was published. De
Candolle (1838) included it in his work, but in
synonymy under E. glaber (= E. ericoides), which
it superficially resembles, but from which it
differs in indumentum and connate marginal
paleae. Despite its obscurity to taxonomists,
Don (1830) mentioned it as an ornamental
known to gardeners in the United Kingdom.
Harvey (1865) did not mention it in his work
on Eriocephalus. Its synonym E. aspala-
thoides , as described by De Candolle, is better
known. It is ironic, however, that almost all
specimens identified with this name, do not
belong to this species but rather to E. ambiguus
(no. 24), E. luederitzianus (no. 25), E. nama-
quensis (no. 31) or E. microphyllus (no. 28), all
taxa distinct from E. decussatus. Acocks
(1975) had a realistic concept of this species
and identified it correctly in most cases. The
main reason for this confusion was Harvey’s
(1865) misinterpretation of De Candolle’s
(1838) description. Harvey (1865) described
the plants as subspinescent while De Candolle
(1838) explicitly described the plants as similar
to E. spinescens but almost without spines
(‘sed rami subinermes’). The spines referred to
by De Candolle are the hardened remains of the
terminal racemose peduncles that sometimes
occur (the capitula are mostly terminal on
brachyblasts).
Another misunderstanding causing confu-
sion and misidentifications was De Candolle’s
(1838) description of the leaves as opposite and
alternate, which should have been opposite with
alternate leaves on flowering shoots. Neither
Burchell (1822) in his original description of E.
decussatus, nor De Candolle (1838) in his
description of E. aspalathoides, mentioned the
connate paleae of the marginal florets, the most
important character distinguishing this taxon
from the closely related E. microphyllus.
Where E. decussatus and E. microphyllus
occur together, they resemble each other in
habit, but can be distinguished by the silvery
sericeous indumentum in E. decussatus, giving
the plants a silvery grey appearance, as opposed
to the blue-green to grey-green to bright green
colour of E. microphyllus.
Common name: kapokbossie.
ASTERACEAE: Eriocephalus
41
Vouchers: Acocks 19486 (PRE); Acocks
19487 (PRE); Leistner 481 (NBG, PRE);
Marloth 3355 (NBG); Miiller 3605 (WIND).
22. Eriocephalus kingesii Merxm. & Eberle
in Mitteilungen der Botanischen Staats-
sammlung, Miinchen 2: 321 (1957); Merxm.:
61 (1967). Type: Namibia: Liideritz, hills across
Nautilus, Kinges 2575 (M, holo.!; PRE!).
Robust, erect to spreading, many-stemmed,
much-branched shrubs, 0.3-0. 6 m high and in
diameter; branches rigid. Old stems displaying
anomalous secondary growth, 10-20 mm in
diameter; young shoots yellow-brown,
appressed sericeous; older shoots brown-grey to
grey-black; brachyblasts opposite, short-lived.
Leaves opposite, decussate, linear or naviculate,
on young shoots 6-12 x 1.0- 1.5 mm, scattered,
on brachyblasts 2-5 x 1.0-2. 5 mm, densely
imbricate, entire, semisucculent, adaxially flat-
tened, concave towards base, abaxially convex,
slightly keeled, surface smooth, blue-green to
yellow-brown, permanently densely sericeous,
apex obtuse, base semi-amplexicaul and with
cushion-like thickenings. Capitula heteroga-
mous disciform, relatively large, 4-8 mm in
diameter, solitary on brachyblasts or racemose,
terminal on young shoots, flowering shoots
rigid, thick; peduncles rigid, appressed
sericeous, 3-12 mm long. Involucral bracts 4 or
5, ovate, obtuse, 4x3 mm, appressed sericeous,
central part thickened, herbaceous, margin
broad, membranous, sometimes red. Paleae :
those of marginal florets mostly free, some-
times connate at base, ovate to oblong, 4-6 mm
long, slightly keeled, firm, apices fringed, abax-
ially lanate, hairs septate; those of central flo-
rets narrowly ovate to oblong, margins fringed,
abaxially long-lanate. Marginal female florets 1
or 2, 5-6 mm long; corolla 2- or 3-lobed, as
long as style, but usually shorter, sometimes
with scattered glands abaxially, white to
creamy. Style branches linear. Ovary (and
cypsela) oblong-obovoid, slightly flattish, trigo-
nous, 2-3 mm long. Seed ovoid, flattish, trigo-
nous, 2. 2-3.0 mm long. Disc florets 5-15, func-
tionally male; ovary sterile; corolla creamy to
yellow, 4.2-6. 5 mm long, 5-lobed. Style undi-
vided, truncate. Stamens 5, well exserted at
maturity. Receptacle after anthesis with dense,
light brown, long-hairy indumentum between
involucral bracts and marginal paleae. Chro-
mosome number. 2 n = 54. Flowering time : al-
most throughout the year, no peak flowering
time determined.
E. kingesii is endemic to Namibia and its dis-
tribution is limited to Liideritz and Diamond
Area No. 1, which falls in the winter-rainfall
region of the Desert and Succulent Steppe
(Giess 1971). Most localities are near the coast
and are subject to fog at night. The average
annual rainfall of this area is less than 50 mm.
Map 11.
These rigid, flat, spreading shrubs with
entire, succulent, silvery sericeous leaves are
fairly common in the areas where they occur.
This is the only Eriocephalus species with a
high seed set (90%) and germination percent-
age. At a temperature of 28°C, 16 of 20 seeds
germinated within 4 days — thus a germination
percentage of 80%.
Since succulence in coastal habitats is often
regarded as being determined by environment,
young and mature plants were cultivated under
uniform conditions at the Botanical Garden of
the University of Stellenbosch. Succulence
remained, leading to the conclusion that it is
genetically determined in this species as is the
case in E. africanus var. africanus (no. 14a),
which also occurs along the coast.
Common name: kapokbos.
Vouchers: Giess & Robinson 13236 (WIND);
Giess & Van Vuuren 686 (BOL, K, PRE,
WIND); Giess & Van Vuuren 707 (BOL, K,
PRE, WIND); Marloth 4764 (NBG); Merx-
miiller & Giess 3069 (M, WIND).
23. Eriocephalus pauperrimus Merxm.
& Eberle in Mitteilungen der Botanischen
Staatssammlung, Miinchen 2: 322 (1957);
42
ASTERACEAE: Eriocephalus
Merxm.: 61 (1967). Type: Namibia: Maltahohe
District, Farm Duwisib MAL 84/Farm Bliitputz
MAL 105/111, Volk 12666 ( M, holo.!; WIND!).
Erect to spreading, many-stemmed, much-
branched shrubs, 350-450 mm tall and in diam-
eter. Older stems displaying anomalous sec-
ondary growth, sometimes twisted and distort-
ed, grey to grey-black; young shoots felty,
glabrescent, whitish, sparsely leafy; brachy-
blasts short-lived, alternate. Leaves alternate,
densely imbricate on brachyblasts, scattered on
young shoots, linear, 4-8 x 0.5 mm, those on
brachyblasts 1-2 x 0.5 mm, entire, adaxially
slightly flattened, concave towards base, abaxi-
ally convex, slightly keeled towards apex, sur-
face pitted with glands in cavities, grey-white,
apex obtuse, base widened, semi-amplexicaul;
leaves at growing point cobwebby/felty,
glabrescent. Capitula heterogamous disciform,
terminal, spicate, 4-6 x 2-3 mm, alternate, ses-
sile, compact. Involucral bracts 4, narrowly
ovate, 4x2 mm, apex obtuse, narrow, herba-
ceous, green, central part with broad membra-
nous margin, glabrous, surface pitted, glands in
cavities. Paleae: those of marginal florets free,
lanceolate, 2.0 x 0.5 mm, membranous, margins
fringed, abaxially long-lanate, hairs septate;
those of disc florets narrowly lanceolate to lin-
ear, margins fringed, abaxially long-lanate.
Marginal female florets 1, 2. 0-2. 5 mm long;
corolla tubular with short, linear to narrow
cuneate lamina, yellow, glandular abaxially,
shorter than style branches but longer than point
of branching of style. Style branches flattened,
linear, 1.2 mm long, apex acute. Ovary (and
cypsela) obovoid, flattened, long-lanate, glan-
dular. Seed narrowly obovoid, slightly flattened,
1.5 mm long. Disc florets 1^4, functionally
male with sterile ovary; corolla 2-3 mm long,
tubular to trumpet-shaped, yellow with red-pur-
ple margin, 5-toothed, abaxially glandular.
Stamens 5, exserted at maturity. Style truncate,
with sweeping hairs. Receptacle after anthesis
with long hairs between involucral bracts and
marginal paleae. Chromosome number. 2n = 1 8.
Flowering time: correlated with rainfall,
January to March and June to September in
summer- and winter-rainfall areas respectively.
Map 12. — Eriocephalus pauperrimus.
The distribution of E. pauperrimus extends
from the southern parts of Namibia through the
Northern Cape to Matjiesfontein in the Western
Cape. The areas where it occurs, receive less
than 200 mm rain per annum. It occurs mainly
in the summer-rainfall area but also partly in the
area that receives both summer and winter rain
at an altitude of 300-600 m. This species is
under-collected in southern Namibia and the
Northern Cape. Map 12.
Although closely related to E. ericoides (no.
26), it can easily be distinguished from that
species by the alternate leaves and sessile ca-
pitula in terminal spikes.
Common name: kapokbos.
Vouchers: Acocks 18254 (PRE); Barker 9300
(NBG); De Winter 3363 (PRE, WIND); Giess
& Muller 12051 (WIND); Goldblatt 6086 (MO,
WIND).
24. Eriocephalus ambiguus (DC.)
M.A.N.Miiller, comb, et stat. nov.
E. aspatalhoides DC. var. ambiguus DC., Prodromus:
148 (1838). Type: Cape Province, precise locality unknown,
Drege 6038 (G-DC, holo.: P!, PRE. photo.!).
E. aspalallwides DC.: 148 (1838), pro parte; Harv.: 203
(1865), pro parte; Merxm.: 60 (1967), pro parte.
ASTERACEAE: Eriocephalus
43
Many-stemmed, erect, much-branched, spi-
nescent shrubs, 0.3-0. 6 m high and up to 450
mm in diameter. Older stems with grey-brown
bark, deeply grooved, eventually displaying
anomalous secondary growth, 10-20 mm in
diameter, breaking up into independent daugh-
ter plants; young branches yellow-brown, later
yellow-grey to grey-brown, irregularly sympo-
dially branched, tips of branches spinescent,
1-18 mm long; brachyblasts 1-10 mm long,
short-lived. Leaves basally semilunate, adaxial-
ly concave, those on dolichoblasts alternate,
linear, 4-15 x 0.5 mm, entire, densely silver-
grey shortly pilose to pilose, apex obtuse;
leaves on brachyblasts scale-like, rosulate, 2-4
x 0.5 mm, entire, apex obtuse. Capitula hetero-
gamous disciform, 4 mm long, solitary on
brachyblasts, rarely in terminal racemes; pedun-
cles 1-11 mm long, shortly pilose. Involucral
bracts 4 or 5, 2.5 x 1 .5 mm, green, with narrow,
sometimes purplish, membranous margin,
ovate, flattened, abaxially appressed sericeous.
Paleae: those of marginal florets free, 2 x 1.5
mm, keeled, lanceolate, apices fringed, abaxial-
ly long-pilose, hairs septate, adaxially smooth,
glabrous; those of disc florets oblong-linear,
membranous, apex fringed, 1 x 0.5 mm, abaxial-
ly long-pilose, adaxially smooth. Marginal
female florets 2-5, yellow, inconspicuous,
2.0-2. 5 mm long; lamina absent; corolla tubular-
filiform, much shorter than branched style. Style
branches flattened, linear, apex acute, 1-2 mm
long. Ovary (and cypsela) oblong-ovoid, slight-
ly flattened, densely lanate at maturity. Seed
1-2 mm long, flattish, trigonous. Disc florets
5-21, yellow, functionally male with sterile
ovary, 2. 5-3.0 mm long; corolla tubular, widen-
ing upwards, 5-toothed; teeth sometimes tinged
red-purple. Stamens 5, exserted at maturity.
Style unbranched, globose, truncate with short,
sweeping hairs. Receptacle after anthesis with
long, soft hairs between involucral bracts and
marginal paleae. Chromosome number. 2n = 18.
Flowering time: January to April.
The distribution of E. ambiguus extends
from the central parts of Namibia to Botswana
and the Northern, Western and Eastern Cape.
Its current known distribution in Botswana is
nus.
limited to a single locality on the border
between the Northern Cape and Botswana. The
largest part of the distribution area receives an
annual rainfall of less than 200 mm; only the
southernmost part of its distribution receives an
average annual rainfall of more than 200 mm.
It is suspected that this species is more com-
mon in the southern parts, but that it has been
under-collected here because of its similarity to
and the resulting confusion with E. karooicus
(no. 17) and E. spine scens (no. 30), both of
which are associated with E. ambiguus in this
area. Like E. karooicus and E. spinescens, E.
ambiguus is found only in low-lying areas, ±
300 m above sea level, mainly in sandy and
clayey soils, never on mountains or hills. Plants
are scattered or at most found in small groups,
but never form dense, dominant communities.
Map 13.
After the description of E. aspalathoides var.
ambiguus by De Candolle (1838), this taxon
was not recognised by later researchers of the
genus Eriocephalus. This can be attributed part-
ly to the fact that later researchers like Harvey
(1865) and Merxmiiller ( 1967) placed it in syn-
onymy under E. aspalathoides and partly to its
poor representation in herbaria. It is only from
1963 onwards that this taxon has become better
represented in herbarium collections.
44
ASTERACEAE: Eriocephalus
E. ambiguus was confused with the earlier
known E. aspalathoides, now E. decussatus
(no. 21). Although E. decussatus has opposite
leaves, they are alternate on flowering shoots,
as in E. ambiguus. The capitula are solitary,
pedunculate on brachyblasts, while the terminal
raceme branches harden to form spinescent tips
after the capitula have been shed. A thorough
investigation of E. ambiguus showed that the
leaves are always alternate, and a true spine is
formed, not merely spinescent branch tips.
Although closely related to E. luederitzianus
(no. 25), it can be easily distinguished from that
species by the spines, the capitula borne mainly
on brachyblasts and the much-branched growth
form. E. luederitzianus , in contrast, has mainly
terminal racemes of which the central flowering
axis becomes hardened and spinescent after
maturity and shedding of capitula.
During good rainy seasons E. ambiguus
tends to form long, drooping shoots without
spines and with long leaves similar to those of
E. luederitzianus instead of the typical spines-
cent habit with rigid branches and very short
leaves. This variation of E. ambiguus can easily
cause confusion with E. luederitzianus, but as
these two species are allopatric, all material can
be separated on distribution alone.
Note: during preparation of the manuscript
and investigation of the taxa, Herman found
three specimens in PRE ( Hafstrom & Acocks
1554, Shearing 558 , Van Rooyen & Bredenkamp
168) in which the paleae of the marginal female
florets were connate. No other differences could
be detected in these three specimens; they fit the
description of E. ambiguus perfectly, except for
the connate paleae.
Common name: doringkapok.
Vouchers: De Winter 3590 (PRE); Giess 14617
(WIND); Giess & Robinson 13253 (WIND);
Muller 791 (WIND); Role Evans 2248 ( BOL).
25. Eriocephalus luederitzianus O.Hoffin.
in Bulletin de I'Hcrbier Boissier I: 86 (1893);
Engl. & Prantl: 270 ( 1894); Merxm.: 62 (1967).
Type: Namibia, ‘Reise von Walfish Bay nord-
ostlich nach Odyitambi, Dec. 1885 bis Febr.
1886’, Liideritz s.n. (Z, holo. !).
E. eenii S. Moore: 351 (1902). Type: Namibia,
Damaraland, Een s.n. (BM. holo.!).
E. squarrosus Muschl. in Dinter: 260 ( 1921 ) nom. nud.
Type: Namibia, Farm Hoffnung WIN 66, Dinter 985 (B,
holo.; SAM!).
E. hirsutus Burtt Davy: 106 (1935). Type: Northern
Province, Bushveld, Klippan. Rehmann 5232 (K, holo.!;
Z!).
E. pubescens sensu Merxm.: 62 ( 1967).
Erect, many-stemmed, sparsely branched
shrubs, 300-500 mm tall. Old stems dark
brown, rough, deeply grooved, displaying
anomalous secondary growth; bark yellow
brown to dark brown; young branches smooth,
yellow-white, densely white sericeous; older
branchlets yellow-brown, superficially grooved,
sparsely hairy; brachyblasts short, up to 10 mm
long. Leaves alternate, densely imbricate, those
on young shoots 4-25 x 0.5- 1.0 mm, those on
brachyblasts linear, 3-6 x 0. 5-1.0 mm, entire,
permanently densely appressed silver-grey
sericeous, adaxially basally concave, abaxially
convex, distally semiterete to cylindrical, apex
obtuse, base semi-amplexicaul. Capitula he-
terogamous disciform, mostly terminal, race-
mose or umbellate racemose, also solitary, ter-
minal on brachyblasts, 4-8 x 4-6 mm; pedun-
cles 2-16 mm long, appressed sericeous.
Involucral bracts 4 or 5, ovate, flattened, ± 3 x
1 .5 mm, central part green, herbaceous with
narrow membranous, colourless to purple mar-
gin, abaxially permanently appressed seri-
ceous. Paleae'. those of marginal florets free,
keeled, lanceolate, 2.5 x 1.0 mm, abaxially
long-pilose/lanate, hairs septate, apex fringed;
those of disc florets oblong to linear, transpar-
ent, membranous, 1-2 x 0.5 mm, abaxially
lanate, adaxially smooth, apex fringed.
Marginal female florets 2-5, indistinct, 2. 5-3.0
mm long; corolla yellow, tubular, narrowed at
throat, sometimes with very short lamina,
scarcely 0.5 mm long, much shorter than
ASTERACEAE: Eriocephalus
45
TABLE 1. — Comparison of morphological characters of Eriocephalus ambiguus and closely related taxa
Character
E. ambiguus
E. luederitzianus
E. microphyllus var.
pubescens
E. merxmuelleri
Leaf arrangement
Alternate
Alternate
Opposite, sometimes
alternate on flowering
shoots
Opposite, sometimes
alternate on flowering
shoots
Leaf length
4—15 mm
4—25 mm
1 .5-4.0 mm
4 — 9(— 14) mm
Indumentum
Permanently
sericeous
Permanently
sericeous
Permanently felty
sericeous
Sericeous, glabrescent
Inflorescence
Solitary on
brachyblasts
Racemose or umbellate-
racemose or solitary
on brachyblasts
Racemose or paniculate
or solitary on brachy-
blasts
Racemose or solitary
on brachyblasts
Involucral bracts
4(5), permanently
sericeous
4(5), permanently
sericeous
4, felty sericeous to
glabrous
4(5), sericeous to
glabrous
Disc florets
5-21, yellow,
sometimes with
red-purple margin
14—28, yellow
(3)4 — 6(— 8), cream-
coloured with red-
purple margin
1— (5) or 6(— 9),
cream-coloured with
red-purple margin
Habit
Much-branched
shrubs with terminal
spines
Many-stemmed shrubs
with long, sparsely
branched shoots
Much-branched shrubs
Much-branched shrubs
with long peduncles
branched style. Style branches flattened, linear,
1-2 mm long, apices acute. Ovary (and
cypsela) oblong-ovoid, slightly fiattish, trigo-
nous, at maturity densely lanate. Seed fiattish,
trigonous, 1-2 mm long. Disc florets 14-28,
functionally male with sterile ovary, yellow, 3
mm long; corolla tube 5-toothed. Stamens (4)5,
well exserted at maturity. Style unbranched,
apex globose, with sweeping hairs. Receptacle
after anthesis densely white or tawny lanate
between paleae of marginal florets and involu-
cral bracts. Chromosome number: 2 n = 36.
Flowering time: October to May with a peak
from January to March.
E. luederitzianus occurs only in the summer-
rainfall area. It extends over the northern half of
Namibia, most of Botswana into the Northern
Province of South Africa. In Namibia it is the
most abundant species with the widest distribu-
tion. Information from Botswana is very defi-
cient, but it seems that with the exception of a
single locality for E. ambiguus (no. 24), this is
the only species occurring in Botswana. It is
also possible that this species is more abundant
in Botswana, but that it has been under-collect-
ed until now. Map 13.
As there is so much confusion between E.
ambiguus (no. 24) [E. aspalathoides pro parte
after De Candolle (1838), Harvey (1865) and
Merxmuller (1967)], E. luederitzianus, E. micro-
phyllus var. pubescens (no. 28b) and E. merx-
muelleri (no. 32), a comparison between these
species is given in Table 1.
During good rainy seasons E. ambiguus pro-
duces water shoots with long leaves and lacking
characteristic spines. This material can easily be
confused with E. luederitzianus. Later growth
shows the characteristic terminal spines. These
two species are allopatric and difficulties
regarding positive identification can be solved
by consulting the distribution map.
In the past, all material of E. luederitzianus
was identified as E. pubescens, the current E.
microphyllus var. pubescens, because Merx-
muller (1967) had put it into synonymy under
E. pubescens. As a result, material of E. merx-
muelleri and E. ambiguus was also identified as
E. pubescens.
Note: during preparation of the manuscript
and investigation of the taxa, Herman found six
specimens in PRE ( Koekemoer 205, Kreulen
46
ASTERACEAE: Eriocephalus
Figure 8. — Eriocephalus ericoides subsp. ericoides: A, flowering shoot with inflorescences, x 1; B. branch with
leaves, x 12; C, capitulum, x 6; Dl, D2, involucral bracts, x 10; E, free marginal palea, x 10; F, central palea, x 15; Gl,
marginal female floret, x 18; G2. branched style, x 25; HI, disc floret x 20; H2, anthers, x 14; H3, truncate style, x 14
(Miiller & Tilson 873, WIND).
ASTERACEAE: Eriocephalus
47
562, Moss & Jacobsen K18, Seydel 1405,
Skarpe S-359 and Vahrmeijer & Steenkamp
3067) in which the paleae of the marginal
female florets were connate. No other differ-
ences could be detected in these six specimens;
they fit the description of E. luederitzianus per-
fectly, except for the connate paleae.
Common name: kapokbos.
Vouchers: De Winter & Leistner 5539 (PRE,
WIND); Giess 13572 (NBG, WIND); Hutchinson
2651 (BOL, PRE); Muller & Kolberg 2119
(WIND); Story 4901 (NBG, PRE).
26. Eriocephalus ericoides (L.f) Druce in
Supplement to Botanical Exchange Club of the
British Islands for 1916: 622 (1917); Merxm.:
61 (1967). Type: Cape Province, exact locality
unknown. Collector unknown (LINN 983.5,
holo., microfiche!; WIND, photo.!).
Tarchonanthus ericoides L.f.: 360 (1782). Type: as
above.
E. glaber Thunb.: 168 (1800); Willd.: 2384 (1803);
Pers.: 497 ( 1 807); Thunb.: 724 (1823); Spreng.: 62 1 ( 1 826);
DC.: 148 (1838); Harv.: 204 (1865). Type: Cape Province,
without exact locality. Thunberg sub Thunberg Herb. nr.
20911 (UPS. holo.; WIND, photo.!).
E. glaber Thunb. var. sessiliflorus Sond. ex Harv.: 204
(1865). Type: Eastern Cape. Graaff-Reinet, Zeyher 23
(MEL. holo.!).
Erect, many-stemmed, relatively sparsely
branched, conical or broom-like shrubs, 0.3-1 .0
m high, 300-400 mm in diameter, not or rarely
spinescent. Old steins displaying anomalous
secondary growth, 15-30 mm in diameter, grey-
brown; dolichoblasts bright green to yellowish,
sparsely to densely white-felty and glandular;
older branchlets grey to grey-brown, delicately
branched, tending to be vertically orientated;
brachyblasts short-lived, sometimes very long
with leaves at apex. Leaves opposite, rarely
alternate on flowering shoots, linear, (0.75-)
1 .0— 3.5(— 7.0) x 0.2-0. 5 mm, entire, adaxially
flattened, concave towards base, abaxially con-
vex, initially densely felty, glabrescent, or per-
manently hairy, when glabrous shiny, bright
green or dull green, pitted with glands in cavi-
ties, apex obtuse to slightly acute, base broad-
ened and semi-amplexicaul; leaves on brachy-
blasts conspicuously decussate, imbricate, ini-
tially shortly felty, glabrescent or permanently
hairy; those on dolichoblasts scattered, sessile
on permanently callous, cushion-like thicken-
ings, much longer than those on brachyblasts.
Capitula heterogamous disciform, spicate race-
mose or racemose or solitary on brachyblasts,
rarely paniculate, 1.5-2. 5 mm long; peduncle
1.0-5. 5 mm long, sometimes almost absent,
felty, glabrescent. lnvolucral bracts 4, ovate to
lanceolate, up to 2 x 1.5 mm, 2 slightly keeled,
other 2 laterally flattened, central part herba-
ceous, green, margin broad, membranous, pur-
ple, abaxially sparsely felty to sericeous,
glabrescent, with permanent, almost sessile
glands in cavities. Paleae'. those of marginal
florets free, 1. 5-2.0 mm long, ovate, keeled and
enveloping florets, abaxially long-lanate, hairs
septate; those of disc florets linear, 2.0 x 0.3
mm, membranous, margins fringed, abaxially
lanate. Marginal female florets ( 1 )2, 2.0-2. 5
mm long; corolla narrowly tubular-filiform;
lamina extending at most to furcation of style,
yellow, cylindrical. Style branches flattened,
linear, apex acute. Ovary oblong to oblanceo-
late, slightly flattened, long-lanate. Seeds
1.2-2. 2 mm long, obovoid, slightly flattened.
Disc florets ( 1— )3— 5(— 7), functionally male with
sterile ovary, 2. 0-2. 5 mm long; corolla tubular
to trumpet-shaped, 5-lobed, red-purple, some-
times yellow towards base. Stamens 5, exserted
at maturity. Style unbranched, cylindrical, apex
globose, with sweeping hairs. Receptacle after
anthesis with dense, white, long hairs between
involucral bracts and marginal paleae. Chromo-
some number: 2 n = 1 8. Figure 8.
E. ericoides occurs from Namibia to the Free
State and the Northern, Western and Eastern
Cape. The plants in Namibia are fairly isolated
from those in South Africa. Except for the indu-
mentum of plants north of the Orange River,
there seem to be no obvious differences be-
tween the plants in Namibia and those south of
the Orange River. They all have a conical to
broom-like growth form. The shape and length
48
ASTERACEAE: Eriocephalus
of the leaves differ very little between the dif-
ferent individuals.
Individuals from a population in the North-
ern Cape, north of the Orange River, have dull
green leaves with permanent, long-felty hairs.
In contrast, other individuals of the species in
South Africa and Namibia have shiny, bright
green, glabrescent leaves. On the grounds of the
dull green leaves with permanent, long-felty
indumentum and geographical isolation, this
group is described as a subspecies of E. eri-
coides.
Two subspecies are recognised:
Leaves glabrescent, shiny, bright green . . .
26a. subsp. ericoides
Leaves permanently long-felty, dull green
26b. subsp. griquensis
26a. subsp. ericoides.
Older leaves glabrescent, shiny, (0.75-)
1.0-3. 0(— 5.0) x 0.3-0.4 mm. Peduncles 1-5 mm
long. Marginal female florets (1)2. Disc florets
( 1— )3— 5( — 7). Flowering time: correlated with
rainfall, January to April in summer-rainfall areas
and July to September in winter-rainfall areas.
E. ericoides subsp. ericoides has the widest
distribution of all Eriocephalus taxa. It extends
from Namibia into the Free State and Northern,
Western and Eastern Cape, being absent only
from the northwestern and western parts. The
species is distributed mostly inland: it does not
extend to the coast and occurs mostly above 300
m altitude. The distribution includes both sum-
mer- and winter-rainfall areas, ranging from
those with an annual rainfall of less than 250 mm
to areas with more than 500 mm. Compared to E.
africanus var. paniculatus (no. 14b), the taxon
with the second-widest distribution, occurring in
coastal areas with higher rainfall, this subspecies
is more representative of the arid karoo region.
The disjunct distribution of E. ericoides subsp.
ericoides is very obvious. Not only does the pop-
ulation in Namibia stand isolated from the rest of
Map 14. — • Eriocephalus ericoides subsp. ericoides; ■ E.
ericoides subsp. griquensis.
the species (both subsp. ericoides and subsp.
griquensis), but there is very little correlation
between the two distribution areas of subsp. eri-
coides. In Namibia, their habitat is high-lying
mountains, 1 000 to 1 700 m in altitude, and
receives summer rainfall of only 250 to 300 mm
rain annually, whereas in South Africa it occurs
both on high-lying parts and near the coast (300
m altitude) and receives both summer and winter
rainfall, with a precipitation of about 500 mm
annually. Both populations of subsp. ericoides
occur in the Karoo-Namib plant geographical
region (Werger 1978). This region is charac-
terised by a wealth of dwarf shrubs belonging to
the Asteraceae. Between this Karoo-Namib plant
geographical region and the Capensis region
where the distribution of subsp. ericoides con-
tinues. there are many floristic similarities. It is
therefore possible that E. ericoides had a much
wider distribution earlier but that it was disrupt-
ed by unknown factors. Map 14.
As this subspecies occurs in so many differ-
ent veld types, it has adapted to various habitats.
This in turn has led to much variation in mor-
phological characters. The most common
growth form is conical or broom-like shrubs
with thin branchlets and delicate, small, bright
green, decussate leaves on brachyblasts. The
leaves at the growing points of young shoots are
ASTERACEAE: Eriocephalus
49
initially felty, but soon become glabrous. The
growing points of the brachyblasts are, how-
ever, felty sericeous.
The arrangement of capitula shows almost as
much variation as there are plants. The most
common is racemose with relatively shortly
pedunculate capitula. Two specimens were col-
lected by Tyson at Murraysburg: Tyson 269
(SAM) has an almost paniculate synflores-
cence, while Tyson 289 (SAM) has sessile ca-
pitula (the so-called var. sessiliflorus Sond. ex
Harv.). The leaves are mostly opposite, but are
often alternate on flowering shoots.
E. ericoides subsp. ericoides is well browsed
compared to E. microphyllus (no. 28). Accord-
ing to Roux (1984), this bush is not unpalatable,
but palatability varies from season to season. It
is best utilised during late autumn and spring.
This subspecies can become invasive in dis-
turbed veld. Roux (1984) mentions that it ham-
pers the establishment of Panicum species in
the Karoo to a certain degree. Common names:
gewone kapokbossie , renosterveldkapok , roos-
maryn , rosemary (Smith 1966); gladdekapok-
bos, regtekapok , gewonekapok , grootkapokbos,
sandveldkapokbos (Roux 1984).
Vouchers: Dahlstrand 2105 (NBG, PRE);
Dyer 427 (GRA, PRE); Hugo 245 (NBG);
Smith 951 (PRE); Walter 1692 (WIND).
26b. subsp. griquensis M.A.N. Miiller, subsp.
nov., E. ericoidi (L.f.) Druce subsp. ericoidi affi-
nis sed foliis permanenter velutinis differt.
Type: Northern Cape, Herbert District: Farm
Eureka, Acocks 8753 (BOL, holo.; PRE).
Older leaves dull green, permanently long-
felty, rarely glabrescent, 1 .5— 3.0(— 7.0) x 0.2-
0.5 mm. Peduncles ( 1 .0— )2.0— 3.5(— 5.5) mm.
Marginal female florets (0)(1)2. Disc florets
(2-)4 or 5(6). Flowering time: correlated with
rain in summer, January to April, occasionally
extending from July to August when the area
receives winter rain.
Subsp. griquensis is restricted to the
Northern Cape, from the Orange River to near
the Botswana border. Map 14.
The habit closely resembles that of subsp.
ericoides , but the leaves of subsp. griquensis
are permanently long-felty, rendering it a dull,
green colour, and are very rarely glabrescent.
Common name: kapokbos.
Vouchers: Coetsee 48 (PRE); Esterhuysen
2295 (BOL, PRE); Kotze 795 (PRE); Leistner
1449 (PRE); Pole Evans 2504 (NBG, PRE).
27. Eriocephalus glandulosus M.A.N. Mid-
ler, sp. nov., E. ericoidi (L.f.) Druce et E. aro-
matico C.A.Sm. affinis sed habitu ramosissimo
spinescenti, foliis saepe ex rubro-purpureis pal-
Iide viridis nitidis; lemma florum marginalium
stylum aequans.
Type: Northern Cape, 5 km E of Williston on
road to Carnarvon, Miiller 3596 (PRE, holo.;
WIND).
Erect to slightly spreading, rounded, many-
stemmed, much-branched, rigid, spinescent
shrubs, 0.2-0. 6 m high and in diameter. Old
stems grey to grey-black, displaying anomalous
secondary growth, growing points sparsely
felty, soon glabrous; dolichoblasts green-yel-
low, glandular, when older yellow-brown,
glabrous, branches rigid. Leaves decussate,
densely imbricate, in 4 rows, obtuse triangular
to linear, those on dolichoblasts 2. 3-6. 2 mm
long, those on brachyblasts 1 .5-2.3 mm long,
entire, shiny, bright green, often with red-purple
tinge, initially sparsely felty, soon glabrous,
with glands in cavities on leaf surface, adaxial-
ly proximally concave, distally flattened, abax-
ially senuterete, keeled, apex obtuse, base semi-
amplexicaul. Capitula heterogamous disciform,
mostly solitary on brachyblasts, sometimes
racemose or umbellate-racemose, 2. 3-2. 6 mm
long; peduncle 3-7 mm long, cylindrical,
glabrous. Involucral bracts 4, broadly ovate,
2.5 x 2.5 mm, with thickened herbaceous cen-
Figure 9.- Eriocephalus glandulosus: A. flowering shoot with inflorescences, x 1; B, branch with leaves, x 4; C, ca-
pitulum. x 3; 1)1, D2, involucrai bracts, x 6; E, free marginal palea, x 6; F, central palea, x 8; Gl, marginal female floret,
' 10: G2, branched style, x 25: II I , disc floret, x 10: H2, anthers, x 20; H3, truncate style, x 25 ( Acocks 548 and 566, PRE).
ASTERACEAE: Eriocephalus
51
tral part and broad membranous margin, mostly
red-purple, sometimes shiny green, glabrous, 2
slightly keeled bracts enveloping 2 flattened
ones. Paleae: those of marginal florets free,
mostly only 2, 2.0-2.4 x 1 .5 mm, keeled with
central hardened part and membranous margin,
apex fringed, abaxially long-lanate, hairs sep-
tate; those of disc florets 2.0-2. 3 x 0.5 mm,
membranous, flattened, narrowly ovate to nar-
rowly lanceolate, margins fringed, abaxially
lanate. Marginal female florets small, 2, white,
3. 6^1. 2 mm, lamina up to 2.2 mm long, strap-
shaped, as long as to slightly longer than style
branches. Style branches strap-shaped, apex
acute. Ovary (and cypsela) oblong to ovoid,
slightly flattened, long-lanate. Seed 1. 4-2.0 mm
long, lanceolate, slightly flattened. Disc florets
10-18, functionally male with sterile ovary,
2. 5-3. 2 mm long; corolla red-purple, trumpet-
shaped to infundibuliform, 5-lobed. Stamens 5,
exserted at maturity. Style unbranched, truncate,
with sweeping hairs. Receptacle after anthesis
densely white, long-pilose between involucral
bracts and marginal paleae. Chromosome num-
ber: 2n - 18. Flowering time : July to October
and February to March according to winter rain-
fall in the western parts and summer rainfall
in the eastern parts of the distribution area.
Figure 9.
The distribution of E. glandulosus extends
over the Northern, Western and Eastern Cape,
in the following Acocks (1975) veld types:
Kalahari Thornveld and Shrub Bushveld,
Kalahari Thornveld invaded by Karoo, Orange
River Broken Veld, Arid Karoo and Desert
False Grassveld, and False Arid Karoo. These
areas receive an average annual rainfall of less
than 400 mm. Although the species occurs over
a large area, it is never found in dense stands,
but rather scattered. Map 15.
Closely related to E. ericoides (no. 26) and
E. aromaticus (no. 8) from which it can be dis-
tinguished by its much-branched, spinescent
habit, shiny, light green leaves, often with a red-
purple tinge, and marginal female florets with
lamina about as long as style branches.
Map 15. — Eriocephalus glandulosus.
E. glandulosus and E. ericoides are closely
related, with some similar characters. Both
species have decussate, initially felty but soon
glabrous leaves and umbellate, racemose or
solitary capitula, terminally on brachyblasts.
They can, however, be distinguished as follows:
the two marginal paleae of E. glandulosus are
relatively broad, 2.0-2. 4 x 1.5 mm, and mem-
branous; the two lateral margins are extended to
touch each other, thus enveloping all the florets,
but they are nonetheless free. In contrast, the
marginal paleae of E. ericoides are narrower,
1. 5-2.0 x 0. 5-1.0 mm, with incurved margins
so that each envelops only one female floret
The lamina of the marginal female floret is
absent in E. ericoides but fairly well developed
in E. glandulosus , as long as to longer than the
style branches.
Common name: kapokbos.
Vouchers: Acocks 548 (BOL, PRE); Acocks
566 (PRE); Hutchinson & Dyer 3124 (BOL,
PRE); Miiller 3590 (WIND);’ Southy in herb
Galpin 5591 (GRA, PRE).
28. Eriocephalus microphyllus DC., Pro-
dromus: 148 (1838). Type: Cape Province, Little
Namaqualand, without exact locality, Drege 2735
(G-DC, holo.; P!, PRE & WIND, photo.!).
52
ASTERACEAE: Eriocephalus
Eigi re 10— Eriocephalus microphyllus: A, flowering shoot with inflorescences, x I; B, capitulum, x 5; C, involucral
brad, x 10; D. marginal palca, x 16; E, central palea, x 16; F, marginal female floret, x 12; G I, disc floret, x 12; G2, anthers,
x 16; G3. truncate style, x 16 (Muller 3564, WIND). E. microcephalus: H. marginal female floret, x 14 (Oliver 3527, NBG).
ASTERACEAE: Eriocephalus
53
E. glaber Thunb. var. pubescens Harv.: 204 (1865) non
E. pubescens DC.: 148 (1838). Type: based on that of E.
microphyllus and the following syntype: Cape Province,
Gariep (without precise locality), Burchell s.n. (G-DC,
PRE, photo.!).
Many-stemmed, markedly dichotomously
but sparsely branched to densely intertwined
shrub, 0.2-0. 4-0. 8 m high, 0.4-1. 2 m in diam-
eter. Old stems displaying anomalous secondary
growth, grey-black; young shoots yellow-
brown, initially felty or felty sericeous, glabres-
cent, glandular; older branches and stems yel-
low-brown to grey-brown to grey-black, 3-5
mm in diameter; brachyblasts short-lived, rela-
tively short, 1-2 mm long. Leaves opposite,
decussate on brachyblasts, densely imbricate,
sometimes alternate on flowering shoots, linear,
those on brachyblasts almost obtuse triangular,
( 1.2—) 1 .5— 4.0(— 7.0) x 0.6-0. 8 mm, entire, blue-
green, bright green to grey-green, felty at grow-
ing points, the rest felty sericeous to glabres-
cent, glandular (subsessile glands in cavities on
leaves), adaxially flattened to slightly concave,
strongly concave towards base, abaxially con-
vex, slightly keeled towards obtuse to slightly
acute apex, base semi-amplexicaul broadened.
Capitula heterogamous disciform, terminal,
racemose or spicate racemose or solitary on
brachyblasts, rarely paniculate, 4-6 mm long;
peduncles 1.5-1 1 .0 mm long, felty to glabrous,
glabrescent. Involucral bracts 4, ovate to broad-
ly lanceolate, 2. 3-4.0 x 1.0-1 ,5(— 2.0) mm, cen-
tral part red-purple to green, herbaceous with
broad transparent membranous margin, ap-
pressed felty sericeous to glabrous, 2 slightly
keeled, 2 laterally flattened, enveloped by the 2
keeled bracts. Paleae: those of marginal florets
free, 3. 2-4. 5 mm long, keeled and enveloping
single floret (mostly female), narrowly ovate to
lanceolate, margins fringed, central part coria-
ceous, rigid with membranous margin, abaxial-
ly long-lanate, hairs septate; those of disc flo-
rets up to 2.5 mm long, broadly to narrowly
lanceolate, outer slightly keeled, inner more
flattened, membranous, margins and adaxially
long-lanate. Marginal female florets (l)2(-4),
2. 5-3. 2 mm long; corolla white, narrowly tubu-
lar, narrowed around style, with short lamina,
narrowly cuneate to oblong, 3-lobed or -toothed.
0.3-0.8 mm long, mostly shorter than point of
style furcation, sometimes as long as style
branches, abaxially with subsessile glands.
Style branches linear, flattened, apex acute.
Ovary oblong to oblanceolate, long-lanate. Seed
2. 0-2. 5 mm long, slightly flattish, trigonous.
Disc florets (3)4— 6(— 8), functionally male with
sterile ovary, 2. 6-3. 5 mm long; corolla tubular
to trumpet-shaped, 5-lobed, proximal half
cream-coloured, distal half gradually darker
red-purple, with subsessile glands abaxially.
Style cylindrical, unbranched, apex globose,
with sweeping hairs. Stamens 5, slightly exsert-
ed at maturity. Receptacle after anthesis with
long hairs between involucral bracts and mar-
ginal paleae. Chromosome number : 2 n = 36.
Figure 10.
Three varieties are recognised:
la Peduncles 1.5-3.0(-4.0f mm long,
permanently felty; leaves blue-
green; branches drooping
28c. var. carnosus
lb Peduncles 5-11 mm long, felty
sericeous to glabrescent; leaves
bright green to grey-green; branch-
es rigid:
2a Leaves initially felty sericeous,
glabrescent; peduncle 6-10 mm
long 28a. var. microphyllus
2b Leaves permanently felty sericeous;
peduncle (5— )7— 9(— 1 1 ) mm long
28b. var. pubescens
28a. var. microphyllus.
Much-branched, rigid shrubs, 0.4-0. 8 m
high, 0.4-1.2 m in diameter, with open branch-
ing. Leaves opposite and decussate, rarely alter-
nate on flowering shoots, green to grey, initial-
ly felty, glabrescent, with subsessile glands in
cavities, those on young shoots 1. 8-4.0 mm
long, those on brachyblasts 1 .5-2.0 mm long.
Capitula terminal, racemose or solitary on
brachyblasts; peduncle 6-10 mm long, densely
felty to glabrous. Involucral bracts 2. 5-4.0 mm
long, sometimes distinct, red-purple to green.
54
ASTERACEAE: Eriocephalus
Map 16. — Eriocephalus microphyllus var. microphyllus.
glabrous. Marginal female florets (1)2. Disc flo-
rets 5-7. Chromosome number. 2 n = 36.
Flowering time: correlated with rainfall (both
summer and winter rainfall), reaching a peak
from February to March and July to August in
the different rainfall areas.
This variety is typical of central Nama-
qualand (Northern Cape) and occurs mainly on
low-lying plateau areas. It is fairly common and
often forms dense stands. Map 16.
Common name: kapokbos.
Vouchers: Hugo 520 (NBG, WIND); Hugo
2884 (NBG); Levyns 5079 (BOL, PRE); Rosch
& Le Roux 921 (KPA-J); Thompson 2398
(NBG, PRE).
28b. var. pubescens (DC.) M. A. N. Muller,
comb, et stat. nov.; non E. glaber Thunb. var.
pubescens Harv.: 204 (1865).
E. pubescens DC., Prodromus: 148 (1838); Harv.: 203
(1865). Type: Western Cape, ‘Bei Mierenkasteel, kaiTooar-
tige Hohe, unter I 000 Fuss’, Drege 6039 (G-DC, holo.:
PRE & WIND, photo.!).
Young shoots felty sericeous, glabrescent.
Leaves always opposite, rarely alternate on flow-
ering shoots, permanently felty sericeous; those
on young shoots 3-7 mm long; those on brachy-
Map 17. — • Eriocephalus microphyllus var. pubescens;
■ E. microphyllus var. carnosus.
blasts 1. 2-2.0 mm. Capitula terminal, racemose,
rarely paniculate; peduncles (5— )7— 9 (-11) mm
long, permanently felty sericeous or glabrescent.
Involucral bracts felty sericeous to glabrous,
2. 3-3. 2 mm long, green. Marginal female florets
2. Disc florets 4-6. Chromosome number: 2 n =
36. Flowering time: mainly July to September.
The distribution of this variety is concentrat-
ed mainly along the west coast. The habitat is
more mountainous than that of var. microphyl-
lus. Map 17.
Common name: kapokbos.
Vouchers: Acocks 16440 (PRE); Acocks
19519 (NBG, PRE); Bolus 9568 (BOL); Muller
4054 (WIND); Rosch & Le Roux 509 (KPA-J,
PRE).
28c. var. carnosus M. A. N. Muller, var. nov.,
E. microphyllo DC. var. microphyllo affinis sed
pedunculis 1.5-3.0(-4.0) mm, permanente velu-
tinis; foliis aeruginosis; ramis cemuis.
Type: Western Cape, ridge NE of Jan de
Boers, Oliver 3474 (PRE, holo.; NBG).
Mostly sparsely branched, compact shrubs,
at most 0.6 m high, normally 200^400 mm tall;
ASTERACEAE: Eriocephalus
55
branches tending to be long, drooping and
sparsely branched, with open branching; young
shoots felty. Leaves opposite to decussate on
brachyblasts, alternate on flowering shoots,
1.6-2. 6 mm long on young shoots, 1.2-1. 8 mm
long on brachyblasts, succulent, felty, glabres-
cent, blue-green, cylindrical distally. Capitula
almost spicate racemose; peduncles relatively
short, 1 .5— 3 ,0(— 4.0) mm long, permanently
felty. Involucral bracts 2. 3-3. 2 mm long, green.
Marginal female florets (1)2 or 3(4). Disc flo-
rets (3)4— 6(— 8). Chromosome number. 2 n - 36.
Flowering time: correlated with winter rainfall,
June to September.
The distribution of this variety is restricted
mainly to the Worcester and Montagu Districts.
It grows on shale and gravel plateaus, forming
relatively dense stands. Map 17.
In contrast to the other two varieties, which
are hardly browsed, this one is palatable and
heavily browsed. Common name: kapokbos.
Vouchers: Compton 2871 (BOL); Miiller
4067 (WIND); Oliver 3473 (NBG); Olivier 220
(NBG, PRE).
29. Eriocephalus microcephalus DC.,
Prodromus: 148 (1838); Harv.: 204 (1865).
Type: Northern Cape, Little Namaqualand,
Modderfontein, Drege 6376 (G-DC, holo.; G!,
K!, PRE, photo.!).
Slender, many-stemmed shrubs, much-
branched from base, 0.4- 1.2 m high. Old stems
dark brown, displaying anomalous secondary
growth; growing points green-purple, felty,
glandular; young shoots red-purple to red-
brown, up to 0.3 mm in diameter, internodes
4. 0-8. 5 mm long; older branches dark brown,
up to 1 mm in diameter, side branches forming
an angle of 70-90° with main axis; brachyblasts
short-lived, up to 1.5 mm long and barely 0.5
mm in diameter. Leaves decussate, grey-green,
relatively small, 0.8-1. 6 x 0.3-0. 5 mm, up to
4.5 mm long on young shoots, entire, scale-like,
obtuse-triangular and sometimes linear-lanceo-
late on young shoots, adaxially flattened, abax-
ially semiterete, distally slightly keeled, leaf
surface with permanent multicellular glands in
cavities, apex obtuse, concave towards base,
leaves at growing point glandular and densely
white-felty, glabrescent. Capitula heteroga-
mous disciform, mostly solitary on brachy-
blasts, also terminal, racemose on dolichoblasts,
in fruiting stage 3x3 mm; peduncle 2. 0-3. 5
(-7.0) mm long, 0. 1-0.2 mm in diameter, slen-
der, initially felty, glabrescent with permanent
glands in cavities. Involucral bracts 4, 2 x 1.2
mm, 2 slightly keeled and margins overlapping
those of other 2, central part slightly thickened,
green to purple, margin broad, membranous,
initially felty, glabrescent. Paleae : those of mar-
ginal florets free, 1.5 x 1.5 mm, slightly keeled,
hard, coriaceous at base, margins membranous,
fringed, abaxially long-lanate, hairs septate;
those of outer disc florets slightly keeled, inner
ones flattened, lanceolate to linear, 1.5 x 0.2
mm, membranous. Marginal female florets 1 or
2, 2.2 mm long; corolla cream-coloured, lamina
obliquely truncate to slightly 3-lobed, shorter
than furcation of style. Style branches linear,
0.5 mm long, apex acute. Ovary oblong, flat-
lish, trigonous, after anthesis long-lanate. Seed
up to 1.0-1. 5 mm long, obovoid, slightly flat-
tened. Disc florets 1 — 4(— 8), functionally male
with sterile ovary, trumpet-shaped, basally yel-
low to cream-coloured, limb 5-lobed, red-pur-
ple. Style truncate, with sweeping hairs. Sta-
mens 5, slightly exserted at maturity. Receptacle
after anthesis with long hairs between involu-
cral bracts and marginal paleae. Chromosome
number : 2 n = 18. Flowering time: mainly June
to September. Figure 10.
E. microcephalus occurs in Namaqualand,
only above 600 in, at the top of mountains or
high on mountain slopes, usually forming
dense, almost impenetrable stands. Where the
closely related E. microphyllus (no. 28) occurs
in the same area, it grows at the foot of moun-
tains or in low-lying areas, but these two are not
habitat-sympatric. Map 18.
Although closely related to E. microphyllus
(no. 28), E. microcephalus can be distinguished
by the thin, small leaves 0.8-1 .6 x 0.3-0. 5 mm,
56
ASTERACEAE: Eriocephalus
scens.
slender, delicate branches and mostly red-pur-
ple young shoots with a diameter of less than 1
mm. The leaves of E. microphyllus are
(1.2-)1.5-4.0(-7.0) x 0.6-0. 8 mm on rigid
branches. Internodes of young shoots of E.
microcepholus are 4. 0-8. 5 mm long as opposed
to the 3-5 mm of those of E. microphyllus. Side
branches are mostly opposite, forming an angle
of 70-90° with the main axis in E. micro -
cephalus, but less than 70° in E. microphyllus.
E. microcephalus is fairly rare and restricted to
a few high mountains, whereas E. microphyllus
is common in Namaqualand.
E. microcephalus did not transplant well and
even cuttings did not survive, but mature E.
microphyllus plants transplanted well and cut-
tings rooted successfully. Common name:
kapokbossie.
Vouchers: Acocks 14980 (PRE); Boucher
3115 (NBG); Esterhuysen 5436 (BOL); Muller
3558 (WIND); Schlechter 11114 (BOL, GRA,
PRE, Z).
30. Eriocephalus spinescens Burch.,
Travels in the interior of southern Africa: 272
(1822); DC.: 147 (1838), pro parte; Harv.: 203
(1865), pro parte. Type: Northern Cape, ‘be-
tween Karree River and Klein Quaggasfontein,
near Frazerburg’, Burchell 1419 (K, holo.!; G-
DC, fragment; PRE & WIND, photo.!).
Robust, many-stemmed, sympodially branch-
ed, spinescent shrubs, 0.5-1. 2 m high and in
diameter. Old stems displaying anomalous sec-
ondary growth, grey to dark grey; young shoots
light yellow-brown, shortly sericeous, glabres-
cent, glandular; older branchlets grey, glabrous.
Leaves opposite on dolichoblasts, decussate
and densely imbricate on brachyblasts, linear,
2.5-3. 5(-5.0) x 0.6-0. 8 mm, entire, perma-
nently densely silver-sericeous, adaxially flat-
tened, slightly concave towards base, abaxially
convex, slightly keeled distally, apex acute.
Capitula heterogamous disciform, solitary, ter-
minal on brachyblasts, 4. 5-5. 2 mm long, ses-
sile or peduncles 1.0-3.5(-5.0) mm long,
densely appressed silver-sericeous. Involucral
bracts 4, oblong-ovate to almost ovate, 4.3 x 2
mm, apex acute, fringed, 2 slightly keeled,
other 2 slightly flattened, central part herba-
ceous, green with purple margin; transparent
membranous margin absent or very narrow.
Paleae : those of marginal florets free, lanceo-
late, up to 4.5 mm long, margins long-pilose,
enveloping floret totally, abaxially long-pilose,
hairs septate, membranous; those of disc florets
lanceolate to oblong, 2-4 x 1-2 mm, membra-
nous, apex acute, margin and abaxially long-
pilose. Marginal female florets 2, cream-
coloured to yellow; corolla tube 5 mm long;
lamina cuneate, 3-lobed, up to 2.2 mm long,
relatively inconspicuous. Style branches flat-
tened, apices acute, 2.5 mm long. Ovary (and
cypsela) oblong, slightly flattened, long-pilose.
Seed 2-3 mm long, oblanceolate, slightly flat-
tened. Disc florets 6-8, functionally male with
sterile ovary, 5 mm long; corolla tubular,
widening upwards, 5-lobed, tubular part yellow
with red-purple margins. Style unbranched,
truncate, apex globose, with sweeping hairs.
Stamens 5, exserted at maturity. Receptacle
after anthesis with abundant white or brown
hairs between involucral bracts and marginal
paleae. Chromosome number: 2n = 36. Flower-
ing time : varying from June to October or
January to March depending on time of rain-
fall.
ASTERACEAE: Eriocephalus
57
The distribution area falls in the transition
zone between winter- and summer-rainfall areas.
E. namciquensis (no. 31), E. spinescens and E.
karooicus (no. 17) are allopatric and succeed
each other from west to east. The distribution of
E. spinescens extends from Calvinia eastwards
in the Northern and Western Cape. This region is
very arid and consists mostly of Arid Karoo and
False Desert Grassveld (Acocks 1975), with an
average annual rainfall of less than 250 mm. E.
spinescens is never found in dense communities,
although it is fairly common in and along water-
courses and seasonal rivulets and in sandy, grav-
elly soil. E. namaquensis, on the other hand,
occurs mostly in high-lying areas on hills and in
stony, clay soil. Map 18.
De Candolle’s (1838) erroneous association
of Burchell’s type material with material from
the current E. karooicus (no. 17), collected by
Drege, resulted in most herbarium material of
E. karooicus being identified as E. spinescens.
Although closely related, there are conspicuous
differences (see discussion under E. karooi-
cus).
E. spinescens is a robust shrub, up to 1 m
high and in diameter, with rigid, strong spines.
E. namaquensis (no. 31), another closely relat-
ed species, is a delicately branched shrub bare-
ly 400 mm tall and in diameter. The capitula of
E. spinescens are relatively large, 4.5-5. 2 mm
long, and fairly shortly pedunculate, 0-3.5
(-5.0) mm long, to almost sessile, borne only
terminally on brachyblasts. It is therefore easy
to distinguish E. spinescens from the closely
related E. karooicus and E. namaquensis , which
are also spinescent, since E. karooicus has a
small, but distinct ray lamina and sessile capit-
ula and E. namaquensis has long-pedunculate
capitula, solitary on brachyblasts, as well as in
racemes borne terminally on dolichoblasts.
Common name: kapokbos.
Vouchers: Maguire 1941 (NBG); Miiller
3599 (WIND); Pole Evans 2248 (BOL); Pole
Evans 2281 (PRE); Van Breda 531 (PRE).
3 1 . Eriocephalus namaquensis M.A.N. Miil-
ler, sp. nov., E. microphyllo DC. affinis sed
indumento argenteo-sericeo, ramisque spinis
terminalibus munitis differt.
Type: Northern Cape, Namaqualand, 29 km
from Loeriesfontein on road to Calvinia, Miiller
3565 (PRE, holo.; K, NBG, WIND).
Many-stemmed, mostly sympodially branch-
ed, spinescent shrubs, 250-450 mm tall and in
diameter. Old stems displaying anomalous sec-
ondary growth, bark grey; young shoots light
brown, often with purple tinge, shortly
sericeous, glabrescent; older branches grey.
Leaves opposite, even on flowering shoots,
decussate and densely imbricate on brachy-
blasts, linear-triangular, 1— 3(— 5 ) x 0.4-0.6 mm,
entire, adaxially flattened and slightly concave
towards base, abaxially convex and keeled dis-
tal ly, permanently densely silver-sericeous,
apex acute. Capitula heterogamous disciform,
solitary on brachyblasts and/or in terminal
racemes, 2. 8-4.0 mm long; peduncles 2.5-12.0
mm long, densely appressed silver-sericeous.
Involucral bracts 4, ovate to narrowly lanceo-
late, 2. 0-3. 5 x 1. 5-2.0 mm, apex acute, 2 slight-
ly keeled, other 2 slightly flattened, central part
herbaceous, green with red-purple tinge, mem-
branous margin narrow or absent. Paleae: those
of marginal florets free, lanceolate, 4-8 mm
long, membranous, margins long-pilose and
enveloping female florets, abaxially long-
pilose, hairs septate; those of disc florets lanceo-
late to narrowly oblong, 2. 5-3.0 mm long,
membranous, margins and abaxial surface long-
lanate, apex acute. Marginal female florets 2,
2. 5-3. 5 mm long, cream-coloured; corolla tube
with short lamina, narrowly cuneate, 3-lobed,
shorter than style furcation. Style branches flat-
tened, apex acute. Ovary (and cypsela) slightly
flattened, long-lanate. Seed 1.5-2. 3 mm long.
Disc florets (3— )5— 8(— 1 0), functionally male
with sterile ovary, 3. 2—4.0 mm long; corolla
tubular, widening upwards, 5-lobed, tubular
part yellow-white with red-purple limb. Style
unbranched, truncate, with sweeping hairs.
Stamens 5, slightly exserted at maturity.
Receptacle after anthesis with dense, white.
58
ASTERACEAE: Eriocephalus
Figijrk 1 1 . Rrioccphalus namaquensis: A I . A2, flowering shoots with inflorescences, x 1 ; B, capitulum, x 6; Cl, C2,
involucral bracts, x 8; D, marginal palea, x 16; E, central palea, x 10; FI, marginal female floret, x 8; F2. branched style,
x 16: Gl, disc floret, x 6: G2, anthers, x 12; G3, truncate style, x 16 (Miiller 3569, WIND).
ASTERACEAE: Eriocephalus
59
Map 19. — • Eriocephalus namaquensis; ■ E. merxmuel-
leri.
long-pilose indumentum between involucral
bracts and marginal paleae. Chromosome num-
ber: 2 n = 18. Flowering time: varying from July
to October and January to March in the differ-
ent rainfall regions. Figure 11.
The western part of the distribution range
falls in the winter-rainfall area, but summer rain
sometimes occurs in the eastern part. The rain-
fall is low, less than 200 mm annually, and the
area is often subject to periodic droughts. This
small shrublet is fairly common in the areas
where it occurs. It is an important component of
the Western Mountain Karoo and the Succulent
Karoo (Acocks 1975). Map 19.
E. namaquensis grows in association with E.
microphyllus var. pubescens (no. 28b), which it
superficially resembles. Especially in the west-
ern part of its distribution range where E.
namaquensis is less spinescent, it is difficult to
distinguish between the two species in their natu-
ral habitat. E. namaquensis has a permanently
silver-sericeous indumentum in contrast to E.
microphyllus var. pubescens which is basically
felty, clearly seen at the growing point. As the
leaves age, the indumentum becomes more
sericeous, slightly wavy, and this can lead to
confusion with E. namaquensis. If there is
doubt, the growing points and peduncles of the
specimen should be examined. In E. microphyl-
lus var. pubescens , the young leaves at the
growing points stick together because of the
intertwined felty indumentum of the overlap-
ping leaves. The leaves of E. namaquensis are
always free.
This species is under-collected, especially in
the southern Karoo (Acocks 1975), partly
because these regions are often subject to
drought and therefore under-collected, and part-
ly because of confusion with species like E.
microphyllus var. pubescens and E. decussatus
(no. 21), which it superficially resembles.
Common name: kapokbos.
Vouchers: Acocks 18489 (PRE); Acocks
19488 (PRE); Hugo 508 (NBG, WIND); Le
Roux 2079 (KPA-J, NBG); Levyns 5032 (BOL).
32. Eriocephalus merxmuelleri M.A.N. Mul-
ler, sp. nov., E. microphyllo DC. affinis sed
habitu ramosissimo, foliis 4— 9(— 15) x 0.5 mm;
lamina florum marginalium femineorum brevis,
0.3-0. 6 mm, interdum brevior sed plerumque
quam stylus longior sed quam stylus cum rami
styli brevior.
Type: Namibia: ‘Aus, an der Strasse nach
Liideritzbucht, 17 August 1963’, Merxmiiller &
Giess 2930 (M, holo.; PRE, WIND).
Erect, many-stemmed, much-branched shrubs,
0.4- 1.2 m high, 0.3-0. 6 m in diameter. Old
stems grey-black, displaying anomalous sec-
ondary growth; young shoots yellow to yellow-
brown; older branches yellow-grey to grey;
branches opposite; brachyblasts short-lived.
Leaves decussate, sometimes alternate on flow-
ering shoots, lanceolate to linear-lanceolate,
obtuse triangular, those on young shoots 4-9
(-14) x 0.5 mm, those on brachyblasts 1-4 x 0.5
mm, entire, very rarely pinnatisect, 3-lobed,
green-grey, indumentum of leaves on growing
point felty sericeous, mature leaves appressed
sericeous to glabrescent (not glabrous), adaxial-
ly more strongly concave from apex to base,
abaxially convex to slightly keeled towards
60
ASTERACEAE: Eriocephalus
Figurf. 12. - Eriocephalus merxmuelleri: A, flowering shoot with inflorescences, x 1; B, branch with leaves, x 10; C,
capitulum. x 5; Dl. D2, involucral bracts, x 10; E, marginal palea, x 16; F, central palea, x 8; Gl. marginal female floret,
x 8; G2. branched style, x 16; 111, disc floret, x 6; H2, anthers, x 20 (Giess 13453, WIND).
ASTERACEAE: Eriocephalus
61
apex, base semi-amplexicaul. Capitula heter-
ogamous disciform, mostly racemose or panicu-
late, rarely solitary on brachyblasts, 3. 5-5.0
mm long; peduncles ( 1 — )2— 7 (— 1 2) mm long,
sparsely felty sericeous, glabrescent. Involucral
bracts 4(5), ovate, 3.2-4.8(-8.0) x 2. 2-4. 8
(-5.5) mm, 2 slightly keeled, other 2 laterally
flattened, central part thickened, herbaceous
with membranous margin, green to red-purple,
sparsely sericeous, glabrescent. Paleae: those
of marginal florets free, slightly keeled, broadly
lanceolate, 3.4-5. 0 x 2. 1-3.0 mm, central part
hard, coriaceous with membranous, fringed
margin, abaxially long-lanate, hairs septate,
enveloping female florets; those of disc florets
narrowly lanceolate to almost linear, 2. 0-3. 5 x
0.6- 1.0 mm, membranous, margins fringed,
abaxially long-lanate. Marginal female florets
( 1 )2— 4, 2.5 mm long; corolla cream-coloured;
lamina short, 0. 3-0.6 mm long, sometimes
shorter than but mostly longer than style furca-
tion, but shorter than style branches, cuneate to
oblong, 3-lobed. Style branches 0.3-1. 5 mm.
Ovary (and cypsela) oblong, slightly flattened,
1.5-2. 5 mm long, long-lanate. Seed oblong-
ovoid, laterally flattened, 1. 5-2.0 mm long.
Disc florets ( 1— )5 or 6(-9), functionally male
with sterile ovary; corolla tubular, widening
upwards, basally cream-coloured to yellow,
limb red-purple, 2. 5-3. 5 mm long. Style
unbranched, apex slightly globose with sweep-
ing hairs. Stamens 5. Receptacle after anthesis
densely white long-pilose between involucral
bracts and marginal paleae. Chromosome num-
ber: 2 n = 54. Flowering time: December to
April and from June to September in the differ-
ent rainfall areas. Figure 12.
The species occurs in both summer- and
winter-rainfall areas and extends over the bor-
der between South Africa and Namibia, but it is
restricted to the Namaqualand Broken Veld
(Acocks 1975). Map 19.
E. merxmuelleri is closely related to E. mi-
crophyllus var. pubescens (no. 28b), which
occurs in Namaqualand.
Common name: kapokbos.
Vouchers: Giess 13454 (WIND); Giess, Volk
& Bleissner 7173 (WIND); Miiller 1380 (PRE);
Pearson 4243 (BOL); Rowland, Scott & Steyn
PRE43673 (PRE).
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LINNAEUS. C. 1760. Plantae rariores africanae. Salvius,
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LINNAEUS. C. (fil.). 1782 ('1781')- Supplementum plan-
tarum systematis vegetabilium. Orphanotrophei,
Brunsvigae.
LOUDON, J.C. 1838. Arboretum et fruticetum brittanicum.
Longman, London.
LOUDON. J.C. 1855. An encyclopedia of plants, new edn.
Longman. London.
MARLOTH. H.W.R. 1932. The flora of South Africa 3,2.
Darter, Cape Town.
MERXMULLER. H. & EBERLE. E. 1957. Compositen
Studien VI. Mitteilungen der Botanischen Staats-
sammlung, Miinchen 2: 321-324.
MERXMULLER. H. 1967. Eriocephalus and Lasiosper-
mum. Prodromus einer Flora von Siidwestafrika
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MOENCH. C. 1794. Supplementum ad methodum plantas.
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MOORE. S. LE M. 1902. A contribution to the composite
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MOORE. S. LE M. 1904. Beitrage zur Kenntnis der afri-
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MULLER. M.A.N. 1988. ’n Morfologiese en taksonomiese
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Journal of South African Botany 30: 49-52.
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um botanicum ... 2.
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Pretoria.
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ASTERACEAE: Eriocephalus
63
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southern Africa, Vol. 1: 147-170.
WILLDENOW, C.L. VON. 1803. Species plantarum 3,3.
Nauk, Berlin.
64
ASTERACEAE: Lasiospermum
9321000 2. LASIOSPERMUM
by M.A.N. Muller, P.P.J. Herman & H.H. Kolberg
(Literature references on p. 72)
Lasiospermum Lag., Genera et species plantarum: 31 (1816) nom. cons, provis.; Trevir.: 205
(1826); Cass.: 304 (1822): Rchb.: 225 (1831); Less.: 250 (1832); DC.: 37 (1838); Endl.: 431,432
(1838); Harv.: 153 (1865); Benth.: 416 (1873); Adamson & T.M. Salter: 803 (1950); Merxm.: 108
(1967); R. A. Dyer: 702 (1975); M.A.N.Muller: 124 (1988); K.Bremer & Humphries: 94 (1993);
K. Bremer: 451 (1994); P.P..I. Herman et al.: 146 (2000); non Lasiospermum Fisch.: 34 (1812).
Type: L. pedimculare Lag. (type cons.).
Eriosphaera F.Dietr.: 221. 222 (1817).
Lanipila Burch.: 259 (1822).
Mataxa Spreng.: 297 (1827).
Eriocarpha Lag. ex DC.: 38 (1838).
Annual or perennial herbs, sometimes decumbent, rooting at nodes. Leaves alternate, long-
pilose to glabrous, pinnatisect to bipinnatisect; lobes narrowly linear, rarely entire. Capitula pedun-
culate, terminal, solitary or paniculate, many-flowered, homogamous discoid or heterogamous
radiate. Involucre broadly saucer-shaped; involucral bracts in 2-4 rows, narrowly oblong to ellip-
tic to almost square, with membranous margin and apex, long-pilose or felted to glabrous.
Receptacle broad, flat to conical with membranous paleae. Ray florets when present, few, female.
Style bifurcate, with linear, truncate branches. Ovary oblong. Pappus absent. Disc florets numer-
ous, campanulate; corolla 5-lobed. Stamens 5, anthers fused, ecaudate and ecalcarate, with lanceo-
late apical appendage; endothecial tissue polarised. Style bifurcate with linear, truncate branches.
Ovary ovoid, slightly triangular, without any appendages, after anthesis with dense, woolly indu-
mentum, hairs often septate. Cypselas oblong-ovoid, slightly flattened, smooth, dark yellow-
brown. Pappus absent. Basic chromosome number, x = 9 (2 n - 18).
The genus name Lasiospermum Lag. (1816) is a later homonym of Lasiospermum Fisch. (1812)
and should be rejected according to Article 64.1 of the ICBN. Cassini (1822) was aware of
Lasiospermum Fisch., but recommended the conservation of the name Lasiospermum for Lagasca’s
genus as Fischer ‘published only the name without a description or indicating diagnostic features;
therefore he (Cassini) felt that the name Lasiospermum should be preserved (retained) for
Lagasca’s genus and Fischer’s genus should receive a different name’. As type species he named
L. pedunculare Lag.
The genus name Scorzonera L. (1735 & 1737) (Asteraceae) was conserved against
Lasiospermum Fisch. (1812). Cassini’s (1822) motivation for the conservation of the genus name
Lasiospermum Lag. is herewith supported, especially as it has been in use for such a long time.
De Candolle ( 1 838) classified the taxa of the genus Lasiospermum in two sections, namely section
Eulasiospermum DC. with discoid capitula and section Lanipila (Burch.) DC. with radiate capitula.
According to Article 21 of the ICBN (Stafleu 1978): ‘The epithet of a subgenus or section is not
to be formed from the name of the genus to which it belongs by adding the ending -oides or -opsis,
or the prefix Eu-.' The implication of this rule means that the section epithet Eulasiospermum as
published by De Candolle ( 1 838) is invalid. The name Lasiospermum is therefore proposed for this
section (Article 22, ICBN).
ASTERACEAE: Lasiospermum
65
The section Lanipila is based on the genus Lanipila described from Burchell 1336 (herbarium
specimen), housed in Kew. Treviranus’s (1826) description of Lasiospermum radiation , based on
the same specimen, does not agree with the Kew specimen, but fits Lasiospermum bipinnatum.
Because of the confusion about the identity of the specimen used by Treviranus for his species
description, the use of the name Lanipila for the section is unacceptable according to Article 22
(ICBN) as the type of Lanipila is in the section Eulasiospermum. The new name Radiation
M. A. N. Muller is proposed for this section.
Key to sections of the genus Lasiospermum
Capitula homogamous discoid; disc florets bisexual, tubular sect. Lasiospermum
Capitula heterogamous radiate; ray florets female; disc florets bisexual, tubular .... sect. Radiation
Section Lasiospermum. Type species; L. pedunculare Lag.
Lasiospermum Lag. sect. Eulasiospermum DC.
Capitula homogamous discoid. Disc florets bisexual.
Section Radiatum M.A.N. Muller , sect. nov. Type species: L. bipinnatum (Thunb.) Druce.
Capitula heterogama radiata. Flosculi radii feminei. Flosculi disci hermaphroditi.
Capitula heterogamous radiate. Ray florets female. Disc florets bisexual.
Key to the species of the genus Lasiospermum
la Capitula heterogamous radiate; rays female, white, pale red-purple, red-purple or purple
with yellow apex (sect. Radiation)'.
2a Ray florets white or pale red-purple, up to 15 x 2.5 mm; perennial herbs . . . 1. L. bipinnatum
2b Ray florets red-purple or red-purple with yellow apex; very small (± 1 mm long, rarely
up to 3.5 mm long), annual herbs 2.L. brachyglossum
lb Capitula homogamous discoid, all florets bisexual, tubular, golden yellow (sect.
Lasiospermum ):
3a Terminal part of peduncle and involucral bracts long-pilose; involucral bracts permanent-
ly hairy and with narrow, inconspicuous, membranous margin 3. L. pedunculare
3b Terminal part of peduncle and involucral bracts densely felted; involucral bracts
glabrescent and with obvious, broad, transparent, membranous margin . . . 4. L. poterioides
Key to Lasiospermum species based on vegetative and geographic features
la Annual herbs 2. L. brachyglossum
lb Perennial herbs:
2a Leaves initially sparsely hairy, soon glabrous; distributed in North-West, Gauteng, the
Free State, Lesotho and the Northern, Western and Eastern Cape 1. L. bipinnatum
2b Leaves initially densely hairy, glabrescent to glabrous; restricted to the Northern and
Western Cape and Roggeveld Mountains;
66
ASTERACEAE: Lasiospermum
Figure 13. — Lasiospermum hipinnatum A, branch with capitula, x I; Bl, ray floret, x 4: B2, branched style, x 20;
Cl bisexual disc floret, x 8; C2, anthers, x 16; C3, branched style, x 16; Dl, D2, paleae, x 8; El. E2, E3, involucral bracts,
' 8; F. cypscla with indumentum, x 4 (Midler 4088, WIND).
ASTERACEAE: Lasiospermum
67
3a Older leaves sparsely long-pilose, rarely glabrous, apex of leaf lobes acute; restricted
to strip parallel with the west coast (Northern and Western Cape), between 300 and
1 500 m altitude 3. L. pedunculare
3b Older leaves glabrescent, apex of leaf lobes obtuse; restricted to Roggeveld
Mountains (Northern Cape), above 1 500 m altitude 4. L. poterioides
1. Lasiospermum bipinnatum (Thunb.)
Druce in Report of the Botanical Exchange
Club of the British Isles for 1916: 631 (1917);
Adamson & T.M. Salter: 803 (1950). Type:
Western Cape, Langkloof, Eselsjagt (Eseljag),
Thunberg 20232 (UPS, holo.; PRE & WIND,
photo.!).
Lidbeckia bipinnata Thunb.: 161 (1800); Willd.: 2165
(1803); Thunb.: 694 (1823). Lancisia bipinnata (Thunb.)
Pers.: 463 (1807). Matricaria bipinnata (Thunb.) Spreng.:
582 (1826).
Mataxa capensis Spreng.: 303 (1827); G.Don: 368
(1839). Type: based on Lasiospermum radiatum.
Perennial, erect to ascending, much-branch-
ed herbs, up to 0.6 m high. Older stems decum-
bent, rooting at some nodes; growing points ini-
tially sparsely long-pilose, soon glabrous.
Leaves alternate, bipinnatisect, lobes linear,
sometimes slightly falcate, yellow-green, 30-80
mm long, smaller on flowering shoots,
mucronate with hard white mucro, margins
irregularly serrate; young leaves sparsely long-
pilose, soon glabrous; petiole semi-amplexicaul
basally, sheathing, margins irregularly serrate.
Capitula heterogamous radiate, terminal, soli-
tary on long glabrous peduncles. Involucral
bracts imbricate, in 3 or 4 rows, enlarging from
outside to inside, up to 3 x 2 mm, with narrow,
transparent margins, glabrous; outer narrowly
oblong to lanceolate, slightly keeled, inner
broadly lanceolate to ovate, more flattened.
Receptacle disc-shaped, flattened. Paleae mem-
branous with green main vein, ovate, 3. 0-3. 5
mm long, marginal paleae keeled, central paleae
flattened, margins irregularly dentate. Ray flo-
rets female, 24^-0, up to 15.0 x 2.5 mm, white
or pale red-purple. Style bifurcate, apices with-
out sweeping hairs. Disc florets 130-150, pale
yellow, bisexual; corolla tubular, 5-lobed (-den-
tate), 3. 5-4. 5 mm long. Style bifurcate with lin-
ear branches, distally truncate, with sweeping
hairs. Stamens 5. Ovary slightly triangular, yel-
low-brown, smooth, lanceolate to oblong-ovoid,
slightly flattened, after anthesis with dense,
white, woolly indumentum, hairs septate.
Cypsela dark yellow-brown, smooth, lanceolate,
flattened triangular. Flowering time : varying
from January to December with a peak from
August to October (winter rainfall) and
November to April (summer rainfall). Figure 13.
L. bipinnatum is widely distributed in North-
West, Gauteng, the Free State, Lesotho and the
Northern, Western and Eastern Cape in both
summer- and winter-rainfall regions. In some
areas it is even regarded as a weed in agricul-
tural land. It grows in dark brown sandy loam,
sandy soil, clay and even dolerite, with prefer-
ence for moist areas like vleis, marshes, river
banks and roadsides where pools of water have
formed. Map 20.
Burchell (1822) described the genus Lani-
pila Burch., without mentioning any species,
from material obtained from the Roggeveld
Map 20. — • Lasiospermum bipinnatum; ■ L. poterioi-
des.
68
ASTERACEAE: Lasiospermum
Mountains as follows: ‘ Lanipila C.G. 1336,
Genus Cotulae affine. Nomen a lana et pila; ob
semina lana involuta, et in capitulo spherico
conglomerata .’ The holotype of the genus
Lanipila (. Burchell 1336 , K!) was collected
between Jackalsfontein and Kuilenberg, near
Sutherland. Treviranus described Lasiosper-
mum radiation in 1826 with 'floribus radiatis’,
i.e. with ray florets, and cited the type as
Burchell 1336 of Lanipila Burch. The descrip-
tion of Lasiospermum radiation agrees totally
with that of Lasiospermum bipinnatum, al-
though the type at Kew on which the descrip-
tion was supposedly based, represents Lasio-
spermum poterioides Hutch., a taxon described
only in 1946. It is concluded that the material of
Burchell used by Treviranus for the description
of Lasiospermum radiation was not the same as
the material at Kew.
Although the plants are eagerly browsed by
sheep, Walsh (1909) reported that it probably
caused animal poisoning. This probability is
confirmed by notes on herbarium specimens
and according to Dr T.F. Adelaar (pers. comm.)
of the Onderstepoort Veterinary Institute, it con-
tains a liver toxin. According to Vahrmeijer
(1981), it causes photosensitivity. Inhabitants of
the Middelburg District (Eastern Cape) make a
decoction of the plant and use it for affection of
the chest. The aromatic nature is apparently
connected to disinfection as it is used as such by
the South-Sotho in an ointment used to disinfect
a sick bay (Phillips 1917).
The ray florets are horizontally orientated in
relation to the capitula during the day, but they
are recurved at night so that the capitulum
resembles a shuttlecock. Ray florets are absent
in some specimens, e.g. Compton 10248 (NBG)
and Schlechter 8961 (Z). It does happen spo-
radically that rays are absent, giving rise to
incorrect identifications, e.g. as L. pedunculare.
Vouchers: Codd 8072 (PRE); Esterhuysen
29688 (BOL); Flanagan 1352 (BOL, SAM);
Jacobsz 2157 (NBG); Jacot-Guillarmod 4756
(PRE).
2. Lasiospermum brachyglossum DC. in
Prodromus: 38 (1838); Harv.: 154 (1865);
Merxm.: 108, 109 ( 1967). Type: Northern Cape,
‘Zilverfontein, auf der Flache’, Drege 2863 (G-
DC, holo.; P!, PRE, photo!, SAM!).
Erect, rarely ascending, annual herbs, 100-
400 mm high. Stems branched or unbranched
from the base, all ending in inflorescences.
Leaves alternate, sometimes rosulate at base;
petioles of basal leaves well developed, with
stipule-like appendages; appendages irregularly
dentate; lamina pinnatisect to bipinnatisect,
entire on flowering shoots, 15-50 x 10-30 mm;
lobes of second order linear, 0.5- 1.0 mm in
diameter, sometimes semifalcate, mucronate
with a hard white mucro. Capitula heteroga-
mous radiate, in panicles or racemes with oldest
capitula terminal and younger ones proximal,
semiglobose when young, up to 5 mm in diam-
eter, in fruiting stage almost globose, up to 15
mm in diameter; distal part of peduncle
glabrous. Involucral bracts in 3 or 4 rows,
imbricate, broadly membranous, 1.5-2. 5 x
1.0-1. 5 mm, glabrous, apex obtuse, outer bracts
relatively narrow, linear, inner ones ovate.
Receptacle conical. Marginal paleae ovate, cen-
tral paleae lanceolate with arista which is the
continuation of the main vein, main vein dis-
tinctly yellow-brown, rest of palea transparent,
membranous, 1.2- 1.8 x 0.8-1. 2 mm. Ray florets
12-20, female, strap-shaped; corolla red-purple
to red-purple with yellow distal part, rarely com-
pletely yellow, very short, 1 mm long, rarely up
to 3.5 x 1.6 mm, 3-dentate, cuneate. Style
branches linear, 0. 1-0.2 mm long. Ovary
oblong-ovoid. Disc florets 130-150, bisexual;
corolla tubular, 5-dentate (-lobed), pure yellow
to yellow with red-purple margin, up to 2.5 mm
long. Stamens 5. Style branches linear, truncate,
with sweeping hairs at apex. Ovary oblong;
ovaries of both ray and disc florets with dense,
woolly indumentum after anthesis, hairs septate.
Flowering time : July to November, with a peak
from July to September.
The distribution of L. brachyglossum var.
brachyglossum extends from Aus in Namibia to
the Oudtshoorn District in the Western Cape
ASTERACEAE: Lasiospermum
69
Map 21. — Lasiospermum brachyglossum.
along the western part of the continent and is
mainly confined to the winter-rainfall area of
southern Africa. Var. sinaicum is confined to the
Sinai Desert. Map 21.
Common names: knoppiesopslag (Nama-
qualand) (Le Roux & Schelpe 1984), knoppies-
stinkkruid (Pofadder, from Conradie 1 , NBG).
Vouchers: Acocks 16908 (PRE); Bolus 392
(BOL, SAM); Giess 14640 (PRE, WIND);
Maguire 1982 (BOL, NBG); Van der Schijjf
8092 (PRE).
3. Lasiospermum pedunculare Lag.,
Genera et species plantarum: 31 (1816); DC.:
38 (1838); Harv.: 154 (1865). Iconotype:
P.Micheli, Nova plantarum genera t. 27
(1729).
Santolina erecta Lam.: t. 671. fig. 4 (1796); Poir.: 508
(1805); non Barr.: 522 (1714); nec 5. erecta Pers.: 407
(1807); nec 5. erecta et S. eriosperma Reichard: 730 (1780);
Desf.: 99 (1804).
IS. pinnata Donn: 107 (1800). Type: ? Western Cape,
collected in 1791, collector unknown.
S. eriosperma Pers.: 407 (1807).
Eriosphaera multifida F.Dietr.: 221 (1817).
S. alpina Bertol.: 43 (1819); Loudon; 694 (1855); non S.
aipina L.: 1180 (1763); L.: 616 (1774); Willd.: 1800(1803);
Guss.: t. 58 (1826); nec Lasiospermum alpinum (L.) Rchb.:
225(1831).
Eriocarpha peduncularis Lag. ex DC.: 38 (1838).
L. eriospermum (Pers.) G.Don: 337 (1839).
L. erectum (Lam.) Druce: 631 (1917).
Erect to ascending, sometimes mat-form-
ing, much-branched, perennial herbs, 150-200
mm tall, up to 1 m in diameter. Older stems
decumbent, sometimes rooting at nodes; stems
cylindrical, permanently white, long-pilose;
young growing points densely white, long-
pilose, but sparsely hairy with age. Leaves:
petiole basally semi-amplexicaul, sheathing
with membranous margins, axils densely long-
pilose, flattened to distal point; lamina bipin-
natisect, 50-150 x 10-30 mm, grey-green
because of long-pilose indumentum; lobes of
first order up to 15 mm long, each with 3 or 4
incisions, lobes of second order linear, 3-6 x
0.4-0. 6 mm, apices acute, mucronate, with
hard white mucro; lobes mostly alternate,
sometimes opposite; older leaves glabrescent
but never glabrous; petiole of leaves on flow-
ering shoots basally with serrate, stipule-like
appendages, petiole of leaves nearer to capitu-
la decreasing in size with reduction in number
of lobes until totally absent. Capitula homoga-
mous discoid, solitary, terminal on long
peduncle; peduncle long-pilose. Involucral
bracts in 3 or 4 rows, imbricate, 3.5 x 2.5 mm,
linear to lanceolate and increasing in size to-
wards the centre to obtuse-triangular, with rel-
atively narrow membranous margin, abaxially
permanently pilose, adaxially glabrous.
Receptacle disc-shaped, flattened. Paleae mem-
branous, transparent, ovate, 1-3-dentate, 3-4
mm long, with main vein, paleae maturing
before florets. Disc florets 130-140, all bisex-
ual; corolla tubular, 5-dentate (-lobed), golden
yellow, turning red-brown with age, 4-5 mm
long. Stamens 5, fused. Style bifurcate with
linear branches and truncate apices, apices
with sweeping hairs. Ovary’ after anthesis with
dense, woolly indumentum. Flowering time:
correlated with the rainy season, August to
October, but a few flowering specimens col-
lected in December and March. Figure 14.
70
ASTERACEAE: Lasiospermum
Flea re 14. -Lasiospermum pedunculare: A. branch with inflorescences, x 1 : B. capitulum, x 2; C 1 . C2, bisexual disc
florets, x 8: D, anthers, x 16; E. branched style, x 20: FI-F5. involucral bracts, x 8; G, palea, x 10; FI. cypsela with woolly
indumentum, x 4 (Muller 4030, WIND).
ASTERACEAE: Lasiospermum
71
Map 22. — Lasiospermum pedunculare.
The distribution of L. pedunculare in south-
ern Africa is, with a few exceptions, restricted
to the winter-rainfall area, all along the west
coast. Until now, it has been found on sandy,
loam and clay soils at altitudes of 300-1 500 m.
Map 22.
The works of earlier researchers like
Linnaeus (1763, 1774), Reichard (1780) and
Desfontaines (1804) caused much confusion
about the identity of Santolina alpina L. and S.
erecla Lam. The type material of both these
species originates from Italy and shows close
relationships. Morphologically S. alpina closely
resembles Lasiospermum pedunculare , except
for the glabrous cypselas of S. alpina in com-
parison to the woolly cover of the cypselas of L.
pedunculare. Another misinterpretation, which
has led to further confusion, is the fact that
Linneaus cited the specimen of Micheli as a
synonym of S. alpina, an error perpetuated by
later researchers.
Only photographs of S. alpina and related
taxa were studied. Another problem encoun-
tered, was locating type material. It is clearly
mentioned that S. maschalantha Spreng. (in
Schrader 1799, Journal fiir die Botanik) was
described from material ‘Aus dem National-
Museum zu Paris’, but no such material could
be located there or in other herbaria. In this spe-
cific case, the description mentioned: ‘paleis
receptaculi lanatis’, i.e. paleae lanate. The prob-
ability does exist that it is the cypselas that are
lanate, meaning that it is a synonym of
Lasiospermum pedunculare. Santolina pinnata ,
put into synonymy under Lasiospermum pedun-
culare by various earlier researchers (Persoon
1807; Bertolini 1819; Don 1839), originated
from southern Africa according to Donn
(1800) — the first indication that the taxon
occurs in southern Africa. In spite of many
attempts, the type material of this taxon could
not be located. A photocopy of material identi-
fied as such from Liverpool [Herbarium,
Merseyside Country Museums (LIV)] does not
agree with southern African material at all.
Italian representatives of the species differ from
southern African material by the presence of
rays (Bertoloni 1819), while southern African
specimens have discoid capitula. Further confu-
sion is added by Reichenbach’s (1831) descrip-
tion, stating that the ray florets are white and
female and that the cypselas are woolly.
The aromatic smell of the plants has led to
the common names laventelkatoen or lavender
cotton by Donn (1800). The lanate indumentum
of the cypselas led to the names veelvertakte
wolbol or vielspaltige Wollkugel (Dietrich
1817) and wolvrug or Wollfrucht (Reichenbach
1831). It is known that the plants are eagerly
browsed by sheep and no poisoning of sheep
has been reported to date.
Vouchers: Bohlmann 202 (NBG); Compton
11544 (NBG); Compton 11792 (NBG); Ester-
huysen 5994 (BOL, PRE); Leipoldt 3531
(BOL).
4. Lasiospermum poterioides Hutch., A
botanist in southern Africa: 140 ( 1946). Type:
Northern Cape, Sutherland, between Matjies-
fontein and Sutherland. Hutchinson 693 (K,
holo.!; BOL!).
Lanipila (sic) Burch.: 259 (1822). Type: Northern Cape,
Roggeveld Mountains, between Jackalsfontein and
Kuilenberg, near Sutherland, Burchett 1336 (K, holo.!:
PRE, photo.!).
72
ASTERACEAE: Lasiospermum
Much-branched, mat-forming, perennial herbs,
rarely erect, rather ascending, 100-300 mm tall,
up to 400 mm in diameter. Older stems decum-
bent, glabrescent, sometimes rooting at nodes;
young growing points densely long-pilose.
Leaves initially rosulate; petiole broadened
basally, semi-amplexicaul, relatively short with
serrate-dentate margins; lamina pinnatisect to
bipinnatisect, 40-75 x 5-15 mm. initially deli-
cately long-pilose, glabrescent; leaves on flow-
ering shoots basally pinnatisect with decreasing
number and size of lobes transending to pedun-
cle; lobes oblanceolate, mucronate with hard,
white mucro. Capitula homogamous discoid,
solitary, terminal, pedunculate, in flower ± 15
mm in diameter, in fruit more than 20 mm in
diameter; distal part of peduncle and involucral
bracts felted, soon glabrescent. Involucre saucer-
shaped, involucral bracts in 3 or 4 rows, imbri-
cate, broadly ovate, rarely obtuse-triangular,
increasing in width from outer to inner, with
conspicuous, broad, transparent, membranous
margin, up to 4 x 3.2 mm. Receptacle flattened.
Paleae broadly ovate to oblong-elliptic, irregu-
larly dentate, transparent, membranous, 3-4 mm
long, outer many-veined, inner with conspicu-
ous main vein. Disc florets ± 250, bisexual;
corolla tubular, 5-dentate (-lobed), golden yel-
low, turning red-brown with age, 4.0-5. 2 mm
long. Stamens 5. Style bifurcate, branches linear,
truncate, with sweeping hairs on distal apices.
Ovary up to 2.5 mm long, after anthesis with
dense, woolly indumentum. Flowering time :
August to October.
This taxon is restricted mainly to the Suther-
land District, the Berg-Roggeveld of the
Beaufort clay series, with a single specimen
from Williston (Northern Cape). Map 20.
L. poterioides can easily be confused with L.
pedunculare . The presence of the felty indu-
mentum at the tip of the peduncle and involu-
cral bracts, the conspicuous, broad, transparent,
membranous margin of the bracts, the leaf lobes
that are almost obtuse and the distribution with
one exception above 1 500 m altitude, distin-
guish L. poterioides from L. pedunculare.
Common names for this taxon are: gansgras,
ganzies gras ( Marloth 9720 , PRE), Reveldsgras
( Hanekom 2124 , PRE). It is called gansgras as
it is one of the first plants turning green after the
rain and is then utilised by geese. Although
eaten by geese without any ill effects, Hanekom
( Hanekom 2124. PRE) mentioned that it caused
'dikkop' in sheep.
Vouchers: Acocks 16931 (PRE); Bayliss 556
(NBG); Hanekom 2124 (PRE); Hutchinson 693
(BOL, K, PRE); Marloth 9720 (PRE).
REFERENCES
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Cape Peninsula. Juta, Cape Town.
BARRELIER. J. 1714. Plantae per Gallium, Hispaniam et
Italiam observatae Iconibus aeneis exhibitae.
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BENTHAM. G. 1873. Ordo 88. Compositae. In G. Bentham
& J.D. Hooker, Genera plantarum 2: 163-533.
Reeve, London.
BERTOLONI, A. 1819. Amoenitates italicae sistentes opus-
cula Typis Annesii de Nobilibus, Bologna.
BREMER. K. & HUMPHRIES. C. 1993. Generic mono-
graph of the Asteraceae-Anthemideae. Bulletin of
the Natural History Museum, London (Botany
series) 23,2.
BREMER. K. 1994. Asteraceae, cladistics and classifica-
tion. Timber Press, Oregon.
BURCHELL. W.J. 1822. Travels in the interior of southern
Africa. Vol. 1: 232, 259, 272. Longman, London.
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DON, G. 1 839. Hotlus brittanicus. edn 2. Ridgeway, London.
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DRUCE. G.C. 1917. Nomenciatorial notes: chiefly African
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631.
ASTERACEAE: Lasiospermum
73
DYER, R.A. 1975. The genera of southern African flower-
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ENDLICHER, S.F.L. 1 838. Genera plantarum , Part 1 . Beck,
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FISCHER, F.E.L. VON. 1812. Catalogue du jardin des
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Razoumoffsky a Gorenki pres de Moscou, edn 2.
N.S. Vsevolojsky, Moscow.
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W.H. Harvey & O.W. Sonder, Flora capensis 3: 153,
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HERMAN, P.P.J., RETIEF, E„ KOEKEMOER. M. & WEL-
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Institute, Pretoria.
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LAGASCA Y SEGURA. M. 1816. Genera et species plan-
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Salvius, Stockholm.
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MERXMULLER, H. 1967. Eriocephalus and Lasiosper-
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Eriocephalus L. (Asteraceae) in suidelike Afrika.
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PERSOON. C.H. 1807. Synopsis plantarum, sen enchiridi-
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SPRENGEL, C.P.J. 1827. Systema vegetabilium 4,2: 297,
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THUNBERG, C.P. 1800. Syngenesia. Prodromus plan-
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Leopoldino-Carolinae Germanicae Naturae Curio-
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VAHRMEIJER, J. 1981. Poisonous plants of southern
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WILLDENOW, C.L. VON. 1803. Species plantarum 3,3.
Nauk, Berlin.
ASTERACEAE: Anthemideae: Eriocephalus, Lasiospermum
75
INDEX*
Eriocarplia Lag. ex. DC., 64
peduncularis Lag. ex DC., 69
Eriocephalus L., 1
ajfinis DC., 39
africanus L., 1, 23
var. africanus, 25
var. paniculatus (Cass.) M.A.N. Miiller, P.P.J. Herman &
H.H.Kolberg, 26
ambiguus (DC.) M.A.N. Miiller, 42
aromaticus C.A.Sm., 16
aspalathoides DC., 39, 42
aspalathoides DC. var. ambiguus DC., 42
brevifolius (DC.) M.A.N. Muller, 21
capitellatus DC., 10
corymbosus Moench, 23
decussatus Burch., 39
dinteri S. Moore, 33
eenii S. Moore, 44
ericoides (L.f.) Druce, 47
subsp. ericoides, 48
subsp. griquensis M.A.N. Muller, 49
eximius DC., 1 1
frutescens R.Br., 23
giessii M.A.N. Muller, 34
glaber Thunb., 47
var. pubescens Harv., 53
var. sessiliflorus Sond. ex Harv., 47
glandulosus M.A.N. Muller, 49
grandiflorus M.A.N. Muller, 29
hirsutus Burtt Davy, 44
karooicus M.A.N. Muller, 31
kingesii Merxm. & Eberle, 41
klinghardtensis M.A.N. Miiller, 19
longifolius M.A.N. Miiller, 11
luederitzianus O.Hoffin., 44
macroglossus B.Nord., 9
merxmuelleri M.A.N. Muller, 59
microcephalus DC., 55
microphyllus DC., 51
var. carnosus M.A.N. Miiller, 54
var. microphyllus, 53
var. pubescens (DC.) M.A.N. Midler, 54
namaquensis M.A.N. Muller, 57
paniculatus Cass., 26
parviflorus Dinter, 33
pauperrimus Merxm. & Eberle, 41
pedicellaris DC., 15
pinnatus O.Hoffin., 8
pteronioides DC., 15
pubescens DC., 54
pubescens sensu Merxm., 44
punctulatus DC., 17
var. tenuifolius (DC.) Harv., 18
var. brevifolius DC., 21
var. pedicellaris (DC.) Harv., 15
purpureus Burch., 13
racemosus Gaertn. non L.. 26
racemosus L., 35
var. affinis (DC.) Harv., 39
var. racemosus, 37
rangei Muschl., 22
scariossisimus S. Moore, 22
scariosus DC., 22
septifer Cass., 23
septulifer DC., 23
sericeus Gaudich. ex DC., 26
simplicifolius Salisb., 35
spicatus Burm. ex DC., 35
spinescens Burch., 56
spinescens sensu DC., 31
squarrosus Muschl. in Dinter, 44
tenuifolius DC., 18
tenuipes C.A.Sm., 30
umbellulatus Cass.. 26
var. argenteus DC., 26
var. glabriusculus DC., 26
variifolius Salisb., 23
virgatus Dinter, 22
xerophilus Schltr., 13
Eriosphaera F.Dietr., 64
multifida F.Dietr., 69
Lancisia bipinnata (Thunb.) Pers., 67
Lanipila Burch., 64, 71
Lasiospermum Lag., 64
sect. Eulasiospermum DC., 65
sect. Lasiospermum, 65
sect. Radiatum M.A.N. Midler, 65
bipinnatum (Thunb.) Druce, 67
brachyglossum DC., 68
erectum (Lam.) Druce, 69
eriospermum (Pers.) G.Don, 69
pedunculare Lag., 64, 69
poterioides Hutch., 71
Lidbeckia bipinnata Thunb., 67
Mataxa Spreng. , 64
capensis Spreng., 67
Matricaria bipinnata (Thunb.) Spreng., 67
Monochlaena racemosus Cass., 26
Santolina alpina Bertol., 69
erecta Lam., 69
eriosperma Pers., 69
? pinnata Donn. 69
Tarchonantlms ericoides L.f., 47
Synonyms are in italics.
A-l
APPENDIX
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. Occasional contributions to the Flora are published in
Bothalia under the title FSA contributions.
Exotic families are marked with an asterisk.
Published volumes and parts are shown in bold.
INTRODUCTORY VOLUMES
The genera of southern African flowering plants, Vols 1 (1975) and 2 (1976). Replaced by Seed plants of southern
Africa: families and genera, published as Strelitzia 10 (2000).
Botanical exploration of southern Africa (1981)
CRYPTOGAM VOLUMES
Charophyta (as Vol. 9 in 1978)
Bryophyta: Part 1: Musci: Fascicle 1: Sphagnaceae, Andreaeaceae, Fissidentaceae, Nanobryaceae, Archidiaceae,
Ditrichaceae, Seligeriaceae, Dicranaceae, Calymperaceae, Encalyptaceae,
Pottiaceae, Bryobartramiaceae, Grimmiaceae (1981)
Fascicle 2: Gigaspermaceae, Ephemeraceae, Funariaceae, Splachnaceae, Bryaceae, Mnia-
ceae, Eustichiaceae, Rhizogoniaceae, Aulacomniaceae, Bartramiaceae (1987)
Fascicle 3: Erpodiaceae, Rhachitheciaceae, Ptychomitriaceae, Orthotrichaceae, Rhabdowei-
siaceae, Racopilaceae, Fontinalaceae, Wardiaceae, Hedwigiaceae, Cryphaeaceae,
Leucodontaceae, Prionodontaceae, Trachypodaceae, Pterobryaceae, Meteoria-
ceae, Leptodontaceae, Neckeraceae, Thamnobryaceae, Hookeriaceae (1998)
Fascicle 4: Fabroniaceae, Leskeaceae, Thuidiaceae, Rigodiaceae, Amblystegiaceae. Brachy-
theciaceae, Entodontaceae, Plagiotheciaceae, Catagoniaceae. Sematophyllaceae,
Hypnaceae, Hylocomiaceae, Polytrichaceae
Hepatophyta: Part 1: Marchantiopsida: Fascicle 1: Targioniaceae, Lunulariaceae, Aytoniaceae, Cleveaceae, Exormo-
thecaceae, Marchantiaceae, Oxymitraceae, Rieciaceae (1999)
Anthocerotophyta
Pteridophyta (1986)
FLOWERING PLANTS VOLUMES
Vol. 1: Stangeriaceae, Zamiaceae, Podocarpaceae, Pinaceae*, Cupressaceae, Welwitschiaceae, Typhaceae, Zoster-
aceae, Potamogetonaceae, Ruppiaceae, Zannichelliaceae, Najadaceae, Aponogetonaceae, Juncaginaceae,
Alismataceae, Hydrocharitaceae (1966)
Vol. 2: Poaceae
Vol. 3: Cyperaceae, Arecaceae, Araceae, Lemnaceae. Flagellariaceae
Vol. 4: Part 1 : Restionaceae
Part 2: Xyridaceae, Eriocaulaceae, Commelinaceae, Pontederiaceae, Juncaceae (1985)
Vol. 5: Part 1: Fascicle 1: Aloaceae (First part): Aloe (2000)
Colchicaceae, Eriospermaceae, Asphodelaceae (Chortolirion, 1995 in Bothalia 25: 31-33; Poellnitzia , 1995
in Bothalia 25: 35, 36)
Part 2: Alliaceae, Liliaceae*. Hyacinthaceae, Agavaceae (1996 in Bothalia 26: 31-35)
Part 3: Dracaenaceae, Asparagaceae, Luzuriagaceae, Smilacaceae (1992)
A-2
Vol. 6: Haemodoraceae, Amaryllidaceae, Hypoxidaceae, Tecophilaeaceae, Velloziaceae, Dioscoreaceae
Vol. 7: Iridaceae: Part 1: Nivenioideae. Iridoideae
Part 2: Ixioideae: Fascicle 1: Ixieae (First part): Ixiinae, Tritoniinae (1999)
Fascicle 2: Syringodea , Romulea (1983)
Vol. 8: Musaceae. Strelitziaceae, Zingiberaceae (1998 in Bothalia 28: 35-39), Cannaceae*, Burmanniaceae. Orchidaceae
( Holothrix , 1996 in Bothalia 26: 125-140)
Vol. 9: Part: Urticaceae (2001)
Casuarinaceae* (2000 in Bothalia 30: 143-146), Piperaceae (2000 in Bothalia 30: 25-30), Salicaceae, Myrica-
ceae, Fagaceae*, Ulmaceae (1999 in Bothalia 29: 239-247), Moraceae, Cannabaceae* (1999 in Bothalia 29:
249-252), Proteaceae
Vol. 10: Parti: Loranthaceae, Viscaceae (1979),
Santalaceae, Grubbiaceae, Opiliaceae, Olacaceae, Balanophoraceae, Aristolochiaceae, Rafflesiaceae, Hydno-
raceae, Polygonaceae, Chenopodiaceae, Amaranthaceae, Nyctaginaceae
Vol. 11: Phytolaccaceae, Aizoaceae, Mesembryanthemaceae
Vol. 12: Portulacaceae, Basellaceae, Caryophyllaceae, Illecebraceae, Cabombaceae, Nymphaeaceae, Ceratophyllaceae
(1997 in Bothalia 27: 125-128). Ranunculaceae, Menispermaceae, Annonaceae. Trimeniaceae. Lauraceae,
Flernandiaceae, Papaveraceae, Fumariaceae
Vol. 13: Brassicaceae, Capparaceae, Resedaceae, Moringaceae, Droseraceae, Roridulaceae, Podostemaceae, Hydro-
stachyaceae (1970)
Vol. 14: Crassulaceae (1985)
Vol. 15: Vahliaceae, Montiniaceae, Escalloniaceae, Pittosporaceae, Cunoniaceae, Myrothamnaceae, Bruniaceae, Hama-
melidaceae, Rosaceae, Connaraceae
Vol. 16:
Fabaceae: Parti: Mimosoideae (1975)
Part 2: Caesalpinioideae (1977)
Part 3: Papilionoideae: Fascicle 1 :
Fascicle 2:
Fascicle 3:
Fascicle 4:
Fascicle 5:
Fascicle 6:
Fascicle 7:
Fascicle 8:
Fascicle 9:
Swartzieae-Robinieae
Indigofereae
Desmodieae, Phaseoleae
Psoraleeae-Galegeae
Loteae-Liparieae
Crotalarieae ( Aspalathus ) (1988)
Crotalarieae ( Bolusia-Lebeckia )
Crotalarieae (Lotononis-Wiborgia)
Crotalarieae ( Pearsonia-Argyrolobium ), Genisteae ( Cytisus-Ulex )
Vol. 17: Geraniaceae, Oxalidaceae
Vol. 18: Part 1: Linaceae, Erythroxylaceae, Zygophyllaceae. Balanitaceae
Part 2: Rutaceae
Part 3: Simaroubaceae, Burseraceae, Ptaeroxylaceae, Meliaceae (Aitoniaeeae), Malpighiaceae (1986)
Vol. 19: Part 1: Polygalaceae. Dichapetalaceae
Part 2: Euphorbiaceae, Callitrichaceae. Buxaceae (1996 in Bothalia 26: 37^10)
Part 3: Anacardiaceae: Fascicle 1: Rhus (1993)
Fascicle 2: remaining genera
Aquifoliaceae (1994 in Bothalia 24: 163-166)
Vol. 20: Celastraceae. Icacinaceae, Sapindaceae, Melianthaceae, Greyiaceae, Balsaminaceae, Rhatnnaceae, Vitaceae
Vol. 21: Part 1: Tiliaceae (1984)
Malvaceae. Bombacaceae, Sterculiaceae
Vol. 22: Ochnaceae, Clusiaceae, Flatinaceae, Frankeniaceae, Tamaricaceae, Canellaceae, Violaceae, Flacourtiaceae,
Tbrneraceae, Passitloraceae, Achariaceae, Loasaceae, Begoniaceae, Cactaceae (1976)
Vol. 23: Geissolomataceae, Penaeaceae, Oliniaceae, Thymelaeaceae, Lythraceae, Lecythidaceae
Vol. 24: Rhizophoraceae, Combretaceae, Myrtaceae, Melastomataceae. Onagraceae (1997 in Bothalia 27: 149-165).
Trapaceae (1998 in Bothalia 28: 11-14), Haloragaceae, Gunneraceae, Araliaceae, Apiaceae, Cornaceae
A-3
Vol. 25: Ericaceae
Vol. 26: Myrsinaceae, Primulaceae, Plumbaginaceae, Sapotaceae, Ebenaceae, Oleaceae, Salvadoraceae, Loganiaeeae,
Gentianaceae, Apocynaceae (1963)
Vol. 27: Part 1 : Periplocaceae, Asclepiadaceae ( Microloma-Xysmalobium )
Part 2: Asclepiadaceae ( Schizoglossum-Woodia )
Part 3: Asclepiadaceae (Asclep ias-A n iso toma)
Part 4: Asclepiadaceae ( Brachystelma , Ceropegia, Riocreuxia) (1980)
Asclepiadaceae (remaining genera)
Vol. 28: Parti: Convolvulaceae (2000)
Part 2: Hydrophyllaceae. Boraginaceae
Part 3: Stilbaceae, Verbenaceae ( Vitex . 1996 in Bothalia 26: 141-151)
Part 4: Lamiaceae (1985)
Part 5: Solanaceae. Retziaceae
Vol. 29: Scrophulariaceae
Vol. 30: Part 1: Bignoniaceae. Pedaliaceae, Martyniaceae, Orobanchaceae
Part 2: Gesneriaceae. Lentibulariaceae
Part 3: Acanthaceae: Fascicle 1: Justiciinae (1995)
Acanthaceae (remaining genera), Myoporaceae
Vol. 31: Part 1: Fascicle 1: Plantaginaceae (1998 in Bothalia 28: 151-157), Rubiaceae (Rubioideae — First part)
Fascicle 2: Rubiaceae (Rubioideae — Second part): Paederieae, Anthospermeae, Rubieae (1986)
Fascicle 3: Ixoroideae, Chinchonoideae
Part 2: Valerianaceae, Dipsacaceae. Cucurbitaceae
Vol. 32:
Vol. 33:
Campanulaceae, Sphenocleaceae (2000 in Bothalia 30: 31-33), Fobeliaceae. Goodeniaceae
Asteraceae: Part 1:
Part 2:
Part 3:
Part 4:
Part 5:
Part 6:
Part 7:
Part 8:
Part 9:
Lactuceae, Mutisieae, 'Tarchonantheae’
Vemonieae. Cardueae
Arctotideae
Anthemideae: Fascicle 1: Eriocephalus , Lasiospennum (2001)
Astereae
Calenduleae
Inuleae: Fascicle 1: Inulinae
Fascicle 2: Gnaphaliinae (First part) (1983)
Heliantheae, Eupatorieae
Senecioneae
FSA CONTRIBUTIONS IN BOTHALIA
FSA contributions 1: Aquifoliaceae. S. ANDREWS. 1994. Bothalia 24: 163-166.
FSA contributions 2: Asphodelaceae/Aloaceae. 1029010 Chortolirion. G.F. SMITH. 1995. Bothalia 25: 31-33.
FSA contributions 3: Asphodelaceae/Aloaceae, 1028010 Poellnitzia. G.F. SMITH. 1995. Bothalia 25: 35, 36.
FSA contributions 4: Agavaceae. G.F. SMITH & M. MOSSMER. 1996. Bothalia 26: 31-35.
FSA contributions 5: Buxaceae. H.F. GLEN. 1996. Bothalia 26: 37-40.
FSA contributions 6: Orchidaceae: Holothrix. K.L. IMMELMAN. 1996. Bothalia 26: 125-140.
FSA contributions 7: Verbenaceae: Vitex. C.L. BREDENKAMP & D.J. BOTHA. 1996. Bothalia 26: 141-151.
FSA contributions 8: Ceratophyllaceae. C.M. WILMOT-DEAR. 1997. Bothalia 27: 125-128.
FSA contributions 9: Onagraceae. P. GOLDBLATT & PH. RAVEN. 1997. Bothalia 27: 149-165.
FSA contributions 10: Trapaceae. B. VERDCOURT. 1998. Bothalia 28: 11-14.
FSA contributions 11: Zingiberaceae. R.M. SMITH. 1998. Bothalia 28: 35-39.
FSA contributions 12: Plantaginaceae. H.F. GLEN. 1998. Bothalia 28: 151-157.
FSA contributions 13: Ulmaceae. C.M. WILMOT-DEAR. 1999. Bothalia 29: 239-247.
FSA contributions 14: Cannabaceae. C.M. WILMOT-DEAR. 1999. Bothalia 29: 249-252.
FSA contributions 15: Piperaceae. K.L. IMMELMAN. 2000. Bothalia 30: 25-30.
FSA contributions 16: Sphenocleaceae. W.G. WELMAN. 2000. Bothalia 30: 31-33.
FSA contributions 17: Casuarinaceae. C.M. WILMOT-DEAR. 2000. Bothalia 30: 143-146.
A-4
FLORA OF SOUTHERN AFRICA
ALPHABETICAL LIST OF PUBLISHED TAXA
* exotic families
Acanthaceae: Justiciinae, Vol. 30. Part 3. Fasc.l (1995)
Achariaceae, Vol. 22 (1976)
Agavaceae (Bothalia 26, 1996)
Alismataceae, Vol. 1 (1966)
Aloaceae (first part): Aloe , Vol. 5, Part 1, Fasc. 1 (2000)
Aloe, Aloaceae (first part), Vol. 5, Part 1, Fasc. 1 (2000)
Anacardiaceae: Rhus, Vol. 19, Part 3, Fasc. 1 (1993)
Andreaeaceae, Bryophyta, Part 1, Fasc. 1 (1981)
Anthemideae, Asteraceae, Vol. 33, Part 4, Fasc. 1 (2001)
Anthospermeae. Rubiaceae: Rubioideae (second part), Vol.
31, Part 1. Fasc. 2 (1986)
Apocynaceae, Vol. 26 (1963)
Aponogetonaceae, Vol. 1 (1966)
Aquit'oliaceae ( Bothalia 24, 1994)
Archidiaceae, Bryophyta, Part 1, Fasc. 1 (1981)
Asclepiadaceae: Brachvstelma-Riocreuxia, Vol. 27, Part 4
(1980)
Aspalathus, Fabaceae: Papilionoideae, Vol. 16, Part 3, Fasc.
6 (1988)
Asparagaceae, Vol. 5 (1992)
Asphodelaceae: Chortolirion, Poellnitzia ( Bothalia 25. 1995)
Asteraceae: Anthemideae: Eriocephalus , Lasiospermum,
Vol. 33, Part 4, Fasc. 1 (2001)
Asteraceae: Inuleae: Gnaphaliinae (first part), Vol. 33, Part
7, Fasc. 2 (1983)
Aulacomniaceae, Bryophyta, Part 1, Fasc. 2 (1987)
Aytoniaceae. Hepatophyta, Part 1, Fasc. 1 (1999)
Bartramiaceae, Bryophyta. Part 1, Fasc. 2 (1987)
Begoniaceae, Vol. 22 (1976)
Brachystelma, Asclepiadaceae, Vol. 27, Part 4 (1980)
Brassicaceae, Vol. 13 (1970)
Bryaceae, Bryophyta, Part 1 , Fasc. 2 ( 1 987)
Bryobartramiaceae, Bryophyta, Part 1, Fasc. 1 (1981)
Bryophyta (three fascicles published 1981. 1987, 1998: see
plan of FSA)
Burseraceae. Vol. 18 (1986)
Buxaceae ( Bothalia 26, 1996)
Cactaceae, Vol. 22 (1976)
Caesalpinioideae, Fabaceae, Vol. 16, Part 2 (1977)
Calymperaceae, Bryophyta, Part 1 , Fasc. 1 (1981)
Canellaceae, Vol. 22 (1976)
Cannabaceae {Bothalia 29, 1999)
Capparaceae, Vol. 13 (1970)
Casuarinaceae (Bothalia 30, 2000)
Ceratophyllaceae (Bothalia 27. 1997)
Ceropegia, Asclepiadaceae, Vol. 27, Part 4 ( 1980)
Charophyta, Cryptogams ‘Vol. 9’ (1978)
Chortolirion, Asphodelaceae (Bothalia 25, 1995)
Cleveaceae. Hepatophyta. Part 1, Fasc. 1 (1999)
Clusiaceae, Vol. 22 (1976)
Commelinaceae, Vol. 4 (1985)
Convolvulaceae, Vol. 28, Part 1 (2000)
Crassulaceae, Vol. 14 (1985)
Crotalarieae, Aspalathus, Fabaceae: Papilionoideae, Vol.
16, Part 3, Fasc. 6 (1988)
Cryphaeaceae, Bryophyta. Part 1, Fasc. 3 (1998)
Cupressaceae, Vol. 1 (1966)
Dicranaceae, Bryophyta, Part 1, Fasc. 1 (1981)
Ditrichaceae, Bryophyta. Part 1, Fasc. 1 (1981)
Dracaenaceae, Vol. 5 (1992)
Droseraceae, Vol. 13 (1970)
Ebenaceae, Vol. 26 (1963)
Elatinaceae, Vol. 22 (1976)
Encalyptaceae. Bryophyta, Part 1, Fasc. 1 (1981)
Ephemeraceae, Bryophyta, Part 1, Fasc. 2 (1987)
Eriocaulaceae, Vol. 4 (1985)
Eriocephalus, Asteraceae: Anthemideae, Vol. 33. Part 4. Fasc.
1 (2001)
Erpodiaceae, Bryophyta, Part 1. Fasc. 3 (1998)
Eustichiaceae, Bryophyta, Part 1, Fasc. 2 (1987)
Exormothecaceae, Hepatophyta, Part 1, Fasc. 1 (1999)
Fabaceae: Caesalpinioideae, Vol. 16, Part 2 ( 1977)
Fabaceae: Mimosoideae, Vol. 16, Part 1 (1975)
Fabaceae: Papilionoideae, Crotalarieae, Aspalathus, Vol. 16,
Part 3, Fasc. 6 (1988)
Fissidentaceae, Bryophyta, Part 1, Fasc. 1 (1981)
Flacourtiaceae, Vol. 22 (1976)
Fontinalaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Frankeniaceae, Vol. 22 (1976)
Funariaceae, Bryophyta, Part 1, Fasc. 2 (1987)
Gentianaceae, Vol. 26 (1963)
Gigaspermaceae, Bryophyta, Part 1, Fasc. 2 (1987)
Gnaphaliinae (first part), Asteraceae: Inuleae, Vol. 33. Part
7, Fasc. 2 (1983)
Grimmiaceae, Bryophyta, Part 1, Fasc. 1 (1981)
Hedwigiaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Hepatophyta, Part I, Fasc. I (1999)
Holothrix, Orchidaceae (Bothalia 26, 1996)
Hookeriaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Hydrocharitaceae, Vol. 1 (1966)
Hydrostachyaceae, Vol. 13 (1970)
Inuleae. Asteraceae: Gnaphaliinae (first part), Vol. 33, Part
7, Fasc. 2 (1983)
Iridaceae: lxieae (first part): Ixiinae. Tritoniinae, Vol. 7, Part
2, Fasc. 1 (1999)
Iridaceae: Syringodea, Romulea , Vol. 7, Part 2, Fasc. 2 (1983)
A-5
Ixieae (first part), Iridaceae: Ixiinae, Trinoniinae, Vol. 7,
Part 2, Fasc. 1 (1999)
Ixiinae, Iridaceae: Ixieae (first part), Vol. 7, Part 2, Fasc. 1
(1999)
Juncaceae, Vol. 4 (1985)
Juncaginaceae, Vol. 1 (1966)
Justiciinae, Acanthaceae, Vol. 30, Part 3, Fasc. 1 (1995)
Lamiaceae, Vol. 28 (1985)
Lasiospermum , Asteraceae: Anthemideae, Vol. 33. Part 4,
Fasc. 1 (2001)
Leptodontaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Leucodontaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Loasaceae, Vol. 22 (1976)
Loganiaceae, Vol. 26 (1963)
Loranthaceae, Vol. 10(1979)
Lunulariaceae, Hepatophyta, Part 1, Fasc. 1 (1999)
Luzuriagaceae, Vol. 5 (1992)
Malpighiaceae, Vol. 18 (1986)
Marchantiaceae, Hepatophyta, Part 1, Fasc. 1 (1999)
Marchantiales, Hepatophyta, Part 1, Fasc. 1 (1999)
Marchantiidae, Hepatophyta, Part 1, Fasc. 1 (1999)
Marchantiopsida, Hepatophyta, Part 1 (1999)
Meliaceae, Vol. 18 (1986)
Meteoriaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Mimosoideae, Fabaceae, Vol. 16, Part 1 (1975)
Mniaceae, Bryophyta, Part 1, Fasc. 2 (1987)
Moringaceae, Vol. 13 (1970)
Myrsinaceae, Vol. 26 (1963)
Nanobryaceae, Bryophyta, Part 1, Fasc. 1 (1981)
Najadaceae, Vol. 1 (1966)
Neckeraceae, Bryophyta, Part 1, Fasc. 3 (1998)
Ochnaceae, Vol. 22 (1976)
Oleaceae, Vol. 26 (1963)
Onagraceae (. Bothalia 27, 1997)
Orchidaceae: Holothrix ( Bothalia 26, 1996)
Orthotrichaceae. Bryophyta. Part 1, Fasc. 3 (1998)
Oxymitraceae, Hepatophyta, Part 1, Fasc. 1 (1999)
Paederieae, Rubiaceae: Rubioideae (second part), Vol. 31,
Part 1, Fasc. 2 (1986)
Passifloraceae, Vol. 22 (1976)
Pinaceae*, Vol. 1 (1966)
Piperaceae ( Bothalia 30, 2000)
Plantaginaceae ( Bothalia 28, 1998)
Plumbaginaceae, Vol. 26 ( 1963)
Podocarpaceae, Vol. 1 (1966)
Podostemaceae, Vol. 13 (1970)
Poellnitzia, Asphodelaceae (Bothalia 25, 1995)
Pontederiaceae, Vol. 4 (1985)
Potamogetonaceae, Vol. 1 (1966)
Pottiaceae, Bryophyta, Part 1, Fasc. 1 (1981)
Primulaceae, Vol. 26 (1963)
Prionodontaceae, Bryophyta. Part 1, Fasc. 3 (1998)
Ptaeroxylaceae, Vol. 18 (1986)
Pteridophyta (1986) (for list of families, see p. v of Pteri-
dophyta volume)
Pterobryaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Ptychomitriaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Racopilaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Resedaceae, Vol. 13 (1970)
Rhabdoweisiaceae. Bryophyta. Part 1, Fasc. 3 (1998)
Rhachitheciaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Rhizogoniaceae, Bryophyta, Part 1, Fasc. 2 (1987)
Rhus, Anacardiaceae, Vol. 19, Part 3, Fasc. 1 (1993)
Ricciaceae, Hepatophyta, Part 1, Fasc. 1 (1999)
Riocreuxia, Asclepiadaceae, Vol. 27, Part 4 (1980)
Romulea, Iridaceae, Vol. 7, Part 2, Fasc. 2 (1983)
Roridulaceae, Vol. 13 (1970)
Rubiaceae: Rubioideae (second part): Paederieae, Antho-
spermeae, Rubieae, Vol. 31, Part 1, Fasc. 2 (1986)
Rubieae, Rubiaceae: Rubioideae (second part), Vol. 31, Part
1, Fasc. 2 (1986)
Rubioideae (second part), Rubiaceae, Vol. 31, Part 1, Fasc.
2 (1986)
Ruppiaceae, Vol. 1 (1966)
Salvadoraceae, Vol. 26 (1963)
Sapotaceae, Vol. 26 (1963)
Seligeriaceae, Bryophyta, Part 1, Fasc. 1 (1981)
Simaroubaceae, Vol. 18 (1986)
Smilacaceae, Vol. 5 (1992)
Sphagnaceae, Bryophyta, Part 1, Fasc. 1 (1981)
Sphenocleaceae (Bothalia 30, 2000)
Splachnaceae. Bryophyta, Part 1, Fasc. 2 (1987)
Stangeriaceae, Vol. 1 (1966)
Syringodea, Iridaceae, Vol. 7, Part 2, Fasc. 2 (1983)
Tamaricaceae, Vol. 22 (1976)
Targioniaceae, Hepatophyta. Part 1, Fasc. 1 (1999)
Thanmobryaceae, Bryophyta, Part 1 , Fasc. 3 ( 1 998)
Tiliaceae, Vol. 21 (1984)
Trachypodaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Trapaceae (Bothalia 28, 1998)
Tritoniinae, Iridaceae: Ixieae (first part), Vol. 7, Part 2. Fasc.
1 (1999)
Turneraceae, Vol. 22 (1976)
Typhaceae, Vol. 1 (1966)
Ulmaceae (Bothalia 29, 1999)
Urticaceae, Vol. 9, Part: Urticaceae (2001)
Verbenaceae: Vitex (Bothalia 26, 1996)
Violaceae, Vol. 22 (1976)
Viscaceae, Vol. 10 (1979)
Vitex, Verbenaceae (Bothalia 26, 1996)
Wardiaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Welwitschiaceae, Vol. 1 (1966)
Xyridaceae, Vol. 4 (1985)
Zamiaceae, Vol. 1 (1966)
Zannichelliaceae, Vol. 1 (1966)
Zosteraceae, Vol. 1 (1966)
"
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