Bothalia

A JOURNAL OF BOTANICAL RESEARCH

Vol. 29,1 May 1999

\ \ I I ' \ V I

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TECHNICAL PUBLICATIONS OF THE NATIONAL BOTANICAL INSTITUTE,

PRETORIA

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

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

BOTHALIA

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

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

ed to the furtherance of botanical science. The main fields covered are taxonomy, ecology, anato-

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

lished annually.

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

STRELITZIA

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

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

MEMOIRS OF THE BOTANICAL SURVEY OF SOUTH AFRICA

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

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

ANNALS OF KIRSTENBOSCH BOTANIC GARDENS

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

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

South African Botany). Discontinued after Vol. 19.

FLOWERING PLANTS OF AFRICA (FPA)

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

mainly by the artists at the National Botanical Institute. Many well known botanical artists have

contributed to the series, such as Cythna Letty (over 700 plates), Kathleen Lansdell, Stella Gower,

Betty Connell, Peter Bally, Fay Anderson, Ellaphie Ward-Hilhorst and Gillian Condy. The Editor

is pleased to receive living plants of general interest or of economic value for illustration.

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

An index to Vols 1-49 is available.

FLORA OF SOUTHERN AFRICA (FSA)

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

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

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

and ecological notes.

Contributions to the FSA also appear in Bothalia.

PALAEOFLORA OF SOUTHERN AFRICA

A palaeollora on a pattern comparable to that of the Flora of southern Africa. Much of the infor-

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

available:

Molteno Eormation (Triassic) Vol. I. Introduction. Dicroidium, by J.M. & H.M. Anderson.

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

Anderson.

Prodromus of South African Megalloras. Devonian to Lower Cretaceous, by J.M. & H.M.

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

BOTHALIA

A JOURNAL OF BOTANICAL RESEARCH

Volume 29,1

Scientific Editor; G. Germishuizen

Technical Editor: B.A. Momberg

NATIONAL

O T A N I C A L

INSTITUTE

2 Cussonia Avenue, Brummeria, Pretoria

Private Bag XlOl, Pretoria 0001

ISSN 0006 8241

May 1999

Editorial Board

D.R Cutler

B.J. Huntley

P.H. Raven

J.P. Rourke

M.J. Werger

Royal Botanic Gardens, Kew, UK

National Botanical Institute, Cape Town, RSA

Missouri Botanical Garden, St Louis, USA

Compton Herbarium, NBI, Cape Town, RSA

University of Utrecht, Utrecht, Netherlands

CONTENTS

Volume 29,1

New species, varieties, status and combinations in Bothalia 29,1 (1999) iv

1 . Studies in the liverwort genus Fossombronia (Metzgeriales) from southern Africa. 7. F. capensis var.

spiralis, a new variety from Western Cape. S.M. PEROLD 1

2. Revision of the genus Calpurnia (Sophoreae: Leguminosae). A.J. BEAUMONT, R.P. BECKETT,

TJ. EDWARDS and C.H. STIRTON 5

3. Studies in the liverwort genus Fossombronia (Metzgeriales) from southern Africa. 8. F elsieae and

F. spinosa, two new Western Cape species with spinose spores. S.M. PEROLD 25

4. Revision of the genus Faucaria (Ruschioideae: Aizoaceae) in South Africa. L.E. GROEN and L.J.G.

VANDERMAESEN 35

5. New species of Sparaxis and Ixia (Iridaceae: Ixioideae) from Western Cape, South Africa, and taxo-

nomic notes on Ixia and Gladiolus. P. GOLDBLATT and J.C. MANNING 59

6. The genus Amphiglossa (Gnaphalieae, Relhaniinae, Asteraceae) in southern Africa. M. KOEKEMOER . . 65

7. Studies in the liverwort genus Fossombronia (Metzgeriales) from southern Africa. 9. A new species

from Mpumalanga and KwaZulu-Natal, with notes on other species. S.M. PEROLD 77

8. Studies in the liverwort genus Fossombronia (Metzgeriales) from southern Africa. 10. Three new

species from Northern and Western Cape. S.M. PEROLD 83

9. Notes on African plants;

Agavaceae. Agave vivipara: the correct name for Agave angustifolia. G.F. SMITH and E.M.A.

STEYN 100

Asteraceae. A note on the genus Philyrophyllum. P.P.J. HERMAN 107

Ericaceae. Two new species of Erica from Western Cape, South Africa. E.G.H. OLIVER and

I.M. OLIVER 95

Ericaceae. Three new species of Erica from Western Cape, South Africa. E.G.H. OLIVER and

I.M. OLIVER 112

Lamiaceae. Salvia thermara, a new species from the Western Cape, South Africa. E.J. VAN

JAARSVELD 100

Ophioglossaceae: Pteridophyta. Two new taxa of Ophioglossiim from tropical Africa. J.E. BUR-

ROWS 109

Polypodiaceae. Polypodium ensiforme, the correct name for Microsorum ensiforme (Polypodioi-

deae). J.P. ROUX 103

Rubiaceae. A new species of Pavetta from the Soutpansberg, South Africa. N. HAHN 107

Solanaceae. Three new records of Solatium section Oliganthes in southern Africa. W.G. WELMAN 98

10. Montane flora of the southern Langeberg, South Africa; a checklist of the flowering plants and ferns.

D.J.MCDONALD 119

11. Flora of the Kap River Reserve, Eastern Cape, South Africa. E.C. CLOETE and R.A. LUBKE 139

12. Vegetation of the Sileza Nature Reserve and neighbouring areas. South Africa, and its importance in

conserving the woody grasslands of the Maputaland Centre of Endemism. W.S. MATTHEWS,

A.E. VAN WYK and N. VAN ROOYEN 151

13. Itinerary and specimen list of M.A. Pocock’s botanical collecting expedition in Zambia and Angola in

1925. M.G. BALARIN, E. BRINK and H.F GLEN 169

14. Miscellaneous notes;

Poaceae. Apomixis in the genus Pentaschistis (Arundinoideae). J.J. SPIES, K.C. KLOPPER and

B. VISSER 203

Progress with the trial phase for registration of new plant names. G.F. SMITH and G. GERMIS-

HUIZEN 207

15. Obituary; EliseBodley (1921-1997). J.P. ROURKE 211

16. Book reviews 215

New species, varieties, status and combinations in Bothalia 29,1 (1999)

Amphiglossa celans Koekemoer, sp. nov., 68

Amphiglossa grisea Koekemoer, sp. nov., 70

Amphiglossa rudolphii Koekemoer, sp. nov., 71

Amphiglossa susannae Koekemoer, sp. nov., 71

Amphiglossa tecta (FBrusse) Koekemoer, comb, nov., 72

Amphiglossa thuja {Merxm.) Koekemoer, comb, nov., 73

Calpurnia reflexus A.J. Beaumont, sp. nov., 13

Erica cubitans E.G.H.Oliv., sp. nov., 112

Erica feminarum E.G.H.Oliv., sp. nov., 97

Erica garciae E.G.//.0//V., sp. nov., 115

Erica hansfordii E.G.H.Oliv., sp. nov., 95

Erica sperata E.G.H.Oliv., sp. nov., 1 15

Faucaria felina (L.) Schwantes subsp. britteniae (L.Bolus) LE.Groen, stat. nov., 41

Faucaria felina (L.) Schwantes subsp. tuberculosa (Rolfe) LE.Groen, stat. nov., 42

Fossombronia capensis S.W.Arnell var. spiralis Perold, var. nov., 1

Fossombronia elsieae Perold, sp. nov., 25

Fossombronia hyalorhiza Perold, sp. nov., 83

Fossombronia marindae Perold, sp. nov., 86

Fossombronia monticola Perold, sp. nov., 87

Fossombronia renateae Perold, sp. nov., 77

Fossombronia spinosa Perold, sp. nov., 29

Gladiolus saxatilis Goldblatt & J.C. Manning, nom. nov., 63

Ixia aurea J.C. Manning & Goldblatt, sp. nov., 61

Ophioglossum richardsiae J.E. Burrows, sp. nov., 109

Ophioglossum vulgatum L. subsp. africanum Pocock ex J.E.Burrows var. taylorianum J.E.Burrows, var. nov., 110

Pavetta tshikondeni N.Hahn, sp. nov., 107

Salvia thermara Van Jaarsv., sp. nov., 100

Section Dichone (Salisb. ex Baker) Goldblatt & J.C. Manning, comb, et stat. nov., 63

Sparaxis auriculata Goldblatt & J.C. Manning, sp. nov., 59

Sparaxis metelerkampiae (L.Bolus) Goldblatt & J.C. Manning, stat. nov., 61

Subgenus Morphixia (Ker Gawl.) Goldblatt & J.C. Manning, comb, et stat. nov., 63

IV

Bothalia29,l: 1-4 (1999)

Studies in the liverwort genus Fossombronia (Metzgeriales) from

southern Africa. 7. F. capensis van spiralis^ a new variety from Western

Cape

S.M. PEROLD*

Keywords: Fossombronia Raddi, F. capensis var. capensis, F. capensis var. spiralis, Hepaticae, Metzgeriales, southern Africa, Western Cape

ABSTRACT

Fossombronia capensis S.W.Amell var. spiralis Perold, var. nov. from Western Cape is described. It is distinguished

from the typical variety by a suite of characters, the most obvious being the strong spirals in the elaters. In F. capensis var.

capensis the spirals are weak except for two Amell collections in which they are quite strong. Subtle differences in spore

ornamentation are then, perhaps, the most definitive character to separate the two varieties.

INTRODUCTION

Perold ( 1997) in an amendment to F. capensis S.W.Ar-

nell, made the following statement: ‘All F. capensis

specimens from the George/Knysna/Brackenfell/Gouna

and Deep Walls Forest and Diep River areas have poor-

ly formed elaters. Collections from the nearby Bloukrans

Pass {Liibenau-Nestle SA 139/2 and S.M. Perold 3534,

3539-3541) have elaters with well-formed spirals, but

have been referred here because the spores and plants are

closely similar’.

I have since come to the conclusion that the collections

from Bloukrans Pass and others like it from elsewhere,

should rather be separated as a variety of F. capensis.

Fossombronia capensis S.W.Amell var. spiralis

Perold, var. nov.

Plantae masculae minores plantis feminis, ut in vari-

etate typica, sed bracteae perigoniales antheridia tegentes

longius dispersae. Sporae 37.5-55.0 pm diametro, super-

ficie distali lamellis 6 ad 8 irregulariter ramosis, inter-

dum debiliter effectis; superficie inter lamellas valde

aspera, granulosa vel transverse grosse striata. Elateres

semper bene evoluti, 75.0-157.5 x 7.5-10.0 pm, omnino

bis vel ter spirales, saepe partim bis et partim ter spirales.

TYPE. — Western Cape, 3118 (Vanrhynsdorp): Matsi-

kama Mountains, Bo-Drift Farm, at Olyfenhoutsrivier,

(-DD), under rocks on island in stream near bridge, S.M.

Perold 3905 (PRE, holo.).

Plants in crowded stands, green; shoots medium-sized

in male plants, up to 10 mm long, 1.4— 2.2 mm high,

2. 8-3. 5 mm wide; female plants rather larger, 10-15 mm

long, 2.0-2. 5 mm high, 3. 3^.5 mm wide, simple or once/

twice furcate, segments moderately to widely divergent,

apically 2-7 mm long. Stems prostrate, slightly or not taper-

ing proximally, dorsally chlorophyllose, ventrally usually

* National Botanical Institute, Private Bag XlOl, Pretoria 0001,

MS, received: 1998-05-13.

purple, plano-convex in cross section, in male plants api-

cally (Figure 10) up to 450 pm (10 cell rows) high, 500

pm wide, basally (Figure IP) 310-350 x 500-550 pm; in

female plants (Figure IM) apically ± 500 pm (10-12 cell

rows) high, 650 pm wide, basally (Figure IN) ± 450 x 600

pm. Rhizoids purple, 12.5-20.0 pm wide. Leaves overlap-

ping, widely spreading, succubously inserted (Figure 2A,

C), smaller apically but soon enlarging, oblong to obovate,

sometimes slightly notched above, lobes equal in size or

not in male plants (Figure lA-E), 1300-2125 x

1100-1925 pm, when sides not parallel narrower below,

1000-1300 pm wide; in female plants (Figure IF-J) most-

ly larger, 1550-2750 x 1550-2575 pm, below 1075-1750

pm; margins (Figure IK) generally almost entire, papillae

if present, few, only 1 or 2. Leaf cells thin-walled, in male

and female plants quite similar, at upper margins sub-

quadrate to rectangular across, 20.0-42.5 x 27.5-40.0

(-57.5) pm, at lateral margins long-rectangular, 65-75 x

20.0- 37.5 pm; upper laminal cells 5- or 6-sided, 35-50 x

40.0- 47.5 pm; middle laminal cells 50-85 x 37.5-62.5

pm; basal cells 62.5-82.5 x 42.5-50.0 pm. Oil bodies

(Figure IL) 7-12 per cell, spherical to ovoid, up to 5 pm

diam., glistening, homogeneous; chloroplasts numerous, ±

5 pm diam.

Dioicous. Antheridia dorsal on stem, in 1 or 2 rows

(Figure 2A, B), short-stalked, globose or ovoid, 220-250

pm diam., each shielded by a perigonial bract (Figure

IQ-T), irregular in shape, 370-450 x 200-300 pm,

sometimes 2 adjacent ones joined together, margins

mostly without papillae, cells in interior 4- or 5-sided,

50-75 X 27.5-37.5 pm. Archegonia in 1 or 2 rows

(Figure 2C, D) along stem, naked; sometimes 2 in

acropetal sequence or 2 adjacent (Figure 2E) on a branch

becoming fertilised. Pseudoperianth (Figures lU, V; 2F)

near apex to rather more proximal, campanulate, project-

ing up to 800 pm above leaves, raised on a short stalk,

then widely flaring above, 1625-2250 pm long, 2(X)0-3125

pm wide across mouth, margin undulating or sometimes

sparsely and shallowly lobed, very rarely with some

angular projections, occasional vertically split along

side, lateral outgrowths quite common; cells at margin

and immediately below comparable in shape and size to

those of leaves, but larger at ± midlength, 100-165 x

32.5-50.0 pm, and at base, 175-195 x 75-80 pm. Cap-

2

Bothalia29,l (1999)

FIGURE I . — Fossomhronia capensis var. spiralis. A-E, male leaves; F-J, female leaves; K, detail of upper leaf margin; L, median leaf cells with

oil bodies (solid lines) and chloroplasts (dotted lines); M, c/s apical part of female stem; N, c/s basal part of female stem; O, c/s apical part

of male stem; P, c/s basal part of male stem; Q-T, perigonial bracts; U, pseudoperianth from side; V, opened pseudoperianth; W, c/s seta;

X, cells in capsule wall. A-K, M-X, S.M. Perold 3905\ L, S.M. Perold 3899. Scale bars: A-J, U, V, 500 pm; K, W, 100 pm; L, X, 50 pm;

M-P, Q-T, 250 pm.

Bothalia 29,1 (1999)

3

FIGURE 2. — Fossombronia capensis var. spiralis. A, male branch with perigonial bracts; B, close-up of bracts; C, female branch with archego-

nia; D, close-up of archegonia; E, 2 pseudoperianths alongside each other, F, single pseudoperianth. A, B, S.M. Perold 39H0\ C, S.M. Perold

3394\ D, S.M. Perold 3S39: E, E, S.M. Perold 3905. A, x 8.3; B, x 25; C, x 9; D, x 36; E, x 13.8; F, x 21.

sales globose, ± 850 |jm diam., wall bistratose, cells in

inner layer irregularly shaped, some trapezoidal, others

triangular or rectangular (Figure IX), 35.0-42.5 x

17.5-37.5 pm, each cell wall with (1)2 or 3 dark brown,

nodular and occasionally semi-annular thickenings. Seta

2.9^. 0 mm long, 290-300 x 240 pm in transverse sec-

tion, 5 or 6 cells across (Figure IW). Spores golden

brown to darker brown, hemispherical, 37.5-55.0 pm

diam., including lamellae projecting at margin; distal

face (Figure 3A-D) convex, with 6-8 irregularly branch-

ing and sometimes poorly developed lamellae, ± 5 pm

high and 5-10 pm apart running across face, a few paral-

lel to each other, others in different directions, occasion-

ally anastomosing and forming some complete or incom-

plete areolae, rarely with a central tubercle, surface

between lamellae highly uneven, granular or mostly

coarsely striated; proximal face (Figure 3E) lacking tri-

radiate mark, flat, covered with irregularly shaped papil-

FIGURE 3. — Fossombronia capensis var. .spiralis. Spores and elaters. A-C, distal face; D, distal surface and lamellae near margin; E, proximal

face; F, elaters. A, S.M. Perold 3541\ B, E, S.M. Perold 3905', C, D, F, S.M. Perold 3814. A, x 835; B, x 867; C, x 789; D, x 1560; E, x

863; F, X 748.

4

Bothalia29,l (1999)

lae and fine granules in between as well as marginally, up

to 20 ‘spines’ projecting around spore periphery, few of

them joined by sections of incomplete wing. Elaters

(Figure 3F) yellow, 75.0-157.5 x 7.5-10.0 pm, tapering

to the tips or not, smooth, entirely 2-spiral or 3-spiral,

often partly 2-spiral and partly 3-spiral, rarely branched.

Fossombronia capensis var. spiralis is restricted to the

winter rainfall area of South Africa, namely Western Cape

(Figure 4). This is referred to as the Fynbos Biome — scle-

rophyllous, microphyllous vascular plant vegetation —

(Cowling et al. 1997; Rutherford 1997) or else the area is

called the Cape Region — phytogeographic region based

on the distribution of vascular plants — (Cowling &

Hilton-Taylor 1997). The plants under discussion grow

on soil at streambanks, on damp vertical earth walls at

roadsides or pathways, near waterfalls, at seepages or at

grotto entrances and are apparently more widespread than

the typical variety, which has now, for the first time, also

been collected at Kirstenbosch (Perold 3955), besides the

localities in the southern Cape previously reported

(Perold 1997). The new variety is distinguished by: the

fairly large, mostly entire leaves; the slightly smaller male

plants bearing 1 or 2 rows of perigonial bracts which are

less crowded than those in var. capensis', the generally

undulating margin of the pseudoperianths; the cells in the

capsule wall with more numerous semi-annular bands;

the spore surface on the distal face with very uneven and

rough, often rather poorly developed lamellae; and the

elaters with well-formed, strong spirals. The latter char-

acter (with rare exceptions) is the most obvious in sepa-

rating the two varieties and the choice of epithet, ‘'spi-

ralis', for this new variety draws attention to this feature.

Flowever, the SEM micrograph of the elaters from F.

capensis var. capensis, Arnell 1477 (G) in Perold (1997:

fig. 3F), shows rather stronger spirals than are usually

encountered in the typical variety, but the spirals appear

mostly quite pale and poorly stained, when examined

under a compound microscope. Two Amell specimens,

1715 and 1716, collected in the Gouna Forest and listed

in the above publication, but not singled out, have quite

strong, well-stained spirals, although the spore ornamen-

tation, probably still the most definitive character, is

clearly that of F. capensis var. capensis. A suite of char-

acters must necessarily be employed for a reliable separa-

tion of the two varieties of F. capensis.

Key to the varieties of F. capensis

la Male plants bearing large, very crowded perigonial bracts

shielding antheridia; leaves with up to 6 scattered papil-

lae at margins and 17-37 oil bodies per median cell;

inner capsule wall with few cells (only ± 2 per field)

having semi-annular thickening bands; distal spore face

with well-formed lamellae, surface between them with

fine cross striations or relatively smooth and appearing

to be cleanly ‘scooped out’; perispore at spore periphery

mostly intact; elaters almost always poorly formed,

(40.0-)62. 5-125.0 pm long, spiral bands pale when

viewed under compound microscope, outer wall collaps-

ing on drying F. capensis var. capensis

lb Male plants bearing fairly large, somewhat spaced perigo-

nial bracts shielding antheridia; leaves generally with

few (1 or 2) or no papillae at margins and with 7-12 oil

bodies per median cell; inner capsule wall with ± half

the cells per field having semi-annular thickening bands;

distal spore face frequently with poorly developed lamel-

lae, surface between them highly uneven and often

coarsely striated; perispore at spore periphery absent

or very incomplete; elaters always well-formed,

(50.0-)75.0-157.5 pm long with strongly de-

veloped spiral bands F. capensis var. spiralis

SPECIMENS EXAMINED

F. capensis var. spiralis

Held at PRE, unless otherwise indicated.

Louwrens CH2881, CH2882. LUbenau-Nestle SA139/2 (private, herb ).

S.M. Perold 2746 pp., 3394, 3407, 3534, 3539-3541, 3814, 3816(b),

3817, 3830, 3858, 3877 (sterile), 3899, 3905 (holotype), 3979, 3980.

ACKNOWLEDGEMENTS

I sincerely thank Dr J.E. Braggins of the University of

Auckland, New Zealand, for kindly refereeing this arti-

cle and his helpful suggestions; also my colleagues at

NBI, i.e. the curator of PRE, Ms M. Koekemoer and Mr

J. Meyer, for their generous help with fieldwork. Dr H.F.

Glen is thanked for translating the diagnosis into Latin as

well as for suggesting the varietal name ‘spiralis' for this

new variety of F. capensis. The curators of BOL, G and

S and Dr Liibenau-Nestle are thanked for the loan of

specimens. I also extend my gratitude to Ms G. Condy

for the drawings, to Mrs A. Romanowski for developing

and printing the photographs and to Ms D. Maree for

typing the manuscript.

REFERENCES

COWLING, R.M. & HILTON-TAYLOR, C. 1997. Phytogeography,

flora and endemism. In R.M. Cowling, D.M. Richardson &

S.M. Pierce, Vegetation of southern Africa. Cambridge

University Press, Cambridge.

COWLING, R.M., RICHARDSON, D M. & MUSTART, RJ. 1997. Fyn-

bos. In R.M. Cowling, D.M. Richardson & S.M. Pierce, Vegetation

of southern Africa. Cambridge University Press, Cambridge.

PEROLD, S.M. 1997. Studies in the liverwort genus Fossombronia

(Metzgeriales) from southern Africa. 2. An amendment to three

species from Western Cape, described by S.W. Amell. Bothalia

27: 29-38.

RUTHERFORD, M.C. 1997. Categorization of biomes. In R.M.

Cowling, D.M. Richardson & S.M. Pierce, Vegetation of south-

ern Africa. Cambridge University Press, Cambridge.

Bothalia 29,1: 5-23 (1999)

Revision of the genus Calpurnia (Sophoreae: Leguminosae)

A.J. BEAUMONT*, R.P. BECKETT,* TJ. EDWARDS* and C.H. STIRTON**

Keywords; Calpurnia E.Mey., Leguminosae, new species, Sophoreae, southern Africa, taxonomy

ABSTRACT

Taxa recognised in this revision are: Calpurnia aurea (Aiton) Benth. subsp. aurea\ C. aurea (Aiton) Benth. subsp. indi-

ca Brummitt; C. floribunda Harv.; C. glabrata Brummitt; C. intrusa (R.Br. in W.T. Aiton) E.Mey.; C. reflexus A.J. Beaumont

sp. nov.; C. sericea Harv. and C. woodii Schinz. A putative hybrid between C. sericea and C. woodii is recorded. With the

exception of C. aurea, all species are restricted to southern Africa. C. reflexus is possibly extinct, and C. woodii is consid-

ered rare. Characters examined in this revision are habitat, habit, vestiture, leaf morphology and anatomy; floral, pollen, fruit

and seed morphology and anatomy; and seedling morphology. Illustrations and a key to taxa are provided.

INTRODUCTION

Calpurnia E.Mey. is a small genus of papilionoid

legumes, and belongs to the Sophora group of the primi-

tive tribe Sophoreae sensu Polhill (1981). According to

Polhill (1981), Calpurnia is closely related to Maackia

Rupr. & Maxim, Cladrastis Raf. and Salweenia Baker f.,

of the Northern Elemisphere. Calpurnia species are slen-

der trees or small shrubs with pinnate leaves and attractive

yellow papilionoid flowers in racemes or panicles. All but

one species are narrow endemics of southern Africa. C.

aurea subsp. aurea extends north into Ethiopia and,

together with C. aurea subsp. indica, into southern India.

Early classifications placed Calpurnia species under

Sophora L., of the tribe Sophoreae (Aiton 1789), Robinia

L., of the tribe Robinieae (L’Heritier 1791), and

Podalyria Willd. (Willdenow 1799) and Virgilia Poir.

(Lamarck 1793), of the tribe Podalyrieae. Meyer (1836)

transferred four species from Virgilia and published a

new species to establish Calpurnia. The genus commem-

orates the Roman poet Calpernicus Siculus who, in the

middle of the First Century A.D., wrote poetry similar to

that of Virgilius. Superficially, species of Calpurnia

resemble those of Virgilia (commemorating Virgilius).

Phillips (1917) recognised seven species of Cal-

purnia. Subsequent workers have contributed taxonomic

notes on species and some have recognised subspecific

taxa (Gillett 1965; Brummitt 1967, 1970). Yakovlev

(1971) provides the most comprehensive recent account

of the genus, notwithstanding the correction of some

erroneous nomenclature by Ross (1976).

MATERIALS AND METHODS

Pollen

Mature buds preserved in the mixture; formalin:

acetic acid: ethanol (FAA), and material from herbarium

* Department of Botany, University of Natal, Pietermaritzburg, Private

Bag XOl, 3209 Scottsville. South Africa.

** The National Botanic Garden of Wales, Middleton Hall, Llanarthne,

Carmarthenshire, SA32 8HN, U.K.

MS. received: 1992-10-14.

sheets were used. Buds from herbarium sheets were

softened by gently boiling in water with a few drops of

‘TeepoP, before the anthers were removed. Anthers were

sonicated in distilled water for one minute to release

pollen. The suspensions were passed through 250 pm

mesh filters to remove anther debris. Two drops of lactic

acid were added to pollen filtrates to prevent excessive

expansion of the pollen grains. Pollen was acetolysed

using a freshly prepared acetic anhydride and concen-

trated sulphuric acid mixture (9:1) for 2 minutes 45 sec-

onds in a water bath at 100°C. For scanning electron

microscopy, acetolysed grains were pipetted onto glass

cover slips and fixed onto brass viewing stubs.

Specimens were viewed using a Hitachi S570 Scanning

Electron Microscope (SEM) at 10 kv. For transmission

electron microscopy, acetolysed grains were embedded

in Spurr’s resin using conventional procedures. Sections

slightly thicker than 0.1 pm were stained and viewed

with a Jeol T200 Transmission Electron Microscope

(TEM) at 80 kv.

Seed morphology and anatomy

Seeds were flash frozen in liquid nitrogen and frac-

tured transmedially through the hilum with a scalpel.

Specimens were coated with gold-paladium and viewed

with a Hitachi S570 SEM at 10 kv.

RESULTS AND DISCUSSION

Pollen

Ferguson & Skvarla (1981) reviewed the pollen of the

Sophoreae and Calpurnia conforms to the general mor-

phology of the tribe. The pollen of all species of

Calpurnia is in monads, tricolporate, spheroidal to pro-

late, and apocolpate (Figure lA, B). Further, colpus mar-

gins are smooth and the texture of the colpus furrow may

be smooth or crustate in all species. Size variation is min-

imal and most grains are about 20 x 15 mm. Compara-

tively fewer columellae and a less continuous tectum dis-

tinguishes C. intrusa pollen (Figure 1C) and C. sericea

(Figure ID) from that of other Calpurnia species. The

pollen wall is between 0.5 mm and 1.0 mm thick in all

species. The pollen of the putative hybrid, C. sericea x C.

6

Bothalia29,I (1999)

FIGURE I. — Pollen features of

Calpurnia. A, C. intrusci pol-

len grain, oblique-polar view;

B, C. sericea pollen grain,

equatorial view; C, C. intrusa

pollen surface, tectum perfo-

rate-reticulate; D, C. sericea

pollen surface, tectum perfo-

rate. A, C, Beautnont 80\ B,

D, Beaumont 53. Scale bars; 1

mm.

woodii, and to a lesser extent C. woodii, have slightly

thicker foot and tectum layers than the remaining

species.

Seeds

Mature seeds of all Calpurnia species except those of

C floribunda, and immature seed only of C. reflexus, were

available for study. Seeds maturing apically and basally

where the pod constricts, are frequently smaller and,

together with seeds maturing next to aborted ovules and

aborted developing seeds, were omitted from this study.

Calpurnia seeds range in size from 3x2 mm to 7 x 4

mm and are oblong-reniform to oblong-ellipsoid, and

slightly compressed laterally. The radicle is horizontal to

slightly deflexed (Figures 5S; 6R; 8S; lOR; IIQ; 13Q).

The testa is uniform in colour, ranging from light yel-

lowish brown, through chestnut to dark brown to brown-

black in all species. A dark brown halo surrounds the

hilum and dark markings extend down the raphe through

the lens to the base of the seed opposite the hilar pole.

The hilum is lateral, and is surrounded by the white rem-

nants of the funiculus that form the rim aril. The hilum of

C. aurea subsp. aurea is elliptic and surrounded by a thin

aril. In C. g lab rata, C. intrusa, C. sericea, C. reflexus, C.

woodii and the putative hybrid C. sericea x woodii, the

hilum is round to obovate, and a little smaller than that

found in C. aurea subsp. aurea. The rim aril is of even

thickness or it may be thickest away from the radicular

lobe, and is coarsely rugulate. The Y-shaped micropyle

lies between the hilum and radicular lobe, next to, or

partly surrounded by the rim aril, and the sculpturing of

the testa attains maximum development closest to the

hilum (Figure 2F). C. aurea subsp. aurea, C. glabrata

and C. sericea have uninterrupted rugose sculpturing of

the testa, in contrast to the testa sculpturing of C. intrusa

which consists of tightly packed rugae in finite groups

interrupted by prominent channels. In all species, an

ovate tracheid bar extends from the micropyle to the

ovule bundle, and tracheids are pitted and orientated with

their long axes perpendicular to the hilar groove.

Calpurnia seeds lack a vascular bundle subtending the

tracheid bar.

Calpurnia E.Mey., Commentariorum de plantis

africae australioris 1: 2 (1836); Benth.: 90 (1837); Walp.:

806 (1842); Harv.: 266 (1862); Benth. & Hook.f.: 456

(1865); Harv.: 88 (1868); Baker: 252 (1871); Hook.f.:

251 (1878); Taub.: 197 (1892); Thonner: 258 (1915);

E.Phillips: 475 (1917); Baker f.: 316 (1926); Hutch.: 329

(1964); J.B.Gillett et al.: 46 (1971); Palmer & Pitman:

898 (1972); J.H.Ross: 196 (1972); Yakovlev: 591 (1972);

R.A.Dyer: 244 (1975); Coates Palgrave: 299 (1977).

Type: C. intrusa (R.Br. in W.T.Aiton) E.Mey.

Slender trees or shrubs, deciduous, unarmed; stems

flexible, old stems glabrescent to glabrous, young stems

usually pubescent. Leaves imparipinnate, pulvinate, petio-

late; stipules triangular to subulate, small, paired, adnate to

base of pulvinus; leaflets 4-28, oblong, ovate to obovate,

symmetrical about the midvein, opposite or sub-opposite,

membranous, flat or slightly conduplicate, base often

slightly asymmetrical, pulvinate, apex acute to retuse, with

or without mucro, terminal leaflet sometimes smaller and

broader; venation brochidodromous, secondary veins in

4—7 pairs. Inflorescence racemose to paniculate, terminal

and axillary; bracts deltoid, ovate or narrowly ovate, ses-

sile; bracteoles deltoid to filiform or rarely foliaceous,

paired or absent, sessile or rarely petiolate; flowers

Bothalia 29,1 (1999)

7

FIGURE 2. — Features of seed surface, tracheid anatomy and testa structure in Calpumia sericea, Beaumont 79. A, hilum; B, tracheid bar;

C, tracheids; D, pits in tracheids; E, seed coat structure; F, seed surface near hilum; G, plan diagram of section through hilar region;

a, rim aril; b, hilar palisade; c, palisade; d, hypodermal cells; e, parenchyma; f, tracheid bar. Scale bars: A, 100 pm; B, E, 30 urn; C,

F, 10 pm; D, 3 pm.

Bothalia29,l (1999)

FIGURE 3.^Calpurnki floribunda: A-O, Bean & Viviers 2306\ P, figure drawn from Batten & Bokelmann (1966: plate 67, fig 2a)^ A,

habit; B, leaf; C, leaflet; D, stipule; E, inflorescence; F, bract; G, hairs at junction of pedicel and hypanthium; H, floww, 1, bud, J,

calyx; Ki, standard petal, lateral view; Kii, standard petal, ventral view; L, wing petal; M, keel petal; N, androecium; O, gynoeci-

urn; P, fniit. A, B, E, P, x 0,8; C, D, F-I, x 4.4; J-O, x 3.5,

Bothalia29,l (1999)

9

10-100, papilionaceous. Hypanthium funnel-shaped,

poorly to moderately developed. Calyx campanulate; tube

cylindrical, base variably intrusive; lobes 5, triangular,

straight, rarely retrorse, upper sinus shallow, rarely all

sinuses equal. Petals exserted, bright sulphur to golden

yellow; standard broadly elliptic, obcordate to broadly

ovate, erect or reflexed, apex emarginate, base clawed,

channelled, glabrous or rarely minutely hairy; pollinator

guides if present with red and brown flecks to larger, solid

patches; calluses absent; wing falcate-ovate to falcate-

obovate, auriculate, apex acute to blunt, base clawed, petal

sculpturing lunate-lamellate or absent; keel boat-shaped,

auriculate, shorter than wings, with lateral pockets, base

clawed, apex acute to blunt, lower margins lightly coher-

ent. Stamens 10, monadelphous from adaxial filaments

fused basally to abaxial filaments fused for up to half of

their lengths, or nearly diadelphous, nine filaments fused

to two thirds of their length, tenth filament fused to one

third of its length, glabrous or rarely basally papillate;

anthers dorsifixed, slightly alternately dimorphic. Pollen

tricolporate; tectum perforate to perforate-reticulate.

Ovary’ oblong-falcate, slightly compressed, stipitate;

ovules 6-16; style filiform, slightly arcuate, glabrous or

with sparse extension of ovary vestiture; stigma capitate,

terminal with elongate peripheral papillae. Fruits linear,

compressed, stipitate, 1-6-seeded, papery or woody, brit-

tle, glabrescent to minutely pubescent; upper suture

winged, dehiscent. Seeds ovoid to unequally reniform,

longer than broad, compressed, pale yellow-brown, chest-

nut to brown-black; hilum small, oval to obovate, subter-

minal, rim aril weakly developed.

Key to species

la Inflorescence paniculate:

2a Stipules up to 1 mm long 1 . C. floribunda

2h Stipules longer than 1 mm:

3a Leaves, pedicels and calyces moderately to densely vil-

lous-tomentose; calyx base truncate; bracts 3-5 mm

long; standard broadly ovate; petal claws and lower

third to one half of the filaments concealed by calyx

lobes 2. C. woodii

3b Leaves, pedicels and calyces glabrescent to puberulous,

minutely tomentose or sericeous; calyx base intru-

sive; bracts to 2 mm long; standard oblong-elliptic

or slightly obovate; petal claws and lower third to

one half of the filaments exposed 3. C. sericea

lb Inflorescences racemose:

4a Calyx lobes reflexed; filament bases densely hairy ... 4. C. reflexus

4b Calyx lobes straight; filament bases glabrous:

5a Seeds not exceeding 3.5 x 2.5 mm 5. C. intrusa

5b Seeds larger than 3.5 x 2.5 mm:

6a Fruits usually larger than 50 x 10 mm:

7a Calyx lobes shorter than calyx tube; ovules 10-16,

native to Africa 6a, C. aurea subsp. aurea

7b Calyx lobes equal to or longer than calyx tube;

ovules 6-7; plants restricted to India

6b. C. aurea subsp. indica

6b Fruits usually less than 50 x 10 mm;

8a Petal claws and filaments exposed in mature flowers

7. C. sericea

8b Petal claws and filaments concealed by calyx in

mature flowers:

9a Plants glabrescent; distribution in Swaziland and

Mpumalanga only 8a. C. glabrata

9b Young stems, leaves, and calyx puberulous; ovary

puberulous-tomentose to sericeous; distribution

in Eastern Cape only 8b. C. floribunda

1. Calpurnia floribunda Harv. in Harv. & Sond.,

Roracapensis 2: 267 (1862); E.Phillips: 480 (1917). Type:

Roadside near Grahamstown, H. Hutton s.n. (TCD, holo.;

K, iso.!).

Shrub or small tree up to 3 m tall; young stems mod-

erately puberulous to glabrescent. Leaves 20-220 mm

long, moderately puberulous; petiole 2-19 mm long;

petiolule 1.0-2. 5 mm long, puberulous; stipules triangu-

lar-ovate, 1 mm long, puberulous, persistent; leaflets

4-20, elliptic-oblong, sometimes slightly ovate or obo-

vate, 6-30 X 4-14 mm, base obtuse to acute, apex obtuse

to retuse, blunt. Inflorescence paniculate, rarely race-

mose, 30-80 mm long, densely puberulous, 5-100-flow-

ered; bracts broadly triangular-ovate, 1 mm long, puberu-

lous, persistent; bracteoles usually absent. Hypanthium

1-2 mm long. Calyx 3-5 mm long, moderately puberu-

lous, base intrusive; lobes 1-2 mm long, straight, shorter

than tube. Petals: standard elliptic, 6-7 x 5-6 mm, erect,

apex reflexed, claw 3-4 mm long, glabrous; wing fal-

cate-oblong to falcate-ovate, 7-10 x 3-5 mm, upper

basal region sculptured, claw 3. 5-5.0 mm long; keel 6-7

X 4-5 mm, apex acute to obtuse, claw 3. 5-5.0 mm long.

Stamens fused at base of staminal sheath, glabrous.

Ovary 5-6 mm long, densely puberulous-tomentose to

sericeous; stipe 3. 0-3. 5 mm long; ovules 7 or 8; style

2.0-2. 5 mm long. Fruit and seed not seen. Figure 3.

The floribunda refers to the profuse flowering

of this species with, as Harvey (1862) observed: ‘pedun-

cles crowded towards the ends of the branches.’

The distribution and small stipules of C. floribunda dis-

tinguish it from C. woodii. The concealed petal claws and

filaments distinguish C. floribunda from C. sericea. The

leaves of C. floribunda resemble those of C. aurea subsp.

aurea but the flowers of C. aurea are considerably larger

than those of C. floribunda, and are borne in racemes.

C. floribunda is endemic to the Eastern Cape (Figure

4), occuring from sea level to 2 000 m. It grows in Valley

Bushveld, Eastern Province Thornveld, Highland

Sourveld and Dohne Sourveld and North Eastern

Mountain Sourveld vegetation types of Acocks (1988),

inhabiting forest margins, open scrub bush and gullies.

Flowering time: January to April.

FIGURE 4. — Known distribution of Calpurnia floribunda, •; C.

woodii and C. sericea x woodii, O; C. reflexus, ■; C. intrusa,

□ and C. glabrata. A, in southern Africa.

10

Bothalia29,l (1999)

HGURE 5 —Calpurnia woodii. Beaumont 90. A, habit; Bi, leaf with terminal leaflet; Bii, leaf posture; C leaflet; D, stipule; E, infloresc^ce,

I bn.« G, bb,c,eole; H. flow.,: 1. bod: J. col,x: K, aandwd p.t* L. w,„| pe,* M, keel ^bd: R »db,.au,n: O P,

immature infructescence; Q, mature mfructescence; R, fruit; S, seed. A, B, E, P-R, x 0.8, C, D, F I, x 4. ,

Bothalia29,l (1999)

11

Vouchers: Bean & Viviers 2306 (NU); Bokelmann 7 (NBG); Britten

6517, s.n. (GRA); Dyer 1235 (GRA); Rycroft 1893 (NBG).

2. Calpurnia woodii Schinz in Bulletin de

I’Herbier Boissier 4: 426 (1896); E.Phillips: 478 (1917).

Type: slopes of Drakensberg, Wood 3516 (Z, holo.; K!,

NH!, PRE, iso.!).

Compact shrubs up to 2 m tall, branched basally;

young stems whitish puberulous-tomentose. Leaves

50-200 mm long, moderately to densely villous-tomen-

tose; petiole 9-26 mm long; petiolule 1-2 mm long,

densely villous-tomentose; stipules subulate to filiform,

3-15 mm long, villous-tomentose, persistent, apex often

curled; leaflets 16-28, oblong-elliptic to ovate, 9^1 x

5-16 mm, base round to acute, apex acute, mucronate.

Inflorescence robust, upright, paniculate, 30-100 mm

long densely villous-tomentose, 10-100-flowered; bracts

shortly oblong to elongate-ovate, 3-5 mm long, densely

villous-tomentose, persistent; bracteoles filiform, often

caducous. Hypanthiiim 1 mm long. Calyx 3-5 mm long,

densely villous-tomentose, base truncate; lobes 0.5-1 .5

mm long, straight, shorter than tube. Petals: standard

suborbicular to very broadly ovate, 4. 0-4. 5 x 3.0-3. 5

mm, erect, apex not reflexed, claw 3. 5-4. 5 mm long,

glabrous; wing falcate-oblong, 6.5-7. 5 x 2. 5-3. 5 mm,

upper basal region pocketed and sculptured, claw 3. 0-3. 5

mm long; keel 3. 5-4.0 x 2.5-3. 5 mm, apex blunt, claw

3. 5^. 5 mm long. Stamens fused basally to one third of

their lengths, glabrous. Ovary 4.5 mm long, densely vil-

lous; stipe 1.5 mm long; ovules 6-8; style 3 mm long.

Fruit 25-40 x 9-12 mm, 1-4-seeded, chartaceous, mod-

erately to sparsely villous, pale brown, venation faint;

upper suture wing to 1 mm broad. Seed 5-6 x 3-4 mm

(Figure 5).

This species commemorates John Medley Wood

(1827-1915), botanist and former curator of the Durban

Botanic Gardens.

Distinguishing characters of C. woodii are: the dense-

ly villous-tomentose leaves and inflorescences which

give shrubs a silver-green to grey-white hue, short,

robust, upright panicles which bear numerous small

flowers, and large, persistent stipules and bracts.

Although similar, the fruits of C. woodii (Figures 5Q, R)

are broader and of thinner texture to those of C. sericea

(Figures 6P, Q).

C. woodii has a very restricted distribution in

KwaZulu-Natal (Figure 4) and occurs at approximately

1 500 m above sea level, in the Underberg region and at

South Downs, near Estcourt, Weenen County. The south-

east-facing grassland slopes overlooking Wagendrift

Dam and the Moor Park Nature Reserve near Estcourt,

support a locally abundant population of this species.

Here, plants form a stand in Southern Tall Grassveld

(Acocks 1988) dominated by Themeda triandra Forssk.

C. woodii occurs on steep, dry slopes in loose, shaley

soils undisturbed by cultivation or grazing. There

appears to be little that is unique to the area where

C. woodii grows that could explain the scarcity of this

species. The flowers are prone to insect herbivory and

beetles of the order Bruchidae eat the seeds. The stems

and leaves however, remain ungrazed even by the local

goats. C. woodii is a very appealing species, especially

when in flower, and it is hoped that an interest in its cul-

tivation could be fostered, which might ensure the sur-

vival of this very restricted and little-known species.

Flowering time: December to May.

Vouchers: Beaumont 72, 90 (NU); Rentz 512 (K); Sidey 124 (MO);

Wood 4377 (BOL, K).

3. Calpurnia sericea Harv. in Harv. & Sond.,

Flora capensis 2: 267 (1862); J.H.Ross: 59 (1976). Type:

Lesotho, ' collector unknown 82' (S, holo.!).

C. obovata Schinz: 426 (1896). Type: KwaZulu-Natal, Ingunga,

Schlechter 6310 (Z, holo.!).

C. mucronulata Harms ex Kuntze: 54 (1898). Type: KwaZulu-Natal,

Klip River Dist., Van Reenen’s Pass, Kuntze s.n. (K, lecto.l, here des-

ignated; NY!).

C. obovata var. pubescens Yakovlev: 183 (1971). Type: KwaZulu-

Natal, Utrecht Dist., Kafir Drift, Tweekloof, Thode A270 (K, holo.!;

PRE, iso.!).

C. intrusa auct., non (R.Br. in W.T.Aiton) E.Mey. sensu stricto.

Virgate shrub to 2 m tall; young stems moderately

puberulous to tomentose or glabrescent to glabrous.

Leaves 30-150 mm long, densely to sparsely pubescent

to puberulous, minutely tomentose to sericeous or

glabrescent to glabrous; petiole 2-20 mm long; petiolule

0.5-2.0 mm long, from puberulous, tomentose, villous or

sericeous to glabrescent to glabrous; stipules narrowly

triangular to subulate, 2-A mm long, from puberulous,

tomentose, villous or sericeous to glabrescent to

glabrous, usually persistent, apex acute to filiform;

leaflets 8-24, shape variable within and among plants,

suborbicular to elliptic or slightly oblong to elliptic-ovate

or obovate-elliptic, 3-25 x 3-17 mm, base subcordate to

obtuse, apex retuse to acute, mucronate to blunt.

Inflorescence upright or pendulous, racemose, rarely

paniculate, 30-200 mm long, densely to sparsely pubes-

cent from puberulous, villous, tomentose or sericeous to

glabrescent to glabrous, 10-100-flowered; bracts

oblong-ovate to triangular, 2 mm long, puberulous,

tomentose, villous or sericeous to glabrescent or

glabrous, persistent, apex irregularly dentate or acute;

bracteoles usually absent. Hypanthium 1.0-1. 5 mm long.

Calyx 2. 0-4. 5 mm long, moderately puberulous, tomen-

tose to shortly sericeous, or glabrescent to glabrous, base

intrusive; lobes 0.5- 1.5 mm long. Petals: standard sub-

orbicular to slightly obovate-oblong, 4-5 x 4—6 mm,

apex not reflexed; pollen guides with red and brown

flecks, rarely with larger, blood-red patches flanking

midvein, or absent, claw 2. 5-^.0 mm long, glabrous;

wing falcate-oblong, 5.0-6. 5 x 2.5-3. 5 mm, upper basal

region sometimes pocketed, sculpturing absent, claw

3.0-4. 5 mm long; keel 4.0-5.0 x 2. 5-3. 5 mm, apex acute

to blunt, claw 2. 5-4.0 mm long. Stamens fused at base of

staminal sheath, glabrous. Ovary 4 mm long, minutely

sericeous along sutures or throughout; stipe 1 mm long;

ovules 5-8; style 2.5 mm long. Fruits 20-50 x 5-9 mm,

1-4-seeded, coriaceous to crustaceous, puberulous or

minutely sericeous to glabrous, mid- to dark brown,

venation moderately prominent; upper suture wing

absent or less than 1 mm broad. Seeds 5. 0-6.0 x 2.5^.0

mm (Figure 6).

12

Bothalia29,l (1999)

FIGURE 6 Calpumiu sericca: A, Biii, Ci, D-G, J-Pii, R, Beaumont & Beckett 46: Bi, Beaumont 68\ Bii, Stirton 12740: Cii, Beaumont & Beckett

s.n.: Hi, li, Beaumont & Beckett 47: Hii. lii, Q. Beaumont 62. A, habit; Bi-Biii, leaf; Ci, li, leaflet; D, stipule; E, inflorescence; F, bract; G, hairs

at base of pedicel and hypanthium; Hi, Hii, flower; li, lii, bud; J, ailyx; K, standard petal; L, wing petal; M, keel petal; N, androecium; O, gynoe-

cium; Pi, immature infractescence; Pii, mature infmctescence; Q, fmit; R, seed. A, B, E, P, Q, x 0.8; C, D, F-l, x 4.4; J-O, x 3.5; R, x 18.

Bothalia29,l (1999)

13

FIGURE 7. — Known distribution of Calpurnia sericea in southern

Africa.

The specific epithet refers to the long, silky, close-

pressed hairs of some specimens.

Ross (1976) provides a comprehensive account of the

correct nomenclature of this species. Individuals of C.

sericea show considerable variation in leaflet number,

shape and size, pubescence length and density of leaflets

and ovaries, and raceme length.

Apart from C. aurea subsp. aurea, C. sericea has the

widest distribution among Calpurnia species (Figure 7). C.

sericea often grows in association with Buddleja species,

Leucosidea sericea and Pteridiiim aquilinum. It favours

riverine habitats, especially river valley flood plains and

stream gullies. Flowering time: November to May.

Vouchers; Beaumont 59, 64. 69 (NU); Hilliard cS: Burn 12580 (K,

NU); Nicholas & Neave 2117 (NH).

3a. Calpurnia sericea x woodii

Figure 8 illustrates the putative hybrid C. sericea x

woodii. This is the first record of possible hybridisation

in Calpurnia. We have only seen two hybrid plants, both

by the river at the picnic site of Moor Park Nature

Reserve in KwaZulu-Natal (Figure 4), with C. sericea

nearby, and C. woodii occurring about 2 km away. The

hybrids are vigorous and express a combination of char-

acters from the parents. The hybrid individuals are taller

than plants of both C. sericea and C. woodii, and com-

prise many close-set, semi-erect, flexible stems which

are reminiscent of those of C. sericea. Vegetative charac-

ters which appear to be derived from C. woodii are the

dense villous-tomentose pubescence of the leaves and

stipules, and the comparatively long stipules. The leaves

of the hybrid plants also resemble those of C. woodii in

shape, they are, however, smaller. The inflorescences of

the hybrid plants are long, lax, many-flowered racemes

concentrated towards the ends of branches. This arrange-

ment is similar to the condition found in C. sericea, com-

pared to the robust, erect, densely-flowered and shortly

paniculate inflorescences of C. woodii. Floral characters

of the hybrid plants which resemble C. sericea are: the

exposed petal claws and stamen bases, and suborbicular-

oblong standard petal. In contrast, the densely villous-

tomentose calyces of hybrid individuals are very similar

to those of C. woodii. Bracts of the hybrid plants are of

intermediate length, between those of C. sericea and C.

woodii. Hybrid fruit length to width ratios are very close

to those of fruits of C. sericea. The slightly longer hilum

of seeds of C. woodii however, is also expressed in

hybrid seeds.

Voucher: Beaumont 99 (NU).

4. Calpurnia reflexus A.J. Beaumont, sp. nov., C.

sericeae habitu fructuque similis sed lobis calycis maturi

revolutis et basi non intrusa, staminibus monadelphibus

et ad basin papillatis puberulisque differt.

TYPE. — Lesotho, 2828 (Bethlehem): Leribe Dist.,

Tsikoane, (-DD), Dieterlen 584 (K, holo.!; MASE!, PRE,

SAM!).

Robinia capensis sensu auct. non Burm.f: 22 (1768). Virgilia robin-

ioides sensu auct. non DC: 98 (1825). Calpurnia robinioides sensu

auct. non (DC.) E.Mey.: 3 (1836). Type: 'Robinia capensis Burm.! cap.

Virgilia robinioides DC.’ (G-DC, lecto.!, here designated).

Shrub or small tree up to 4 m tall; young stems puberu-

lous. Leaves 30-100 mm long, moderately puberulous to

glabrescent; petiole 4-23 mm long; petiolule 1-2 mm

long, puberulous; stipules triangular-ovate, 1-2 mm long,

puberulous, caducous; leaflets 4-12, elliptic, or slightly

ovate- or obovate-elliptic, 4-40 x 4—20 mm, base round to

acute, apex round to truncate or rarely slightly retuse,

mucronate to blunt. Inflorescence racemose, 40-80 mm

long, moderately to densely puberulous, 10-30- flowered;

bracts narrowly ovate-triangular, 1.0-1. 5 mm long,

puberulous, caducous; bracteoles filiform, longer than

bracts, caducous. Hypanthium 1-2 mm long. Calyx 5-7

mm long, puberulous, base campanulate; lobes 2-3 mm

long, revolute, shorter than or equal to tube; anterior lobe

shorter than other lobes. Petals: standard obovate, 6-9 x

5-7 mm, erect, apex reflexed, claw 3-4 mm long, papil-

late; wing falcate-oblong, 9-1 1 x 3-5 mm, upper basal and

rarely upper central regions lightly sculptured, claw

3.0-3. 5 mm long; keel 6-8 x 4—5 mm, inner anterior mar-

gins puberulous-tomentose, apex round to obtuse, claw

3-4 mm long. Stamens monadelphous, nine fused to two-

thirds of their lengths, the tenth fused to one third of its

length, base densely papillate-puberulous. Ovary 5 mm

long, margins tomentose-sericeous, sides sparsely papil-

late towards the stipe; stipe 3 mm long; ovules 6-8; style

3 mm long, filiform, glabrous. Immature fruits 30-45 x

8-10 mm, 1-3-seeded, ligneus, glabrescent to glabrous,

straw-coloured, venation prominent; upper suture wing to

0.5 mm broad; mature fruits not known. Mature seed not

known (Eigure 9).

Burman (1768) briefly described a plant which he

named Robinia capensis, although he made no reference

to the specimen upon which he based his description. De

Candolle (1825) listed seven species under Virgilia. He

transferred Burman ’s Robinia capensis, renaming it

Virgilia robinioides (and adopting Virgilia capensis for

Sophora capensis Burm.f). De Candolle (1825) appar-

ently was the first person to have referred to a specimen

upon which Burman (1768) based his description of

14

Bothalia29,l (1999)

nCURE 8 — Calnurnia sericeu x woodii, Beaumont 99. A, habit; B, leaf; C, leaflet; D, stipule; E, inflorescence; F, bract, hairs at base of

pedicel and hypanthium; H, flower; 1. bud; J, calyx; K, standard petal; L, wing petal; M, keel petal; N. androeaum; O, ^noKium; P,

immature infructescence; Q. mature infructescence; R, fruit; S, seed. A, B, E, P-R, x 0.8; C, D, F-1, x 4.4; J-O, x 3.5, S, x

AJB ’90

Bothalia29,l (1999)

15

FIGURE 9, — Calpumia reflexus: A-E, Hi-Hiv, J-Q, Dieterlen 5H4', F, G, 1, Archibald 509. A, habit; B, leaf; C, leaflet; D, stipule; E, inflores-

cence, F, bract; G, bracteole; Hi-Hiv, successive floral maturation; I, bud; J, calyx; K, standard petal; L, wing petal; M, keel petal; N,

androecium; O, base of staminal sheath; P, gynoecium; Q, immature infructescence. A, B, E, Q, x 0.8; C, D, F-I, x 4.4; J-N, P, x 3.5;

O, scale bar, 0.5 mm.

16

Bothalia29,l (1999)

Bothalia29,l (1999)

17

Robinia capensis. He reported this specimen to be

housed in the Delessert Herbarium, Conservatoire et

Jardin botaniques de la Ville de Geneve (G-DEL). Dr A.

Charpin (Geneve) has investigated at our request, the

whereabouts of this specimen, but unfortunately he

reports that the specimen is no longer in the Delessert

Herbarium. Dr Charpin however, did find a specimen in

the De Candolle Herbarium (G-DC), labelled "Robinia

capensis Burm.! cap. Virgilia robinioides DC.’. This pre-

sumably is part of Burman’s original specimen of

Robinia capensis, and possibly the last surviving rem-

nant of this collection. This specimen bears metacentric

trichomes on the calyx, and the dorsal surface of the stan-

dard petal is densely hairy throughout. Metacentric hairs

are absent from Calpurnia, but are a characteristic of a

number of legume genera including Indigofera. Dr B.D.

Schrire (Royal Botanic Gardens, Kew) examined a

microfiche of the specimen (which shows a leaf but no

flowers) and confirmed it to be Indigofera fnitescens.

Accordingly, we reject all Calpurnia synonyms associat-

ed with the Burman fragment in the De Candolle

Herbarium, Geneva (G-DC), and propose the new name

Calpurnia reflexus. The specific epithet refers to the rev-

olute nature of the calyx teeth, which is unique in

Calpurnia (Figure 9).

In addition to the revolute calyx teeth, the monadel-

phous stamens and densely papillate base of the staminal

sheath are unique characters for this species. Yakovlev

(1971), citing calycine and staminal characters, suggest-

ed that this taxon might be worthy of recognition as a

new genus. We prefer to recognise this taxon as a

Calpurnia species because of the similarity of the

remaining vegetative and floral characters between it and

the other members of this genus. There is a paucity of

representative herbarium material of C. reflexus. This

plant appears to be very scarce and might be extinct. The

first two authors visited the recorded Lesotho localities

of the Leribe District, and west of the Thaba-Putsoa

Range near Ramabantas between Roma and Nyakosoba,

in May 1990, but were unable to find the plant. The west-

ern lowlands of Lesotho are heavily overgrazed, the flat

land is cultivated or badly eroded, and consequently,

indigenous flora survives only on isolated koppies.

C. reflexus is an upland species, occuring in mountain

ravines around 1 800 m.

Vouchers: Archibald 509 (GRA); Barber 813 (K); Drege 6402. I, a,

45 (P); Muller 731 (GRA, PRE).

5. Calpurnia intrusa (R.Br. in W.T.Aiton) E.Mey.,

Commentariorum de plantis Africae australioris 1,2: 4

(1836). Type: Hort. Kew 1796 (BM, lecto.!, here desig-

nated).

Virgilia intru.sa R.Br. in W.T.Aiton: 4(1811); DC.: 98 (1825).

C. villo.ia Harv.: 268 (1862); E.Phillips: 475 (1917). Type: Mund &

Maire s.n. sub Herb. Reg. Berol. (K, holo.!).

C. intrusa \ni. glabrataYakovlew 195 (1971). Type: Western Cape,

Oudtshoom Div. Meiringspoort, Acocks 18292 (PRE, holo.!).

Shrub or tree up to 6 m tall; young stems puberulous-

tomentose to glabrescent. Leaves 20-110 mm long;

leaflets 4-24, oblong, sometimes slightly ovate- or obo-

vate-oblong, 7-21 x 4-9 mm, base round to obtuse, apex

round to obtuse, mucronulate or blunt, abaxial surface

villous-tomentose to tomentose-sericeous rarely glabres-

cent to glabrous, adaxial surface glabrescent to glabrous,

rarely villous-tomentose to tomentose-sericeous; petiole

5-15 mm long, pubescent; petiolule 0.5-1. 5 mm long,

pubescent; stipules filiform, 3-5 mm long, villous-

tomentose to glabrescent, often caducous. Inflorescence

pendulous, racemose, 30-100 mm long, densely villous-

tomentose to sericeous or glabrescent, peduncle often

bearing trifoliolate and unifoliolate leaves, 6-30-flow-

ered; bracts narrowly triangular, 3 mm long, sparsely

hairy, often caducous; bracteoles filiform to foliaceous,

often persistent. Hypanthium 2^ mm long. Calyx 5-8

mm long, moderately villous-tomentose to glabrescent,

base intrusive; lobes 1. 5-5.0 mm long, straight, abaxial

and wing lobes equal to or longer than tube, adaxial lobes

shorter than tube. Petals: standard broadly obovate, 6-10

X 7-11 mm, erect, apex reflexed, claw 3-6 mm long,

glabrous; wing falcate-elliptic to falcate-obovate, 8-15 x

3-8 mm, upper central region lightly sculptured, claw

3-5 mm long; keel 6.5-10.0 x 4.0-6. 0 mm, apex obtuse,

claw 3-8 mm long. Stamens fused basally for nearly half

their lengths, glabrous. Ovary 5-10 mm long, villous-

tomentose to sericeous usually along margins, rarely

hairy throughout or glabrous; stipe 1.5-2. 5 mm long;

ovules 9-14; style 2. 5-4.0 mm long. Fruits 16^3 x 5-6

mm, 1^-seeded, chartaceous, glabrescent to glabrous,

pale sandy-brown, minutely flecked grey, venation faint;

upper suture wing absent. Seeds 2. 5-3. 5 x 2. 0-2. 5 mm

(Figure 10).

The specific epithet refers to the intrusive nature of

the calyx base.

Many workers have misapplied the name Calpurnia

intrusa to specimens of C. sericea Harv. The confusion

surrounding the identity of C. intrusa, has arisen because

successive workers perpetuated a misunderstanding of

the original concept of this taxon. Meyer’s (1836) con-

cept of C. intrusa is in fact, C. sericea Harv., and not the

Western Cape endemic which Brown described (Aiton

1811). This taxonomic mistake was perpetuated (Phillips

1917; Burtt-Davy 1932; Henkel 1934) until Brummitt

(1967) and Ross (1976) clarified the nomenclature.

Brummitt (1967) describes two specimens at the British

Museum of Natural History mounted on one sheet; Hort.

Kew 1796 and Hort. Kew 1797, and identified as Virgilia

intrusa. We examined these specimens and, although

they do not bear Brown’s handwriting, we believe that

they represent the plant which he had in mind when he

described Virgilia intrusa. Brummitt (1967) also notes

that the plant illustrated in tablet 2617 in Curtis’s

Botanical Magazine 53 (1826), is in fact, C aurea subsp.

sylvatica, although Brown verified the illustration to be

of Virgilia intrusa. Plants of C. aurea subsp. aurea

resemble those of C. intrusa in some respects. Both have

large flowers, and smaller specimens of C. aurea tend to

be found in southern Africa, where they resemble C.

intrusa with respect to organ size. Tablet 2617 in Curtis’s

Botanical Magazine 53 (1826) clearly portrays the inflat-

ed base of the calyx above the hypanthium, typical of C.

aurea, and unlike the deeply intrusive calyx of C.

intrusa. Meyer (1836) in transferring V. intrusa to Cal-

purnia, cites a Drege specimen (which is a specimen of

Bothalia29,l (1999)

Fir.URF 1 1 -Calnurniu ciurea subsp aurea. Beaumont 26. A, habit; B, leaf; C, leaflet; D, stipule; E, inflorescence; F, bract; G hairs at base

“r Mi H%owen I, bud; J. uulyx. K. uBudurd puiut U wing pert M ku.l peM N^nd»ec,um: O gy.oec.um,

F infructescence hairs at base of pedicel and hypanthium; Q, seed. A, B, E, F x 0.8, C, D, , X . , , - - , Q-

Bothalia 29,1 ( 1999)

19

C. sericea Harv.), and describes the flowers of the new

combination thus: 'in genere minimi, vix tres lineas longi

liitei’. Flowers of C. sericea are small and generally of

this length (3 lines being equivalent to 6 mm), whereas

flowers of C intrusa sensu Brown are usually about

twice this length. Meyer (1836) makes no mention of the

collections Hort. Kew 1796 and Hort. Kew 1797 in the

British Museum of Natural History.

Apart from C. reflexus, plants of C. intrusa tend to

have fewer leaflets per leaf among Calpurnia species.

The leaflets are characteristically oblong and usually

glabrescent above and pubescent below. Yakovlev (1971)

established C. intrusa van glabrata to distinguish speci-

mens with ‘totally glabrous leaflets’. However, the

leaflets of the type specimen of C. intrusa van glabrata

are not glabrous. The upper surfaces of the leaflets are

usually glabrous but the rachis, petioles and lower leaflet

surfaces are sparsely to moderately villous. Some speci-

mens have leaflets which are equally pubescent on both

surfaces (e.g. Hort. Kirstenbosch, sub BOL470050).

Ovary pubescence, both in terms of distribution and den-

sity, is also highly variable, ranging from hairy through-

out, hairs confined to the sutures, to glabrous. We do not

support Yakovlev’s establishment of varieties in C.

intrusa based on variation of pubescence of leaflets or of

ovaries because these characters are so variable and char-

acter states are not clearly defined.

C. intrusa is endemic to the Western Cape, and occurs

south of 33“ S and between 20“ and 23“ E in the Groot

Swartberge region (Figure 4). It occupies an island of

alluvial sand and calcrete surrounded by conglomerate

sandstone, in turn surrounded by quartzitic sandstone

with shale and tillite. C. intrusa is found in gorges in

relict forest patches. Flowering time: December to June.

Vouchers: Beaumont 80 (NU); Goldblatt 7450 (MO, PRE); Middle-

most 607 (NBG); Muir 1968 (BOL, PRE); Stokoe 8779 (SAM).

6. Calpurnia aurea (Aiton) Benth., Commenta-

tiones de leguminosarum generibus: 26 (1837); Baker:

252 (1871); Bedd.: 89, t.l2 (1874); Hook.f.: 251 (1878);

Fiori: 175 (1912); J.S.Henkel: 207 (1934); J.B.Gillett:

421 (1965); Brummitt:123 (1967); J.B.Gillett et al.: 46

(1971); Compton: 247 (1976); Coates Palgrave: 299 (1977);

Britto: 469 (1986). Type: ‘Nat. of Africa.’ ‘Sophora foliis

pinnatis: foliolis numerosis oblongo-ovalibus supra

glaberrimis, caule fruticoso.’ 'Hort. Kew. 1778.' (BM,

lecto.!).

Sophora aurea Aiton: 44 (1789). Virgilia aurea (Aiton) Lam,: 454, t.

326 (1793). Podalyria aurea (Aiton) Willd.: 502 (1799).

Robinia subdecandra L’Herit.: 157, t. 75 (1791); C. subdecandra

(L’Herit.) Schweick; 237 (1937); Brenan: 410 (1949); Eggeling &

Dale: 298 (1952); L.Touss.: 45 (1953); Dale & Greenway; 355 (1961);

F.White; 146 (1962). Type; grown at Paris from seed obtained in

Ethiopia by Bruce s.n. : (G, holo.; K, microfiche!).

Sophora sylvatica Burch.; 146 (1824). Virgilia sylvatica (Burch.)

DC,; 98 (1825). Calpurnia sylvatica (Burch.) E.Mey.; 2 (1836); Harv.;

267 (1862); Calpurnia aurea subsp. sylvatica (Burch.) Brummitt: 123

(1967). Type: forests of the Boschberg, Burchell 3138 (K, holo.l).

C. lasiogyne E.Mey.: 3 (1836); Harv,: 267 (1862); F.Muell.; 31

(1866); J.M.Wood & M.S.Evans; 6 (1899); E.Phillips: 477 (1917);

Burtt Davy; 354 (1932). Type: ‘Zwischen Omtata und Omsamwubo,’

Drege Vb. 47 (MOI, PI, K, isosyn.l); ‘prope Port Natal, 3000 feet,’

Drege V.c, 34 (P, syn.l).

C. aurea var. major Oliv. & Baker f.: 339 (1886); Baker f; 594

(1926). Type; Kilimanjaro 6000 feet, H.H. Johnston s.n. (K, holo.!).

6a. subsp. aurea.

Shrub or slender tree to 15 m tall; young stems dense-

ly puberulous-tomentose to glabrescent. Leaves 40-250

mm long, moderately puberulous-tomentose to glabres-

cent-glabrous; petiole 10-30 mm long, densely puberu-

lous-tomentose to glabrous; petiolule 1^ mm long,

densely puberulous-tomentose to glabrous; stipules trian-

gular, 1. 5-3.0 mm long, puberulous to glabrous, often

persistent; leaflets 8-30, oblong-elliptic to ovate or obo-

vate, 10-55 x 5-25 mm, base subcordate to acute, apex

retuse to obtuse, blunt to mucronulate. Inflorescence pen-

dulous, racemose, 40-250 mm long, densely puberulous-

tomentose to glabrescent, 8^0-flowered; bracts broadly

triangular, 0. 5-3.0 mm long, puberulous-tomentose to

glabrous, persistent; bracteoles usually absent.

Hypanthium 2-5 mm long. Calyx 5-13 mm long, moder-

ately puberulous to glabrous, base campanulate; lobes

2-5 mm long, straight, shorter than tube. Petals: standard

suborbicular to obovate, 9-20 x 8-20 mm, erect, apex

reflexed, claw 5-13 mm long, glabrous; wing falcate-

oblong, falcate-ovate or falcate-obovate, 6-25 x 4-12

mm, sculpturing present in upper basal and central

regions or absent, claw 4—9 mm long; keel 7-20 x 5-10

mm, apex broadly acute to obtuse, claw 4-9 mm long.

Stamens fused basally less than one fifth of their lengths,

glabrous. Ovary 7-15 mm long, densely sericeous to

glabrous; stipe 6-10 mm long; ovules 10-16; style 6-9

mm long. Fruits 40-140 x 9-23 mm, 3-8-seeded, char-

taceous, puberulous to glabrous, pale sandy-brown to

brown-black, venation moderately prominent to faint;

upper suture wing to 3 mm broad. Seeds 5-8 x 3-5 mm

(Figure 11).

The epithet aurea refers to the golden-yellow flowers.

In South Africa it is known as Natal laburnum.

C. aurea subsp. aurea, the best known Calpurnia spe-

cies, is a frequent entry in African floras and botanical

checklists. It is cultivated as an ornamental and as a

shade tree in tea and coffee plantations (Brummitt 1967).

C. aurea subsp. aurea has the largest vegetative, flo-

ral and fruit characters in Calpurnia. It is the tallest

species, often growing as a slender tree in forest clear-

ings and forest margins. In drier, open scrub and grass-

land habitats it grows as a shorter, much branched shrub

(Figure 12). Another characteristic of this species is the

base of the calyx tube which is slightly inflated around

the apex of the hypanthium. When viewed laterally, the

base of the calyx tube does not obscure the hypanthium,

but the calyx tube and teeth conceal the petal claws and

lower portions of the stamens (Figure IIH). Flowering

time: throughout the year when mild to hot weather pre-

vails, usually during spring and summer.

We recognise two subspecies in Calpurnia aurea:

subsp. aurea and subsp. indica. Subsp. indica occurs in

India; however, specimens from Indian herbaria were

20

Bothalia 29,1 (1999)

FIGURE 12. — Known distribution of Calpurnia aurea subsp. aurea in

southern Africa.

unavailable to us for study. Although limited material from

other herbaria was examined, the present authors feel that

more consultation of specimens of subsp. indica is needed

in order to present a full descriptive account of this taxon.

Britto (1986) provides a clear account of the two sub-

species in India. He uses ovule number per pod to distin-

guish subsp. indica (with six to seven ovules) from subsp.

aurea with 12 to 15 ovules. Brummitt (1967) cites differ-

ences in the relative lengths of the teeth to the campanu-

late part of the calyx, to distinguish the two subspecies.

We reject previous decisions by other authors to

recognise subsp. sylvatica within C. aurea, or to recog-

nise it as a distinct species. Former taxonomic treatments

distinguished such specimens by their glabrous leaflets

and ovaries. Vestiture of leaves and ovaries is notorious-

ly variable among other members of this genus as well as

among specimens of C. aurea sensu lato. The infraspe-

cific discrimination in C. aurea based upon degree of

pubescence of these features has been quite arbitrary.

Previous workers did not consider the pubescence of

other structures. Vestiture on stipules, petioles, rachides,

and petiolules do not necessarily correlate with the

pubescent or glabrous nature of the leaflets of specimens.

Features of the inflorescence (the peduncles, bracts,

pedicels and calyx) may also be variously pubescent to

glabrous, independent of the condition of the ovary.

Plants in this genus are rarely, if ever, glabrous, and usu-

ally some pubescence is present, at least at the junctions

of structures for example, bracts and pedicels and peti-

oles. Plants of C. aurea vary greatly in the distribution

and density of their pubescence, and classification at the

specific or subspecific levels using these criteria is

impractical and controversial with respect to defining the

limits between character states.

Vouchers: CodJ & De Winter 3267 (K, PRE); Edwards 1014 (NU);

Galpin 1214 (BOL, PRE, SAM); Hilliard 2517 (NU); Slirlon 5201

(MO, PRE, SRGH).

7. Calpurnia glabrata Brummitt in Kew Bulletin

24: 71 (1970). Type: Swaziland, Mbabane Dist., Sheba’s

Veil, Compton 27133 (K, holo.!; NBG!, PRE, iso.).

Shrub or small tree to 3 m tall; young stems glabrescent

to glabrous. Leaves 70-360 mm long, glabrescent to

glabrous; petiole 11^8 mm long; petiolule 2-6 mm long,

glabrescent to glabrous; stipules triangular, 1-2 mm long,

glabrescent to glabrous, persistent; leaflets 8-26, oblong-

obovate to obovate, 14—38 x 5-16 mm, base acute and

sometimes attenuate towards petiolule, apex blunt to

rounded and sometimes slightly retuse, mucronulate to

blunt. Inflorescence flexible, racemose, 80-210 mm long;

glabrescent to glabrous, 15^0- flowered; bracts triangular,

0.5-1. 0 mm long, glabrescent to glabrous; bracteoles

minute to absent. Hypanthium 2-3 mm long. Calyx base

intrusive; lobes 0.5-2. 5 mm long, triangular, straight,

shorter than tube. Petals: standard suborbicular to broadly

elliptic, sometimes slightly obovate, 4—6 x 5-7 mm, erect,

apex reflexed, claw 3-^ mm long, glabrous; wing falcate-

elliptic to falcate-ovate, 4—9 x 2. 5-5.0 mm, upper basal

region lightly pocketed or sculptured, claw 3-6 mm long;

keel 3.0-7. 5 x 2. 5-5.0 mm, apex acute to obtuse, claw 3-5

mm long. Stamens fused basally to half their lengths,

glabrous. Ovary 3-8 mm long, glabrous or upper margin

sparsely puberulous; stipe 2.5-7. 5 mm long; ovules 6 or 7;

style 2-3 mm long. Fruits 29-45 x 10-12 mm, 1-3-seed-

ed, ligneous, glabrous, deep brown, venation prominent;

upper suture wing 0. 5-2.0 mm broad. Seeds 5.0-7.0 x

3. 5-5.0 mm (Figure 13).

The epithet glabrata refers to the glabrescent nature

of plants.

The most striking feature of C. glabrata is the almost

glabrous nature of all parts. Presence or absence of hairs,

however, is a character to be used with great caution

when identifying Calpurnia specimens. In the absence of

floral or fruit characters, vegetative specimens of C.

glabrata may be confused with glabrescent specimens of

C. aurea subsp. aurea occurring in the southern Cape.

Flower size varies in C. glabrata (Figure 13E). Flowers

may he similar in size to those of C. sericea (Figure 6E)

or larger, resembling those of C. floribunda (Figure 3E).

The calyx of C. glabrata conceals the wing and keel

petal claws, and the filament bases. These features are

exposed in flowers of C. sericea, which C. glabrata oth-

erwise resembles. In general outline and size, flowers of

C. glabrata may resemble those of C. floribunda, but the

pubescence of the leaves, calyx and ovary of the latter is

denser than in C. glabrata.

The fruits of C. glabrata most closely resemble those

of C. sericea and C. reflexus, although the upper suture

wing of C. glabrata fruits is slightly broader.

C. glabrata (Figure 4) is concentrated in Swaziland

but it also occurs in Mpumalanga between 30° and 31° E

and in KwaZulu-Natal to 29° S. It occurs between 200 to

1 500 m. Flowering time: October to April.

This species occurs predominantly in North Eastern

Mountain Sourveld (Acocks 1988) in hillside grassland.

Acacia caffra scrub, roadsides and riverbanks. In

Swaziland it grows in disturbed roadside grassland with

the weeds Pteridium aquilinum (bracken) and Psidiurn

guajava (wild guava).

Vouchers: Beaumont 96 (NU). Compton 63 (NBG); 28152 (NBG,

PRE); 29869 (K, NBG, PRE, SRGH); 32203 (K, NBG, PRE).

Bothalia 29,1 (1999)

21

FIGURE n.—Calpurnia glabrata, A-O, Beaumont 96\ P, Q, Compton 28152. A, habit; B, leaf; C, leaflet; D, stipule; E, inflorescence; F,

bract, G, flower, H, buds, 1, calyx; J, standard petal; K, wing petal; L, keel petal; M, androecium; N, gynoecium; O immature

infructescence; P, fruits; Q, seed. A, B, E, O, P, x 0.8; C, D, F-H, x 4.4; 1-N, x 3.5; Q, x 18.

22

Bothalia 29,1 (1999)

SPECIMENS EXAMINED

Abbott 1351 (6a) NH; 2464 (6a) UNITRA. Acheson 25 (6a) SRGH.

Acocks 9861 (3) NH; 18292 (5) PRE. Anon. 732 (5) B; ,<?S97(6a) BOL;

17315 (5) NBG, Archibald 509 (4) GRA. Ash 232 (6a) MO.

Balkwill, Manning & Meyer 794 (3) NU. Bants 5 (6a) NY. Barber 813

(4) K. Barbosa 9430 (6a) LISC. Barbosa & Moreno 9739 (6a) DISC.

Barker 5219 (6a) NBG. Battiscombe 514 (6a) K. Baur 575 ( 3) SAM.

Bayliss 4443 (6a) MO, NBG, NY; 8762 (6a) MO, NY. Bean & Viviers

2306 (1) NU. Beattie 30 (3) NH, NU. Beaumont 72. 73. 90 (2) NU; 2,

55, 57, 58. 59. 60. 64. 65, 66, 69 (3) NU; 99 (3a) NU; 80 (5) NU; 25,

26, 29, 31. 32, 35, 49, 50, 52 (6a) NU; 96 (7) NU; 100 (3) ?; Beaumont

& Beckett s.n. (3) NU. Bews 1299 (3) NU. Bie gel 1550 (6a) SRGH.

Bokelmann 7(1) NBG. Bourquin s.n.. 36, 159 (6a) NH. Brassnett 116

(6a) K. Britten s.n., 6517 (1) GRA; 1 (6a) BOL. Brown & Shapiro 275

(6a) U. Bruce s.n. (6) K. Bullock 2817 (6a) B, K, LISC. Burchell 3138

(6) K; 3138. 3233 (6a) K. Buthelezi 372 (3) NH.

Cadman, Edwards & Norris 3319 (3) NU. Cawe 473 (6a) UNITRA.

Chase 381 (6a) BM, NU, SRGH; 4221 (6a) BM. Codd 9750 (6a) UPS.

Codd & De Winter 3267 (6a) K. Coetz.ee 938 (6a) SRGH. Compton

22471 (3) NBG; 28027 (6a) K, NBG; 29270 (6a) NBG; 63, 24703,

28152 Cl) NBG; 27133, 32203 (7) K, NBG; 29869 (7) K, NBG, SRGH.

Corby 1637, 1994 (6a) SRGH. Couto 107 (6a) LISC, SRGH. Crewe 43

(6a) NU.

Dahlstrand 2662 (6a) MO. Dahlstrand & Mogg 33 (6a) J. De Menezes

1622 (6a) BM, K, LISC, SRGH. Deschamps, Murta & Da Silva 1184,

1186, 1188, 1233, 1234, 1303 (6a) LISC. Dieterlen 37(3) K, P, SAM;

584 (4) K, MASE, SAM. Drege l.a. 45 , 6402 (4) P; .s.n. (6a) K, MO;

s.n. (6a) K; s.n. (6a) P; V.b, 47 (6) MO, P, K; V.c, 34 (6) P. Duckworth

163 (6a) D. Killick Herb. Dutton & Tinley 8 (6a) NH, NU. Dyer 1235

( 1) GRA. Dyer & Verdoorn 4220 (6a) K.

Ecklon & Zeyher 1142 (6a) MO. Edwards 2694 (2) NH, NU; 32 (3)

NU; 725, 804, 934, 1014, 1121, 2316, 2374 (6a) NU; 25270 (6a) J.

Evans 456 (3) NH; 537 (6a) NH. Exell & Mendonca 2481 (6a) BM.

Eyles 6159 (6a) SRGH.

Faden 74 (6a) MO. Feinauer s.n. (3) NBG. Ferreira 224 (3) K.

Flanagan 127 (6a) SAM; 808 ( I ) GRA, SAM; s.n. (3) SAM.. Forrester

& Gooyer 186 (6a) J. Fry 2776 (6a) GRA.

Galpin 1214 (6a) BOL, SAM; s.n. (6a) BOL; s.n. (7) BOL. Ger.stner

4090 (3) NH; 4092 (7) NH. Gillett 14244 (6a) BOL. Goldblatt 7450 (5)

MO. Goldsmith 36 (6a) SRGH. Goodier & Phipps 150 (6a) SRGH.

Go.ssweiler 12678 (6a) LISC. Greenhow s.n. (6a) NY, SRGH.

Greenway 7596 (6a) K, Greenway & Kanuri 12061, 12473 (6a) K.

Guillarmod 9878 (3) GRA. Guy 72 (6a) NU.

Henderson 400 (6a) MO. Henriques 1059 (6a) LISC. Herb. Le Testu

(sheets 1 and 11) 4249 (6a) BM. Hilliard 2780, .s.n. (3) NU; 1306 (3)

NH, NU. 1040, 2112, 2517 (6a) NU. Hilliard & Burn 8501 (7) NU;

10064 (7) MO, NU; 12580 (3) K, NU. Hort. Kew 1778 (6) BM; 1796

(5) BM. Huntley 768 (3) NU. H. Hutton s.n. (1) K.

Johnson 62 (6a) NBG; 696 (6a) B, BOL. H.H. Johnston s.n. (6) K.

Johnston s.n. (6a) BM.

Kemp 990 (6a) MO. Kuntz.e s.n. (3) K, NY.

Lawn 37 (6a) NH. Leach 9658 (6a) SRGH. Lebrun 3912 (6a) BM, K;

5498 (6a) P. Leistner 3028 (3) SRGH. Levyns 6933 (6a) BOL, P. Lubke,

Everard & Avis 2697 (6a) GRA.

Maas Geesteranus 4976 (6a) G, MO. Maguire 157 (6a) J. McClean 37

(3) MO, PRE; s.n. (6a) NH. Mendes 923 (6a) ?. Middlemo.st 607 (5)

NBG. Mogg 7546 (3) NH; 25374 (3) MO; 27864, 31071 (6a) J. Moll

828 (6a) NU,- 1896, 3358 (6a) K, NU; 4448 (6a) SRGH; 5459 (3) NH.

Moore s.n. (6a) NBG. Moreno 27 (6a) LISC. Mo.s.s 27753 (7) BOL, J.

Muir 1968 (5) BOL. Muller 731 (4) GRA. Mund & Mairc s.n. (5) K.

Nel 234 (6a) NBG. Nelson s.n. (4) K. Nel.shiungani s.n. (6a) J. Nicholas

& Neave 2117 (3) NH. Nicholson 276 (6a) NH.

Obermeyer, Schweickerdt & Verdoorn 353 (6a) NH.

Pappe s.n. (6a) K. Paterson s.n. (6a) BOL. Paulo (sheets 1 and II) 750

(6a) MO. Pegler 79 (3) GRA; 155 (6a) BOL, K, SAM. Phillips .s.n. (6a)

NU. Pienaar 128 (6a) MO, SRGH. Platt s.n. (3) NH. Polhill & Paulo

2304 (6a) B, LISC, Powell-Cotton 667 (6a) BM, MO, NY.

Ranales 89 (3) NU. Rauh & Schlieben 9642 (6a) B, K, SRGH. Rentz512

(2) K. Retief 1004 (3) K. Rudatis 2056 (3) NH. Rycroft 1893 ( 1 ) NBG,

Scheepers 1024 (6a) MO, SRGH. Schimper 200 (6a) K. Schlechter

6310 (3) Z. Schlieben 4499 (6a) B, BM, G, LISC. Schrire 246 (6a) NH.

Semsei 2785 (6a) K. Sidey 124 (2) MO. Sim 19071 (3) NU. Stalmans

69 (6a) K, Stirton 5201 (6a) MO, SRGH. Stokoe 8779 (5) SAM. Story

3407 (6a) MO. Strey 6394 (6a) NH, NU; 8606 (6a) K, NH, SRGH.

Swynnerton 79 (6a) BM, K, SAM.

Taylor 1752 (6a) NY. Teixeira 1642, 2158, 3409 (6a) LISC. Teixeira &

Andrade 4389 (6a) LISC, SRGH; 5172, 5218 (6a) LISC. Thode A270

(3) K, PRE; s.n. (3) K; s.n. (6a) NH. Tinley 696 (6a) NH. Torre &

Pereira 13 (6a) LISC. Tyson 1248 (3) UPS; 1355 (3) BOL, SAM;

2595, 3062 (6a) SAM.

Van Wyk 7139 (3) NH, UNITRA.

Ward 518 (6a) NH; 3238 (7) NU. Werger 243 (3) SRGH. Wild 2103 ,

2171 (6a) SRGH; 4286 (6a) K, LISC, MO, SRGH; 4335 (6a) LISC,

MO, SRGH. Wilson 771 (6a) K. Wood 3516 (2) K, NH; 4377(2) BOL,

K; s.n. (3) MO; 11153 (3) NH, UPS. Wright 1975 (3) NU.

Zeyher 777 (6a) K, P.

ACKNOWLEDGEMENTS

Dr R.K. Brummitt (Royal Botanic Gardens, Kew,

UK) and an anonymous referee of an earlier manuscript

are thanked for their constructive and valuable com-

ments. We thank cited herbaria for the loan of specimens;

Dr A. Charpin of Conservatoire et Jardin botaniques de

la Ville de Geneve for finding the type of Robinia capen-

sis\ and Mrs J. Browning of the Department of Botany,

University of Natal, Pietermaritzburg for discussion and

support. We also thank Mr R. Scott-Shaw of the Natal

Parks Board who first drew our attention to the putative

hybrid C. sericea x C. woodii. The authors gratefully

acknowledge the Foundation for Research Development

(FRD) and the University of Natal Research Fund for

financial support.

REFERENCES

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Bothalia 29.1: 25-33 (1999)

Studies in the liverwort genus F ossombronia (Metzgeriales) from

southern Africa. 8. F. elsieae and F. spinosa, two new Western Cape

species with spinose spores

S.M. PEROLD*

Keywords: Fossombronia Raddi, F. elsiecie Perold, F. spinosa Perold, Hepaticae, Metzgeriales, southern Africa, Western Cape

ABSTRACT

Two new species with spinose spores are described from the Western Cape. F. elsieae Perold is quite a large plant, gener-

ally with almost entire leaves, its spores on the distal face having relatively few, rather coarse spines, which often appear bro-

ken and are occasionally linked to form abbreviated ridges. F. spinosa Perold is a smallish plant with irregularly lobed leaves,

its spores on the distal face having numerous, fine spines, which sometimes coalesce to fonn short ridges. A table which com-

pares F. glenii Perold, F. leucoxantha Lehm. and F. montaguensis S.W.Amell with the two new species, is supplied.

1. Fossombronia elsieae Perold, sp. nov.

Plantae dense coarctatae, apicem versus virides, cito

albescentes et translucentes. Folia late patentia, imbrica-

ta, obovata vel suboblonga. Dioicae. Antheridia bracteis

forma irregulari subtensa. Pseiidoperianthium campanu-

latum; ore undulato cum lobis ± 6 rotundatis. Sporae

47.5- 57.5 pm diametro, superficie distali convexa, cum

9 vel 10 seriebus spinarum grossarum conicarum, inter-

dum in cristis irregularibus concatenatarum; superficie

proximali sine nota triradiata, cum cristis brevibus et spi-

nis nonnullis. Elateres 125-165 pm longi, 7.5-12.5 pm

lati, omnino ter spirales vel alii bis spirales, alii apicem

versus bis spirales, alii ter, raro quater apices versus ter,

spirales.

TYPE. — Western Cape, 3318 (Cape Town); Table

Mountain, top of Nursery Gorge, (-CD), E. Esterhiiysen

24885 (BOL, holo.).

Plants in crowded stands, apically green, soon

bleached and translucent, rather fragile; shoots medium-

sized to quite large, 6-12 mm long, ± 2.5 mm high,

3. 5- 4. 5 mm wide; simple (Figure 2A) or apically furcate,

segments 2-3 mm long, moderately divergent, some-

times with lateral buds toward base. Stems prostrate,

fresh apical shoots often arising from somewhat with-

ered base (Figure 2B), in recent collections dorsally

chlorophyllose, ventrally purple, in cross section plano-

convex, in male plants at apices (Figure IN) 300-350

pm ( 10 cell rows) high, 500-600 pm wide and at bases

(Figure 10) 220-350 x 300^30 pm; in female plants at

apices (Figure IP) 300-400 pm (12 cell rows) high,

450-500 pm wide, tapering proximally and basally

(Figure K^), 250-300 x 400-450 pm. Leaves spreading

widely, overlapping, succubously inserted on stem, quite

large, except for young apical ones, plane and rounded

above, mostly longer than wide and narrowing toward

base, obovate to almost oblong, in male plants (Figure

lA-E) 1125-2000 X 1125-1500 pm, in female plants

(Figure IF-K) somewhat larger, 1700-2125 x 1625-2125

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

MS. received: 1998-09-23.

pm, margins nearly entire, with few (1-3) papillae, most-

ly at slight angulations. Leaf cells thin-walled, in male

plants tending to be slightly smaller, but not significant-

ly so, in female plants at upper margin (Figure IL) sub-

quadrate to rectangular across, 45-50 x 55.0-62.5 pm, at

lower lateral margins long-rectangular, 90.0-112.5 x

27.5- 47.5 pm; upper laminal cells 5- or 6-sided,

62.5- 80.0 X 50-55 pm; middle laminal cells 87.5-132.5

X 57.5-62.5 pm; basal cells 75-125 x 37.5-50.0 pm. Oil

bodies present in fresh material only (Figure IM), up to

34 per cell, shiny, globular to slightly irregular in shape,

± 5 pm diam.; chloroplasts similar in size.

Dioicous. Antheridia dorsal on stem, in 1 or 2 rows,

short-stalked, globose, ± 270 pm diam., subtended by

perigonial bracts (Figure IR-U), single or paired

(Figure 2C, D), shape irregular, body 400-500(-600) x

1 80-280(-3 10) pm, upper margin with low, blunt or tall,

spinous projections, up to 300 pm long and 160 pm wide

at base, tapering to a pointed tip topped with a papilla,

cells in interior 5- or 6-sided, 60.0-82.5 x 37.5^5.0 pm.

Archegonia (Figure 2E) in 1, 2 or more rows, dorsally

along stem. Pseudoperianth (Figure IX, Y; Figure 2F)

campanulate, near stem apices, 1500-1800 pm long,

sometimes slit along the side, one margin overlapping

the other, base narrow, ± 875 pm wide, upwardly widely

flaring, 2250-2625 pm across undulating mouth, with ±

6 rounded lobes, marginal papillae few, cells comparable

in shape and size to those of leaves. Capsules globose, ±

625 pm diam., cells in inner layer of bistratose capsule

wall (Figure IV) ± rectangular, roughly triangular or

irregularly shaped, 52.5-65.0 x 30-50 pm, each cell wall

with 1-3 nodular and some semi-annular thickenings.

Seta from very short to 3.25 mm long, 150 pm or 9 cell

rows wide (Figure IW). Spores brown to dark brown,

hemispherical, 47.5-57.5 pm diam. including spines pro-

jecting around periphery; distal face convex (Figure

3A-D), covered with 9 or 10 rows of coarse, often coni-

cal spines (or a total of ± 39), ± 5 pm long, basally joined

by fine lines, sides grooved, apices sometimes ‘broken’,

apparently hollow inside, occasionally linked to form

irregular ridges; proximal face (Figure 3E) almost flat,

lacking a distinct triradiate mark, with short, slightly sin-

uous or occasionally branched ridges and some blunt

26

Bothalia 29,1 (1999)

FIGURE 1. — Fossomhronia elsieae. A-E, leaves of male plant; F-K„ leaves of female plant; L, detail of marginal area of leaf; M, median leaf

cells with oil bodies (solid lines) and chloroplasts (dotted lines); N, c/s apical part of male stem; O, c/s basal part of male stem; P, c/s api-

cal part of female stem; Q, c/s basal part of female stem; R-U, perigonial bracts; V, cells in capsule wall with thickenings; W, c/s seta; X,

pseudoperianth from side; Y, opened pseudoperianth. A-L, N-V, X, Y, Esterhuysen 24885; M. W, Bra^gins 97/356A. Scale bars: A-K, X.

Y, 5(K) pm; L, W, 100 pm; M, V, 50 pm; N-Q, R-U, 250 pm.

Bothalia29,l (1999)

27

FIGURE 2. — Fossombronia elsieae. A, simple shoot; B, fresh apical shoot from rather withered base; C, D, male stem with perigonial bracts; E,

female stem with archegonia from above; F, female stem with pseudoperianth and capsule from above. A, E, S.M. Perold 3475\ B-D, F.

Esterhuysen 24885. A, x 7.8; B, x 12.8; C, x 10.5; D, x 16; E, x 17; F, x 17.

spines in between, 21-25 spines projecting around

periphery. Elaters (Figure 3F) yellow, smooth, 125-165

pm long, 7.5-12.5 pm wide in centre, often tapered

toward tips and ending in a loop, 5 pm wide; 3-spiral

along entire length, occasionally 2-spiral only at tips,

remainder 3-spiral; some wholly 2-spiral; rarely loosely

4-spiral, tapering tips 3-spiral, or only exceptionally

branched into 2 near apex, both tips 2-spiral, the lower,

unbranched part 3-spiral.

Fossombronia elsieae has been named in honour of Ms

Elsie Esterhuysen, the doyenne of southern African plant

collectors, who collected it in October 1951 at the top of

Nursery Gorge, Table Mountain. This specimen was

FIGURE 3. — Fossombronia elsieae. Spores and elaters. A, B, distal face; C, side view of distal face; D, detail of spines and lamellae at margin of

distal face; E, proximal face; F, elaters. A, S.M. Perold 3475; B-F, Esterhuysen 24885. A, x 761; B, x 71 1; C, x 876; D, x 1560; E, x 715;

F, X 1013.

28

Bothalia29,l (1999)

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misidentified by Amell as F. crispa. In my paper re-exam-

ining F. leucoxantha Lehm. (Perold 1997c), I referred to

this collection, which I have now selected as the holotype

of this new species, stating that it has ‘almost entire

leaves’. I thus distinguished it from F leucoxantha, which

has ‘denticulate to incised-dentate leaves’, in spite of both

species having spinose spores. The spores in F. leucoxan-

tha almost invariably have papillae between the at least 90

spines covering the distal face, whereas in F. elsieae there

are no papillae and only up to 39, rather coarse, often bro-

ken spines on the distal face. Clearly, there are subtle dif-

ferences between these two species (and F. spinosa) in the

spore ornamentation, but much more obvious differences

vegetatively. In the recently described F. glenii (Perold

1997a) from the northern summer rainfall region, the

spines on the distal face of the spores are often double and

the ± undulating leaf margins have shallow angular lobes.

In Table 1 , some characters of F elsieae and F. spinosa are

compared to those of F. glenii Perold, F leucoxantha and

F. montaguensis S.W.Arnell.

In addition to the methodology employed by Scott &

Pike (1988), and others, who relied ‘almost completely

on spore characters — surface sculpturing, often colour

and, much less, spore size — for discriminating species’,

I have tried to use vegetative characters also. Extensive

use of LM and SEM micrographs of the spores and thal-

li has, of course, made comparison between specimens

easier. At times, however, it remains extremely difficult

to establish species parameters in this genus. Accordingly,

I am still undecided where to place the other specimen,

Arnell 616, which I also excluded from F. leucoxantha in

my treatment of that species (Perold 1997b). Other

species with coarse spines on the distal spore face have

been described from elsewhere in the world, notably F.

caespitiformis De Not., which has a total of only ± 25

spines on the distal face and ± 29 around the periphery.

Vegetatively it does not appear to be closely related to F.

elsieae, nor do the Australian species with coarsely spin-

ose spores, F. cultrifomiis, F magnaspora and F. trunca-

ta, which were described by Scott & Pike (1987).

Fossombronia elsieae (and F spinosa), with spores

ripening in spring (September/October), are hitherto

known from only a few localities in the Western Cape

(Figure 4), which is in the winter rainfall area of south-

ern Africa. This is referred to as the Fynbos Biome (scle-

rophyllous, microphyllous vascular plant vegetation)

(Cowling et al. 1997; Rutherford 1997), or else the area

is called the Cape Region — phytogeographic region

based on tbe distribution of vascular plants (Cowling &

Hilton-Taylor 1997). The holotype specimen of F

elsieae, as already mentioned, was collected on Table

Mountain. It grew on black, humus-rich soil containing

quartz grains derived from Table Mountain sandstone.

Other specimens referred here are Braggins 97/354 from

Silvermine Reserve, near Kalk Bay as well as Braggins

97/356A from Stinkwood Trail, Kirstenbosch. Their

spore ornamentation is closely similar to that of the holo-

type specimen, but some of the leaves are apically divid-

ed into 2 ± triangular lobes. Schelpe 6368 is from Bain’s

Kloof, Witte River and Perold & Koekemoer 3475 is from

Tradouw Pass. F. elsieae is distinguished by its fairly

large size, by its obovate to oblong, almost entire leaves,

as well as by its spores with the distal face having rela-

IMGURE 4. — Distribution of F. elsieae. □: and F. spinosa. •. in south-

ern Afnca.

tively few, rather coarse spines which are occasionally

linked to form short ridges.

2. Fossombronia spinosa Perold, sp. nov.

Plantae dense coarctatae, laete virides, apicibus vel

solum marginibus foliorum apicalium purpureis. Folia

suberecta, imbricata, plicata, supra irregulariter lobata.

Dioicae. Antheridia bracteis obtecta, marginibus superi-

oribus cum processibus acutis. Pseudoperianthium

infundibuliforme; ore convoluto, cum processibus angu-

latis vel spinosis. Sporae 35-45 pm diametro, superficie

distali convexa, cum 11-15 seriebus spinarum tenuium,

interdum in cristis brevibus concatenatarum; superficie

proximali sine nota triradiata, subtiliter vel grosse granu-

lar!, cum lamellis tenuibus nonnullis. Elateres 100-150

pm longi, medio ± 7.5 pm lati, laxe bis vel ter spirales,

vel medio ter et apices versus bis spirales.

TYPE. — Western Cape, 3320 (Montagu): Kogman’s

Kloof, 5 km from Montagu, immediately beyond tunnel

on road to Ashton, at Loftus Bridge, along disused road

along the left side of the river; on soil on vertical rock

face, (-CC), S.M. Perold 3835 (PRE, holo.).

Plants in dense colonies, bright green, apical tips occa-

sionally entirely purple or else only margins of young

leaves deeply stained dark red; shoots smallish, up to 9 mm

long, (0.9-)l.l-l .6 mm high, 1.4— 2.4 mm wide; rarely

simple (Figure 5A), mostly apically furcate, terminal seg-

ments 2-A mm long, moderately to widely divergent, side

branches 1-3, apparently arising from lateral buds. Stems

prostrate, sometimes arched, apices tuberous, occasionally

slightly raised, plano-convex in cross section, in male

plants (Figure 6F) ± 220 pm (10 cell rows) high, ± 420 pm

wide, basally (Figure 6G), ± 210 x 400 pm; in female

plants apices (Figure 70) ± 280 pm (11 cell rows) high, ±

450 pm wide, tapering proximally and basally (Figure 7P),

± 150 X 250 pm; dorsally chlorophyllose, ventrally purple.

Rhizoids purple, 10-15 pm wide, often with internal myc-

orrhizal strands. Leaves suberect, overlapping, decurrent,

succubously inserted, apically small, soon becoming larg-

er, shape irregular, often shorter than wide, in male plants

(Figure 6A-E) 625-825 x 825-1200 pm, in female plants

(Figure 7A-L) 8(X)-1375 x 825-1650 pm, mostly some-

30

Bothalia 29,1 (1999)

FIGURE 5. — Fossomhmnia spinosa. A, simple shoot seen partly from side; B, male stem with row of perigonial bracts from above, antheridium

indicated by arrow; C, female stem with row of archegonia from above; D, shoot with pseudoperianth; E, close-up view of pseudoperi-

anth; F, pseudoperianth with capsule (wall disintegrating) from above. A, C-F, S.M. Perold 3834', B, 5. W Arnell 738. A, x 9.6; B, x 19.7;

C, X 18; D, x8.2;E, x22;F, x2l.

what narrower below, ‘mched’ and irregularly lobed above,

with 3 or 4(5) rather angular lobes, 400-500 x 450-500

|jm, margins with (4)5-16 papillae, mostly sessile, some-

times raised on 1 or 2 basal cells. Leaf cells thin-walled, in

male plants not appreciably different from those of female

plants, at upper margin (Figure 7M) subquadrate to rectan-

gular across, 20.0-37.5 x 27.5-42.5 pm, at lateral margins

long-rectangular, 40.0-87.5 x 22.5-32.5 pm; upper lami-

nal cells 5- or 6-sided, 37.5-47.5 x 30-45 pm; middle 1am-

inal cells 42.5-62.5 x 30.0-37.5 pm; basal cells 55-75 x

42.5-52.5 pm. Oil bodies (Figure 7N) 6-27 per cell, shiny,

globose, ± 2.5 pm diam.; chloroplasts numerous, rounded,

± 5 pm diam.

Dioicous. Male plants rare. Antlieridia dorsal on stem,

in 1 or 2 rows (Figure 5B), short-stalked, globose, ± 230

pm diam., each one shielded by a bract (Figure 6H-K),

230-340 X ± 330 pm, sometimes 2 adjacent ones Joined

together, upper margins with pointed projections up to

180 pm long, marginal cells above 27.5-37.5 x

25.0-32.5 pm, cells in interior 4- or 5-sided, 25-45 x

30-35 pm. Archegonia in a row dorsally along stem

(Figure 5C). Pseudoperianth (Figures 5D, E; 7Q, R)

proximal to stem apex, sometimes 2(or 3) close together,

funnel-shaped, raised on a short stalk, ± 500 x 675 pm,

widely flaring above, 1250-1500 pm high, up to 2500

pm wide across convoluted mouth, divided into ± 15

irregularly-shaped projections, some spinous, ± 120 pm

long, others angular, up to 450 pm wide, cells compara-

ble in shape and size to those of leaves. Capsules globose

(Figure 5F), 625-875 pm diam., cells in inner layer of

bistratose capsule wall roughly quadrangular, rectangu-

FIGURE 6. — Fossomhmnia spinosa. A-E, leaves of male plants; F, c/s apex of stein of male plant; G, c/s base of stem of male plant; H-K, perigo-

nial bracts. A-K, 5. W Arnell 735. Scale bars; A-E, .SOO pm; F-K, 2.S0 pm.

Bothalia29,l (1999)

31

FIGURE 7. — Fossombronia spinosa. A-L, leaves of female plant; M, detail of marginal area of leaf, with papilla; N, median leaf cells with oil

bodies (solid lines) and chloroplasts (dotted lines); O, c/s apex of stem of female plant; P, c/s toward base of stem of female plant; Q,

pseudoperianth from side; R, opened pseudoperianth; S, cells in capsule wall with thickenings. A, B, N, Q, R, S.M. Perold 3834\ C-J, O,

S, SM. Perold 3835', K, L, P, Schelpe s.n. Scale bars: A-L, Q, R, 500 pm; M, 100 pm; N, S, 50 pm; O, P, 250 pm.

32

Bothalia29,l (1999)

FIGURE 8. — Fossombroniu spinosa. Spores and elaters. A-C, distal face; D, detail of spines at margin of distal face; E, proximal face; F, elaters.

A, F, S.M. Perold 3H35\ B, S.W. Arnell 735\ C-E, Schelpe s.n. A, x 959; B, x 986; C, x 734; D, x 2294; E, x 674; F, x 863.

lar or irregularly shaped, 32.5-75.0 x 20.0-37.5 pm,

each cell wall with 1 or 2(-4) nodular and only rarely

semi-annular thickenings (Figure 7S). Seta 1. 0-2.9 mm

long, 200-350 pm diam. Spores light brown to darker

brown, hemispherical, 35-45 pm diam., including spines

projecting around periphery; distal face convex (Figure

8 A-C), densely covered with 1 1-15 rows of fine spines,

(or a total of ± 95), almost smooth or faintly grooved, ±

5 pm long, apices quite often truncate (Figure 8D), occa-

sionally ‘broken’ and apparently hollow inside, some-

times linked to form several short ridges; proximal face

(Figure 8E) slightly concave or almost flat, lacking a dis-

tinct triradiate mark, finely to coarsely granular and with

some thin, irregular lamellae, numerous (± 40) spines

projecting around periphery. Elaters (Figure 8F) yellow,

mostly smooth, 100-150 pm long, ± 7.5 pm wide in the

centre and tapering slightly toward tips, ending in loops,

loosely 3- or 2-spiral throughout, or 3-spiral in the mid-

dle and 2-spiral at one or both tips.

The name Fossomhronia spinosa refers to its finely

spinose spores and the species is so far known from only

three localities in the Western Cape (Figure 4), which is

in the winter rainfall area of southern Africa. The holo-

type specimen of F. spinosa and Perold 3834 grow on

pockets of sandy, rather coarse soil derived from sand-

stone, on a vertical rock face, kept damp by seepage

water. They are mixed with Targionia hypophylla, Riccia

species, Gonioniitrium africanum and Bryum species.

Some of Arnell’s collections of this newly isolated

species, 5. Vk Arnell CH4037, 793, from Bath Kloof and

Kogman’s Kloof, Montagu, as well as my own sterile

specimens, S.M. Perold 3453 and 3454 p.p., from Bath

Kloof have regrettably been wrongly identified as F.

montaguensis (.see Table 11). In Perold (1997b: figs 8C,

D; 9A, B), the figures purporting to illustrate F. mon-

taguensis, are actually those of F. spinosa. Arnell 735

had been identified as F. leucoxantha, no doubt on

account of its spinose spores. The Schelpe specimen was

collected at Malmesbury Common on clayey soil more

than 40 years ago. I returned there in 1997, but failed to

find more material of this species. Koekenioer 1415 p.p.

is from the outskirts of Worcester.

Fossonibronia spinosa can be recognised by its rather

small size, frilly, angularly lobed leaves and finely spin-

ose spores, of which some spines coalesce to form short

ridges. F. leucoxantha, also from the Western Cape, has

similar finely spinose spores, but mostly with papillae

interspersed between the spines, and has ‘incised-den-

tate’ leaves (Table 1).

SPECIMENS EXAMINED

Held at PRE, unless otherwise indicated. Bracketed

numbers after collectors’ name and number refer to the

species in the text in alphabetical order, namely: F.

elsiei ( 1 ) and F. spinosa (2).

Arnell 735 (2) BOL; 785 p.p. (2); 793 (= CH4033) (2); 793 (= CH4035) (2)-

CH4037 (2).

Bnt^pns 971354, 97I356A ( 1 ).

Esterhuysen 24885 ( I) BOL (holotype); s.n. (1) BOL.

Koekenioer 1415 p.p. (2).

S.M. Perold 3834 (2); 3835 (2) (holotype). Perold & Koekenioer 3475 ( 1 ).

Schelpe 6368 (\) BOL; s.n. (2) BOL.

ACKNOWLEDGEMENTS

I sincerely thank Dr E.O. Campbell for refereeing this

article and for her helpful suggestions; also my col-

leagues at NBI, i.e. the curator of PRE, Ms M.

Koekemoer for help with fieldwork. Dr H.E. Glen for the

Latin diagnoses, the artist, Ms G. Condy for the draw-

Bothalia 29,1 (1999)

33

ings, the photographer, Mrs A. Romanowski for devel-

oping and printing the photographs and Ms D. Maree for

typing the manuscript. Dr J. Braggins of the University

of Auckland, New Zealand, is kindly thanked for making

available to me the specimens which he collected locally

during his visit here in 1997. The curator of BOL is

thanked for the loan of specimens.

REFERENCES

COWLING, R.M. & HILTON-TAYLOR, C. 1997. Phytogeography,

flora and endemism. In R.M. Cowling, D M. Richardson &

S.M. Pierce, Vegetation of southern Africa. Cambridge

University Press, Cambridge.

COWLING, R.M., RICHARDSON, D.M. & MUSTART, P.J. 1997.

Fynbos. In R.M. Cowling, D.M. Richardson & S.M. Pierce,

Vegetation of southern Africa. Cambridge University Press,

Cambridge.

PEROLD, S.M. 1997a. Studies in the liverwort genus Fossombronia

(Metzgeriales) from southern Africa. 1 . Three new species from

Northern Province, Gauteng and Mpumalanga. Bothalia 27:

17-27.

PEROLD, S.M. 1997b. Studies in the liverwort genus Fossombronia

(Metzgeriales) from southern Africa. 2. An amendment to three

species from Western Cape, described by S.W. Amell. Bothalia

21: 28-38.

PEROLD, S.M. 1997c. Studies in the liverwort genus Fossombronia

(Metzgeriales) from southern Africa. 4. A re-examination of F.

crispa, F. leucoxantha and F. tumida. Bothalia 27: 105-115.

RUTHERFORD, M.C. 1997. Categorization of biomes. In R.M. Cow-

ling, D.M. Richardson & S.M. Pierce, Vegetation of southern

Africa. Cambridge University Press, Cambridge.

SCOTT, G A M. & PIKE, D.C. 1987. Studies on Fossombronia in

Australia II. Fourteen more new species. Journal of the Hattori

Botanical Laboratory 62: 367-386.

SCOTT, G A M. cfe PIKE, D.C. 1988. Revisionary notes on Fossom-

bronia. The Bryologist 9\ : 193-201.

Bothalia 29,1 ; 35-58 ( 1999)

Revision of the genus Faucaria (Ruschioideae: Aizoaceae) in South

Africa

L.E. GROEN* and L.J.G. VAN DER MAESEN*

Keywords: Aizoaceae, distribution. Eastern Cape, ecology, Faucaria, morphology, phylogeny. South Africa, stat. nov., taxonomy

ABSTRACT

Faucaria Schwantes is a small genus of succulents, concentrated in south-central Eastern Cape. Distinguishing features

of Faucaria are the characteristically dotted and toothed leaves and the structure of the seed capsule. A taxonomic review

of the genus is presented, including relationship, diagnostic features, variation, distribution and ecology. The revision

reduces the number of species from 33 to six; F bosscheana, F. felina, F. gratiae, F nemorosa, F. subintegra, and F. tigri-

na. Two new subspecific combinations, F. felina subsp. britteniae and F felina subsp. tuberculosa have been made. A

cladogram is included. A key to the taxa is provided and each species is described and illustrated.

CONTENTS

Abstract 35

Introduction 35

Material and methods 36

Taxonomy 36

Faucaria Schwantes 36

Key to species 37

1. fcoyyc/teanrz (A. Berger) Schwantes 37

2. /e/ma (L.) Schwantes 39

2a. snhsTp. felina 40

2b. subsp. britteniae (L. Bolus) L.E.Groen 41

2c. subsp. tuberculosa (Rolfe) L.E.Groen 42

3. gratiae L. Bolus 42

4. nemorosa L. Bolus ex L.E.Groen 43

5. subintegra L. Bolus 44

6. tigrina (Haw.) Schwantes 45

Cytotaxonomy 46

Character assessment 46

Morphology 46

Stems 46

Leaves 46

Idioblasts 47

Flowers 47

Fruits 49

Seeds 49

Anatomy of epidermis 50

Anatomy of leaves 51

Germination and development 51

Pollination biology 51

Fruit and seed dispersal 51

Cultivation 52

Ecology 52

Distribution and habitat 52

Conservation status 53

Discussion 53

Relationship with other genera 53

Phylogeny 54

Conclusions 55

Acknowledgements 56

References 56

Index 58

* Department of Plant Taxonomy, Botanical Gardens of Wageningen

Agricultural University, P.O. Box 8010, Wageningen, The Netherlands.

MS. received: 1996-02-07.

INTRODUCTION

The earliest record of a Faucaria dates from before

1707, and consists of a watercolour of F. felina subsp.

felina in the Moninckx Atlas (Wijnands 1983). The

Moninckx Atlas is a collection of nine volumes of water-

colours, painted from plants in the Hortus Medicus

Amstelodamensis. Material of this specimen was proba-

bly collected at a location near Algoa Bay, [the present-

day Port Elizabeth region], during a search for fresh

water by a Dutch vessel. Herbarium material from this

period could not be traced. The earliest known herbarium

sheet of F. felina s. 1. was made by Thunberg, who visit-

ed the Eastern Cape on his second journey, from

September 1773 to January 1774 (fuel 1918; Dyer 1937).

After Thunberg, only a few herbarium specimens of this

succulent genus were deposited in herbaria until ± 1925.

Even Ecklon and Zeyher, who collected extensively in

the Eastern Cape between 1831 and 1832, left only a few

herbarium specimens. More herbarium collections were

made between 1930 and 1938 by Mmes W. van

Ryneveld, G.V. Britten, L.L. Britten, I.E. King, Messrs

H.W. James, G.G. Smith, and other (often amateur) col-

lectors. Plants from these collectors were often intro-

duced into living collections, especially those of the then

acknowledged expert on the Mesembryanthema, Louisa

Bolus in Kirstenbosch and Rondebosch.

Haworth (1803) subdivided Mesembryanthemum and

placed the two species known at that time, M. tigrinum

and M. felinum, later considered to be species of

Faucaria, in the section Denticulata. Both species recur

in the new section Ringentia (Haworth 1812) together

with some species now placed in the genus Stomatium.

Schwantes (1926) erected Faucaria, accommodating M.

bosscheanum, M. felinum, M. lupinum, M. tigrinum and

M. tuberculosum. Field collections during the nineteen-

thirties enabled Louisa Bolus to describe 25 species

between 1933 and 1938 from living material. Her species

concept was very narrow, and often based on minor dif-

ferences. With the exception of F. subintegra these

species turned out to be synonyms of already known

Faucaria species. Hartmann placed Faucaria in the

Stomatium group, one of the 1 1 groups in the

Ruschioideae (Hartmann 1991). This group is charac-

36

Bothalia29,l (1999)

terised by homocellular xeromorphic leaves with crystals

in irregular zones, flowers with five separate nectaries,

capsule with or without covering membranes, without

closing bodies, and with valve wings broad to reduced.

MATERIAL AND METHODS

Live material of more than 60 field collections were

studied over the period 1985-1992. An extensive field

study in 1991 completed the taxonomic research. SEM

studies of epidermis, pollen and seed of fresh material were

carried out with a JEOL JSM-5200. This material was

dried using a Balzers Union CPD 020 critical point dryer

with carbon dioxide and subsequently sputter-coated with

platinum using a Polaron Sputter Coating Unit E5100. The

morphological studies were made with a Zeiss Axiophot

photomicroscope. Germination experiments with seeds of

1 1 accessions harvested in the wild were carried out in Petri

dishes, placed in a germination incubator for 12 hours at

25°C, and 12 hours at 20°C. The material for microtome

sectioning was embedded in Kulzer’s Technovit 7100 and

stained with 1 % Toluidine blue in 1 NHCl. A 4% solution

of formalin was used to examine the idioblasts (Hartmann

1977). The presence of tannins was established by blue

colouring of tannin with a 1% EeC13 solution (Hartmann

1977). Cytotaxonomic studies were done on root tips of

germinating seeds, stained with Feulgen reagent. Besides

the living accessions, 300 herbarium sheets were studied

from B, BOL, G, GRA, HBG, K, MO, NBG, PRE, SAM

and WAG. A cladistic analysis was carried out with all

eight taxa of Faucaria using Orthopterum L. Bolus as the

outgroup. Thirteen characters were investigated (Table 1 ).

The most parsimonious cladogram was determined using

Hennig86 (Farris 1988).

TAXONOMY

Faucaria Schwantes in Zeitschrift fur Sukkulenten-

kunde 2: 176 (1926); Schwantes; 105 (1927); N.E.Br. et

al.: 212 (1931); L.Bolus: 22 (1933a); H.Jacobsen: 129

(1933); Poelln.: 41 (1933); L.Bolus: 109 (1937); L.Bolus;

1 14 (1938); E.Phillips: 320 (1951); Schwantes: 20 (1952);

H.Jacobsen: 1362 (1955); Schwantes: 129 (1957);

H.Jacobsen; 422 (1970); Herre: 156 (1971); H.Jacobsen:

1141 (1978); H.Jacobsen: 454 (1981); F.MeIntosh: 167

(1989). Type: Faucaria tigrina (Haw.) Schwantes 3: 105

(1927).

Mesembryanthemum L.: 487 (1753) p.p.; Haw.: 161

(1794/1795); DC.: t. 152 (1805); Haw.: 216 (1812); Haw.:

89 (1821); Haw.: 109 (1824); DC.: 419 (1828); Salm-Dyck;

t. 5, 6 (1836); Salm-Dyck; t. 2 (1854); Sond.: 388 (1862);

A.Berger: 265 (1908).

Prelinnean: Mesembryanthemum rictum felinum reprae-

sentans, Dill.: 240, t. 187, fig. 230 (1732); L.: 218, No.

12 (1738)[1737].

Plants perennial, forming clumps or tufts, 20-100

(-150) mm high and up to 200 mm across; stems some-

times developing with age; thick, turnip-shaped taproots

sometimes present; isophyllous. Leaves succulent, 2-4

crowded pairs at top of stem, fused at base, covering

short intemodes, upper side flat, smooth or rough, ovate-

rhomboid to linear-spatulate, underside rounded in lower

part, keeled and 3-angled in cross section towards tip,

leaf margins in upper half with 0-12 cartilaginous, often

bristle-tipped teeth on each leaf margin; epidermis

homocellular xeromorphic, often with whitish dots.

Flowers solitary, terminal; pedicel flattened, (0-)l-4(-6)

X 2. 5^.0 mm, 0.4-1. 5 mm thick, not scented; receptacle

2-keeled, sometimes slightly constricted at apex, base

broadly bell-shaped, often flattened, top with 5 or 10

grooves, in middle usually raised; bracts absent. Sepals

(4)5(6), unequal, 2 outer sepals keeled with small dorsal

subapical appendage, 3 inner sepals not keeled, all free to

base. Petals many, free, in 3 or 4 series, linear, narrowed

to base, on inside shining yellow, outer petals dull yel-

low, rarely white or reddish. Nectaries free, crest-like.

TABLE 1 . — The characters (1-13), character states and coding in the cladistic analysis of Faucaria. The first state of a character is coded 0, the

second 1, the third state 2

1 = Some plants stem-forming with age/stem-forming absent

2 = Keel of leaves sharp/rounded at top

3 = Whitish leaf margins not present/not continuously/continuously

4 = Teeth less than 4/usually more than 5

5 = Bristles shorter than 2 mm/sometimes/always longer than 2 mm

6 = Bristles not papillate/sometimes/always papillate

7 = Whitish dots on upper surface absent except on margins/dots always present

8 = Whitish dots on upper surface small/large dots

9 = Epidermis smootb/somewhat uneven/rough

10 = Cuticular folds absent/not very developed/well-developed cuticular folds

1 1 = Petals fewer than 1 OO/sometimes/always more than 100

12 = Nectaries greenish or ochreous/brownish

13 = Tumble fruits sometimes present/absent

Bothalia 29,1 (1999)

37

(4)5(6), yellow, greenish to dark brown. Staminodes

absent. Stamens numerous, 7-11 mm long; filaments

sparsely papillate at base, yellowish, at base whitish;

pollen yellow. Gynoecium inferior, (4)5(6) fused carpels

with parietal placentation. Style absent; stigmas (4)5(6),

filiform, erect, in medium-aged flowers about as long as

stamens, top usually curved, yellowish, with elongated

blunt papillae ± 10-16 pm wide on inside. Capsule

hygrochastic, woody, long-persistent, sometimes break-

ing off and forming tumble fruits, brownish, obovate,

usually laterally slightly compressed; forming a flattened

to nearly semiglobose top, flattened to globose in lower

part, often flattened on one side; valves when closed,

with small slits between them, each concave, ascending

to form a central peak, expanding keels surrounded by

broad, expanding sheets, ending in a short awn and

extending into locules; covering membranes absent;

closing bodies absent; stiff, small valve wings present;

fruit stalk flattened, 0-4(-6) x 2. 5^.0 mm, 0.4-1. 5 mm

thick. Seeds 60-100(-200) per capsule, broadly ovate to

tetrahedral, dark-brown, 1.4-1. 7 x 0.9-1. 4 mm, covered

with warts. Chromosome number: 2n = 18 (Moore

1973).

The generic name is derived from faux (= jaw, throat);

Faucaria - a collection of jaws, likening the toothed

leaves to the open jaws of an animal.

Diagnostic features

Leaves usually toothed, rarely toothless, dispersed

calcium oxalate crystal concentrations usually visible as

small dots. Flowers opening during late afternoon, clos-

ing at sunset; petals 1.0-1.25 mm wide. Fruit locules

hidden by split septa (resembling covering membranes).

Covering membranes and closing bodies absent. Valve

wings stiff, straight and standing erect over locules in

open fruits.

Key to species

la Upper leaf surface shiny green, usually without white dots, with conspicuous, often continuous, white cartilagi-

nous margins and keel, with 0-3 white cartilaginous teeth without papillae 1 . fi bosscheana

lb Upper leaf surface rarely shiny green, with white dots, without continuous, white cartilaginous margins, teeth

absent to many, usually partly greenish:

2a Leaves with a blunt keel, only present at tip, dots minute, rare or absent, teeth and bristles very reduced to absent;

fruits often firmly attached:

3a Leaves ovate-rhomboid in upper part, epidermis hard, somewhat glossy, green to yellowish, teeth absent or coni-

cal, 1-2 mm long, usually not bristle-tipped; sepals thicker than wide 5. K subintegra

3b Leaves oblong-lanceolate in upper part, epidermis soft, dull greyish green, with (0-) 2-4 teeth, reduced to

small whitish tubercles, shorter than 1 mm; sepals longer than wide A. F. nemorosa

2b Leaves often with sharp keel, present from halfway up, distinct dots usually present, teeth and bristles often

well developed; fruits often easily detached (tumble-fruits);

4a Stem at base obscured by leaves which have numerous white distinct dots, confluent into patches, teeth well

developed, ending in long bristles 6. F. tigrina

4b Stem often leafless at base, dots on leaves present, not confluent into patches, teeth poorly to well developed,

with or without long bristles:

5a Leaves olive-green to reddish, margins irregularly undulate, large-lobed or obscurely toothed with up to 5

small teeth, usually ending in short bristles, 1-3 mm long 3. F. gratiae

5b Leaves mostly greyish green, margins regularly few- to many-toothed, very rarely absent, teeth well devel-

oped, usually ending in short or long bristles;

6a Leaves with some to many whitish tubercles on upper surface 2c. F. felitia subsp. tuberculosa

6b Leaves without tubercles on upper surface, some tubercle-like teeth on margins may be present:

7a Leaves fresh green to greyish green but not glaucous, dots clearly visible, minute dots with (0-)3-8(-10)

teeth 2a. F. felina sahsp. felina

7b Leaves partially or completely whitish or glaucous, caused by very numerous, barely visible dots, with

6-10(-13) teeth 2b. F. felina subsp. britteniae

1. Faucaria bosscheana (A.Berger) Schwantes in

Zeitschrift fiir Sukkulentenkunde 2: 111 (1926);

L. Bolus: 153 (1930); L. Bolus: 22 (1933a); H. Jacobsen:

129 (1933); L.Bolus: 110 (1937); H.Jacobsen: 1363

(1955); Schwantes: 131 (1957); H.Jacobsen: 423 (1970);

H.Jacobsen: 1141 (1978); H.Jacobsen: 454 (1981);

F.McIntosh: 167 (1989); S. A. Hammer: 1, 3 (1991).

Type: De Wildeman: 108, fig. 230 (1908) (neo.!, here

designated).

Mesembryanthemum bosscheanum A.Berger; 269 (1908).

F albidens N.E.Br.; 349 (1927); L.Bolus: 153 (1930); H.Jacobsen;

1362 (1955); H.Jacobsen: 423 (1970); H.Jacobsen: 1141 (1978);

H.Jacobsen: 454 (1981); F.McIntosh: 166 (1989). Type: J. Frith s.n. (K,

lecto.!, here designated).

F. haagei Tischer: 20 (1929); G.D.Duursma; 128 (1930). F. boss-

cheana var. haagei (Tischer) H.Jacobsen in H.Jacobsen & G.D. Rowley:

80 (1955); H.Jacobsen: 1363 (1955); H.Jacobsen: 423 (1970);

H.Jacobsen; 1142 (1978); H.Jacobsen: 455 (1981). Type: Tischer: 21

(1929), unnumbered plate (lecto.!, here designated).

F. paucidens N.E.Br. et ah: 210 (1931). Type: N.E.Br. et ah: 210, fig.

100 (1931) (holo.!).

F. kendrewensis L.Bolus: 469 (1934). Type: Kendrew, Van der Berg

NBG1619/34, fl. 28 April 1934 (BOL, holo.l).

F. peersii L.Bolus: 108 (1937). Type: near Klipplaat, Peers BOL2I979,

fl. 15 March 1937 (BOL, holo.!).

Plants small, clumped or tufted (Figure lA). Leaves

4-8 per rosette, crowded, thick ovate-lanceolate or lin-

ear-spatulate, 15^0(-50) x 5-15(-25) mm, margins

undulate or with 0-3(-5) short whitish stout teeth on

each leaf margin, without or with short stout smooth bris-

tles; margins and keel with almost semiglobose epider-

mal cells; leaf surface smooth, often without obvious

white dots, numerous small, dark green subepidermal

idioblasts visible, wax rugose; leaf margins and keel at

least partially whitish cartilaginous, glossy or shiny fresh

green, turning reddish in very sunny conditions. Flower

with 70-90 petals; stamens 120-180. Nectaries largely

38

Bothalia29,l (1999)

FIGURE I. — Faucaria species. A, F. bosscheana. Van Jaarsveld 1 1 120, B, F. felina subsp. britteniae. Van Jaarsveld 11106, pi. culta; C, F.

ffratiae. Van Jaarsveld IIIIO, pi. culta; D, F subintef>ra, Groen I082\ E, F. tif>rina, Groen & Marx I052b\ F, F. nemorosa. Van

Jaarsveld NBG87H/H5. Scale bars: A-F, 10 nun.

Bothalia 29,1 (1999)

39

greenish. Fruit stalk mostly absent or (0.5-)2.8-5.8 x

2. 0- 2. 6 mm, 0.7- 1.4 mm thick. Fruits often easily

falling off (tumble fruits), 7.0-11.3 x 6.5-12.0 mm,

7.0- 10.5 mm thick; lower half of fruit mostly flattened at

one side, or more rarely convex on 2 sides, valves on top

of fruit bent downwards in the middle forming a depres-

sion there. Seeds 1.35-1.6 x 0.9-1.45 mm. Flowering

period'. Feb.-July, main period March.

Distribution', western part of the Eastern Cape, wide-

spread in scattered populations (Figure 2A).

Ecology, restricted to Karoo Biome, altitude 750 m

and up, open vegetation, flat terrain, north-facing

foothills, in flat cavities of rock plates, between stones on

slopes. Rainfall: 200 mm (arid Karoo) to about 350 mm

(False Karroid Broken Veld), mainly in March and

November. Geological formation: often sandstone of

Beaufort Group, Adelaide Subgroup; Dwyka tillite,

pH(H20) 7.5.

F. bosscheana was named after Van den Bossche of

Tirlemont (Belgium), in whose garden the first plants

were grown from seed received from H. Bolus, Cape

Town in December 1904.

The descriptions of Berger and De Wildeman have been

based on two different plants: De Wildeman sent seedlings

to Berger in la Mortola (Italy), whose description (Berger

1908) predates the description of De Wildeman (1908) by a

month. Berger did not conserve herbarium specimens of F.

bosscheana. Both descriptions indicate a form with narrow

leaves, F. albidens, which occurs around Graaff-Reinet and

Kendrew. The type specimen of F. albidens is very similar

to the picture of the De Wildeman specimen. The confusion

of what should be regarded as F. albidens is already noted

by Bolus (1930). She noted that the chief difference

between F. albidens and F. bosscheana seemed to be the

length of the stamens. The broad- leaved form is common in

cultivation and has therefore been regarded by growers as

the real F. bosscheana. All listed synonyms of F. boss-

cheana are based on characters such as leaf form and num-

ber of teeth. These characters are quite variable and may

vary from koppie to koppie according to Hammer (1991).

They have in common the smooth epidermis, often without

conspicuous white dots, the few-toothed, uniform green to

brownish coloured leaves and (not) continuous white mar-

gins. F. haagei was a cultivated variant of F. bosscheana,

characterised by dark green broad leaves, undulating or

often entire leaf margins. F. paucidens, like F. albidens, is a

narrow-leaved form of F. bosscheana, described from a

plant in cultivation. According to Hammer this form varies

considerably around Graaff-Reinet. F. peersii is a normal

variant of F. bosscheana, not distinct at specific level.

Specimens examined

WESTERN CAPE. — 3222 (Beaufort West): Beaufort West, (-BC),

Perold NBG2657/35 (BOL); ibid.. Grant 1374ft (PRE); Otzen

NBGI153/36 (BOL). 3322 (Oudtshoom): Prince Albert, (-AA), J.A.C.

Kaorts BOL46534 (BOL).

EASTERN CAPE. — 3123 (Victoria West): Murraysburg, (-DD),

Hart. Stell. Univ. Garden s.n. (BOL). 3224 (Graaff-Reinet): Graaff-

Reinet, (-BC) Van Jaarsveld 11170 (NBG); 30 km from Pearston and

Jansenville, Smith 7251 (BOL); Graaff-Reinet Dist., (-BC),

Barkhuizen BOL46594 (BOL); Graaff-Reinet, (-BC), Luckhaff

BOL46596 (BOL); near Oatland Railway station, 24 km north of

Klipplaat, (-CD) Marx 202 (WAG); Kendrew, (-DA), Peers BOL46595

(BOL); ibid.. Cook & Long s.n. (BOL); dry plains, near Graaff-Reinet,

H. Bolus 574 (BOL); Karoo Nat. Park, Graaff-Reinet, Groen 1025,

1027, 1035 (GRA, WAG). 3323 (Willowmore): Willowmore, (-AD),

Hort. Stell. Univ. Garden 9337, 9861 (BOL); 29 km from Willowmore,

on road between Willowmore and Uniondale, (-CA), Mrs N.R. Urton

s.n. (BOL). 3324 (Steytlerville): Klipplaat, between Miller and Mount

Steward, (-AB), Long s.n. (BOL); Klipplaat Dist., Peers BOL46537

(BOL); Baroe station, near Klipplaat, (-AB), Stayner BOL22191

(BOL); Stevtlerville, (-AD), Hort. Stell. Univ. Garden 8659 (BOL).

2. Faucaria felina (L.) Schwantes in Zeitschrift fur

Sukkulentenkunde 2: 177 (1926); L. Bolus: 22 (1933a);

H. Jacobsen: 129 (1933); L. Bolus: 110 (1937);

H. Jacobsen: 1364 (1955); Schwantes; 131 (1957);

H. Jacobsen: 423 (1970); H. Jacobsen: 1143 (1978);

H. Jacobsen: 455 (1981); Wijnands: 148 (1983); Van

Jaarsv.: 53 (1984). Type: Dill.: 240, t. 187, fig. 230

(1732) (lecto.!, here designated).

FIGURE 2. — Distribution of Faucaria in South Africa: A, F. bosscheana', B, F. felina s.l. : F felina subsp./e/inrt, •; F. felina subsp. britteniae, O.

C, F. gratiae', D, F. nemorosa', E, F. subintegrw, F, F. tigrina.

40

Bothalia29,l (1999)

2a. subsp. felina

Mesembrycinthernum ringens L. H felinum L.: 487 (1753). M. felimim

(L.) Weston: 172 (1770); Lam.: 486 (1788); Haw,: 161 (1794/1795);

Haw.; 31 (1803); DC.: t, 152 (1805); W.T.Aiton: 218(1811); Haw.: 216

(1812); Haw.: 89 (1821); DC.: 419 (1828); G.Don: 128 (1834); Eckl. &

Zeyh.: 308 (1834-1837); Salm-Dyck: t. 6 (1836); Sond.; 397 (1862);

A.Berger: 267 (1908); N.E.Br.: 76 (1920).

M. lupinum Haw.: Ill (1824). F. lupina (Haw.) Schwantes: 176

(1926). Type: watercolour, Bowie s.n., in 1823 (K, lecto.!, here desig-

nated).

F. militaris Tischer: 280 (1932). Type: Tischer: 280 (1932), unnum-

bered plate (lecto!, here designated).

F. duncunii L. Bolus; 22 (1933a). Type: Port Elizabeth Dist., Coega

Kop. W Duncan BOL19645 (BOL, holo.!).

F felina (L.) Schwantes var. jamesii L. Bolus; 395 (1933b). Type:

Somerset East Dist., near Halesowen, alt.l 200-1 525 m, James 8, fl.

3-02-1933, (BOL, lecto.!, here designated).

F. aciitipetala L. Bolus: 443 (1934). Type: Somerset East Dist.,

Bruintjeshoogte, W. van Ryneveld NBG686/34, fl. 21 Feb. 1934 (BOL,

holo.!).

F. cradockensis L. Bolus; 448 (1934). Type: Somerset East Dist.,

Cradock, B. Rogers NBG687/34 (BOL, holo.!).

F. crassisepala L. Bolus: 442 (1934). Type: Somerset East Dist.,

Bruintjeshoogte, W. van Ryneveld NBG2352/33, fl. 22 April 1934

(BOL, holo.!).

F. jarnesii L. Bolus ex Tischer: 99 (1934) nom. nud.

F. latipetala L. Bolus: 443 (1934). Type: Fort Beaufort Dist., W. van

Ryneveld NBG686/34. fl. 14 April 1934 (BOL, holo.!).

F. laxipetala L. Bolus: 444 (1934). Type: Grahamstown Dist., near

Grahamstown, comm. Long & Cook NBG286/33, (BOL, holo.!).

F. longidens L. Bolus; 470 (1934). Type: Somerset East Dist., Yester

Farm, W. van Ryneveld NBG2355/33, fl. 7 July 1934 (BOL, holo.!).

F longifolia L. Bolus: 471 (1934). Type; Fort Beaufort Dist., W van

Ryneveld NBG690/34, fl. 3 May 1934 (BOL, holo.!).

F. montana L. Bolus: 448 (1934). Type: Somerset East Dist., hill near

Cradock, H.W. James NBG685/34, fl. 2 April 1934 (BOL, holo,!).

F multidens L. Bolus: 450 (1934). Type: Fort Beaufort Dist., W. van

Ryneveld NBG141 2/33, fl. 5 April 1934 (BOL, holo.!).

F. ryneveldiae L. Bolus; 446 (1934). Type: Fort Beaufort Dist., W. van

Ryneveld NBG688/34, fl. 31 March 1934 (BOL, holo.!).

F. Candida L.Bolus: 105 (1937). Type: Somerset East Dist., Scanlan,

near Cradock, Mrs C.H. Mclane NBG65I/37 (BOL, holo.!).

F. kingiae L.Bolus: 106 (1937). Type; Grahamstown Dist., near

Alicedale, I.E. King 15 (BOL, holo.!).

F. multidens L.Bolus var. paardepoortensis L.Bolus: 107 (1937).

Type: Port Elizabeth Dist., Klein Winterhoek Mountains at

Paardepoort, I.E. King 4, fl. 4 May 1937 (BOL, holo.!).

F. plana L.Bolus: 107 (1937). Type: collector and locality unknown,

BOL2I978, fl. 1 May 1937 (BOL, holo.!).

F. uniondalensis L.Bolus: 106 (1937). Type: Willowmore Dist., near

Uniondale, E. Markoetter SUG 10662, fl. 24 March 1936 (BOL,

holo.!).

Plants clump-forming or shrubby, short-stemmed

with age. Leaves spreading, crowded, 4-8, variable,

ovate-rhomboid to linear-lanceolate, 15-70 x 5-15(-20)

mm, keeled; keel at an oblique angle with tip of leaf,

with (0-)3-8(-10) teeth on each margin; teeth 0.5-3. 0 x

1—4 mm, usually bearing more or less papillate bristles of

l-5(-6) mm long, sometimes one or more small teeth

present on keel, rarely teeth with very broad base up to 4

mm broad, between teeth furrows often visible; leaf sur-

face nearly smooth to rough from a variable number of

small epidermal whitish dots (Figure 3), emerging from

subepidermal crystals, sometimes confluent into larger

flecks present on leaf margins and keel; epidermis with

cuticular folds, wax cover thin; light green-olive, dark

green, grey-green, or glaucous. Flowers yellow, white-

flowering plants occasionally found; petals 100-130,

(1.0-)1.25 mm wide, acute; stamens 200-250. Nectaries

variable in colour, greenish, yellowish, yellow-brown,

brownish. Fruit stalk absent or 1-4 x 2.5— 4.0 mm,

0.4-1 .5 mm thick. Fruits easily falling off, top of capsule

variable in shape, lower part of capsule often flattened on

one side, (5.0-)9.0-13.5 x (5.0-)7. 5-12.0 mm and

5. 0-8. 5 mm thick. Seeds 1.4-1. 6 x 1.0-1. 3 mm.

Flowering period: March-August, main flowering peri-

od March to June. Differences in flowering date between

locations in the large area of distribution are small. In

several cases it has been noticed that the plants start to

flower after rain.

Distribution: Eastern Cape; the area between Graaff-

Reinet, Cradock, Bedford, Uitenhage, Oudtshoorn,

Uniondale and the coast. Large populations occur near

Cradock, in the valleys of the Sundays, Swartkops and

Fish Rivers, around Port Elizabeth and Addo (Figure 2B).

Ecology: (open) Grassland, Karoo, Valley Bushveld.

Rainfall: from about 200 mm or less (Karoo) to 700 mm

(Valley Bushveld, Grahamstown). Geological fonnation;

quartzite sandstone of Witteberg Series, Kommadagga

Subgroup (Cape system); shale Karoo Succession,

Beaufort Group; tillite Dwyka formation; sandstone of

Uitenhage Group, pH (H2O) 6. 3-7. 5.

Faucaria felina s.l. is variable and is distributed over

a vast region in the centre of the area of the genus. The

majority of the diagnostics between Faucaria segregates

were based on local, variable forms of this species, as

already mentioned. Judging from the available field data

it is impossible to maintain them as separate taxa. Plants

FIGURE 3. — Faucaria felina subsp. felina. Van Jaarsveld 1 1120, pi.

culta, papillae and calcium oxalate dots above subepidermal

idioblasts clearly visible. Scale bar: 5 mm.

Bothalia29,l (1999)

41

with long leaves and many thin teeth grow in more

humid conditions, usually within the boundaries of the

Tongaland-Pondoland phytochorological regions. Plants

with shorter, stout leaves and teeth, described as F. felina

s.stK, grow in the harsher conditions of the dry Karoo-

Namib phytochorological region.

After the publication of a watercolour of F. felina in

the Moninckx Atlas before 1707 (Wijnands 1983),

Boerhaave (1719) published a short description that

Martyn (1728) used in his more detailed paper together

with an excellent watercolour of F felina subsp. felina.

Linnaeus (1738, 1753) published the species referring to

Dillenius (1732). Very variable in their vegetative char-

acters, most described Faucaria ‘species’ are to be con-

sidered as mere populations and fall into synonymy with

F felina s.l. Details are discussed with the synonyms.

Faucaria felina subsp. tuberculosa has been regarded

as a distinct species up till now. However, this taxon dif-

fers from F felina only in the tubercles on the leaves. As

such F. tuberculosa can not be classified as a species on

the tubercles only and should be regarded as a subspecies

of F felina. Some succulent lovers will have problems

with the rejection of F ryneveldiae and F. longifolia as

separate species. The long, narrow, few-toothed leaves

seem distinct. However, this rare leaf form occurs on

widely diverging localities, e.g. in Addo {Liebenberg

7697). As genetically isolated populations develop the

same leaf type, it is still hard to describe them as a single

taxon, as F. longifolia and F. ryneveldiae differ in other

characters. The ovary top of the type specimen of F.

longifolia is hollow, whereas that of F. ryneveldiae is

acute. Intermediates with rather long leaves and more

teeth also occur. Further research must indicate whether

F. ryneveldiae should remain a synonym of F. felina s.l.

or regain species status. With some hesitation F. lupina is

regarded here as conspecific with F. felina. The differ-

ences are small, the leaves are greener, slender, more

glossy, and the teeth normally have rather long bristles.

The number of teeth is variable, from 0-10. In the notes

of Haworth (1824), F. lupina was described as having

longer and narrower glaucous leaves, and with longer

and twice as many teeth as F. felina. Salm-Dyck (1836)

gives a more detailed description and says that this plant

differs from F. felina by having greener, shiny leaves,

without points and more teeth with bristles. The teeth on

the keels described for F. militaris occur occasionally in

F. felina.

F. felina subsp. felina grows together with F. boss-

cheana in a large area. However, they do not grow sym-

patrically, F. bosscheana prefers a more karroid area than

F. felina. Intermediate populations are short-leaved, and

the leaves are partly white margined and have a smooth

epidermis. The leaves bear 3-6, long-needled teeth.

Specimens examined

WITHOUT LOCALITY : Cape of Good Hope, Thunberg sheet 12009

(UPS-Thunb., microfiche).

EASTERN CAPE. — 3224 (Graaff-Reinet): between Graaff-Reinet

and Beaufort, Ecklon & Zeyher 1983 (SAM); Halesowen, James

BOL22728, BOL24959 (BOL); near Jansenville, (-CC), King 56

(BOL). 3225 (Somerset East): Somerset East, (-BA), Ferguson s.n.

(BOL); Somerset East, near Cradock, (-BA), James BOL21977,

BOL22725 (BOL); between Cradock & Bedford, (-BC), James

BOL22193, BOL22194. BOL46646. BOL46647 (BOL); Mortimer,

(-BC), Groen 1106 (WAG). 3226 (Fort Beaufort): Bedford, (-CA), Van

Ryneveld s.n. (BOL); Farm Richfontein, near Yester, (-DC), Van

Ryneveld BOL46708 (BOL). 3322 (Oudtshoorn): (-CA), s.c.,

BOL46579 (BOL). 3323 (Willowmore): 8 km NE Uniondale, (-CA),

Markoetter s.n. (BOL). 3324 (Steytlerville): Brakfontein, (-BB), Van

Ryneveld BOL46711 (BOL); Sandpoort, Springbokvlakte, (-BD), Marx

207 (WAG). 3325 (Port Elizabeth): 4 km south of Lake Mentz, (-AA),

Marx 210 (WAG); road to Jansenville, about 8 km beyond Glenconnor,

(-AC), King 3 (BOL); ± 32 km from Uitenhage on mountain road to

Steytlerville, (-AD), King 12 (BOL); near Bluecliff Station, (-AD),

King 62 (BOL); Addo Bush, (-BC), King 6 (BOL); Addo Drift, (-BC),

King 8 (BOL); 48 km from Port Elizabeth on Uitenhage-Steytlerville

road, (-BC), Stayner BOL22189 (BOL); hills ± 8 km south of Selbome,

in Sundays River Valley, (-CA), King 9a. b (BOL); ‘am

Zwartskoprivier Uitenhage’, (-CB), Ecklon & Zeyher 1981 (SAM);

Swartskoprivier, (-CB), Zeyher 2580 (B, G); hills north of Uitenhage,

(-CB), Muir 3999 (K); 19 km out of Port Elizabeth on Uitenhage road,

(-CD), King 2 (BOL); Soutpoort, near Addo, (-DA), Groen 1092

(WAG); near Addo, (-DA), Long & Cook NBG282/33 (BOL);

Coegakop, (-DA), King 1 (BOL); Coega, G.V Britten s.n. (BOL);

Perseverance, (-DC), Groen 1119 (WAG); New Brighton, (-DC),

LBolus BOL46580, BOL46581 (BOL, HBG); Port Elizabeth, (-DC),

Dahlstrand 1884 (STE, MO); near Port Elizabeth, (-DC), King 52

(BOL). 3326 (Grahamstown): near Swartwaterpoort, (-AA), King 41,

42 (BOL); ibid.. Van Jaarsveld 11142 (NBG); near Middleton, (-AB),

Van Ryneveld BOL22205 (BOL); Tootabie, (-AC), Groen 1080

(WAG); near Alicedale, (-AC), King 16, 17, 39, 43 (BOL); Eagles

Craig, near Alicedale, (-AC), King 35 (BOL); near Grahamstown,

(-BC), G.V. Britten BOL21986 (BOL).

2b. subsp. britteniae (LBolus) L.E.Groen, stat. nov.

Faucaria britteniae L. Bolus in South African Gardening & Country

Life 23: 22 (1933a); H.Jacobsen: 1363 (1955); H.Jacobsen: 423 (1970);

H.Jacobsen: 1142 (1978); H.Jacobsen: 455 (1981); F.McIntosh: 167

(1989). Type: Grahamstown Dist., on the road to Committees, 20 miles

from Gr^amstown, LL. Britten 5395, coll. 12 July 1926 (BOL,

holo.!).

F. coronata L. Bolus: 446 (1934). Type: Grahamstown Dist., between

Carlisle Bridge and Riebeeck East, G.V. Britten 34, fl. 15 April 1934

(BOL, holo.!).

F. speciosa L. Bolus: 445 (1934). Type: Grahamstown Dist., near

Grahamstown, LL. Britten & A.H. Harcourt-Wood NBG691/34, fl. 10

April 1934 (BOL, holo.!).

F. smithii L. Bolus: 32 (1936). Type: Albany Dist., Fish River valley,

near the foot of Pluto’s Vale, about 20 miles E of Grahamstown, G.G.

Smith 90, fl. 15 March 1936, (BOL, holo.!).

F. grandis L. Bolus: 125 (1938). Type: Albany Dist., exact locality

unknown, probably coll, by G.V. Britten BOL21986, fl. 28 March 1936,

(BOL, holo.!).

Plants clump-forming or shrubby, stem-forming with

age; stems up to 120 mm long (Figure IB). Leaves spread-

ing, ovate-rhomboid, 30-55(-70) x 12-33 mm, keeled;

keel sharp, forming a pronounced chin in almost rectan-

gular arrangement as seen from side, toothed; 6-10(-13)

teeth on each margin, 3-5 mm broad at base, 2.5-5. 0 mm

long normally with long, papillate bristles; epidermis with

well-developed wax cover, thicker than in subsp. felina]

uniform or partially whitish green with numerous small

greyish white dots, or dull green with white clearly visible

dots. Flowers large for genus, 60-80 mm diam.; petals

125-160; stamens 260-350. Nectaries mostly yellowish.

Fruit stalk absent or (0.5-)2.2-3.8(-6.5) x 2.3-6. 5 mm,

0.50-1.4 mm thick. Fruits easily falling off (tumble

fruits), 7-13 X 10.5-13.0(-15.0) mm, 7-10 mm thick.

Seeds 1.4—1. 6 X 1.0-1. 3 mm. Flowering period: March-

August, occasionally throughout the year.

42

Bothalia29,l (1999)

Distribution: northeast of Grahamstown, Fish River

Ridge, northern hills on foot of mountain range at

Riebeeck East (Figure 2B). It occurs near the areas

where F. felina subsp. felina and F. tigrina are found.

Intermediate populations with F felina subsp. felina are

found at Fish River Ridge, north of Grahamstown. These

plants are robust, have few but large teeth with bristles,

and the whitish sheen of F. felina subsp. britteniae.

Ecology: (dense) Valley Bushveld, Noorsveld, Arid

Savanna, open karroid Valley Bushveld, on slopes

between stones. F. felina subsp. britteniae is found in

open rocky spots in dense thicket, in open grassy spots or

under small bushes with a preference for north-facing

slopes. Rainfall; 350-500(-700) mm. Geological forma-

tion: Karoo System, Ecca Series, tillite; Dwyka Series;

Bokkeveld Series, pH (H2O) 6. 7-7. 7.

Faucaria felina subsp. britteniae is unique in the

genus because of the (partially) whitish grey epidermis.

This taxon deserves the rank of subspecies by the pres-

ence of numerous very small dots on the leaves, papillate

bristles and restricted area of distribution. Additional fea-

tures are the large number of teeth and the size. The most

characteristic specimens with many, but rather small

teeth are found at Ecca Pass, north of Grahamstown.

Specimens examined

EASTERN CAPE.— 3226 (Fort Beaufort): Kat River hills, 15.5 km

south of Fort Beaufort, (-DC), Marx 196 (WAG). 3326

(Grahamstown): Signalkop, near Carliste Bridge, (-AA), Van

Jaarsveld 11090 (NBG); Buffelsdrif, (-AA), Van Jaarsveld 11095

(NBG); Fish River Ridge, (-AB), G.V Britten BOL22120, BOL46640,

BOL46641 (BOL); Hellskloof, (-AB), G.V Britten s.n. (BOL); Hell’s

Poort, (-AB), R.A. Dyer s.n. (K); Pigots Bridge, (-AB), Smith 1992

(BOL); Clifton Farm, (-BA), Groen 1061 (WAG); Botha’s Hill, Ecklon

& Zeyher 1982 (SAM); near Grahamstown, L.L. Britten s.n. (BOL);

Ecca’s Pass Nature Reserve, (-BA), Gwen 1057 (GRA, WAG); near

foot of Pluto’s Vale, (-BA), Smith 598 (BOL, K); Committees Flats,

G. V. Britten s.n. (BOL); Uplands, Kransdrift, (-BA), Groen 1071

(WAG); Fort Brown, (-BA), A. Walton BOL22195, BOU6557 (hOL):

32 km from Grahamstown on road to Committees, (-BB), G.V. Britten

s.n. (BOL); near Committees Hotel, (-BB), Smith 5456, 5457 (BOL).

3327 (Peddie): between Chalumna and Committees, (-AB), Herre s.n.

(BOL).

2c. subsp. tuberculosa (Rolfe) L.E.Groen, stat. nov.

F. tuberculosa (Rolfe) Schwantes in 21eitschrift fiir Sukkulentenkunde

2: 177 (1926); L.Bolus: 382 (1930); N.E.Br. et al.: 212 (1931);

H. Jacobsen: 130(1933);L.Bolus; 110(1937); H.Jacobsen: 1369(1955);

Schwantes: 131 (1957); H.Jacobsen: 424 (1970); H.Jacobsen: 1146

(1978); H.Jacobsen: 456 (1981); F.McIntosh: 167 (1989). Type: Rolfe

s.n., aquaprint Curtis’s Botanical Magazine t. 8674b (1916), (holo.!).

Mesemhryanthemum tuherculosum Rolfe: t. 8674b cum fig. (1916).

Plants clumped or single. Leaves spreading, 4—8 per

rosette, crowded, thick, ovate-rhomboid, 15-20 x 15-20

mm, keeled, with 3-6 short, whitish stout teeth on each

margin, almost without bristles, some teeth may be rudi-

mentarily developed; surface with many epidermal

whitish dots, sometimes confluent on leaf margins and

keel; upper surface with blunt, whitish tooth-like tubercles

(this may vary from a single tubercle, to, very rarely, a

complete tuberculate surface). Flowers yellow, as F. feli-

na. Stigmas 5, filiform, free to the base. Nectaries yellow-

brown, large. Fruits and seed as in F. felina subsp. felina.

Distribution: Bedford surroundings; field data very

scarce (Figure 4).

Ecology: grassland, often camouflaged by grasses in

association with many herbs. Rainfall: 493 mm.

Geological formation; Beaufort group, Adelaide sub-

group (Karoo Succession).

As Bolus (1930) already noticed, the aquaprint of the

united stigmas in Curtis’ Botanical Magazine is not cor-

rect, in Faucaria these are free to the base.

At a glance F. felina subsp. tuberculosa can be identi-

fied by the tubercles on the leaves which are unique in

the genus. A typical plant is compact, with whitish dots

and patches and resembles F tigrina. Cultivated hybrid-

ised plants often show tubercles more prominent than in

wild material.

Specimens examined

WITHOUT LOCALITY: photo original plant, Oct 1921 (BM); N.S.

Pillans 281 sin. loc. (SAM); Hort. Pillans s.n.. May 1915 (BOL); N.E.

Brown s.n., Oct. 6 1913 (K).

EASTERN CAPE. — 3226 (Fort Beaufort): Bedford, (-CA), Van

Ryneveld BOL46630 (BOL); Normandale, south of Bedford, Van

Jaarsveld 11115 (NBG); ibid., Groen 1118 (WAG); Adelaide, (-CB),

Van Ryneveld s.n. (BOL); (-CB), W.E. Armstrong BOL46691 .

BOL22190 (BOL); Kat River Hills, (-CD), ± 64 km from

Grahamstown and 5 km from the Kat River, G.G. Smith 10 (BOL).

3326 (Grahamstown): 32 km west of Grahamstown (-AA), on road to

Bedford, G.G. Smith 710 (BOL); between Fort Beaufort and

Grahamstown, A.P.J. Kluth BOL46628 (BOL).

3. Faucaria gratiae L.Bolus in Notes on Mesem-

bryanthemum and allied genera 2: 394 (1933b);

H.Jacobsen: 1365 (1955); H.Jacobsen: 423 (1970);

H.Jacobsen: 1 143 (1978); H.Jacobsen: 455 ( 1981 ). Type:

Albany Dist., Mitford Park, near Riebeeck East, G.V.

Britten 35, fl. 23-3-1933, (BOL, holo.!).

E hooleae L.Bolus: 447 (1934), Type: Albany Dist,, Mitford Park,

near Riebeeck East, Hoole & G.V. Britten NBG684/34, fl. 10 April

1934 (BOL, holo.!).

Plants clump-forming, very short-stemmed (Figure

1C). Leaves erect, ovate-lanceolate, 21.5-35.0 x

10.0-16.5(-20.0) mm; leaf margins irregular, sometimes

undulating or with (0-)2-5(-7) obscure, normal to

FIGURE 4. — Distribution of Faucaria felina subsp. tuberculosa.

Bothalia 29,1 (1999)

43

coarse teeth, without or with few bristles of 0. 1-^.0 mm

long; shiny green, often with a reddish lustre, base red-

dish, with many small white crowded dots, margins and

keel whitish. Flowers yellow; sepals 5, 8-10 mm long,

base 2. 5-5.0 mm broad, unequal; petals 70-84, emar-

ginate to acuminate, 15.0-21.0 x 1.0-1.75 mm; stamens

8-9 mm long; filaments white. Nectaries brown-orange.

Stigmas 8-11 mm long, shorter than longest stamens.

Fruit stalk absent or up to 1 x 3 mm wide, 1 mm thick.

Fruits sessile, when closed with rounded top, 0.65-9.0 x

0.5-0. 8 mm, 0.6-0. 9 mm thick. Seeds 0.85-1.0 x

1.15-1.30 mm. Flowering period: March-May.

Distribution: restricted to a radius of 3 km around

Riebeeck East (Figure 2C).

Ecology: Grassy Fynbos with forest groups. The ecol-

ogy of the plant is peculiar: without shelter in compact

clumps in sandy soil in cracks and hollows of flat sand-

stone plates. Rainfall: about 380 mm per annum.

Geological formation: sandstone. Karoo System,

Witteberg Series, Kommadagga Subgroup.

F gratiae and F. hooleae are known only from three

herbarium sheets. The descriptions do not differ signifi-

cantly, the type specimens originate from the same pop-

ulation. The plants are very small with short leaves and

irregular margins, rather different from those of F.felina.

Van Jaarsveld (1990) visited the locality and rediscov-

ered these plants. I examined this rare species at the same

location in 1992, and collected it, 3 km further away.

Specimens examined

EASTERN CAPE. — 3326 (Grahamstown): Mitford Park, near

Riebeeck East, (-AA), G.V. Britten s.n. (BOL); ibid.. Van Jaarsveld

lino (NBG); Palmietfontein, near Riebeeck East, Groen, Brink &

Marx 1068 (WAG).

4. Faucaria nemorosa L.Bolus ex L.E.Groen in

Succulenta 73: 39 (1994). Type: Fort Beaufort Dist., near

Alicedale, Swartwaterpoort, Archibald 960, 29-7-1937

(BOL, holo.!).

Plants stem-forming, branching from base, forming

small clumps, (Figure IF). Stems up to 150 x 7-13 mm,

internodes 5-10 mm long. Leaves spreading, 3-5 leaf

pairs together per branch, crowded, connate at base,

oblong-lanceolate, 25^5 x 9-14 mm, slightly triangular

in upper part, with leaf sides often unequal, keeled

towards apex; keel often asymmetric, flat on face, mar-

gin each with (0-)2-4 reduced teeth, smaller than 1 mm,

bristles 0. 1-1.0 mm long or absent; epidermis dull grey-

green, soft, without any white dots or flecks, numerous

small subepidermal idioblasts visible as dark green dots.

Flowers yellow; sepals 5, 8-12 mm long, at base

0.3-0.65 mm wide; petals 70-90, conspicuously nar-

rowed downwards, some of innermost row very small,

linear, (15-)20-25 x (1.0-)2.0-2.3 mm in upper part, at

base (O.l-)0. 2-0.5 mm wide, yellow, top emarginate or

obtuse, some very small ones acute. Receptacle 8-9 x

7-9 mm. Stamens 150-200, 5-8 mm long; anthers yel-

low, 1 mm long. Nectaries olive-green, smooth. Ovary

top 5-lobed, almost flat but very slightly raised in mid-

dle, stigmas 5, filiform, 10 mm long, in young flowers

shorter or as long as stamens, in old flowers longer. Fruit

stalk 1-2 X up to 4 mm, 1 mm thick. Fruits tightly

attached, dark brown, 9.5 x 7-8 mm, 5-7 mm thick,

valves bend to a central depression. Seeds 1.3- 1.5 x

0. 9-1. 0 mm. Flowering period: June-September.

Distribution: from a single locality near Alicedale

(Figure 2D) in the Eastern Cape.

Ecology: hills, (300-)600-900 m altitude. Fish River

Scrub in which Portucalaria afra and small succulents

are dominant. F. nemorosa is solitary or grows in small

groups among a F felina subsp. felina population on a

hill (E. van Jaarsveld pers. comm.). Rainfall: 350-400

mm, mainly in summer months. Geological formation:

quartzite sandstone, Dwyka Series, Karoo System.

When verifying the Faucaria sheets from BOL, three

sheets could not be determined satisfactorily. The speci-

mens looked similar to F. subintegra but their leaves

were different. They originate from a single locality near

Alicedale, and have been collected between 1937 and

1939 by Mrs E.E.A. Archibald and Grace Britten, both

from Grahamstown. Louisa Bolus grew field accessions

in her garden, and recognised these specimens as some-

what different. She made an additional note on the sheet:

‘may be only a variety of F subintegra'. She referred to

the habitat by her note on the sheet Faucaria nemorosa,

1. e. Faucaria from the forest. She did not publish this

name. The live collection of the Kirstenbosch Botanic

Garden includes a plant, collected by Ernst van Jaarsveld

in 1985, very much like the Archibald specimen in the

Bolus Herbarium. This peculiar plant bears soft dull

greyish green leaves without any sign of whitish dots,

and the teeth are small and indistinct. Like the herbarium

specimens of F nemorosa the leaves are similar to those

of F. subintegra, but longer and narrower. Moreover this

plant has been collected in the same area as the original

F. nemorosa. The scarce, conserved fruiting material was

of the normal Faucaria type, but not like those of

Orthopterum. Unfortunately the plant could not be traced

again in 1991, even after repeated efforts.

Study of the living material confirmed some relation-

ship between F. nemorosa and F. subintegra. The reduc-

tion of the teeth in F. nemorosa is almost complete, being

reduced to small tubercles. The fruits are similar, rather

small, tightly attached on the plants and often broader

than long. The scarce ecological data on F. subintegra

indicate differences in habitat between the two taxa. F.

subintegra is restricted to shallow hollows or crevices in

bare sandstone rocks in full sun under very dry circum-

stances. F. nemorosa grows in soil in the understorey of

shrubs in partial shade. SEM micrographs of F. nemorosa

show cuticular folds which are also found in F. subinte-

gra (Figure 5D), a common feature for Faucaria, that is

missing in Orthopterum (Figure 5H).

Specimens examined

EASTERN CAPE. — 3326 (Grahamstown): near Alicedale,

Swartwaterpoort, (-AA), G. Britten .v.n. (BOL); ibid.. Van Jaarsveld

NBG878/85 (WAG!).

44

Bothalia29,l (1999)

FIGURE 5, — Scanning electron micrographs of leaf epidermis. A, B, F. bosxcheana, Bayer 4965: A, epidermis above showing rather even

surface with small dispersed wax particles; B, under surface with a ribbed stnicture of the cuticle and few wax particles. C, F. feli-

nu. Van Jaarsveld 6H97, under surface with cuticular folds. D, F. suhintegra. Van Jaarsveld 6950, under surface with coarse ribbed

structure. E, F, F. ligrina, Mullins s.n. : E, under surface with raised dots and stomata between them; F, upper surface with well-devel-

oped cuticular folds. G, F. hosscheana, Bayer 201 1 h, margin with semi-globose epidennis cells. H, Orthopterum waltaniae. Hammer

s.n., under surface without cuticular folds and with large flakes of wax. Scale bars; A, D, E, 100 pm; B, C, E G, 50 pm; H, 10 pm.

5. Faucaria subintegra L.Bolus, Notes on

Mesembryanthemum and allied genera 2: 449 (1934);

H. Jacobsen; 1368 (1955); H. Jacobsen: 424 (1970);

H. Jacobsen: 1146 (1978); H. Jacobsen: 456 (1981);

F.McIntosh: 167 (1989). Type; Peddie Dist., Chalumna

River Bank, J.I. Browlee s.n., fl. 12 April 1934 (BOL,

lecto.!, here designated).

F. suhindurata L.Bolus: 444 (1934). Type; Peddie Dist., near Peddie,

G.G. Smith 102, 0. 12-4-1934 (BOL, holo.!).

Bothalia29,l (1999)

45

Plants clump-forming, in older stages stem-forming;

stems 4-7 mm diam. (Figure ID). Leaves 4-8, crowded,

very thick, spreading, ovate-rhomboid, 20-35 x 10-15

mm, bluntly keeled at top, margins with 0-8 very short

conical teeth having short bristles on each side that dis-

appear in older leaves; often shiny, glaucous, sometimes

with a violet lustre, with minute white dots, margins and

keels indistinctly whitish. Flowers with 80-92 linear

petals; stamens 150-180. Nectaries yellowish green.

Fruit stalk absent or 2-A x 2-4 mm, 1.0-1. 4 mm thick.

Fruits tightly attached on plant, 7-10 x 9-12 mm, 6-9

mm thick; top of capsule flat, or in most fruits a raised

centre. Seeds dark brown, 1.1-1. 4 x 1.0-1. 1 mm.

Flowering period'. March-May.

Distribution', the most south-central part of the

Eastern Cape (Figure 2E).

Ecology. Subtropical Coast, Open Valley Bushveld

with grassveld in between, overgrazed, in full sun on

open places on top of north-facing river bank, cavities in

rock plates in flat sandstone rock under very dry circum-

stances. Rainfall: 500-900 mm (Valley Bushveld) or

more (Subtropical Coast). Geological formation:

Sandstone of the Beaufort Group, Adelaide Subgroup

(Karoo Succession).

F. subintegra possesses leaves with indistinct teeth

and often rounded leaf margins and keel. The epidermis

is smooth with indistinct small whitish dots. F.

subindurata was described from the same area in the

same publication, and is very clearly conspecific. The

two names therefore have the same priority but the name

F. subintegra is chosen here because it better describes

the almost entire leaf margins in older leaves.

Intermediate plants have been found between F. felina

subsp. britteniae and F. subintegra. The leaf shape was

that of F. subintegra, the number and shape of the teeth

those of F. felina subsp. britteniae. It is unknown if these

intermediate populations are the result of natural hybridi-

sation.

Specimens examined

EASTERN CAPE. —3327 (Peddie): Wooldridge, (-AB), King 76

(BOL); a couple of km west of Wooldridge, (-AB) Smith 3146 (BOL);

Chalumna River, (-BA), G.V. Britten s.n. (BOL); East London, (-BB),

collector unknown, SUG10855 (BOL); Farm Kamaskraal, King 53

(BOL); Chalumna River bank, (-BA), Gwen 1082 (WAG); Chalumna

River bank, near King Williams Town, (-CD), Henderson 2041 (NBG).

6. Faucaria tigrina (Haw.) Schwantes in Zeit-

schrift fiir Sukkulentenkunde 2: 177 (1926); L. Bolus: t.

267 (1927); G.D.Duursma: 128 (1930); N.E.Br. et al.:

211 (1931); L.Bolus: 22 (1933a); H.Jacobsen: 130

(1933); L.Bolus; 110 (1937); H.Jacobsen; 1369 (1955);

Schwantes: 129 (1957); H.Jacobsen; 424 (1970); Herre:

156 (1971); H.Jacobsen: 1146 (1978); B.RBarkhuizen:

126 (1978); H.Jacobsen; 456 (1981); J.A.Retief; 23

(1989); F.McIntosh: 167 (1989); S.A. Hammer: 4 (1991).

Type; Edwards, 1817, t. 260, unnumbered plate, (1817)

(neo.!, here designated).

Mesembryanthemum tigrinum Haw.: 164 (1794/1795); Haw.: 31

(1803); W.T. Alton: 218(1811); Haw.: 216(1812); Edwards: t. 260 cum

fig. (1817); Haw.: 89 (1821); DC.: 419 (1828); G.Don: 128 (1834);

Salm-Dyck: t. 5 (1836); Sond.: 397 (1862); Eckl. & Zeyh.: 308

(1834-1837); A.Berger: 266, fig. 57 (1908).

F. tigrina (Haw.) Schwantes forma splendens H.Jacobsen & G.D.

Rowley: 81 (1955); H.Jacobsen: 1369 (1955); H.Jacobsen: 424 (1970);

H.Jacobsen: 1146 ( 1978); H.Jacobsen: 456 (1981). Type: not designated.

F. tigrina ‘Superba’ nom. nud., Weber: 12 (1968).

Plants forming compact clumps, stem-forming absent

(Figure IE). Leaves very crowded, pressed against each

other, more or less erect, ovate-rhomboid in upper half,

lowest half of leaves square, sharply keeled at top,

30-40(-50) X 15-20(-25) mm, toothed; teeth on leaf mar-

gins (5-)9-10(-12) with long bristles, pointed backwards;

grey-green, turning to bluish purple, margins and keel

whitish; epidermis with numerous clearly visible white

dots, confluent into larger flecks, often arranged in curved

lines towards keel, minute white dots absent. Flowers with

100-120 petals, linear, acute, 1.0-1.25 mm wide; stamens

250-300. Nectaries brownish, surface often 3-ribbed.

.Fruit stalk absent or (0.5-)2.5-3.8 x 3. 0-4.0 mm, 0.9-1. 6

mm thick. Fruits coming loose rather easily, top of fruit

flattened to slightly spherical, 7. 0-9. 5 x 8.0-10.0(-12.0)

and (6-)8-9 mm thick. Seeds 1.15-1.50 x 1.0-1.25 mm.

Flowering period: March-May.

The species was discovered as early as 1790 (Bolus

1927), probably because of its accessibility. Faucaria

tigrina is a compact, crowded, distinctly flecked plant

having 9-10 distinct teeth with long bristles on the short

leaves. The many white-coloured flecks on the often red-

dish leaves resemble the lichens and reddish rocks in its

natural surroundings. Some populations of F. felina

subsp. britteniae at Hell’s Poort, 35 km west of

Grahamstown and near Fort Beaufort, look like F. tigri-

na because of the numerous large flecks on the leaves

and the long bristles. However, the leaves of F. felina

subsp. britteniae are larger, not pressed against each

other and the number of teeth is reduced to 5 or 6. Forma

splendens, published by Jacobsen & Rowley in 1955,

was recognised on the basis of its reddish leaves, typical

for some populations. F. tigrina ‘Superba’ is a superflu-

ous synonym, used in seed catalogues to differentiate

good plants and seed, reintroduced by Mrs Dudley

Ryder, from the usually inferior hybrids named F. tigrina

(Weber 1968).

Distribution: narrow endemic at the edge of

Grahamstown (Figure 2F). These populations are endan-

gered because of cultivation and urbanisation (Retief

1989; Van Jaarsveld 1990; Groen 1991; Chan 1992).

Ecology: Fynbos, Renosterbosveld on open rocky

patches. Rainfall: 681 mm per annum (Grahamstown).

Geological formation; sandstone, Dwyka Series, Karoo

System, pH(H20) 4. 7-5. 7.

Specimens examined

EASTERN CAPE. — 3326 (Grahamstown): Gowieskloof, near

Grahamstown, (-BC), Gowie s.n. (K); ibid. G.V. Britten 36 (BOL);

ibid.. Van Jaarsveld 11104 (NBG); ibid., A.J. Mullins comm. E. Brink

(GRA); Burnt Kraal, near Grahamstown, (-BC), Groen & Marx 1052b,

1054 (GRA, WAG).

46

Bothalia29,l (1999)

CYTOTAXONOMY

The chromosome number of Faucaria is usually list-

ed as 2n = 18 (Moore 1973). The chromosome number of

the studied material of all species was also 2n = 18. The

chromosomes are however difficult to study, and this

number is therefore based on a few plates only.

CHARACTER ASSESSMENT

The classification of Faucaria is based on a combina-

tion of variable characters (Table 1 ). There could be even

more variation within taxa than between taxa. These diffi-

culties are reflected in the key; it proved quite difficult to

construct a usable one. In this treatment, considerable

importance is awarded to epidermis characters, being one

of the few fairly usable characters. Other characters such as

number of teeth or leaf shape proved to be too variable for

making reliable taxonomic decisions. Cladistic analysis is

also hampered by this lack of usable information and must

be regarded as uninformative. Despite these shortcomings,

the authors find this monograph important as the first tax-

onomic treatment of this horticulturally important genus.

Morphology

Stems

Stems of Faucaria taxa are short, erect or creeping, or

may be virtually absent. The stems increase in length, to

a maximum of about 150 mm long and up to 10 mm

thick. The woody stems show a limited secondary anom-

alous growth and cork development. The internal struc-

ture consists of successive rings of meristem in the

phloem or pericycle. Cork develops in the inner part of

the cortex or inside the endodermis. The stems are

branched, so that small clumps develop. The leaves at the

top of the branches cover the stem completely, whereas

older parts of the stems are mostly leafless.

Leaves

The leaves of Faucaria show a distinct interspecific

variation, whereas the variation in separate populations of

one species is usually smaller. In the pre-juvenile stage,

all species lack flecks and teeth, are fresh green, often

with white margins. In the juvenile stage the subsequent

leaves gradually develop the characteristics of the adult

leaves (Figure 6A). In the adult stage the successive

leaves are similar. In conditions of poor light the plants

stay longer in the pre-juvenile and juvenile stage than in

full sunlight. The ontogeny of F. hosscheana is very short,

compared to the other species; it reaches the flowering

stage while still in possession of juvenile leaves; the

neoteny of this species could be initiated by the very dry

circumstances of its habitat, which tend to induce a short-

ening of its development. It has been noticed that extreme

conditions promote the development of taxa with

neotenic character states (Ihlenfeldt 1971a).

The shape of adult leaves may vary from ovate-rhom-

boid to ovate-lanceolate, depending on species, popula-

tion and environment. Individual populations could be

recognised in some cases on the basis of leaf characters.

FIGURE 6. — A, Seedling of F. bosscheana, Marx 202, cotyledons still

visible and 2 pairs of leaves developed, 252 days after sowing;

B, F. felina subsp. britteniae, Groen 1058, fruit, central part

raised; C, F. tif’rina, Groen 1054, central part of fruit not raised;

D, F felina subsp. felina, 84BG4I408, flowerbud just before

anthesis; E, F. felina subsp. britteniae, Groen 1057, epidermis

of under surface of leaf, superficial stoma surrounded by cutic-

ular folds, wax particles dispersed. Scale bars: A, D, 1 mm; B,

C, 5 mm; E, 10 pm.

Leaves exposed to full sun are shorter, wider, more

intensely coloured and flecked than leaves sheltered

from the sun. Old withered leaves turn brownish, red-

dish, greyish to blackish. Plants of Faucaria are isophyl-

lous. All leaves are narrow at the base, flat on the upper

side, with the lower side keeled and triangular in cross

section towards the top. The keel becomes rounded

lower down so that towards the base the leaf is hemi-

spherical. Towards the base the leaves are fused for a

short distance and consequently the internodes are hid-

den by a leaf sheath. The leaf margins are often sharp at

the apex, a mucro is absent on the ± rounded leaf tip. The

differences in leaf form can be remarkable (Huber 1943;

Bolus 1937), this taxonomically variable character

proves, however, to be useful in the key.

The leaves of Faucaria very often bear teeth on the

leaf margins in the upper part. Their development is

simultaneous with that of the leaf, but very young leaves

bear relatively large teeth. These teeth develop succes-

sively, the differences in size could be large in young

stages; e.g. in a leaf primordium of 1 .5 mm length with

the largest tooth at 1 .3 mm, the ratio with two other teeth

Bothalia29,l (1999)

47

is 16:3; 1. In later stages the differences between the teeth

become gradually smaller. The teeth are emergences of

the leaf because deeper tissue layers contribute to the for-

mation of the projections. The teeth consist of epidermal

and hypodermal cell layers that are characterised by

strongly thickened collenchymatic or sclerenchymatic

cell walls. Some populations have leaves with small,

poorly developed teeth on the upper side, the so-called

tubercles. These usually functionless tubercles may be

present on the margins or elsewhere on the upper side.

Plants with tubercles on the upper surface have often

been cultivated under the name F. tuberculosa. In the

ontogeny, teeth appear in the 3rd or 4th leaf pair. In adult

leaves the teeth are most prominent in the young stage,

normally decreasing in size with age. The number and

size of teeth vary with each taxon, growing circum-

stances, and even the population.

The teeth often bear bristles, depending on the taxon.

Certain taxa like F. tigrina always possess bristles,

whereas others, like F. bosscheana, usually lack them. A

strong development of bristles seems to be correlated

with well-developed oxalate dots in the epidermis. The

presence of papillae on the teeth is also variable, they are

only numerous on the teeth of F. tigrina. In F. felina s.l.

the teeth are smooth to moderately papillate, some acces-

sions are very papillate, whereas other populations are

hardly or not papillate. F. bosscheana, F. gratiae, F.

nemorosa and F. subintegra show no or only a few papil-

lae.

Idioblasts

Idioblasts occur on the stem, leaves and receptacle of

Faucaria (Figure 7A, B). They are roundish, spherical or

elongated. The elongated idioblasts in the lower part of

the leaves and the central tissue are orientated in the

growth direction of the leaves. The subhypodermal

idioblasts are visible as longitudinal or roundish dots in

the epidermis. They are very often obscured by calcium-

oxalate dots that develop above the idioblasts. They may

or may not contain bundles of needle-like calcium oxalate

raphides. The idioblasts vary considerably in size: small

roundish raphide idioblasts are about 80-185 pm where-

as only liquid-containing idioblasts may vary between

100 and 650 pm. The most and largest idioblasts may be

found at the strongly cutinized edges. Subepidermal

idioblasts contain mostly tannin-like substances,

idioblasts in the central tissue are often tannin-free. Dark

brown-coloured tannin idioblasts occur especially in old

leaves. From the outside they can be localised as small

dark brown or blackish dots. It could not be established

whether the percentage of tannin-like substances in

decayed leaves is higher than in younger leaves.

Flowers

Flower shape and colour within Faucaria is not very

variable, and only some differences in petal and stamen

number can be observed between species. Variation in

colour in the nectaries has some taxonomic value as was

suggested by Schwantes (1957). But even within popula-

tions the colour can vary between green and brown. The

single flower (Figure 6D) is a termination of the shoot.

produced by the more developed of the two buds of the

distal leaf pair. Sometimes both buds become fertile or

stay sterile. The flower is produced on the topmost inter-

nodium (homologous to the flower pedicel), resulting in

(almost) sessile flowers. The study of Troll & Weberling

(1981) of F. felina proves that the reduction of the stem

region is important, resulting in the absence of bracts.

The free sepals often show membraneous margins.

The calyx consists of five sepals of which the two outer

sepals are leaf-like (Ihlenfeldt 1960, 1971a; Haas 1976).

The three-keeled inner sepals are produced by the edge of

the receptacle. Difference in thickness of the sepals is of

some taxonomic importance. The two keeled outer sepals

are slightly longer than the inner three and bear small dor-

sal subapical appendages, homologous with the protrud-

ing keel of the leaf (Kaussmann & Schiewer 1989).

The yellow linear-lanceolate petals are whitish at the

base, dull yellow below, sometimes reddish at the tip,

outer surface reddish, especially in fading flowers. Very

occasionally a white-flowering mutant can be found

among many yellow-flowering ones. One of these

mutants has been described as F. Candida. The yellow

colour originates from 90-95% dopaxanthin and free

lemon-yellow betalamic acid (Hofmann 1973; Reznik et

al. 1988). The petal number depends on the species and

size of the flower and varies between 70 and 160 (Figure

8). Staminodes are absent. The 100-350 erect stamens

fill the centre of the flower, providing abundant pollen.

The presentation of large amounts of pollen and the hid-

den nectaries characterise the pollination type of

FIGURE 7. — Subepidermal idioblasts in leaf section. A, B F. subinte-

gra, Van Jaarsveld 6950. The dark coloured tannin contents

have disappeared in some cases during preparation. Scale bar;

160 pm.

48

Bothalia29,l (1999)

0 30 60 90 120 150 1 80

numbers of petals

FIGURE 8. — Variation in numbers of petals of Faucaria species.

Faucaria as melittophilous (Vogel (1954). Because of

the flat, open flower and the stamen-filled centre, the

flower subtype could be classified as stamen carpet

flower (Vogel 1954; Hartmann 1991).

The anthers consist of thin-walled cells, usually weak-

ly papillate at the base. The papillae are hair-like projec-

tions from the basal cells, which may occur higher up.

Variation in papillae occurs and in some flowers almost

epapillate stamens have been found. The stamens vary in

length from 7 to 1 1 mm, depending on their position, the

longest are placed at the outside of the cone, the inner-

most are the shortest. The stamens are colourless except

for a somewhat yellowish apex. Pollen grains are spheri-

cal-subprolate, 28-32 pm diam., 3-colpate (Figure 9A,

B); with microreticulate, microspinulose tectum. Our

FIGURE 9. — Scanning electron

micrographs of F. felina

subsp. felina. Van Jaarsveld

6897. A, B, pollen grain: A,

polar view; B, equatoral

view. C, valve wings have

been dissected from valves in

background for a view of

open fruit; split septa with

their hollows clearly visible.

Scale bars: A, B, 10 pm; C, I

mm.

Bothalia 29,1 (1999)

49

observations do not indicate any differences between

taxa. The style is absent, usually five stigmas are present,

free to the base. On the inner side they bear elongated

blunt papillae of about 10-16 pm wide, which may form

small bunches. The stigmas continue to grow during

flowering, at anthesis they are shorter than or as long as

the stamens, in old flowers they overtop them. The stig-

mas can be curled or recurved, especially in old stages.

The green to brown nectaries of Faucaria vary in size,

and form, even within species. They may be button-like,

elongated, crest-like, smooth or ribbed. Mostly they are

slightly less then 1 mm long. Nectaries in F. bosscheatia

and F. subintegra are button-like, 1 mm or less long and 0.5

mm wide, with a smooth or slightly uneven surface. Those

of F. hooleae are smooth, and hutton-like to elongated. The

nectaries of F. felina s.l. vary from smooth button-like to

elongated or crest-like. The surface may be smooth to

ribbed. The variation is clearly depicted in the watercolour

drawings of F ryneveldiae. One drawing shows smooth,

crest-like nectaries, the other drawing, button-like, smooth

ones. The nectaries of F tigrina are ribbed and elongated.

The receptacle is 2-keeled and sometimes slightly

constricted at the apex, and may form a short tube up to

1 mm long. The top of the inferior ovary is flattened to

conical, with 5 grooves, sometimes with another 5 alter-

nating less prominent grooves. Bolus (1934, 1937)

emphasised especially the number of these grooves in

Faucaria and the slope of the tissue between them to the

centre. The middle of the ovary itself is usually elevated

for 1-2 mm. A considerable variation exists in the con-

struction of the top of the ovary, and may even vary

between flowers of the same plant.

Fruits

The top of the ovary, and immature and mature

Faucaria fruits vary in shape between the taxa (Bolus

1937). The variation in shape of the top of the ovary and

the immature fruits is so large that this character is not

used in this study. The mature fruits show less variation

among the taxa and is included only in the species

description.

In the developing fruit, parenchyma tissue develops

from the valves downwards, and splits the lamellae of

the five septa. The top of the ripening fruit is often coni-

cally swollen by this developing parenchyma tissue, the

grooves are well developed at this stage. In ripe capsules

the parenchyma has disappeared and only five empty

hollows remain of the pulp masses, so that, together with

the real locules, the fruit seems to be 10-celled (Figure

9C). At first sight the capsule seems empty, because only

the hollows are visible, but in fact the locules are hidden

by the bent lamellae. The ripe capsule is 5-(6)-locular,

woody, persistent, light brown to dark brown, with a flat-

tened or subglobose base, and a flat to globose or raised

top (Figure 6B, C). The closed valves may be bent

upwards or show inwardly folded margins, leaving small

fissures between them. The deep locules are covered by

the split and bent cell lamellae (Figure lOB). Ordinary

covering membranes and closing bodies are absent. The

combined closing mechanism of deep locules and cell

lamellae is very effective, since very old fruits normally

still contain some seed for late distribution. The expand-

ing keels end in an awn. The two expanding keels of one

valve are lateral on each side of the valve, parallel with

those of the neighbouring valve. The valve wings are

stiff and straight, and when the fruits are fully opened,

they stand over the locules. The capsules of Faucaria are

variable in opening, some fruits open rapidly within min-

utes, whereas others need half an hour or more.

Seeds

A capsule contains 60-100(-200), broadly ovate, dark

brown, warty seeds of 1.0-1. 5 mm diam. Taxonomically

important differences between the seeds could not be

observed. The exotesta is more or less uniform, however,

a longitudinal row of elongated cells on the rear is

absent. The warts on the seeds exist on the papillae of the

exotesta and are common in the Ruschioideae. Ehler &

Barthlott (1978) found with the aid of the scanning elec-

tron microscope that the warts of Faucaria are covered

with an epicuticular dimorphic layer of erect rodlets. One

FIGURE 10, — Fruit of A, Orthopterum waltoniae, Gwen 1090; B, Faucaria tigrina, Groen 1054. cb, closing body; EK, expanding keel; L, lamel-

la; LO; locule; S, seeds (lined); V, valve; VR, valve membrane; VW, valve wing. Scale bars: A-C, 1 mm.

50

Bothalia 29,1 (1999)

layer consists of rodlets of about 6 x 1 |um. On and

between these rodlets other small, often bent rodlets are

present that measure 1^ x 0.4 pm. They also found

small differences in the shape of these rodlets between F.

tigrina and F. felina subsp. tuberculosa. From SEM

preparations it could only be concluded that slight differ-

ences between the species exist, but these do not seem to

be of any taxonomic use (Figure llA-F).

Anatomy of epidermis

The wax cover on the epidermis may occur as small

particles of different size (Figure 5A, B), but is obscured

by the cuticular folds of the epidermis. Form, structure and

occurrence of the cuticular folds of 21-36 pm wide

depends on the position on the leaf and on the taxon. The

folds may form cobweb-like structures in combination

with small epidermal elevations of the whitish dots

(Figure 5E). A more ribbed structure can be found in

species without pronounced elevated dots such as F. subin-

tegra or F bosscheana. The most developed cuticular fold

structures can be found in species with strongly oxalate-

encrusted epidermis, like F. tigrina. The cuticular folds are

generally less developed on the upper surface than the

lower surface of the leaves. A flake-like wax cover is

strongly developed in the related Orthopterum waltoniae,

where the cuticular folds are absent (Figure 5H).

The epidermis of Faucaria is of the xeromorphic type

(Ihlenfeldt & Hartmann 1982; Ihlenfeldt 1983) and is

encrusted with calcium-oxalate crystals in the outer

walls. Strongly encrusted parts of the epidermis are visi-

ble as whitish dots and flecks of 0.15-5.0 mm or more

diam., very often developed above subhypodermal tan-

nin idioblasts. The outer walls of the epidermal cells in

these dots are dome-like and 16-26 pm thick with a thick

layer of oxalate crystals, compared to the 8-13 pm thick

outer epidermis of the surrounding epidermal cells.

These dome-like epidermal structures may be further

developed in papillae, of 20-150 x 20-50 pm that are

located on margins, keels and teeth of the leaves (Figure

5G). Their presence is however variable (see description

of leaf). Besides these large papillae, small club-shaped

papillae occur on the epidermis, which are about 7. 1-9. 1

X 2. 8-3.9 pm in size. The outer wall of the strongly

encrusted epidermis cells reaches to about half-way

down the tangential walls, the thickness may vary from

2-4 pm. The inner walls of the epidermis of these cells

are 3-6 pm thick, in less encrusted cells 2 pm.

The development of the epidermal calcium-oxalate

crystal dots, flecks or zones differs from species to

species and may be restricted to the leaf margins, as in F.

bosscheana, or even absent in some populations of F.

felina subsp. /e/ma. The crystal concentrations are slight-

ly elevated, giving the leaves a pattern of depressions

and elevations. These elevations are also partially due to

the presence of subepidermal idioblasts in the mesophyll

(Figure 3).

The stomata on upper and lower leaf surfaces gener-

ally occur in relatively less oxalate-encrusted parts

between the strongly oxalate-encrusted elevated dots.

FIGURE 1 1 . — Scanning electron mi-

crographs of seeds. A, B, F.

lif’rinu, G.V. Britten .t.n.

(BOL); C, F. felina subsp.

felina, James R (BOL); D, F.

felina subsp. felina, N.S.

Pillans NBG446/I6 (BOL);

E, F. bosscheana, Bayer 2395

(BOL); F, F. subintegra,

NBGW46/33 (BOL). Scale

bars: 100 pm.

Bothalia 29,1 (1999)

51

The stomata occur regularly in these areas, whereas their

density is very variable, ranging from 6-128 per mm^

The anisocytic and paracytic stomata are level with the

surrounding epidermal cells. These superficial stomata

(Figure 6E) occur also in other genera with a xeromor-

phic epidermis e.g. Nananthiis and Rhombophyllum

(Ihlenfeldt & Hartmann 1982). The guard cells measure

22-37 X 6-12 pm. The stomata may be surrounded par-

tially or completely by cuticular folds of the neighbour-

ing cells. These projections partially form a secondary

outer surface. The epidermal protection of the stomata of

Faucaria may be absent or almost sunken in the cuticu-

lar folds and/or protected by the elevated leaf surface

(Figure 5A-F). This type of differentiation was described

earlier from the genera Odontophorus, Octopoma and

some species of Cheiridopsis (Ihlenfeldt & Hartmann

1982). Instead of surface sculpturing in the form of

papillate epidermis cells like these taxa, Faucaria has

cuticular folds.

Anatomy of leaves

The leaves possess a large central vascular bundle,

occasionally accompanied by some parallel lateral vas-

cular bundles. A ring of lateral vascular bundles is also

present. The distinction between assimilation parenchy-

ma and chloroplast-free, central, water-storing, spongy

parenchyma is not very sharp. Assimilation parenchyma

is absent in the dorsally lower part. A faint ring of later-

al vessels could be observed in the transition zone

between the two tissues.

Germination and development

Fresh seed of Faucaria will germinate within a few

days after sowing. The germination curve is bimodal like

that of Trichodiadema spp. (Ihlenfeldt 1971b). The first

seedlings appeared three days after sowing and the first

germination peak appeared after six days, the second one

after 12 days. New seedlings appeared up to 33 days after

sowing. The percentage germination varied from 34 to

94%, with an average of 67%. In the first stage the ger-

mination is by means of a true operculum, which breaks

off before the embryo emerges. Bregman & Bouman

(1983), who studied seeds of numerous Cactaceae and a

few Portulacaceae, found that all operculate species stud-

ied were succulents, whereas the inoperculate species

were not. The cracks may be dorsal or lateral near the

micropyle. The germination does not depend on disinte-

gration of the tough testa. The empty testa may still be

attached to the cotyledons. It was noticed in the present

study that with germination, the primary root stayed

enclosed in the inner integument in some seeds. The pri-

mary root in these seeds may break finally through the

inner integument after days or even several weeks. This

delay in germination is advantageous if only a single

short rain shower has fallen.

The succulent green cotyledons of the seedlings are

slightly broader than long (5:4), with a clear slit between

them. With their fused base they form a flat semicircular

or elliptic top of the seedling. After germination they

grow to about 5 mm in length and width. Dupont (1968)

has observed seedlings of 79 genera in the Aizoaceae sd.

and found that the cotyledons of Faucaria are about

average in their degree of succulence. Differences

between species of this genus in size, form and colour are

hardly noticeable in the seedling stages. Within some

months after sowing, the seedlings develop their first leaf

pair. These leaves are uniformly shiny green and possess

a white, cartilaginous margin without teeth. At this stage

differences between species are not yet noticeable. After

a vegetative development of ± seven months to three

years the lirst Bowers develop. The flowers appear in

autumn, mostly around April/May in the southern hemi-

sphere and in September/November in the northern hemi-

sphere.

Pollination biology

Faucaria sets seed easily after cross-pollination. The

main flowering period is in autumn, from March till

June, but occasionally flowering occurs outside this peri-

od. Differences in the flowering time between species are

usually small; F bosscheana flowers one month earlier

than F felina sd. in culture in the southern hemisphere.

Rain may be important to induce flower development, in

several cases the plants start flowering after the onset of

the rains. Nectar does not seem to play an important role

in pollination. The nectaries are not so easy to reach,

insects have to pass the stamen brush. This unspecialised

floral type is a representative of the cantharophilous syn-

drome, where pollination takes place by small malachiid

beetles. Gess & Gess (1989) found small wasps

(Hymenoptera, Masaridae) as one of the main pollinators

of the Aizoaceae. Liede et al. ( 199 1 ) found in a similar-

ly structured flower of Conophytum subfene stratum

Schwantes ( = C. pillansii) that both insects play a role in

the pollination, but that the beetles outnumbered the

wasps. Wind pollination is unlikely, as the stigmas are

not exposed and dry pollen is not present (Chan 1992).

The flowers of Faucaria open around 15:00-17:00, and

close at sunset. It has been observed in the field that

mixed populations of Faucaria, Bergeranthus and

Orthopterum flower simultaneously. This could indicate

a combined pollinator attraction by flower synchrony.

Fruit and seed dispersal

Faucaria has variable fruit dispersal. Some capsules

become detached easily, others are persistent. The fruits

can either be detached actively from the plant by the next

developing leaf pair or passively when the very short

pedicel withers. The seed is not specially adapted for dis-

persal and combined with the effective closing mecha-

nism seed dispersal of Faucaria depends on capsule dis-

persal. There is a close correlation between hygroscopic

opening, ombrochory and germination (Hartmann 1988).

In Faucaria, ejection of seeds is prevented by a combi-

nation of deep locules and overgrown cell lamellae. In

old capsules most seeds are usually still present and only

after complete deterioration, which could be after years,

these seeds come free from the fruits. This is an inge-

nious method for these plants to overcome drought con-

ditions and re-establish populations after long periods of

low rainfall. Observations and small experiments with

seed dispersal also indicated that seeds of Faucaria are

difficult to wash out from the capsules. This means that

52

Bothalia29,l (1999)

ombrochory in Faucaria is not very important. Fruit and

seeds of Faucaria are not wind-dispersed (anemo-

chorous) like nearly half of the species of the central

lower Karoo community (Cowling & Roux 1987). The

relatively large and heavy capsules usually stay within a

few centimetres of the plants. The re-establishment on

locations after unpredictable long periods of drought in

the south-central Eastern Cape is probably more impor-

tant than dispersal. The establishment of new popula-

tions is difficult, because of the absence of vegetative

dispersal and the poor seed dispersal.

Cultivation

Like many other species of the Eastern Cape,

Faucaria is rather easy to grow in cultivation. In nature,

leaf succulents in the Eastern Cape grow throughout the

year, but a summer peak is apparent (Hoffman 1989). In

cultivation it is attempted to restrict growth to spring and

summer. The best results in the Netherlands were

obtained by growing plants in a sunny place in a heated

winter frame, protected from rain in the growing season

by glass, watered generously in summer and early

autumn. Species of Faucaria overwinter without water-

ing at a minimum of 5°C. Under these conditions plants

grow well and remain compact. If grown in a warm

greenhouse, plants at low light intensity produce long

etiolated leaves. Soil is not very critical; in nature

Faucaria grows in several soils. An appropriate potting

mixture is 1 part compost to 2 parts sharp sand and a

small amount of clay. Pots of 70-120 mm diam. are suit-

able. The pH can probably be below 7 according to field

data (see species descriptions). Propagation is easy by

seed or cuttings. Faucaria species hybridise easily in

cultivation, causing frequent confusion (Hammer 1991).

To obtain unhybridised seed it is necessary to isolate

species when in flower. Plants sometimes suffer from

(root) mealy bugs or they may be attacked by the vine

weevil, Otiorhynchus sulcatus (R). Watering during cold

or moist periods may result in rotting of the leaves, ulti-

mately killing off the plant.

Ecology

Faucaria is found very often under the shelter of open

bushes. This ‘nurse plant’ phenomenon whereby

seedlings readily establish themselves beneath the

canopy of certain plants is common in most arid and

semi-arid areas, including the Karoo (Cowling & Roux

1987). Faucaria usually grows on rocky hill slopes, on

top of koppies, on open spots in the bush or more rarely

in cracks of rocks. The plant has a preference for sun-fac-

ing north slopes, but is not restricted to that position.

Field data recorded plants growing in gravel, sand, tillite,

sandstone, shale, loam, and clay. The plants are probably

not sensitive to low pH.

In this genus a very compact growth is not an adaption

against aridity; F. suhintegra grows in very dry circum-

stances and develops a rather long stem. Correlation

between location factors and species seems small. The

genus occurs in different rainfall regimes, the southern-

most populations of F. felina subsp. felina thrive exclu-

sively in a maximum winter rainfall area. F. hosscheana

occurs in regions with mainly autumn and summer rain-

fall. The remaining taxa grow in areas with mainly spring

and autumn rainfall. Faucaria does not grow at very high

altitudes, and is mostly recorded between 600-850 m,

rarely up to 1 500 m. A comparison of the distribution area

with the vegetation maps of Lubke et al. (1986) indicates

that Faucaria is for the greater part an element of the

Tongaland-Pondoland and Karoo-Namib phytochorologi-

cal regions, which contain many succulent species from

karroid origin, distributed in the thicket (Lubke et al.

1986). The distribution area of F tigriua is within the

boundaries of the southern phytogeographical centre of

the Cape Floral Kingdom as defined by Olivier et al.

(1983); Bond & Goldblatt (1984); Ellis (1988). The dry

thickets of these river valleys were defined by Cowling

(1983) as Kaffrarian Succulent Thicket, and as Valley

Bushveld Thicket by Lubke etal. Like Noorsveld (Acocks

No. 24) and Spekboomveld (Acocks No. 25) these thick-

ets are classified by Lubke et al. (1986) as Subtropical

Thicket. Faucaria has been reported in all three.

Distribution and habitat

The genus is found south of 31° S and between 22°

and 28° E. The main distribution area is within the south-

central region of the Eastern Cape, but it extends slight-

ly into the Western Cape. The genus occurs from the

southeastern coast at about sea level to 1 500 m in the

interior of the Namib-Karoo region. Few locations are

reported from the dry northwestern districts of the area,

but these have been botanically poorly explored (Gibbs

Russell et al. 1984). Faucaria shows a patchy distribu-

tion; the plants are scattered in populations of different

size. Temperature and rainfall change markedly from the

coast towards the interior. The more extreme tempera-

tures occur in the north and west of the area, with maxi-

mum air temperatures of more than 43°C in summer, and

winter minimum falling below -3°C at Graaff-Reinet

(Palmer 1989). The unpredictable rainfall diminishes

gradually from south to north, the river valleys receive

500-900 mm per annum and the area at Graaff-Reinet

has recorded extreme annual precipitation between 144

and 664 mm (Acocks 1988; Palmer 1989). Part of the

distribution area of Faucaria is situated in the ‘stem suc-

culent zone’ as defined by Jurgens (1986). This vegeta-

tion type, characterised by tall long-lived succulents,

only exists in regions with a moderate variability in rain-

fall. The diagram of Ellenberg (1981) indicates that

Faucaria can grow in regions with a rainfall of 400-700

mm and a rainfall variability index of 2. 5-3. 5.

The present distribution of Faucaria partially fits the

hypothesis of Cowling (1983). He postulated that the

semi-arid river valleys of the south-central Eastern Cape,

where Faucaria is concentrated, comprise an endemic

centre and a relatively ancient centre of karroid taxa. He

suggested that karroid species in the south-central

Eastern Cape river valleys were more widespread during

the last glacial era. Thicket taxa of tropical origin e.g.

Euphorbia were established in the Holocene (White et

al. 1941). The present-day succulent thicket communi-

ties consist of these immigrants and newly evolved

species. The semi-arid climate promotes the division of

large or medium-sized populations into smaller ones.

Bothalia29,l (1999)

53

These small populations become isolated from each

other, resulting in small, but often distinct differences

between the populations of Faucaria.

Conservation status

Only Hall et al. (1980) listed a conservation status for

two Faucaria taxa, F. Candida and F longidens syn-

onyms of Ffelina subsp./^/ma. The conservation classi-

fication was listed as indeterminate resp. uncertain,

which means that insufficient data are available. As for

other plant taxa it seems to be more important to protect

the habitat than that of Faucaria itself. Faucaria fe Una

sJ. prefers Eastern Cape transitional thicket, comprised

of Valley Bushveld, Noorsveld and Spekboomveld. This

subtropical thicket is extremely poorly conserved

(Noorsveld = 0%, Spekboomveld = 1.8%, Valley

Bushveld = 1.2%) (Hoffman & Everard 1987). More

than 30% has been converted to wheat cultivation or

degenerated to wasteland by overgrazing. The southern

form of Ffelina subsp./e/ma (published as F. lupina), is

threatened by human activities in its habitat around

Algoa Bay.

It is preferable to assign the lUCN code ‘Vulnerable’

to F tigrina, as this species is restricted to the surround-

ings of Grahamstown, where it is threatened by human

activity such as housing development. There are about

1 700 plants in seven colonies in the wild, according to

a field study of Chan (1992). According to Hilton-

Taylor (1996), F. tigrina is also listed as Vulnerable.

Two other species with a very limited distribution, F.

nemorosa and F. gratiae, are not threatened at the

moment, but must be regarded as Vulnerable. About 800

plants of F gratiae exist in two populations according to

Chan and the size of the population of F. nemorosa is

small but unknown.

DISCUSSION

Classification of species of Faucaria and the genus

itself is problematic. On species level, taxa with striking

but taxonomically difficult characters such as F. rynevel-

diae, remain a subject for discussion. Hartmann (1988)

described eight fruit types in the Ruschioideae. Faucaria

had been regarded in this scheme as being of uncertain

position. In Hartmann (1993) Faucaria fruits had been

classified as similar to the Leipoldtia type but lacking

tubercles. However, Orthopterum, the close relative of

Faucaria, with its small tubercles, does not fit properly

in the Leipoldtia type, as will be discussed later.

Relationship with other genera

Faucaria is included in the subfamily Ruschioideae of

the Mesembryanthema, a monophyletic branch without tax-

onomic rank within the Aizoaceae (Caryophyllales). The

Ruschioideae comprise 107 genera (Bittrich & Hartmann

1988) and are distributed mainly in southern Africa.

The Ruschioideae are characterised by: ovary almost

always with parietal to basal placentation; nectaries

crest-shaped, either as separate nectaries or in a ring,

rarely flat and inconspicuous, very rarely absent; fruits

with expanding keels of mainly valvular origin, never

reaching to the centre of the fruit, if not hygrochastic,

xerochastic, or breaking into mericarps.

The Ruschioideae are divided into 1 1 groups, mainly

on epidermal, flower and fruit characters by Hartmann

(1991). The epidermal characters employed are: epider-

mis mesomorphic or xeromorphic and homocellular or

heterocellular. Flower features used are form and struc-

ture of the nectaries and the presence/absence of fila-

mentous staminodes. Fruit characters are very important

for this classification. Hartmann (1991) placed Faucaria

in the Stomatium Group, mainly on the basis of epider-

mis features. The problems in classifying Faucaria fruits

will be discussed later (see phylogeny).

The Stomatium Group comprises nine genera with the

following diagnostic characteristics:

Plants are perennial. Leaves two to three pairs to a

branch, homocellular xeromorphic, the crystals in irreg-

ular zones; surface rough or warty, from elevations

above subhypodermal tannin idioblasts, often denticu-

late. Flowers with five separate nectaries. Capsule with

or without covering membranes; closing bodies absent;

valve wings broad to reduced. Genera: Chasmatophyl-

luni, Faucaria, Frithia, Mossia, Neohenricia, Ortho-

pterum, Rabiea, Rhinephyllurn, Stomatium.

Frithia, Mossia and Neohenricia have morphological-

ly little in common with Faucaria (Table 2). Rabiea with

its different floral structure and 7-10-locular Titanopsis-

type capsule is also not close to Faucaria. Faucaria and

Stomatium differ mainly in flower, fruit and seed fea-

tures; morphologically the latter often looks like a small

Faucaria with compact growth and toothed leaves.

Flower types are, however, different, the recess flower of

Stomatium with a shallow hypanthium contrasts with the

large carpet flower of Faucaria (Table 2). The scented

flowers of Stomatium with small petals 0.25-0.5(-1.0)

mm wide can readily be distinguished from the scentless

flowers of Faucaria where the petals are 1.0-1.25 mm

wide. The often club-shaped stigmas of Stomatium, 1-3

mm long, differ markedly from the 8-11 mm long fili-

form stigmas of Faucaria. The light brown, non-woody

Delospemia-type fruit of Stomatium is not clearly dis-

tinct from the dark-coloured woody Faucaria capsule

(Figure lOA, B). The parallel expanding keels and seed

pockets are in contrast to the diverging keels and the

seed-retaining syndrome of Faucaria by means of bent,

split septa. The seeds of the two genera are also different,

the light brown Stomatium seeds have a smooth or papil-

late brown testa, those of Faucaria are dark brown and

tuberculate.

The similarities between Chasmatophyllum and

Rhinephyllurn and Faucaria are restricted to compact

growth and similar flower shape. Teeth on the often

rounded leaves of the former two are, however, general-

ly indistinct or even missing, whereas in Faucaria the

teeth are usually well developed and often the leaves

have white margins. In Rhinephyllurn, furthermore, the

nectaries are often hardly separated, but in

Chasmatophyllum and Faucaria they are clearly distant.

54

Bothalia29,l (1999)

TABLE 2. — Morphological similarities in the Stomatium Group

+ character states present; - character states absent. Fa, Faucaricr, Or, Orthopterunv, Rh, Rhinephyllunr, Ch, Chasmatophyllum\ St, Stomatium;

Ra, Rabieu; Fr, Frithia; Mo, Mossia; Ne, Neohenricia.

The fruit of Chasmatophyllum and Rhinephyllum can not

be misidentified as Faucaria fruits, mainly because of

the absence of the unique stiff, upright valve wings of the

latter.

Vegetatively Orthopterum is closer to Faucaria than

to any other genus. Non-fruiting plants of Orthopterum

with their smooth, more or less shiny appearance resem-

ble forms of Faucaria felina, described as F lupina, even

though the leaves of Orthopterum usually have fewer

teeth and white dots.

The flowers are almost identical and those of

Orthopterum also open in the late afternoon. The real dif-

ference between the two genera lies in the fruits.

Nevertheless the fruits of Orthopterum are the only ones

within the Ruschioideae that have close similarities to

those of Faucaria fruits (see Phylogeny).

Hybridisation between Faucaria and other members

of the Stomatium group is unknown. Up to now only one

natural intergeneric hybrid has been reported by Hammer

& Liede (1990) between Faucaria and Rhombophyllum.

Phylogeny

The Stomatium Group is polyphyletic, a cladogram

could not be constructed. The artificial character of the

Stomatium Group is reflected by the absence of any diag-

nostic feature common to all taxa. The diversity of the

group is also shown by the presence of three of the eight

described Ruschioideae fruit types (Hartmann 1991).

Orthopterum, Stomatium, Rhinephyllum and Chasmato-

phyllum resemble Faucaria as mentioned earlier (Table

2). Only the fruits are considered suitable for a meaning-

ful examination of relationships within the Stomatium

Group. Five out of the nine genera in the group possess

Delosperma-type fruits (Hartmann 1988). The fruits of

this type have parallel expanding keels, probably an apo-

morphic character (Hartmann 1988), quite different from

the diverging expanding keels of Faucaria and

Orthopterum. The fruits of Stomatium and Rhinephyllum

belong to the Delosperma type, those of Chasmato-

phyllum to the Drosanthemum type. This last-mentioned

type has translucent covering membranes which are

absent in Faucaria. Out of the genera involved, only the

fruits of Orthopterum show similarities with those of

Faucaria (Figure lOB, C).

Close examination reveals that the fruits of Faucaria

might be derived from those of Orthopterum by reduction

and lignification. They share split septa, which bend over

the locules to resemble covering membranes. The fruits of

both genera lack real covering membranes. The upper

part of the split septa of Orthopterum is translucent, flex-

ible and raised, the lower part is partially lignified. This

expanded part has disappeared in Faucaria and the lamel-

lae are completely lignified. Both fruits have divergent,

non-broadened expanding keels. The expanding keel of

Orthopterum is long, that of Faucaria short. The long

awn of the expanding keel of Orthopterum is reduced to

a short subulate point in Faucaria. The large, broad,

reflexed, flexible valve wings of Orthopterum have been

reduced to small upright wings in Faucaria, standing

rigidly over the locules. The valve wings of Orthopterum

are reflexed, away from the opening fissure of the locules.

In spite of differences in lamellae, expanding keels and

valve wings, the basic fruit structure of the genera is the

same. The common features indicate a common ancestor,

confirming the hypothesis of Schwantes (1957) that they

are related. The presence of closing bodies in Ortho-

pterum has no influence on this relationship, as the taxon-

omic importance of these structures cannot be determined

(Hartmann 1988). The ontogeny of the fruits also sug-

gests that Orthopterum and Faucaria are close relatives.

Mature fruits of Orthopterum are conical and acute, pro-

viding space for the raised septa. The ripening fruits of

Faucaria also develop this conical top (Figure 6C), but

this plesiomorphic feature disappears in later develop-

mental stages, as the septa are lower, and the ripe fruits

are flat- or low-topped.

Bothalia 29,1 (1999)

55

Hartmann (1993) assumed that Faucaria fruits are

similar to the Leipoldtia type, the striking feature of the

genus being firm, stout valve wings fused to the valves

in their basal part only and standing rigidly above the

locules after the capsule has opened. Faucaria shares

with the Leipoldtia type the distinct, non-broadened radi-

al awn and expanding keels. Other fruit types (Dro-

santhemum, Titanopsis) show broadened expanding

keels. However, the two apomorphic characters of the

Leipoldtia type (Hartmann 1983); large closing bodies

and the elaborate covering membranes, are missing. The

large valve wings of Orthopterum and the small valve

wings of Faucaria both occur in the Leipoldtia type. The

large valve wings are common, the reduced valve wings

of Faucaria are only shared with Antiruima. Ortho-

pterum fits even better into this fruit type, as it has small

closing bodies similar to those in Jordaaniella and

Fenestraria. The presence of these small closing bodies

does not prevent classification of Orthopterum in the

Leipoldtia fruit type. The pentamerous fruits of

Orthopterum and Faucaria are also represented in this

fruit type, e.g. in Antimima fruits. The absence of cover-

ing membranes in Faucaria and Orthopterum is the sin-

gle, but essential feature, preventing them from being

readily classified in tbe Leipoldtia type. If the presence

of this character prevents classification in this type, then

Faucaria and Orthopterum fit into none of the fruit types

described by Hartmann (1988). It must be concluded that

in the present classification Faucaria cannot be satisfac-

torily placed. Only Orthopterum is closely related to

Faucaria and together they appear to form an isolated

group within the Ruschioideae. Therefore it is proposed

to re-introduce the Faucaria fruit type of Schwantes

(1952). This will fit into the fruit scheme of Hartmann

(1988) with the following characters:

Faucaria type of fruit (Schwantes 1952)

Expanding keels stout, divergent with distinct radial

awn, not broadened, connate in their basal part only,

extending rigidly above the locules, expanding sheet pre-

sent. Covering membranes absent. Closing body absent

or small. Genera: Faucaria Schwantes, Orthopterum

Schwantes.

For a cladistic analysis of the genus Faucaria itself

only a few characters are usable, such as crystal dots and

teeth. Four Faucaria species show good visible crystal

dots dispersed over the leaves in the outer epidermal

layer. The three remaining species, F. bosscheana, F

nemorosa and F. subintegra, share with the outgroup

Orthopterum the presence of, mostly only a few, concen-

trations of crystals around the teeth and margins on the

leaves. The size and number of crystal dots and teeth are

dictated by the taxa and are influenced in a minor way by

the habitat. In culture all species turn more greenish than

in nature.

The development of hypodermal crystal concentra-

tions, teeth or bristles reflects the phylogeny in the

ontogeny, and indicates the polarity of these characters.

The ontogenetic transformations are an invaluable tool in

the recognition of characters (sets of homologous states).

The absence of oxalic crystal dot concentration in the

leaves in the juvenile stage of Faucaria indicates that

this feature is an advanced character. Because of the

neotenic character of F. bosscheana, this species is con-

sidered basal in the genus by the phylogenetic analysis

(Figure 12). The differences with the other species are,

however, not so large as the diagram suggests. The dia-

gram reflects a close relation between F nemorosa and F

subintegra. Both show a reduction in tooth development.

The fruits are similar in form and attachment on the

plant, and the two species have the same, somewhat

shrubby growth. The ecology of the two species, howev-

er, differs markedly: F. subintegra grows in the open, on

very dry river banks. F. nemorosa is found under shrubs.

A close relationship exists between F. felina s.l. and F.

gratiae. F tigrina is also related to the F felina Group,

but is distinct in morphological and epidermal features.

CONCLUSIONS

Faucaria together with Orthopterum forms an isolat-

ed group within the Ruschioideae. The genus consists of

six species. Two species, F bosscheana and F felina s.l.

occur over a large area, four are much more localised.

The most common and widely distributed species, F. feli-

na S.L, covers most of the diversity in Faucaria, more

than three quarters of the names in the genus were based

on forms of this species. The variation is continuous,

only three forms could be recognised as separate taxa,

giving rise to subsp. britteniae, suhsp. felina and subsp.

tuberculosa. Macro-morphological information was suf-

ficient for a proper taxonomic analysis in the genus.

Orthopterum

F. nemoro.'ui

F. subintegra

F. bosscheana

F. gratiae

F. felina subsp. tuberculosa

F. felina suhsp. felina

F. felina subsp. britteniae

F. tigrina

FIGURE 12. — Cladogram for the species in Faucaria. Characters, represented by numbers in the cladogram are given in Table I. Symbols: O-I

character state changement, ■; 1-0 homoplasy, □; 1-2 character state changement, •. Consistency index 67, retention index 72.

56

Bothalia 29,1 (1999)

Epidermal characters proved important in this genus.

Neither seed surface, nor pollen morphology provide

taxonomically important data; similarly the structure of

flowers and fruits is of limited taxonomic value.

The cladistic analysis is based on only 13 characters

(Table 1) and must be regarded as preliminary. The

cladogram reflects the impression that Faucaria has

arisen from a toothless or few-toothed ancestor. F. boss-

cheana, with its neotenic tendencies, is probably the less

advanced species with a relatively simple epidermis and

leaf structure, with little ecological specialisation in kar-

roid circumstances. F tigrina, on the other hand, shows

a complicated epidermis structure, is ecologically spe-

cialised, even showing mimicry with reddish lichens.

ACKNOWLEDGEMENTS

I thank the directors of the herbaria of B, BOL, G,

GRA, HBG, K, MO, PRE, SAM, NBG and WAG for the

loan of herbarium sheets. In particular I am thankful to

the herbaria of BOL and K for the loan of type specimens.

I acknowledge with gratitude the help of the National

Botanic Gardens and the National Botanical Institute of

South Africa, for providing assistance, help and facilities.

Drs O.A. Leistner, S.P. Cowling and S.A. Hammer are

thanked for refereeing and advice. Mrs Chan, Mrs E.

Brink, S.A. Hammer, G. Marx, E. van Jaarsveld and Dr &

Mrs P.J. Vorster kindly provided help during field trips

and extensive information about wild populations. A col-

lection permit was obtained from the Chief Directorate of

Nature and Environmental Conservation, Cape Town.

Thanks are due to the Department of Plant Taxonomy at

Wageningen Agricultural University, particularly to the

late Dr D.O. Wijnands. Financial support for research in

South Africa was obtained by the Dutch Eoundation of

Scientific Research, the Eoundation Neerlandia and the

Van Ewijck-Stigting.

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bryanthenti 1 : t. 5, 6. Henry & Cohen, Dusseldorf & Bonn.

SALM-REIFFERSCHEID-DYCK, PRINCE J.M.F.A.H.I. VON J.

FURST ZU, 1854. Monographia generum Aloes et Mesem-

bryanthemi 6: t. 2. Henry & Cohen, Dusseldorf & Bonn.

SCHWANTES, G. 1926. Zur Systematik der Mesembrianthemen.

Zeitschrift fiir Sukkulentenkunde 2, 11: 176, 177.

SCITWANTES, G. 1927. Zur Systematik der Mesembrianthemen.

Nachtrag Zeitschrift fiir Sukkulentenkunde 23: 105.

SCHWANTES, G. 1952. Die Friichte der Mesembryanthemaceen.

Vierteljahrsschrift der Naturfdrschenden Gesellschafft Zurich

97, Beiheft 2: 1-38.

SCHWANTES, G. 1957. Flowering stones and midday flowers:

129-131, 340. Benn, London.

SONDER, O. W. 1862. Mesembryaceae Lindl. In W.H. Harvey & O.W.

Sender, Flora capensis 2: 388, 389, 396, 397. Dublin.

TISCHER, A. 1929. Faucaria haagei Tisch. spec. nov. Monatsschrift

der Deutschen Kakteen-Gesellschaft 1: 20, 21.

TISCHER, A. 1932. Einige neue Mesembryanthema. Monatsschrift der

Deutschen Kakteen-Gesellschaft 3: 280.

TISCHER, A. 1934. Faucaria jamesii L. Bolus ex Tischer. Kakteen-

kunde: 99.

TROLL, W. & WEBERLING, F. 1981. Infloreszenzstudien an Aizo-

aceen, Mesembryanthemaceen und Tetragoniaceen. Tropische

und Subtropische Pflanz.enwelt 35: 195-289.

VAN JAARSVELD, E. 1984. The family Mesembryanthemaceae with

special reference to Faucaria felina and Glottiphyllum muirii.

Cactus and Succulent Journal (USA) 56: 53.

VAN JAARSVELD, E. 1990. Kougadam en Oos Kaap versameltog.

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VOGEL, S. 1954. Bliitenbiologische Typen als Elemente der Sip-

pengliederung. In W. Troll & H. von Guttenberg, Botanische

Studien 1 . Fischer Verlag, Jena.

WEBER, C. 1968. Notes on the nomenclature of some Aizoaceae.

Baileya 16: 12.

WESTON, R. 1770. Botanicus Universalis et Hortulanus (Weston’s

Nurseryman) I: 172. London.

WHITE, A., DYER, R.A. & SLOANE, B.L. 1941. The succulent

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Rotterdam.

58

Bothalia29.1 (1999)

INDEX

Bergeranthus Schwantes, 31

Chasmatophylluin Dinter & Schwantes, 53, 54

Cheiridopsis N.E.Br., 51

Conophytum Lavis, 5 1

subfenestratum Schwantes, 51

pillansii Lavis, 5 1

Delosperma N.E.Br., 53

Drosanthemum Schwantes, 54

Faucaria Schwantes, 36

acutipetala L. Bolus, 40

albidens N.E.Br., 37

bosscheana (A.Berger) Schwantes, 37

var. haagei (Tischer) H. Jacobsen, 37

britteniae L. Bolus, 41

Candida L. Bolus, 40

coronata L. Bolus, 41

cradockensis L. Bolus, 40

crassipetala L. Bolus, 40

duncanii L. Bolus, 40

felina (L.) Schwantes, 39

subsp. britteniae (L.Bolus) L.E.Groen, 41

subsp. felina, 40

subsp. tuberculosa (Rolfe) L.E.Groen, 42

war. jamesii L.Bolus, 40

grandis L.Bolus, 41

gratiae L.Bolus, 42

haagei Tischer, 37

hooleae L.Bolus, 42

jamesii L.Bolus, 40

kendrewensis L.Bolus, 37

kingiae L.Bolus, 40

latipetala L.Bolus, 40

laxipetala L.Bolus, 40

longidens L.Bolus, 40

longifolia L.Bolus, 40

lupina (Haw.) Schwantes, 40

militaris Tischer, 40

rnontana L. Bolus, 40

rnultidens L.Bolus, 40

var. pardepoortensis L.Bolus, 40

neinorosa L.Bolus ex L.E.Groen, 43

paucidens N.E.Br., 37

peersii L.Bolus, 37

plana L.Bolus, 40

ryneveldiae L.Bolus, 40

smithii L.Bolus, 41

speciosa L.Bolus, 41

subindurata L.Bolus, 44

subintegra L.Bolus, 44

tigrina ‘Superba’, 45

tigrina (Haw.) Schwantes, 45

forma splendens H. Jacobsen & G.D. Rowley, 45

tuberculosa (Rolfe) Schwantes, 42

uniondalensis L.Bolus, 40

FrithiaiV.E.Br, 53, 54

Jordaaniella H.E.K. Hartmann, 55

Mesembryanthemum L., 36

bosscheanum A. Berger, 37

felinum ‘(L.)’ Weston, 40

lupinum Haw., 40

ringens L. (J felinum L., 40

tigrinum Haw., 45

tuberculosum Rolfe, 42

Mossia N.E.Br, 53, 54

Nananthus N.E.Br., 51

Neohenricia L.Sr;/i(.v, 53, 54

Octopoma A.E.Br., 51

Odontophorus N.E.Br, 51

Onhopterum L.Bolus, 51, 53-55

RabieaA.E.B/:, 53, 54

Rhinephyllum N.E.Br, 53, 54

Rhombophyllum (Schwantes) Schwantes, 51

Ruschioideae Schwantes in Ihlenf., Schwantes & Straka emend.

Bittrich & H.E.K. Hartmann, 53

Stomatium Schwantes, 53, 54

Stomatium Group, 53

Titanopsis Schwantes, 55

Trichodiadema Schwantes, 5 1

Bothalia 29,1: 59-63 (1999)

New species of Sparaxis and Ixia (Iridaceae: Ixioideae) from Western

Cape, South Africa, and taxonomic notes on Ixia and Gladiolus

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

Keywords: comb, nov., Gladiolus scixatilis, Iridaceae, Ixia aurea. Ixioideae. South Africa, sp. nov., stat. nov., taxonomy. Western Cape

ABSTRACT

Sparaxis auriculata is a new species of this western Cape and western Karoo genus. It resembles S. villosa (Burm.f.)

Goldblatt in its flowers but is probably allied to S. galeata (Jacq.) Ker Gawl. Sparaxis metelerkampiae, currently a sub-

species of S. variegata Sweet is raised to species rank. We propose changes to the infrageneric classification of the southern

African winter rainfall genus Ixia, and describe a new species, I. aurea. It is related to I. odorata Ker Gawl. but differs in the

larger deep yellow or orange flowers which are unscented. In Gladiolus, we propose the new name, G. saxatilis, for the

Mpumalanga endemic, originally described as G. litliicola Goldblatt & J.C. Manning, a homonym for an Ethiopian species.

1. Sparaxis auriculata Goldblatt & J.C. Manning,

sp. nov.

Plantae 150-400 mm altae, cormo conico 12-14 mm

diam., tunicis fibrosis, foliis 5-8 anguste lanceolatis

acutis 10-15 mm latis, caule plerumque 1- vel 2-ramoso,

spica subsecunda 5-7-flora, bracteis scariosis 15-20 mm

longis, floribus zygomorphis bilabiatis luteis violaceis

notatis, tubo oblique-infundibuliforme ± 24 mm longo,

tepalis inaequalibus, tepalo dorsale ± 25 x 18-20 mm,

inferioribus auriculatis, filamentis ± 18 mm exsertis,

antheris 7-8 mm longis.

TYPE. — Western Cape, Farm Koorlandshoek (Collins-

hoek), slopes of the Gifberg, Aug. 1998, Goldblatt &

Manning 10966 (NBG, holo.; K, MO, PR£, S, iso.).

Plants 150^00 mm high. Conn globose-conical,

12-14 mm diam., tunics of medium-textured fibres most-

ly oriented vertically. Leaves 5-8, mainly basal, upper-

most inserted in middle of stem, sheathing lower part of

stem, reaching to between middle of stem and base of

spike, narrowly lanceolate, acute, mostly 10-15 mm

wide, with a moderately prominent submarginal vein

making margin appear somewhat thickened, sheathing

parts of leaves speckled with purple (sometimes fading

when dry). Stem usually with one or two branches;

branches produced in axil of cauline leaf, usually inclined

and slightly flexed above leaves. Spike mostly 5-7-flow-

ered, weakly flexuose, more or less secund; bracts pale

below with whitish veins, becoming streaked with light

brown above, 15-20 mm long, lightly lacerate and with

prominent central cusp, inner slightly shorter and with 2

apical cusps. Flowers zygomorphic, bilabiate, predomi-

nantly cream-coloured, dorsal tepal pale mauve, lower

tepals deep yellow fading to cream-coloured at apices,

abaxial half of throat obscurely lined with dull grey-pur-

ple, unscented; perianth tube obliquely funnel-shaped,

upper part curving outwards, ± 18 mm long, lower tepals

united for a further 6-7 mm, basal cylindrical portion nar-

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

RO. Box 299, St. Louis, Missouri 63166, USA.

** National Botanical Institute, Private Bag XI, Claremont 7735, Cape

Town.

MS. received: 1998-12-17.

row and filled by style, without nectar; tepals unequal,

dorsal largest, broadly ovate, erect, ± 25 x 18-20 mm,

upper laterals oblong-ovate, ± 20 x 10-12 mm, spreading

in upper half, lower three abruptly deflexed above and

pinched together at bases, directed downward, ±18x8

mm, strongly channelled, lower laterals with prominent

auriculate basal lobes. Filaments unilateral and arcuate,

whitish, ± 18 mm long; anthers cream-coloured, 7-8 mm

long, exserted completely; pollen cream-coloured. Ovary’

3.5 mm long, oblong-trigonous; style arching over sta-

mens, white, ± 4 mm long, dividing just below apex of

anthers, branches filiform, diverging and recurved in

upper half, slightly wider toward apex. Capsules and

seeds unknown. Chromosome number unknown. Flower-

ing time: August. Figure 1 .

Distribution and habitat: the species is so far known

only from the lower slopes of the Gilberg Mountains in the

northernmost part of the Western Cape (Figure 2). Plants

were found to be very local but fairly common among rocks

and in cleared places on bush-covered, sandstone slopes.

Apparently first recorded in 1979 by I.S. Walters on

the Farm Vleikraal (or Veekraal) on the slopes of the

Gifberg east of Klawer, Sparaxis auriculata was initially

associated with the superficially similar S. villosa

(Burm.f.) Goldblatt, then classified in the genus Synnotia

Sweet. A second gathering was made in 1997 on Gifberg

Pass east of Vleikraal and we re-collected the species

there on the middle slopes of the Pass on the Farm

Koorlandshoek in early August, 1998. Examination of

living plants, in particular the underground organs, con-

firmed our expectation that the two prior collections rep-

resented an undescribed species. S. auriculata can be dis-

tinguished mainly on vegetative features. The corms are

narrowly conical and have tunics of medium-textured

fibres, whereas the leaves have the sheaths liberally spot-

ted with dark purple, blades with acute apices and, espe-

cially unusual for Sparaxis, a prominent submarginal

vein that makes the margins appear thickened.

Narrow, conical corms with soft corm tunics are char-

acteristic of only two species of Sparaxis, S. galeata

(Jacq.) Ker Gawl. and 5. roxburghii (Baker) Goldblatt,

60

Bothalia 29,1 (1999)

both northwestern Cape species (Lewis 1956; Goldblatt

1992). The underground part of the stem is enclosed by a

thick neck of fine fibres and leaves with speckled

sheaths, but both species differ in lloral details from S.

auriculata. Sparaxis galeata has short-tubed, extremely

sweetly fragrant flowers with the dorsal tepal rcllexed,

the leaves are prostrate, oblong and obtuse, and the

species favours dry, stony, clay flats. The rare S. rox-

hurgliii has long-tubed flowers (the tube 20-25 mm

long), otherwise resembling those of S. galeata, and is

known from a single population in the mid Olifants River

valley on stony slopes. Neither species has the submar-

ginal vein of 5. auriculata.

The llowers of Sparaxi.s auriculata are much like those

of the common southwestern Cape S. villosa in general

FIGURE 1. — Sparaxis auriculala. A,

corm and leaves; B, flowering

spike; C, D, tlower: C, front

view; D, side view. Scale bar;

A-D, 12 mm. Artist: John

Manning.

appearance, but they are substantially larger in all respects.

In S. auriculata the dorsal tepal is ± 25 x 18-20 mm and the

anthers are 7-8 mm long, compared with 14—19 x 10-12

mm for the dorsal tepal and 3-^ mm for the anthers in S. vil-

losa. Furthermore, the style in the latter species divides near

the anther bases, the style branches are ± 3 mm long, rarely

reaching beyond the middle of the anthers, and the lower

tepals are inclined slightly below the horizontal and without

auriculate basal lobes. In S. auriculata the style branches are

± 4.5 mm long and exceed the anther apices, and the lower

tepals are directed downwards and are prominently lobed

near the base. The corms and leaves of the two species dif-

fer substantially and in S. villosa the corms are globose with

coarse, netted fibres, the vertical elements of which are

thickened and claw-like below, while the leaves lack the

unusual spotting so marked in S. auriculata.

Bothalia29,l (1999)

61

FIGURE 2. — Distribution of Sparaxis auriculutu, and Ixia aiirea,

•, in Western Cape.

Specimens examined

WESTERN CAPE. — 3118 (Vanrhynsdorp); Farm Vleikraal east of

Klawer, on east-facing lower slopes of the Gifberg, (-DC), Aug. 1979,

Walters 15 (NBC); Gifberg Pass, (-DD), 25 Aug. 1997, Van Rooyen.

Steyn & De Villiers 587 (NBC, PRU).

2. Sparaxis metelerkampiae (L.Bolus) Goldblatt

& J.C. Manning, stat. nov.

Synnotia metelerkampiae L.Bolus, in Annals of the Bolus Herbarium

3: 77 (1923). Synnotia variegata Sweet var. metelerkampiae (L.Bolus)

G.J.Lewis: 148 (1956). Sparaxis variegata (Sweet) Goldblatt subsp.

metelerkampiae (L.Bolus) Goldblatt: 158 (1992). Type: Western Cape,

Clanwilliam Div., near Eendekuil, F. Metelerkamp BOL16039 (BOL,

holo.).

Goldblatt (1992) recognised two subspecies of Sparaxis

variegata with subsp. metelerkampiae (L.Bolus) Goldblatt

being distinguished from the typical subspecies by its small-

er, longer-tubed flowers which are largely purple (including

the anthers), a style which usually divides below the middle

of the anthers, and short style branches only 2-A mm long.

The two were maintained as subspecies because of the pres-

ence of some apparently intermediate populations.

Field observations show that populations of Sparaxis

variegata and S. metelerkampiae consistently differ and

do not intergrade. This is particularly significant as their

geographic ranges overlap. Apparent intermediates in

herbaria may either be depauperate individuals or unusu-

al forms of either taxon. The morphological differences

between the two reflect differences in their pollination

biology. S. metelerkampiae is pollinated by the long-pro-

boscid flies Prosoeca peringueyi and P. sp. (Diptera:

Nemestrinidae) (Manning & Goldblatt 1996), whereas S.

variegata is pollinated by the large anthophorine bee,

Anthophora diversipes (Hymenoptera: Apidae) (unpub-

lished observations). These differences in morphology

and reproductive biology suggest that it is more appro-

priate to recognise S. metelerkampiae as a species.

The two species can be distinguished as follows.

Sparaxis variegata has flowers with a perianth tube 37-44

mm long with the lower part 25-32 mm long and the dor-

sal tepal 25-30 mm long. The style divides at or near the

anther apices, the style branches are 4-5 mm long and the

cream-coloured anthers are 6-7 mm long. The perianth is

usually predominantly yellow with the dorsal tepal and the

tips of the others coloured violet whereas the lower part of

the throat is streaked with dark violet lines. In S. metel-

erkampiae the perianth tube is 45-50 mm long with the

lower part 35-40 mm long and the dorsal tepal is 14-18

mm long. The style usually divides below the middle of

the anthers, the style branches are 2-4 mm long and the

purple anthers are 2-4 mm long. The perianth is uniform-

ly purple in colour except for the lower tepals which are

marked with narrow white longitudinal streaks.

Greenhouse-grown plants of S. metelerkampiae are self-

compatible and autogamous, whereas S. variegata is self-

incompatible (Goldblatt 1992).

Specimens examined

NORTHERN CAPE. — 3119 (Calvinia): Nieuwoudtville-Oorlogs-

kloof, (-AC), Leipoldt 872 (BOL, K).

WESTERN CAPE. — 3118 (Vanrhynsdorp): Zyferfontein, NE of

Rietvleiberg, (-DC), Oliver 3812 (K). 3218 (Clanwilliam): Graafwater,

(-BA), Compton 220 (NBG); between Clanwilliam and Graafwater,

(-BA, -BB), Van Breda 1267 (PRE, Z); near Paleisheuwel, (-BC),

Levyns 10154 (BOL); Eann Klein Remhoogte, west of Algeria, stony

bank in sand, (-BD), Snijman 887 (MO, NBG); 10 km S of

Clanwilliam, Lewis 4284 (K, PRE, SAM); Hex River Farm N of

Citrusdal, sandstone slopes. Goldblatt 6148 (MO); Grey’s Pass, (-DB),

Barker 1476 (NBG), Salter 2733 (K). 3219 (Wuppertal): top of Pakhuis

Pass, recently burned slopes in sandy soil, (-AA), Goldblatt 6404

(MO); Pakhuis Pass, Salter 3641 (BOL, K), Leipoldt BOL21277,

Gillett 4062 (BOL, PRE), Esterhuysen 8032 (PRE); foot of Pakhuis

Pass, Goldblatt 5657A (MO); Algeria campsite, 700 m, (-AC), Bos 503

(PRE); Kriedouw Krantz Pass, Cederberg, Gillett 4114 (MO); slopes of

Middelberg, Elandskloof, (-CA), Hafstrdm & Acocks 331 (PRE);

Elandskloof, Leighton 1341 (NBG); slopes behind warm baths,

Stephens 7086 (BOL, K).

3. Ixia aurea J.C. Manning & Goldblatt, sp. nov.

Plantae 200-400 mm altae, cormo globoso 10-16 mm

diam., tunicis fibrosis reticulatis, foliis 5 vel 6 lanceolatis

(8-) 10- 16 mm latis supra leviter contortis, caule eramoso,

spica leviter flexuosa 6-20-flora, bracteis scariosis 8-9

mm longis, floribus hypocrateriformibus luteis, tubo

anguste infundibuliforme 8-11 mm longo, tepalis subae-

qualibus ellipticis patentibus 16-23 x 7-9 mm, filamentis

decurrentibus 5-6 mm exsertis, antheris 6-7 mm longis.

TYPE. — Western Cape, Darling District, Farm Wolwe-

fontein, rocky slope among granite boulders, 22 Sept.

1998, Goldblatt & Manning 11029 (NBG, holo.; K, MO,

PRE, S, WAG, iso.).

Plants 200-4(X) mm high. Comis globose, ± 10 mm

diam., tunics of fairly fine reticulate fibres, often with few

small cormlets at base. Cataphylls membranous, upper one

reaching shortly above ground level. Leaves 5 or 6, all basal,

narrowly lanceolate, (8-) 10-16 mm wide, usually reaching

to about middle of stem, uppermost leaf usually entirely

sheathing, margins and midribs not thickened, lightly twist-

ed in upper half Stem erect, unbranched, ± 0.8 mm diam.

below base of spike. Spike crowded, secund and spirally

twisted, (6-)12-20-flowered; bracts scarious, translucent,

outer ± 7 mm long, with prominent dark central vein,

obscurely three-dentate, central tooth cusp-like and much

exceeding lateral teeth, inner bract about as long as outer,

62

Bothalia 29,1 (1999)

FIGURE 3. — Ixia aurea. A, corm and leaves; B, flowering spike; C,

floral bracts; D, androecium; E, gynoecium. Scale bars: A-E,

10 mm. Artist: John Manning.

two-kceled, each keel with a prominent dark vein, bicuspi-

date. Flowers hypocrateriform, actinomorphic, uniformly

deep yellow or orange, unscented; perianth tube straight and

narrowly funnel-shaped, 8-11 mm long. Filiform in lower

half to two-thirds, widening gradually above, basal part

tightly enveloping style, containing nectar in upper third;

tepals obovate-elliptic, subcqual, spreading, margins slightly

upeurved, 16-23 x 7-9 mm. Filaments in.serted ± 1 mm

below apex of tube, decurrent for ± 1 mm, 5-6 mm long.

straight, weakly diverging; anthers 5-6 mm long (before

anthesis), diverging, yellow; pollen yellow. Ovary ovoid, ± 3

mm long; style straight and erect, dividing opposite lower

third of anthers, branches arching outward, 4.5-7 .0 mm

long, reaching to shortly above base of the anthers. Capsules

barrel-shaped-trigonous, retuse, showing outline of seeds, ±

7x5 mm. Seeds subglobose, reddish brown, ± 1 .2 mm long,

testa sculpturing scalariform. Flowering time: September,

occasionally into early October. Figure 3.

Distribution and habitat: Ixia aurea is known only

from the granitic slopes above the Darling- Yzerfontein

road (Figure 2) and is conspicuous when in flower, the

brilliant yellow-orange spikes making a bright splash of

colour among the late spring flora.

Despite occurring alongside the road in such a well-

visited region as Darling, the species has rarely been col-

lected. Plants from the type locality were originally mis-

takenly identified as a form of I. dubia Vent, of section

Ixia by Goldblatt et al. (1998) because of the general

resemblance to that species, despite the lack of a dark

central mark to the flowers. More critical examination

shows that I. aurea does not belong in section Ixia as cur-

rently circumscribed, for it has flowers with a narrowly

funnel-shaped tube, tightly enclosing the style in the

cylindrical lower half but flaring above and containing

nectar in the upper third. In addition the filaments

diverge from the base and do not occlude the mouth of

the tube. In section Ixia, to which I. dubia belongs, the

perianth tube is filiform-cylindrical almost to the apex,

closely envelops the style and does not contain more than

minute traces of nectar. The filaments are also coherent

or connate from the base and thus close off the mouth of

the tube. We suspect that records of this species were

never made because even from a short distance the plants

may be confused with two other common species from

the Darling area, I. maculata L. and I. dubia. Apart from

the floral differences, I. aurea may be distinguished from

I. dubia by the leaves which lack thickened margins and

midrib and are lightly coiled in the upper half. The blades

of I. dubia and its allies have thickened margins and

midribs and are plane. We suspect that the relationships

of I. aurea lie with the two other yellow-flowered, nar-

row-tubed species, I. tenuifolia Vent. (= I. framesii

L. Bolus) and /. odorata Ker Gawl. Of these, /. odorata

may be its closest ally, based on general appearance, the

broad, twisted leaf blades, and the many-flowered

spikes. The flowers of I. odorata are in general smaller

than those of I. aurea, with tepals up to 15 mm long, are

white to lemon-yellow with a darker tube and never deep

golden yellow to orange and are always sweetly scented

of magnolia. The flowers are visited by a variety of bees

for nectar and pollen.

The flowers of I. aurea are pollinated by a range of

insects, including the hopbine beetles Heterochelus

arthriticus, Lepithrix ornatella and Pachycnema cras-

sipes (Coleoptera: Scarabidae) and the horsefly Meso-

myia edentula (Diptera: Tabanidae). The beetles use the

flowers as sites for assembly and mating while the flies

are attracted to the flowers as a source of nectar. Both the

beetle species and the horsefly become covered in pollen

after visiting several flowers. Ixia aurea thus joins the

long list of species of this genus, mostly in section Ixia,

Bothalia29,l (1999)

63

which are pollinated in part at least by hopliine beetles

(Goldblatt et al. 1998).

INFRAGENERIC CLASSIFICATION OF IXIA

The following currently accepted infrageneric classi-

fication of Ixia, was proposed by Lewis (1962):

Subgenus Ixia

Section Ixia

Section Morphixia (Ker Gawl.) Pax

Section Hyalis (Salisb. ex Baker) Diels

Subgenus Dichone (Salisb. ex Baker) G.J. Lewis. .

Subgenus Dichone and section Ixia (subgenus Ixia)

share the synapomorphy of a fdiform perianth tube which

tightly clasps the style for its entire length. The tube also

contains little or no nectar and the filaments are contigu-

ous or connate from the base, effectively closing the

mouth of the tube. We assume that this character arose

once, for there seems no evidence to the contrary.

Observations on the pollination biology of the genus are

also consistent with this assumption and suggest that the

species with filiform tubes have adopted derived pollina-

tion strategies which offer pollinator rewards other than

nectar. The two taxa have flowers which are modified in

different ways for different pollination strategies. In

Dichone the flowers are uniformly pink and the anthers

are short, somewhat inflated and dehisce tardily or only

from the base by short slits. The flowers are buzz-polli-

nated by anthophorine bees (Apidae: Anthophorinae)

which vibrate the flowers to extract the pollen (unpub-

lished observations). In contrast, flowers of section Ixia

are variously coloured, often with dark centres, and the

anthers are large and longitudinally dehiscent. These flow-

ers are adapted for pollination by hopliine beetles

(Scarabidae) or by bees foraging for pollen. In tbe remain-

ing sections Hyalis (Salisb. ex Baker) Diels and

Morphixia (Ker Gawl.) Pax, the flowers have hollow peri-

anth tubes which normally contain significant quantities of

nectar; they are largely pollinated by nectar-foraging

insects, particularly bees and long-tongued flies. Because

of the shared synapomorphy we believe that Dichone,

with section rank, and section Ixia should together consti-

tute the subgenus Ixia. This decision is reinforced by the

close resemblance between 7. micrandra (subgenus

Dichone) and I. flexuosa (section Ixia). Sections Hyalis

and Morphixia are placed in a second subgenus for which

we propose the name subgenus Morphixia.

We, therefore, put forward the following infrageneric

classification for Ixia:

Subgenus Ixia. Type: Ixia polystachya L.

Section Ixia

Section Dichone (Salisb. ex Baker) Goldblatt &

J.C. Manning, comb, et stat. nov.

Tritonia subgenus Dichone Salisb. ex Baker, Hand-

book of the Irideae: 190 (1892). Ixia subgenus Dichone

(Salisb. ex Baker) G.J. Lewis: 159 (1962). Type: 7. eru-

bescens Goldblatt (= 7. crispa L.f.).

Subgenus Morphixia (Ker Gawl.) Goldblatt &

J.C. Manning, comb, et stat. nov.

Morphixia Ker Gawl., Iridearum genera: 105 (1827).

Ixia section Morphixia (Ker Gawl.) Pax: 154 (1888).

G.J.Lewis: 66 (1962). Type: 7. latifolia D.Delaroche.

Ixia subgenus Hyalis Salisb. ex Baker: 161 (1892) (?

in part?). Ixia section Hyalis (Salisb. ex Baker) Diels:

486 (1930). Type: 7. paniculata D.Delaroche.

NOMENCLATURAL NOTE IN GLADIOLUS

The name G. lithicola, used for a local endemic restrict-

ed to the Mpumalanga escarpment. South Africa, described

by Goldblatt & Manning (1998) is a homonym for another

species from Ethiopia (Goldblatt 1996). We propose the

new name, G. saxatilis for the Mpumalanga plant. The epi-

thet alludes to the mostly lithophytic habit of the species.

Gladiolus saxatilis Goldblatt & J.C. Manning, nom.

nov., pro G. lithicola Goldblatt & J.C.Manning, Gladiolus

in southern Africa 82 (1998), horn, illeg. non G. lithicola

Goldblatt, Gladiolus in tropical Africa 284 (1996).

REFERENCES

BAKER, J.G. 1892. Handbook of the Irideae. Bell, London.

BOLUS, H.M.L. 1923. Novitates Africanae. Annals of the Bolus Her-

barium 3: 70-85.

DIELS, F.L.E. 1930. Iridaceae. Die naturlichen Pflanzenfamilien, edn

2, 15a: 463-505.

GOLDBLATT, R 1992. Phylogenetic analysis of the South African

genus SparcLxis (including Synnotia) (Iridaceae: Ixioideae), with

two new species and a review of the genus. Annals of the

Missouri Botanical Garden 79: 143-159.

GOLDBLATT, P. 1996. Gladiolus in tropical Africa. Timber Press,

Portland, Oregon.

GOLDBLATT, P. & MANNING. J.C. 1998. Gladiolus in southern

Africa. Femwood Press, Cape Town.

GOLDBLATT, P, BERNHARDT, P. & MANNING, J.C. 1998. Polli-

nation of petaloid geophytes by monkey beetles (Scarabaeidae:

Rutelinae: Hopliini) in southern Africa. Annals of the Missouri

Botanical Garden 85: 215-230.

KER GAWLER, J.B. 1827. Iridearum genera. Brussels.

LEWIS, G.J. 1956. A revision of the genus Synnotia. Annals of the

South African Museum 50: 1 37- 151.

LEWIS, G.J. 1962. South African Iridaceae. The genus Ixia. Journal <f

South African Botany 28: 45-195.

MANNING, J.C. & GOLDBLATT, P. 1996. The Prosoeca peringueyi

{Diptera: Nemestrinidae) pollination guild in southern Africa:

long-tongued flies and their tubular flowers. Annals of the Mis-

souri Botanical Garden 83: 67-86.

PAX, FA 1888. Iridaceae, Die naturlichen Pflanzenfamilien, edn 2,5:

137-158.

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Bothalia 29,1: 65-75 (1999)

The genus Amphiglossa (Gnaphalieae, Relhaniinae, Asteraceae) in

southern Africa

M. KOEKEMOER*

Keywords; Amphiglossa DC., Asteraceae, comb, nov., Gnaphalieae, Pterothrix DC., Relhaniinae, southern Africa, sp. nov., taxonomy

ABSTRACT

It is proposed that the generie concept of Amphiglossa DC. be broadened to include the genus Pterothrix DC. The two gen-

era were previously separated on the presence or absence of ray florets. A thorough study of various characters showed that

the type of florets was overemphasised in the past and that the pappus and cypselas provide sufficient evidence to unite the two

genera. The genus consists of 1 1 species of which Amphiglossa celans Koekemoer, A. grisea Koekemoer, A. rudolphii

Koekemoer and A. susaimae Koekemoer are newly described. A key to the species as well as detailed descriptions and distri-

bution maps are provided. Photographs are included for each species and key characters for the genus are also illustrated.

INTRODUCTION

Formerly the two genera, Amphiglossa DC. and

Pterothrix DC., were poorly known and distinguished by

the absence or presence of ray florets. Even De Candolle

(1838) was not always sure to which genus certain

species belonged, hence his description of A. perotri-

choides DC. for a species without ray florets. A thorough

study of various characters showed that the type of flo-

rets was overemphasised in the past and that the pappus

and cypsela provide sufficient evidence to unite the two

genera. The genus consists of 1 1 species, four of which

are newly described here.

The extensive fieldwork I undertook over a period of

six years during different seasons, to cover the different

flowering times, was very valuable in revising this

group. Not only were four new species discovered but

another three, that were mainly known from type mater-

ial, were re-discovered after earlier collections by Ecklon

& Zeyher and Drege.

Nine species in the genus are characterised by very

restricted distributions in Namibia and the Northern,

Western and Eastern Cape of South Africa. Two other

species, A. tomentosa (Thunb.) Harv. and A. triflora DC.

have wider distributions in the other South African

provinces and Namibia.

Specimens were studied in BM, BOL, BR, E, G, G-DC,

K, LINN, NBG, P, PRE, S, SAM, SBT, TCD, TUB, UPS

and WAG (abbreviations as in Holmgren et al. 1981).

Many of the older collections are of a very poor quality

and were of limited value in the revision.

Amphiglossa DC., Prodromus 6: 258 (1838);

Benth; 324 (1873); E. Phillips: 799 (1951). Type:

Amphiglossa corrudifolia DC.

* National Herbarium, National Botanical Institute, Private Bag XlOl,

0001 Pretoria.

MS. received; 1998-10-19.

Pterothrix DC.: 279 (1838); Benth.: 324 (1873);

E. Phillips: 798 (1951). Type: Pterothrix spinescens DC.

Perennial, woody, much-branched, wiry shrubs or

dwarf shrubs, erect or decumbent, stoloniferous or rhi-

zomatous. Branches glabrous or with a thin layer of

arachnoid hairs, tips rarely spinescent. Leaves small,

imbricate or widely spaced, often in fascicles, sessile or

inconspicuously stalked, ericoid, convex, abaxial surface

arachnoid or glabrous, adaxial surface woolly, margins

smooth, tips mucronate, acute or rounded. Capitula radi-

ate or discoid, terminal, sessile, solitary, or in small

groups of distinct units, homogamous or heterogamous;

ray florets 3-12, bilabiate or pseudobilabiate florets

female, disc florets bisexual, 5-lobed. Involucral bracts

in several series, outermost chaffy or foliaceous, inner-

most longer, narrower. Receptacle discoid, nude or with

small extensions around cypsela bases. Anthers 5, synge-

nesious, apically acuminate, basally tailed. Style bifid,

stigmatic area in two strips along margins of style arms,

inflated cells at branch tips short, rounded. Nectary pre-

sent in all florets, a small disc between style base and

cypsela. Cypselas small, asymmetrical, glabrous, papil-

lose. Pappus bristles 1 8-32, densely plumose from tip to

base, in one series, free or rarely fused at the very base.

Flowering time', mainly November-March, some species

in spring-summer and others in summer-autumn.

Amphiglossa is recognised by its small, sessile capit-

ula that are terminally borne, solitary or in small groups,

a chaffy involucre, papillose cypselas (Figure lA, B) and

pappus of many fine bristles (Figure 1C, D) that are pro-

fusely plumose and interlocking in the central parts.

Amphiglossa also has a germination strategy unique in

the Relhaniinae. Fresh, mature cypselas germinate with-

in 24 hours by means of a basal hygroscopic substance

that absorbs water very quickly, causing the seedcoat to

burst and thereby facilitating germination. A gelatinous

blob (Figure IE, F), about the size of the swollen

cypsela, then surrounds the root tip.

Amphiglossa belongs to the subtribe Relhaniinae with

which it shares adaxially tomentose leaves. The

Relhaniinae are subdivided further by flower colour: the

66

Bothalia29,l (1999)

FIGURE 1. — Diagnostic characters of Amphif’lossa. A, cypsela of A. rudolphii, x 45; B, papillose cypsela surface of A. corrudifolia, x 820; C,

plumose pappus tips of A. griseu, x 40; D, interlocking pappus bristles of A. rudolphii, x 25; E, F, gell substance released to aid germina-

tion in A. rud(dphii and A. tomentosa respectively, x 20.

Bothalia29,l (1999)

67

yellow-flowered genera are referred to the Relhania

group and the white/pink-flowered genera to the

Metalasia group. In Anderberg’s (1991) cladistic analy-

sis, Amphiglossa groups terminally with Disparago

Gaertn., Stoebe L. and Elytwpappus Cass. The other

related genera in the Metalasia group are: Hydroidea

Karis, Lachnospermum Willd., Phaenocoma D.Don,

Planea Karis, Atrichantha Hilliard & Burtt, Bry’omorphe

Harv., Calotesta Karis, Dolicothrix Hilliard & Burtt and

Metalasia R.Br. (Anderberg 1991).

Diagnostic characters: pappus of 18-32 very fine

bristles, densely plumose from close to base to tip, inter-

locking and released en masse with seeds; cypselas

papillose.

Distribution and habitat: Namibia, Western, Eastern

and Northern Cape, Free State, Lesotho, North-West,

Gauteng and Mpumalanga. Dry, rocky and sandy areas,

often in deep sand or on granite outcrops.

Conservation status: most species not threatened.

Key to the species of Amphiglossa

la Ray florets present:

2a Brachyblasts numerous, very obvious, to 15 mm long;

leaves imbricate;

3a Ray florets with 1 or 2 small lobes in sinus A. callunoides

3b Ray florets without lobe in sinus A. cornidifolia

2b Brachyblasts, if present, short; leaves in fascicles, not imbri-

cate:

4a Ray floret lamina curled like a watch-spring, strap-

shaped; capitula 6. 5-7. 5 x 2. 0-3.0 mm A. grisea

4b Ray flower lamina open and spreading, ovate; capitula

8.5-10.0 X 2. 5-3. 5 mm A, tomentosa

lb Ray florets absent;

5a Capitula with more than 10 florets; restricted to white sand

dunes in the Witsand complex near Postmasburg .... A. tecta

5b Capitula with 4 or 5 florets; distribution not as above;

6a Some terminal branches modified into spines; distributed

outside the Western Cape A. triflora

6b Spines absent; distributed in Namibia and in the Western

Cape:

7a Leaves minute, adpressed; plants known from the

Kovis Mountain near Liideritz (Namibia) and pos-

sibly also granite mountains further south A. thuja

7b Leaves longer than 5 mm, usually spreading; plants

distributed in Western Cape (A. celans in the

Caries area):

8a Plants wiry, sparsely branched and sparsely leaved,

often scrambling; known from a small area west

of Caries A. celans

8b Plants cushion-like shrubs or dwarf shrubs; distrib-

uted in the Western Cape:

9a Leaf tips rounded; involucre with glossy, yellow

sheen and small tomentose patches on abaxial

surface; restricted to the Ceres Karoo (Kagga

Kamma and Katbakkies areas north of Ceres)

A. susannae

9b Leaf tips acuminate; involucre chaffy or greenish

with tomentose abaxial surface; distributed to

the north, west and south of the Ceres Karoo:

10a Shrubs taller than 0.3 m; leaves widely spaced,

fascicles rare; involucre tips acute A. rudolphii

10b Shrublets smaller than 0.3 m; leaves not so

widely spaced, in fascicles; involucre tips

aristate A. perotrichoides

Amphiglossa callunoides DC., Prodromus 6: 259

(1838). Type: Eastern Cape, 3325 (Port Elizabeth):

District Uitenhage, in the stony channel of the Zwartkops

River, fl. Jan. 1835, Ecklon 51 (G-DC, lecto.!, here des-

ignated). Figure 2B.

Stoloniferous, woody, multistemmed, blue-grey

shrubs, to 0.5 m tall. Branches slender and very flexible,

erect at first, bending over and becoming stoloniferous,

rooting where they touch the ground; older plants form-

ing well-anchored bushes, covering an area of up to 6 m^;

young active shoots up to 3 mm diam., older, main

branches 1. 5-2.0 mm diam. Primary brachyblasts devel-

oping in every leaf axil on main stems, numerous sec-

ondary brachyblasts developing in leaf axils of primary

brachyblasts; stems of brachyblasts and young branches

often white-tomentose; brachyblasts mostly negatively

geotropic, whereas main branches positively geotropic,

thus creating the impression that secondary branching is

restricted to one side of the main branch. Leaves persis-

tent, sessile, adpressed, narrowly triangular, margins

entire, slightly inrolled, midrib prominent, adaxial surface

woolly, abaxial surface smooth, tips shortly mucronate,

young leaves yellowish green, older leaves fading to grey,

giving a characteristic blue-grey sheen to plants; subtend-

ing leaves on main stems alternate, up to 10 mm long and

2 mm wide at base, about a leaf length apart, leaving

branch exposed; brachyblast leaves minute, up to 2 mm

long, imbricate, subtending leaves on brachyblasts not

much longer than ordinary leaves. Capitula radiate, het-

erogamous, 7-8 x 2-3 mm, 8-10-flowered with the same

number of ray and disc florets; single head terminally

borne on brachyblast, linear when young, involucre

spreading at maturity to release pappus and seeds. Outer

involucral bracts 10-15, foliaceous in upper part and

along midvein, remaining green around midrib, in 3-6

series, ovate, 2. 5-4. 5 mm long, increasing in length from

outer to inner series, the outer about half the length of

innermost bracts. Inner involucral bracts scarious along

margins, with multicellular glands, 10-15, in 4 or 5

series, 6.0-7. 5 mm long, linear to narrowly oblong, hya-

line towards apex, those in innermost series hyaline in

upper half, mucronate, as long as pappus. Receptacle flat-

tened, less than 2 mm diam., honeycombed, ridges short-

er than 1 mm. Bilabiate or pseudobilabiate florets white,

female, 4 or 5; lamina ± 1.5 x 0.5 mm, 3-lobed; tube

4. 5-5.0 mm long. Disc florets white, bisexual, 5-lobed,

5. 0-5. 5 mm long. Cypselas subterete, ± 1 mm long,

chestnut-coloured, inconspicuously ribbed, papillose,

developing in 3-5 florets, more often in bilabiate or

pseudobilabiate florets; released in a cluster together with

pappus at maturity. Pappus bristles 18-23, ± 5 mm long,

barbed in lower fifth, plumose in upper part, seta up to 1.5

mm long. Elowering time: January-April.

Diagnostic characters: stoloniferous growth form,

numerous brachyblasts; leaves very small; capitula

small, terminal on brachyblasts; small white florets just

visible at tip of involucre.

Distribution and habitat: known from two localities

only: a short section of the Kwa-Zunga River (inland part

of Swartkops River), NW of Uitenhage between

Springfontein and Rooikrantz Farms, and the Farm

Hoeree in the Kouga Mountains, NW of Joubertina

(Figure 3). Observed growing in abundance amongst

boulders in the bed of the Kwa-Zunga River.

68

Bothalia 29,1 (1999)

FIGURE 2. — Branches, leaves and

capitula of selected species. A,

Amphiglossa celans; B, A.

callutioides', C, A. grisew, D,

A. rudolphiv, E, A. susannae;

F, A. tecta; G, A. thuja; H, A.

tomentosa; 1, A. triflora. Scale

bars: 10 mm.

Conservation status; rare, but locally abundant from

two localities only; threatened by sandmining activities in

the riverbed near Uitenhage and also by natural flooding.

N umber of collections known: 25.

Vouchers: Ecklon 45; Ecklon ct Zeyher 2916; Koekemoer 1148;

Oelofsen 171.

Amphiglossa celans Koekemoer, sp. nov., suffru-

tex scandens multicaulis leniter lignosus, vel volubilis,

ad 0.3 m altus. Capitula homogama, solitaria, terminalia,

quinqueflora. Bracteae involucrales 12-17, stereoma in

parte centrali supero perspicue manifestum. Flosculi

tubulosi albi vel rosei, hermaphroditi. Cypselae sub-

teretes, in floribus omnibus perfecte evolutae. Pappi

setae 18-26, in fundo connatae, basin versus hamatae,

hamulis implicantibus. Florescentia Februarii-Aprilis.

Adhuc ex area parva inter pagos Garies et Kotzesrus, in

provincia Capensi Boreali nota.

TYPE. — Northern Cape, 3017 (Hondeklipbaai): 35 km

from Garies to Kotzesrus, (-DD), 1995-11-20, Koekemoer

1099 (PRE, holo.; MO, NBG, S, iso.). Figure 2A.

The specific epithet ‘celans’ (= hiding) refers to the

unobtrusive manner in which this plant grows amongst

other plants.

Scrambling, multistemmed, weakly woody subshrub

or scrambler, up to 0.3 m tall. Stems up to 2 mm diam.,

branches bare or very sparsely leaved in lower part, grey-

Bothalia29,l (1999)

69

ish or brick-red with a shiny appearance, often with

zigzag branching pattern. Leaves sessile, ascending,

elliptic to linear, margins entire, inrolled, twisted, adaxi-

al surface woolly, tips shortly mucronate, up to 8.0 x 1.5

mm, seemingly caducous after growing season, greyish

green, very sparse on most branches, ± half a leaf length

apart on terminal branches; young, active shoots up to 3

mm diam., large-leaved, leaves flat, up to 16 x 6 mm.

Capitiila discoid, homogamous, solitary in last 3 or 4 ter-

minal leaf axils, 7-8 x 2-3 mm, 5-flowered. Involucral

bracts 12-17, up to 8.0 x 1.3 mm, stereome clearly visi-

ble in central upper part, outermost bracts foliaceous,

joined together by hairs on outside, innermost bracts

scarious along margins, tinged dark wine-red towards

slightly recurved, mucronate apex. Receptacle conical,

less than 1 mm diam., honeycombed, ridges shorter than

1 mm. Florets white or pink, bisexual, 5-lobed, 4. 5-5.0

mm long. Cypselas subterete, ± 1 mm long, inconspicu-

ously ribbed, papillose, annulus not distinct. Pappus

bristles 18-26, connate at very base, ± 4 mm long,

barbed in lowermost part with barbs interlocking, short-

ly plumose along shaft ending in tufted tip, seta up to 0.2

mm long. Flowering time: February-April.

Diagnostic characters: wiry branches; small solitary

capitula; dark-tipped involucre.

Distribution and habitat: poorly known, currently

only from a small area between Garies and Kotzesrus

(Figure 3). ‘Strandveld’, deep sand, with sparse vegeta-

tion, of predominantly mesembryanthemous and astera-

ceous elements that seldom grow taller than 500 mm.

Conservation status: rare.

Number of collections known: 2.

Vouchers: Koekemoer 1099, 1100.

Amphiglossa corrudifolia DC., Prodromus 6: 258.

(1838). Type: in Africa australi ad Zwarte Camisberge et

Onder Bol^eweld, alt. 2-300 ped, Drege 2723 (G-DC,

lecto.!, here designated, BM!; P!; PRE, photo.!).

Erect, rhizomatous, chaffy-green shrubs, up to 0.4 m

tall. Branches rigid, up to 4 mm diam., secondary

branches ascending, to 2 mm diam., developing in each

leaf axil on main stem; older plants forming well-

anchored bushes; stems becoming rhizomatous and pro-

ducing copious shoots when covered by sand deposits

during floods. Brachyblasts developing in each leaf axil

of secondary branches, generally short, up to 5 mm

long, bearing capitula close to stems; secondary brachy-

blasts rare. Young branches and stems of brachyblasts

often white-tomentose. Leaves bright green, sessile,

adpressed, triangular, margins entire, slightly inrolled,

midrib prominent, adaxial surface woolly; subtending

leaves to secondary branches on main stems alternate,

up to 11 mm long, 4 mm wide at base, ± a leaf length

apart, leaving branch exposed, browning with age and

then often reflexed; subtending leaves on secondary

stems much shorter, up to 5 mm long, 1.5 mm wide at

base, becoming brown in older parts, ascending to

inclined; leaves on brachyblasts minute, up to 2 mm

FIGURE 3. — Geographical distribution of A. calluru)ide.<:, •; and -4.

cekms, ♦.

long, imbricate. Capitula radiate, heterogamous, 7-8 x

2-3 mm, 8-10-flowered, with same number of ray and

disc florets; single heads terminally borne on short

brachyblasts, linear when young, involucre spreading at

maturity to release pappus and seeds. Outer involucral

bracts 10-15, 2. 5-4. 5 mm long, foliaceous in upper part

and along midvein, in 3-6 series, ovate, increasing in

length from outer to inner series, the outer about half the

length of innermost bracts. Inner involucral bracts scar-

ious along margins, 10-15, 6. 0-7. 5 mm long, in 4 or 5

series, linear to narrowly oblong, those in outer series

remaining green around midrib, hyaline towards apex,

those in innermost series hyaline in upper half,

mucronate, as long as pappus. Receptacle flattened, less

than 2 mm diam., honeycombed, ridges shorter than 1

mm. Ray florets white, female, 4 or 5, lamina shorter

than 1 mm, narrower than 0.5 mm, tapering towards tip,

1-3-lobed; lobes equal or very uneven; tube 4.5-5.0 mm

long. Disc florets white; tube often tinged with pink

below lobes, bisexual, 5. 0-5. 5 mm long, 5-lobed; lobes

often asymmetrical. Cypselas subterete, ± 1 mm long,

greyish, inconspicuously ribbed, papillose, developing

in 3-5 florets, more often in ray florets, released in a

cluster at maturity, with pappus interlocking. Pappus

bristles 18-23, ± 5 mm long, barbed in lower fifth,

plumose in upper part, seta up to 1.5 mm long.

Flowering time: January-April.

Diagnostic characters: rhizomatous; growing in

sandy riverbeds; leaves small, triangular; brachyblasts

numerous; capitula small, solitary; pappus fluffy.

Distribution and habitat: known from the Farm Klein

Sleutelfontein in a dry riverbed in the Houtrivier, a trib-

utary of the Sand River, near Prince Albert and from an

unknown locality of Drege in the ‘Bockland’ (Figure 4).

In deep shale-derived soils, in river or sand beds amongst

grassy tufts. This species was recollected in September

1998 in the same area where Drege originally found it.

Although it appears to be quite rare, an extensive search

in the area is needed to establish the extent of the distri-

bution.

Conservation status: rare, but well-conserved on pro-

tected farmland; could be endangered by flooding.

70

Bothalia 29,1 (1999)

FIGURE 4. — Geographical distribution of A. corrudifolia, •; and A.

grisea, ♦.

Number of collections known: 11.

Vouchers: Drege 2723: Kuekemoer 1291, 1380.

Amphiglossa grisea Koekemoer, sp. nov., frutex

compactus multicaulis lignosus caesiellus, ad 0.3 m

altus. Folia juvenia glauca, seniora brunnescentia.

Capitula heterogama, 9-flora, cum flosculis 3 ligulatis et

6 tubulosis. Bracteae involucrales exteriores 8-14, in

parte superiori et secus nervum medium foliaceae.

Flosculi radii albi, feminei, lamina ±1.5 mm longa,

plerumque in cochleam voluta. Flosculi tubulosi albi,

hermaphroditi. Cypselae subteretes, ectocarpio libero.

Pappi setae 15-20, in quinta parte basali hamatae, supra

plumosae. Florescentia Januarii-Martii. Inter pagos

Nieuwoudtville et Clanwilliam, in provincia Capensi

Occidentali nota.

TYPE. — Western Cape, 3118 (Vanrhynsdorp):

Nardouwsberg, ‘Brand se Berg’, turn-off from R363

road, top of pass, about 4 km after crossing the

Doringrivier, (-DC), 1995-11-21, Koekemoer 1106

(PRE, holo.; MO, NBG, S, iso.). Figure 2C.

The specific epithet 'grisea’ refers to the blue-grey

colour of these shrubs.

Compact, multistemmed, woody blue-grey shrub, up

to 0.3 m tall. Branches slender, up to 2 mm diam., entan-

gled, bare in lower parts, new growth branching copi-

ously at short intervals; secondary branches often white-

tomentose, leafy, often with zigzag branching pattern;

tertiary branches frequently at right angles but soon

bending upwards, giving a candelabrum appearance.

Brachyblasts absent or reduced to leafy clusters. Leaves

up to 6 X 1 mm, sessile, adpressed, linear, margins entire,

slightly inrolled, midrib prominent, adaxial surface

woolly, young leaves greyish green, older leaves brown-

ish. Capitula radiate, heterogamous, 6. 5-7. 5 x 2-3 mm,

9-flowered, with 3 ray and 6 disc florets; single heads

terminally borne on tertiary branches, linear when

young, involucre spreading at maturity to release pappus

and seeds. Outer involucral bracts 8-14, 2.5— 4.5 mm

long, foliaceous in upper part and along midvein, ovate,

increasing in length from outer to inner series, outer

about half the length of innermost bracts. Inner involu-

cral bracts scarious, 15-20, 5. 0-6.0 mm long, linear to

narrowly oblong. Receptacle flattened, less than 2 mm

diam., honeycombed, ridges shorter than 1 mm. Ray flo-

rets white, female, lamina ±1.5 mm long, characteristi-

cally rolled like a watch-spring, 3-lobed. Disc florets

white, bisexual, 5-lobed, up to 4 mm long with few

glands on outer surface of lobes. Anthers apically trun-

cate. Cypselas subterete, ± 1 mm long, greyish, incon-

spicuously ribbed, papillose, with loose ectocarp, but

ecto- and endocarp seemingly attached at cypsela base.

Pappus bristles 15-20, ± 3.5 mm long, barbed in lower

fifth, plumose in upper part, seta up to 1 mm long, bris-

tles free, truncate and tufted at apex. Flowering time:

January-March.

Diagnostic characters: compact grey bush; small,

unobtrusive capitula; lamina of ray florets rolled like a

watch-spring.

Distribution and habitat: Western Cape between

Nieuwoudtville and Clanwilliam (Figure 4). Sandy flats

and mountain foothills.

Conservation status: not threatened; thrives on distur-

bances.

Number of collections known: 6.

Vouchers: Hugo 1027: Koekemoer 588, 1106, 1111, 1215.

Amphiglossa perotrichoides DC., Prodromus 6:

258. (1838). Type: Western Cape, Swellendam Div.,

Burchell 7499 (K, lecto.!, here designated; PRE!).

Pterothrix perotrichoides (DC.) Harv.: 275 (1894), syn. nov.

Woody, multistemmed grey shrub, up to 0.3 m tall.

Main branches up to 1.5 mm diam., long, slender and

very flexible, erect at first, bending towards ground, pos-

sibly becoming stoloniferous where they touch soil; sec-

ondary branches developing from base upwards, creating

impression that branching is restricted to only one side of

main branch. Brachyblasts reduced to short leafy clus-

ters. Leaves sessile, adpressed or ascending, linear, up to

10 mm long, margins entire, slightly inrolled, adaxial

surface woolly; subtending leaves 8-16 x 1 mm.

Capitula discoid, homogamous, 8-9 x 2-3 mm, 4-5-

flowered, borne individually in the 4-8 terminal leaf

axils of secondary branches. Involucral bracts 16-20,

outer bracts ± 3.5 mm long, inner bracts up to 8 mm long,

foliaceous bracts absent or in some florets restricted to 1

or 2 outermost bracts, basally ovate with aristate tips,

upper part hyaline and spreading somewhat at maturity.

Receptacle flattened, less than 2 mm diam., honey-

combed, ridges shorter than 1 mm. Disc florets white,

bisexual, 5-lobed, often asymmetrical, 5. 0-5. 5 mm long.

Anther tails short and stout. Cypselas terete, ± 1 mm

long, without visible ribs, papillose, annulus fairly well

developed. Pappus bristles 25-30, ± 5 mm long, barbed

in lower fifth, plumose in upper part, seta up to 1.5 mm

long. Flowering time: December-January.

Bothalia29,l (1999)

71

FIGURE 5. — Geographical distribution of A. pewtrichoides, •.

Diagnostic characters', homogamous heads; involucre

reflexed in upper third, tips aristate.

Distribution and habitat, known from Swellendam

area and Cederberg (Figure 5). Coastal and Mountain

Rhenosterveld.

Conservation status: probably endangered, together

with the threatened Renosterveld.

Number of collections known: 8.

Vouchers: Acocks 23237\ Burchell 7499', Koekenwer 602, 608.

Amphiglossa rudolphii Koekemoer, sp. nov., fru-

tex virgatus robustus ad 1 .2 m altus et 1 m diametro. Rami

laeves perfragiles, maxime implexi, fruticem densum,

sparse foliatum formantes. Folia breve petiolata.

Capitula homogama, semper quinqueflora. Bracteae

involucrales 30-35, plerumque in pagina adaxiali tomen-

tosae. Flosculi tubulosi albi, hermaphroditi. Cypselae

solum in uno e 5 flosculis evolutae, subteretes, asymmet-

ricae, griseolae, ectocarpio libero. Pappi setae 20-28,

basin versus perbreviter hamatae, supra plumosae.

Florescentia Februarii-Junii. Solum ex areis paucis in

provincia Capensi Occidentali nota: Botterkloof, ad meri-

diem pagi Nieuwoudtville, ad Loeriesfontein et prope

Worcester.

TYPE. — Western Cape, 3119 (Worcester): 50 km S of

Nieuwoudtville on R364 road to Clanwilliam, 1 km N of

turn-off to Farm Lasvlei, (-CD), 1996-03-20, Koekemoer

1210 (PRE, holo.; NBG, S, MO, BOL, iso.). Figure 2D.

The specific epithet of this plant is to commemorate

my father, Rudolph, in gratitude for his patient assistance

in collecting specimens and seed of this species.

Wiry, robust shrub up to 1.2 m tall and 1 m wide.

Branches smooth and very brittle, rarely more than 5 mm

diam., very much entangled to form dense, sparsely-

leaved bushes; older branches bare with conspicuous leaf

scars. Leaves very shortly petiolate, ascending, linear to

narrowly oblong, twisted ± 270°-360°, greyish green,

margins entire, slightly inrolled, adaxial surface woolly.

generally 6-10 x 1-2 mm, on new shoots to 30 x 5 mm,

probably persistent for one season only. Brachyblasts

generally absent, except on new, actively growing

branches and then only 2-^-leaved shoots. Capitula dis-

coid, homogamous, 7-10 mm long, 1-2 mm wide, con-

sistently 5-flowered; heads developing in the 2-7 termi-

nal leaf axils of a branch, linear when young, involucre

spreading in upper half at maturity, lower part remaining

tubular. Involucre with 30-35 bracts, bearing character-

istically tomentose markings on adaxial surfaces, fairly

uniform in texture and appearance, becoming progres-

sively longer from outer to inner bracts. Outer involucral

bracts scale-like, 1. 5-2.0 mm long and almost as wide,

thus very widely ovate in shape, following series widely

ovate, up to 2.5 mm long, next two series ovate, up to 4

mm long and 5 mm long respectively, innermost series

lanceolate to linear, up to 8 mm long. Inner involucral

bracts 10-15, in 4 or 5 series, scarious along margins,

with multicellular glands, 6.0-7. 5 mm long, linear to nar-

rowly oblong, those in outer series remaining green

around midrib, hyaline towards apex, those in innermost

series hyaline in upper half, mucronate, as long as pap-

pus. Receptacle flattened, less than 2 mm diam., honey-

combed, ridges shorter than 1 mm. Disc florets white,

bisexual, 5-lobed, 6-7 mm long; lobes 0.8-1. 0 mm long.

Anthers basally feathery-tailed. Cypselas subterete,

asymmetric, ± 1 mm long, greyish coloured, papillose,

indistinctly ribbed, with 3 ribs visible, developing in only

1 of the 5 florets; ectocarp loose, easily removable, 1 cell

layer thick; endocarp darker and thicker, forming a cavi-

ty filled with a hygroscopic gel-like substance, which

expands rapidly when it comes into contact with water,

causing the seedcoat to burst; cotyledons dark purplish;

released in a cluster together with entangled pappus at

maturity. Pappus bristles 20-28, ± 5.5 mm long, barbed

for a very short distance at base, plumose in upper part,

seta up to 1 mm long. Flowering time: February-June.

Diagnostic characters: large bush; branches brittle;

leaves widely spaced; capitula small, terminal on branch-

es; involucre spreading to release fluffy pappus; usually

only one fairly large, subterete seed per capitulum.

Distribution and habitat: known from a few areas

only: Botterkloof, south of Nieuwoudtville, Loeries-

fontein and Worcester area (Figure 6). Rocky terrain in

Mountain Renosterveld.

Conservation status: not threatened.

Number of collections known: 5.

Vouchers: Cunradie s.n.\ Hartman 12947\ Koekemoer 1210, 1211,

1214.

Amphiglossa susannae Koekemoer, sp. nov., fru-

tex virgatus multicaulis lignosus fumosus, ad 0.5 m altus.

Apices foliorum rotundati. Capitula homogama, quin-

queflora, nitore claro flavo-viridi. Bracteae involucrales

20-28, cum strictura circa quinta parte infra apicem;

bracteae intimae apicibus cochlearibus. Flosculi tubulosi

albi, hermaphroditi. Cypselae subteretes, badiae, ectocar-

pio ut videtur libero. Pappi setae 20-26, in quinta parte

basali hamatae, supra plumosae. Florescentia Novembris

72

Bothalia29,l (1999)

FIGURE 6. — Geographical distribution of A. rudolphii, •; and A.

tecta, ♦.

Decembrisque. Crescit in provincia Capensi Occidentali,

in planitie alta mentis Swartruggens.

TYPE. — Western Cape, 3219 (Wuppertal); Kagga

Kamma Nature Reserve, SE of chalets in red sand,

(-DA), 1995-11-24, Koekemoer 1115 (PRE, holo.; MO,

NBG, BOL, K, S, iso.). Figure 2E.

The specific epithet of this plant is to commemorate

my mother, Susan, who very enthusiastically helped to

collect specimens at Kagga Kamma.

Wiry, erect, multistemmed, woody, yellowish grey

shrub, up to 0.5 m tall. Stems up to 2 mm diam., bare in

lower parts. Leaves 3-7 mm long, sessile, mostly soli-

tary, but sometimes in small fascicles in young growth,

ascending, linear, margins entire, slightly inrolled, adax-

ial surface woolly, leaf tips rounded; young leaves yel-

lowish green, older leaves fading to dark grey and drop-

ping off easily. Capitula discoid, homogamous, 6-7 x 2

mm, 5-flowered, terminal, solitary or in groups of up to

4, their bright, greenish yellow colour contrasting with

the darker leaves. Involucre of 20-28 bracts, increasing

in length from outer to inner series, 5. 5-6.0 mm long,

with a constriction at about one fifth from tip and yel-

lowish below constriction. Outer involucral bracts

broadly ovate, with acute hyaline tips, not foliaceous,

following series with hyaline part above constriction,

gradually increasing in width and resulting in a membra-

nous, spoon-shaped tip in innermost bracts. Receptacle

conical, less than 2 mm diam., honeycombed, ridges

shorter than 1 mm. Disc florets white, bisexual, 5-lobed,

4. 5-5.0 mm long, outer lobes of closed florets very glan-

dular. Cypselas subterete, ± 1 mm long, chestnut-

coloured, inconspicuously ribbed, papillose, appearing to

have a loose ectocarp, with a small attachment area

between endo- and ectocarp at the base. Pappus bristles

20-26, ± 4.5 mm long, barbed in lower fifth, plumose in

upper part, seta up to 1.5 mm long. Flowering time:

November and December.

Diagnostic characters: yellowish sheen to leaves and

capitula; terminal heads small; involucre yellowish in

lower parts with tips hyaline; innermost involucre with

spoon-shaped tips.

Distribution and habitat: on plateau on Swartruggens

Mountain in the Ceres-Karoo (Figure 7). Deep red or

white sand in level areas between TMS outcrops.

Conservation status: not threatened.

Number of collections known: 5.

Vouchers: Esterhuysen 29338: Koekemoer 609, 1115, 1116: Taylor

12106.

Amphiglossa tecta (FBrusse) Koekemoer, comb.

nov.

Pterothrix tecta F.Brusse in Bothalia 20: 67 (1990), syn. nov. Type:

Northern Cape, 2822 (Glen Lyon): Hay Dist., Witsand, some 70 km

SW of Postmasburg, (-CB), 1989-11-26, Brusse 5629 (PRE, holo.!;

AD, B, BAF, BH, BM, BOL, BR, BRI, C, CAN, CANB, COI, E, EA,

G, GH, GRA, HBG, J, Kl, L, LD, LG, LlSU, LMA, M, MEL, MO,

NBGI, NH, NSW, NU, O, P, R, SI, SRGH, U, UC, UPS!, US, W,

WAG!, WIND, Z, iso.). Figure 2F.

Erect, robust, single or multistemmed shrub, to 0.6 m

tall, aromatic. Stems gnarled, dark grey, up to 40 mm

diam., often buried in sand towards base; secondary

branches up to 5 mm thick, bark thin, golden chestnut-

brown to reddish brown or reddish in places, glabrous or

with golden flakes pealing off; tertiary branches white-

tomentose when young, becoming golden-yellow,

glabrous and peeling when older. Leaves alternate, ses-

sile, linear, 1.0-11.0 x 0.7-2. 2 mm, broadest at base,

ascending but spreading in upper half, margins involute;

upper surface canaliculate, woolly-white; abaxially

glabrous, epidermis sloughing off and becoming follicu-

late, especially at base; midrib raised, particularly at

base, apex obtuse, mucro very small, inconspicuous.

Capitula discoid, solitary, terminal, sessile, homoga-

mous, discoid, cylindrical, 7. 0-8.0 x 2. 5-3. 5 mm,

10-12-flowered. Involucral bracts 20-26, becoming

progressively longer inwardly, tips hooded, pellucid,

bronze, innermost bracts glabrous, lanceolate, up to 6.0 x

O. 8-1 .0 mm, remaining bracts ovate, 1. 5-6.0 x 1. 5-2.0

mm, stereomes entire, with conspicuous white-tomen-

tose patches at apices. Receptacle honeycombed, ridges

between cypsela bases shorter than 1 mm. Disc florets

FIGURE 7. — Geographical di.stribution of A. susannae, •; and A.

thuja, ♦.

Bothalia 29,1 (1999)

73

bisexual and fertile, ± 7 mm long; corolla white or pale

mauve from above, ± 5.5 mm long, 5-lobed; tube cylin-

drical and widening towards apex, 5 mm long, purplish

above, becoming pale greenish below; lobes deltoid,

acute, spreading. Nectary present. Cypsela subterete,

1.0-1. 8 mm long, glabrous, papillose, cream-coloured,

5-ribbed, apex with thickened ring, does not seem to

have a loose ectocarp or cavity filled with gel-hke sub-

stance. Pappus bristles free, plumose, but base barbed,

3. 5-4. 5 mm long. Flowering time: mainly November,

but also recorded in January, March and July.

Diagnostic characters: capitula 10-1 2-flowered;

involucre with conspicuous white tomentose patches

abaxially; innermost bracts with tips hooded; restricted

to Witsand.

Distribution and habitat: it has a restricted distribu-

tion in the northwestern area of the Witsand dune com-

plex, 70 km southwest of Postmasburg in the Northern

Cape (Figure 6). White sand dunes of Aeolian origin,

deposited during the Middle and Upper Pleistocene, as

were the red Kalahari sands. The water table in the dune

complex is high and acts as a reservoir to support vege-

tation in the basin.

Conservation status: not threatened. The Witsand area

is now being conserved by Cape Nature Conservation

and as long as the habitat is protected, there is no threat

of extinction. Although it is localised, it is abundant in

the area.

Number of collections known: 10.

Vouchers: Brusse 5629: Koekemoer 1066: LeisUier 1170, 1390, 1614:

MacDonald 76/43.

Amphiglossa thuja (Merxm.) Koekemoer, comb.

nov.

Pterothrix thuja Merxm. in Mitteilungen der Botanischen

Staatssammlung. Miinchen. Heft 2: 36 (1954). Type: Karasburg,

Rotkop, Dinter6008 (M, holo.; BM!, PRE!, K!, S!, G!, E!, iso.). Figure

2G.

P. thuja Merxm.: 407 (1954), nomen.

P. trifurcata C.A.Sm. ex Merxm. in sched. herb. Kew, nomen. Cited

specimen: Namibia, Rote Kuppe, alt. 800 m, Dinter 1025 (K!).

Erect woody, single-stemmed, brownish grey, strong-

ly aromatic, compact dwarf shrub, up to 0.2 m tall; very

old plants up to 0.45 m tall but then breaking up, with

main branches prostrate. Bark of older stems becoming

greyish and flaky with age, tearing off in strips to uncov-

er a chestnut-coloured stem; older main stems up to 30

mm diam., tending to split and appearing very uneven

when seen in cross section, dead wood persistent on

plant for a long time. Main branches up to 5 mm diam.,

profusely branching into very fine shoots, less than 1 mm

diam., with active growth restricted to very tips of fine

branches, which are covered in minute leaves. Leaves up

to 1.5 mm long, scale-like, adpressed, sessile, greyish

green, alternate, about a leaf length apart, margins entire,

slightly inrolled, adaxial surface woolly. Capitula dis-

coid, small, up to 7 X 2 mm, homogamous, 3-4-flow-

ered, terminally borne on branchlets, linear when young,

involucre spreading at maturity to release pappus and

seeds. Outer involucral bracts 22-28, 1-6 mm long,

ovate with acuminate tips, becoming progressively

longer inwards, outermost scale-like. Inner involucral

bracts linear, membranous in upper fifth, this part slight-

ly ascending, apex shortly mucronate. Receptacle coni-

cal, less than 2 mm diam., honeycombed, ridges shorter

than 1 mm. Disc florets white, bisexual, 5-lobed, 5. 0-5. 5

mm long. Cypselas subterete, ± 1 mm long, greyish,

inconspicuously ribbed, papillose, often only one per

capitulum, appearing to have a loose ectocarp, but cavi-

ty with gel-like substance not obvious as in A. rudolphii.

Pappus bristles 18-23, ± 5 mm long, barbed in lower

fifth, plumose in upper part, seta up to 1.5 mm long.

Flowering time: January-March.

Diagnostic characters: compact shrub with branches

copiously branched into very fine, secondary branches;

leaves and twigs pleasantly aromatic when crushed.

Distribution and habitat: restricted to Kovis Moun-

tain near Liideritz in Namibia, possibly also on other

mountains in the area (Figure 7). In sand on granite

mountains.

Consetyation status: not threatened.

Number of collections known: 7.

Vouchers: De Winter & Hardy 7898: Dinter 6008: Koekemoer 1200.

Amphiglossa tomentosa (Thunb.) Haiw. in Harv.

& Sond., Flora capensis 3: 276 (1894). Type; Thunberg

(sheet 20103 UPS!). Figure 2H.

Relhania tomentosa Thunb.: 640 (1823). Athrixia tomentosa Less.:

364. (1832). Pterothrix tomentosa DC. 280. (1838).

Amphiglossa nitidula DC.: 259. ( 1 838). syn. nov. Type: in Africa aus-

trali ad Zwarte Onder Bokkeveld et Zeederbergen, alt. 2-300 ped.

1835. Drege 2781 (G-DC!, P!).

Pterothrix flaccida Schlechter in sched. Cited specimen: Brandvlei,

1896-01-08, 5di/et7i?er 1655 (K!, NBG!, PRE!, SI).

Amphiglossa tomentosa (Thunb.) Harv. var. breviligulata Merxm.,

syn. nov., nomen. Cited specimen: Brandvlei, 1897-01-08, Schlechter

9923 (M, holo.; PI, El, G! (4x), NBG!, K!, iso.).

Pteronia engleriana Muschl.: 98 (1911). Pterothrix engleriana

(Muschl.) Hutch. &E.Phillips: 277 (1917). Type: Aus.G.S.W.A, 1 400 m,

Dinter 7707 (K!, SAM!).

Pterothrix flaccida Schltr. ex Hutch. & E. Phillips: 379 (1917). syn.

nov., nomen.

Woody, compact, multistemmed dwarf shrubs, up to

0.6 m tall, but very often with growth stunted and then ±

0.2 m tall. Stems up to 3 mm diam., smooth, wiry and

brittle, produced from a well-developed rootstock, older

dead stems persistent; stems bare for largest part, leaf

cluster scars prominent, profusely and very finely

branched in upper parts, young growth often with zigzag

appearance. Leaves sessile, in fascicles; subtending

leaves up to 15 x 1 mm, about twice as long as others in

fascicle, ascending, linear, straight or twisted, margins

entire, slightly inrolled, adaxial surface woolly; fascicles

widely spaced, at least a leaf length apart, leaving

74

Bothalia29,l (1999)

12 14 16 18 20 22 24 26 28 30 32 34 36

FIGURE 8. — Geographical distribution of A. tomentosa, •.

branches exposed. Capitiila radiate, heterogamous,

8.5-10.0 X 2. 5-3. 5 mm, 8-9-flowered, with more disc

than ray florets; one to three heads terminally borne on

each of the last 2-4 nodes of branches, linear when

young, involucre spreading at maturity. Involucml bracts

30-35, scarious, often tinged wine-red, with very distinct

white-tomentose patches on abaxial side, all very similar

except for difference in size, foliaceous bracts limited to

2-5, outer bracts 2. 5-3.0 x ± 1.5 mm, tomentose patch

fairly large in upper half, inner bracts to 8 x ± 1 mm,

tomentose patch becoming increasingly smaller and

almost disappearing in innermost bracts. Receptacle flat-

tened, less than 2 mm diam., honeycombed, ridges short-

er than 1 mm. Ray florets white or pink, female, 3-4 (-5),

lamina 2. 8-3. 5 x 1.8 mm, 3-lobed; tube 5. 2-5. 4 mm long.

Disc florets white or pink, bisexual, 5-lobed, 5. 5-6. 5 mm

long. Cypselas subterete, ± 1 mm long, greyish coloured,

inconspicuously ribbed, papillose, with distinct annular

ridge. Pappus bristles 20-28, ± 5 mm long, barbed in

lower fifth, plumose in upper part, seta up to 1.5 mm

long. Flowering time: September-December.

Diagnostic characters: compact shrub; 3-5 ray flo-

rets; wiry, brittle branches; involucre with conspicuous

tomentose patches abaxially.

Distribution and habitat: from Liideritz southwards to

the Western Cape and inland to Oudtshoorn (Figure 8).

Sandy soils, often in deep red sand and on granite out-

crops.

Conservation status: not threatened.

Number of collections known: 75.

Vouchers: Acocks 85S3\ Compton 3201: Esterhuysen 19291,

Koekemoer 1097, 1104: Merxmuller & Giess 32122.

Amphiglossa triflora DC., Prodromus 6; 258.

(1838). Type; C.B.S. ad Zwart Kei auf der Flache, alt.

4-5000 ped, Drege 3679 [G-DC!, K!, S!, P (4x)!]. Figure

21.

Helichrysum armutum Dinter ex Merxm. nom. nud.

Erect woody, densely entangled, greyish shrub, up to

0.4 m tall, with many stems developing from a well-

developed, underground rootstock. Stems rigid, up to 2

mm diam., old, dry stems persistent; young active shoots

to 2.5 mm diam., often white-tomentose; terminal

branches frequently spinescent; young spines leafy, older

spines bare, 8-25 mm long. Leaves sessile, up to 8 mm

long, basally adpressed, ascending in upper part, at least

half a leaf length apart, narrowly triangular, margins

entire, inrolled, midrib prominent, adaxial surface wool-

ly, persistent for at least two seasons, young leaves yel-

lowish green, older leaves fading to grey. Capitula dis-

coid, homogamous, 8-9 x 1.5-2. 5 mm, 4-5-flowered;

heads terminally borne, solitary or in groups of up to 8.

Involucral bracts 20-25, up to 7.5 mm long, increasing

progressively in length from outer to inner series, outer

broadly ovate, inner linear, hyaline in upper part, tips

aristate, tomentose patch on abaxial side. Receptacle flat-

tened, less than 2 mm diam., honeycombed, ridges short-

er than 1 mm. Disc florets white, bisexual, 5-lobed,

5. 0-5. 5 mm long. Cypselas subterete, ± 1 mm long,

papillose, with a distinct annulus. Pappus bristles free,

caducous, 25-32, ± 5 mm long, barbed in lower fifth,

plumose in upper part, seta up to 1.5 mm long.

Flowering time: mainly October-March, occasionally in

September, April and June.

Diagnostic characters: compact, dense, spiny shrub

with small, linear capitula.

Distribution and habitat: Namibia, Northern Cape,

North-West, Lesotho, Free State, Gauteng and

Mpumalanga (Figure 9). Sandy soils, most often on lime-

stone outcrops or sandstone koppies, frequently in dis-

turbed or overgrazed veld.

Conservation status: not threatened.

Number of collections known: 80.

Vouchers; Edwards 4175: Giess 12566: Hilliard & Burn 10669:

Koekemoer 1057, 1064.

Pterothrix spine.scens DC.: 280. (1838), syn. nov. Type:, C.B.S. in

regionibus interioribus inter flumen Gariep et urbcin Litakun detexit,

Burchell 1845 (G-DC!, PREI, K!).

FIGURE 9. — Geographical distribution of A. triflora, •.

Bothalia29,l (1999)

75

NAMES EXCLUDED

Amphiglossa alopecuroides Sch.Bip. = Stoebe alopecu-

roides (Lam.) Less.

Amphiglossa kolbei Bolus = Disparage kolbei (Bolus)

Hutch.

Pterothrix cymbaefolia Harv. Does not belong to

Amphiglossa or any other related genus. An almost leaf-

less twig with two old capitula was seen in S. The leaves

are 10-20 x 4-5 mm, shortly petiolate, flat, not involute

and slightly tomentose on the abaxial surface.

ACKNOWLEDGEMENTS

This revision is part of a larger research project on the

systematics of the Metalasia group (Gnaphalieae,

Relhaniinae, Asteraceae) and the support of my super-

visor, Prof. B-E. van Wyk of the Rand Afrikaans Uni-

versity, is acknowledged. The financial support and use

of facilities at the NBI is acknowledged in appreciation.

Fieldwork assistance of my parents and generous overall

support are valued dearly. Adela Romanowski is thanked

for the miracles in the darkroom. Dr H.F. Glen for the

Latin diagnosis. Dr S.M. Perold for proofreading the

manuscript, the editors of the journal and the referees for

valuable comments.

REFERENCES

ANDERBERG, A. A. 1991. Taxonomy and phylogeny of the tribe

Gnaphalieae (Asteraceae). Opera Botanka 104.

BENTHAM, G. 1873. Compositae. In G. Bentham & J.D. Hooker,

Genera plantarum 2,1: 163-533. London.

BRUSSE, F. 1990. A new species of Pterothrix (Gnaphalieae) from the

northern Cape. Bothalia 20: 67-70.

DE CANDOLLE, A.R 1838. Compositae. Prodromus systematis natu-

ralis regni vegetabilis 6: 1-687. Treuttel & Wiirtz, Paris.

HARVEY, W.H. 1894. Compositae. In W.H. Harvey & O.W. Sonder,

Flora capensis 3. Hodges & Smith, Dublin.

HOLMGREN, P.K., KEUKEN, W. & SCHONFIELD, E.K. 1981.

Index Herbariorum, edn 7. Regnitm Vegetabile 106.

HUTCHINSON, J. & PHILLIPS, E.P. 1917. A revision of the genus

Pteronia (Compositae). Annals of the South African Museum 9:

277-327.

LESSING, C.F. 1832. Synopsis generum compositarum. Berolini.

MERXMULLER, H. 1954. Taxa nova vel critica: Compositae. Mit-

teilungen der Botanischen Staatssammlung, Munchen 2: 31^1.

MUSCHLER, R.C. 1911. Compositae africanae novae. 1. Botanischer

Jahrbiicher d6: 94—124.

PHILLIPS, E.P. 1951. Compositae. The genera of South African flow-

ering plants, edn 2. Memoirs of the Botanical Survey of South

Africa No. 25.

SCHLECHTER, R. 1900. Plantae Schlechterianae novae vel minus

cognitae describuntur. II. Botanische Jahrbiicher 27: 86-220.

THUNBERG, C.P 1800. Prodromus plantarum capensium. Edman,

Uppsala.

THUNBERG, C.P 1823. Flora capensis. Stuttgart.

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Bothalia 29,1: 77-82 (1999)

Studies in the liverwort genus F ossombronia (Metzgeriales) from

southern Africa. 9. A new species from Mpumalanga and KwaZulu-

Natal, with notes on other species

S.M. PEROLD*

Keywords; Fossombronia, F. renateae, liverworts, Metzgeriales, new species, southern Africa, tropical Africa

ABSTRACT

A new species of Fossombronia, F renateae, from Mpumalanga and KwaZulu-Natal in southern Africa is described. It is

distinguished by its reflexed, undulating leaves and often stipitate pseudoperianth with a highly convoluted mouth, as well

as by completely or incompletely reticulate spores and usually by short, blunt elaters with two spirals which often form rings

or coils. Notes are provided on two unnamed species from Mpumalanga which cannot be fully described as yet, since ripe

spores are not available for study. Brief references are made to some tropical African species.

Fossombronia renateae Perold, sp. nov.

Plantae repentes, dense congestae vel cum muscis

intercrescentes. Folia late patentia, undulata, oblonga vel

breviora quam lata, supra leve lobata. Dioicae. Antheridia

in plantis masculis bracteis obtecta. Pseudoperianthium

campanulatum; ore perirregulariter lobato et maxime

convolute. Sporae 37.5^7.5 pm diametro, cum 8 vel 9

cristis irregularibus, aliquando anastomosantibus, aliquot

areolas formantibus. Elateres plerumque bis spirales,

annulares vel spirales, 25-75 x 12.5-17.5 pm.

TYPE. — Mpumalanga, 2530 (Lydenburg): Lone

Creek Falls, on soil, at margin of pond below falls,

(-BB), Perold & Koekemoer 4073 (P^, holo.).

Plants smallish to medium-sized to rather large,

creeping, in dense, crowded stands or intimately to

loosely intergrown with mosses, green; shoots often sim-

ple, in male plants 5-13 x 3—4 mm, 1.3-2. 5 mm high; in

female plants 9-15 x 3. 5^.0 mm, 1.6-2. 5 mm high,

sometimes furcate or with short, lateral branches near

apex and/or toward base. Stems prostrate, in some popu-

lations distally markedly fleshy, tapering proximally, in

living material from above, hardly distinct from adjoin-

ing leaf bases, sometimes faintly longitudinally striate, in

cross section plano-convex, in male plants at apex

(Figure IL) up to 600 pm (12-14 cell rows) high, ± 630

pm wide, at base (Figure IM) ± 570 x 540 pm; in female

plants at apex (Figure IN) up to 680 pm (15 cell rows)

high, ± 1000 pm wide, at base (Figure 10) ± 420 x 700

pm. Rhizoids purple, 12.5-25.0 pm wide. Leaves over-

lapping by about V3 of their width, spreading widely,

undulating, succubously inserted, often slightly decur-

rent, apically smaller, soon becoming larger, oblong or

shorter than wide, shallowly lobed above; in male plants

(Figure lA-F) 1200-2500 x 1300-2375 pm; in female

plants mostly larger (Figure IG-I), 1250-2500 x

1350-3500 pm, sometimes narrower above or below;

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

MS. received: 1999-01-18.

margins with up to 13 well-spaced, sessile papillae, but

sometimes fewer, only 2 toward leaf base, occasionally

raised on a basal cell. Leaf cells thin-walled, in male

plants not appreciably different from those of female

plants, at upper margins (Figure IJ) rectangular across,

occasionally isodiametric, 25.0-37.5 x 27.5-52.5 pm, at

lateral margins long-rectangular, 50.0-87.5 x 15.0-22.5

pm; upper laminal cells 5- or 6-sided, 35.0-52.5 x

27.5-37.5 pm; middle laminal cells 47.5-75.0 x

32.5^5.0 pm; basal cells 75.0-107.5 x 42.5-62.5 pm.

Oil bodies few (Figure IK), only 4-6 per cell, 5 pm

diam., or a little larger, round, but some slightly irregular

and with a central indentation; chloroplasts numerous,

mostly rounded, ± 5 pm diam.

Dioicous. Antheridia dorsal on stem, in 1 or 2 crowd-

ed rows, short-stalked, globose, distal ones often larger,

220-270 pm diam., white, proximal ones smaller, ± 160

pm diam., yellow, each shielded by a perigonial bract

(Figures IP-S; 2A, B), 460-580 x 200-380 pm, some-

times 2 adjacent ones joined together and then 2- or 3-

lobed, mostly with apical processes topped by a papilla,

marginal cells ± rectangular, 40-75 x 27.5-40.0 pm,

inner cells usually 5-sided, 65.0-87.5 x 22-35 pm.

Archegonia in 1 or 2 rows dorsally along stem (Figure

2C), distally and proximally, naked, sometimes 2 per

branch, at intervals or 2 adjacent or even 3 in a cluster,

becoming fertilised. Pseudoperianth (Figures IT, U;

2D-F) campanulate, about as tall as leaves, raised on a

short stalk, then widely flaring above, up to 1625 pm

long, ± 3500 pm wide across mouth, very irregularly

lobed and highly convoluted, lobes 300-400 x 400-450

pm, with papillae at angulations, some sessile, others

raised on a basal cell; cells comparable in shape and size

to those of leaves, except for basal ones which are larger,

100.0-112.5 X 37.5-45.0 pm. Capsules globose, ± 750

pm diam., walls bistratose, cells in inner layer irregularly

shaped (Figure IV), subquadrate, long-rectangular or tri-

angular, 25-50 X 27.5-40.0 pm, each cell wall with 1 or

2 dark brown, nodular and sometimes semi-annular thick-

enings. Seta 1.1 -2. 5 mm long, in cross section (Figure

IW) 200-300 pm diam., 8 cells across. Spores light

brown, hemispherical, 37.5^7.5 pm diam., including

78

Bothalia29,l (1999)

FIGURE I . — Fossomhronia renateae. A-F, male leaves; G-I, female leaves; J, upper leaf margin; K, median leaf cells with oil bodies (solid lines)

and chloroplasts (dotted lines); L, c/s male stem apex; M, c/s male stem base; N, c/s female stem apex; O, c/s female stem base; P-S,

perigonial bracts; T, pseudoperianth from side; U, opened pseudoperianth; V, cells in inner capsule wall; W, c/s seta. A-F, K-M, P-S,

Perold & Koekemoer 4071', G-I, N, O, Perold & Koekemoer 4073', J, Lubenau-Nestle SA265', T, U, Lubenau-Nestle SA264 p.p.; V, Doidge

CH3564, W, Perold & Koekemoer 4078. Scale bars: A-I, T, U, 500 pm; J, W, 100 pm; K, V, 50 pm; L-S, 250 pm. Artist: G. Condy.

Bothalia29,l (1999)

79

FIGURE 2. — Fossombronia renateae. A, male plant with perigonial bracts; B, close-up of perigonial bracts; C, female plant with archegonia; D,

female plant with pseudoperianths (indicated by arrows); E, female plant with pseudoperianth near apex of branch; F, close-up of

pseudoperianth. A, B, E, F, Perold & Kaekemoer 407I \ C, Perold & Koekemoer 4073\ D, Liibenau-Nestle SA265. A, x 8,7; B, x 20; C, D,

X 8; E, X 8; F, x 25.6.

odges projecting at margin; distal face (Figure 3A-D)

convex, with a tendency to form 8 or 9 well-marked are-

olae across face, 5. 0-7. 5 pm wide, occasionally ridges

running parallel to each other or else in different direc-

tions, interconnected by faint or distinct cross walls or

anastomosing and forming incomplete areolae, some-

times with small inclusions, surrounding ridges 2. 5-4.0

pm high, crenulate above; proximal face (Figure 3E)

lacking triradiate mark, mostly flat, covered with coarse,

very irregular spicules or spikes, 20 or 21 ‘spines’ pro-

jecting up to 2.5 pm high around spore periphery and

completely or somewhat incompletely joined by a ± 5 pm

wide, membranous wing or perispore, its margin crenu-

late. Elaters in most populations relatively few, with 2

FIGURE 3. — Fossombronia renateae. A-E, spores; F, elater. A, B, distal face; C, side view of distal face; D, detail of part of distal face; E, prox-

imal face. A, Perold & Koekemoer 4073\ B-E, Liibenau-Nestle SA265\ F, Liibenau-Nestle SA264 p.p. A, x 1046; B, x 908; C, x 881; D,

F, X 1744; E, x 752.

80

Bothalia29,l (1999)

spirals (rarely with 3 spirals partly), light brown or green-

ish, often in rings or coils (Figure 3F), rather short and

stout, blunt at both ends, or ± wedge-shaped, 25-75 x

12.5-17.5 pm, sometimes sticking to spores.

Fossombronia renateae is so far known from only two

localities, both in the Afromontane, summer rainfall region

of southern Africa, from Lone Creek Falls, near Sabie in

Mpumalanga and Xumeni Forest Reserve, near Donny-

brook in KwaZulu-Natal (Figure 4), but it must surely be

more widespread. At Lone Creek Falls, the plants grow on

soil between rocks and are kept moist by spray from the

falls. LFnfortunately, there are no details about the substrate

and conditions on the label of the Doidge collection from

Xumeni Forest Reserve. Bryum alpinum Huds. ex With,

and Funaria limbata (C.Miill.) Broth, are mosses that grow

together with the Lone Creek plants.

This newly isolated species has been named F. renateae

in honour of Dr Renate Lubenau-Nestle of Kempten,

Germany, who collected it when on a visit to South Africa

in 1992 and kindly lent her specimens to PRE for exami-

nation. In 1998, accompanied by Ms M. Koekemoer, cura-

tor of PRE herbarium, I visited Lone Creek Falls (and the

vicinity) three times. In April we failed to find fertile spec-

imens of this species, but in June we were successful,

although the material only had a few dehisced capsules

retained from the previous season, and fortunately still

with some unshed spores. These spores and elaters are

closely similar to those of the Lubenau-Nestle specimens.

Samples of the fresh collections with young capsules were

cultivated for two months (until August) before ripe spores

were obtained. Many of these spores were malformed, but

the ornamentation was fairly similar to that of the earlier

collections. The elaters, however, were more numerous,

usually with 2 (rarely 3) darker and thicker spirals, 45-110

(-132.5) X 10.0-17.5 pm; only a few elaters were identi-

cal to the Lubenau-Nestle and Doidge ones.

Scott & Pike (1988) refer to the elaters of F. cale-

donica Steph. as ‘bispiral, with the spirals breaking down

into rings’ (translated by them from Latin). They found

similar elaters in F papillata (and several other species),

where capsules with elaters of this sort and capsules with

normal elaters occurred in the same colony. Accordingly,

they considered the production of such elaters a mere

abnormality not worthy of specific recognition.

However, in a genus where there are so few reliable tax-

onomic characters to discriminate between species, I

would hesitate to discard this one entirely.

The plants collected by Perold & Koekemoer are

decidedly more robust and the stems fleshier than in the

earlier collections. The species can be distinguished by

its undulating leaves, by its often stipitate pseudoperi-

anths, of which the mouth is highly convoluted, by com-

pletely or incompletely reticulate spores and generally by

short and relatively wide elaters, the two spirals often

forming rings or coils.

Fossombronia capensis S.W.Arnell from the winter

rainfall area in the southern Cape, also has short and

stout elaters, usually with loose, pale-coloured spirals

(Perold 1997a), but in spore ornamentation it differs

completely from F renateae.

FIGURE 4. — The distribution of Fossombronia renateae in southern

Africa.

NOTES ON TWO UNNAMED SPECIES FROM

MPUMALANGA

A new, unnamed species was found beside the

Malieveld River, 9 km East of Sabie on the road to

Hazyview, Mpumalanga. It has been collected three

times, Perold & Koekemoer 3573, 4053, 4082, in the

past few years. This species grows on the concrete sup-

ports of the bridge, just above or at the water level, as

well as on the soil of the streambank. Unfortunately, ripe

spores have not yet been obtained and it cannot be

described at this stage.

This species can be recognised by almost entire,

rounded, slightly concave and rather stiff, imbricate

leaves, 1250-1750 x 1500-2000 pm. They overlap by ±

'A of their width and are not decurrently inserted on the

stem. The shoots are up to 19 mm long, including the

apical branches which are 5-7 mm long. The pseudope-

rianth is cupulate, also rather stiff and the margin is

undulating. The perigonial bracts shielding the antheridia

in the male plants, are conspicuous.

Another species of Fossombronia collected at Lone

Creek Falls, Perold & Koekemoer 4059, 4060, is very

large, with up to 50 mm long shoots. It is an erect-grow-

ing, bottle-green, aquatic plant. Its leaves are sub-

quadrate to rectangular, 2800-3375 x 2725-3200 pm,

and only very slightly decurrent at the ‘trailing’ edge,

rather stiff, 4-stratose at the base and bistratose up to ±

midlength. Unfortunately, only specimens with archego-

nia were found, and the plants kept in cultivation deteri-

orated rapidly. It cannot, therefore, be described at this

stage, but it is still worth mentioning, in case fruiting

material turns up later.

Several more, large, aquatic specimens, without

spores have been collected elsewhere in southern Africa.

It would appear that fertilisation of these plants in quite

rapidly flowing streams, seldom takes place.

NOTES ON SOME TROPICAL AFRICAN SPECIES

It is unlikely that the above-mentioned plants belong

to F. pulvinata, a sterile species described by Stephani

Bothalia29,l (1999)

81

(1911, 1917) from Rwanda, leg. Mildbraed 22182 (G!).

The leaves of F. pulvinata, in contrast to the above-

mentioned South African plants, are strongly decurrent,

much wider than long (‘7 mm lata, medio 4 mm longa’)

and rather lax. Scott & Pike (1988) think that this species

may be F. australis.

Jones (1990) reports Noteroclada porphyrorhiza

(Nees) Steph. with purple rhizoids, as once recorded

from Kilimanjaro. The pseudoperianth is said to be later-

ally compressed and bilabiate, but spores are not men-

tioned. This species had previously been transferred to

Fossombronia porhyrorhiza (Nees) Prosk. by Proskauer

(1955). I have not seen this specimen and am unable to

comment on it. I have, however, examined a sterile spec-

imen from Lydenburg, Wilms 024671(G), which had

been referred to Fossombronia (= Androcryphia) por-

phyrorhiza Nees. This plant has surely been misidenti-

fied, as the type species is from South America, which

Scott & Pike (1988) think may also be F. australis.

Another Fossombronia species described from tropi-

cal Africa on Mt Kilimanjaro in Kenya, is F. grandis

Steph., leg. Volkens 1226 (G18524) holotype (G!). It is a

large plant, as the specific epithet is intended to convey.

The shoots are 20 mm long and simple, the leaves large

and broadly rounded, the margins with some short, longi-

tudinal folds. Stephani (1900) reported the spores of this

dioicous species to be 34 pm diam.; my measurements

are 32.5-37.5 pm diam. The distal face is coarsely retic-

ulate, with rather thin, 2. 5-5.0 pm high ridges delimiting

4 or 5(-6) angular areolae, 7.5-10.0 pm wide, across the

diameter and with ± 12 spinous projections connected by

a perispore, at the periphery; the proximal face is gene-

rally concave in the centre, with fine, irregular, sinuating

ridges, sometimes forming small areolae, partly or com-

pletely surrounded and often a little obscured by several

large, angular areolae, seemingly ‘folded’ inward from

the outer row on the distal face. The elaters are 242-280

X 7.5 pm and often 2-spiral. I have also examined a

specimen, Adam & Jager 9211/X from Mt Elgon

National Park, which was kindly lent by Dr Chuah-

Petiot. It, and a sterile specimen, Poes 92 14/ AC had been

assigned to F. grandis by Prof T. Poes. Chuah-Petiot

(1995) reports F. grandis (Poes 9236/AA) from Naro

Mom. A specimen, Hedberg 1747h (BOL) from Mount

Kenya (at the stream below Tyndall glacier), was deter-

mined by Arnell (1956) as F. grandis. It is sterile and the

determination is doubtful.

The spore ornamentation in F. grandis is so closely

similar to that of F. angulosa (Dicks.) Raddi, [said to be a

salt-tolerant (Smith 1990), mediterranean-atlantic species

(Jovet-Ast 1946)], and also large as well as dioicous, that

I am surprised no one else has remarked on it. Scott &

Pike (1988) regard both F. angulosa and F. grandis as

good species and do not comment on the spore ornamen-

tation. I found the leaves in F. angulosa to be mostly

longer than wide and shortly lobed above. In F. grandis

the leaves are generally wider than long, the apex round-

ed and, at the base, several cell layers thick, the latter also

found to be the case in F. angulosa. In both species the

pseudoperianths are stipitate. Despite the similarities

between these two species, it seems phytogeographically

unlikely that F. grandis would be conspecific with

F. angulosa. Moreover, I have examined only two sporu-

lating specimens of F. grandis, the type specimen and the

Adam & Jager collection, which makes me hesitate to

place this species in synonymy under F. angulosa.

Krauss (1846) collected a specimen of the so-called

F. angulosa from southern Africa ‘in mpibus rivulor natalen-

sium’, but this has not been confirmed and is thought to

have been a misidentification of the commonly occurring

F. crispa Nees (= F. zeyheri Steph.) (Perold 1997b, c).

SPECIMENS EXAMINED

Held at PRE, unless otherwise stated. Species listed in

alphabetical order.

F. angulosa (duplicates in PRE)

T. Arts (Portugal and Madeira) 15519, 15567, 15605, 15732, 15778,

15912, 16277, 16329, 19815, 19819, 19883, 19921, 19929, 20003,

20190. Bryhn NH31448. Cumow CH656. Piquet CH536B.

F. grandis

Adam & Jager 9211/X. Poes (sterile) 9214/AC (University of Nairobi).

Hedberg (sterile) 1747h (BOL). Volkens 1226 = (018524) (G).

Lone Creek specimens (very large)

Perold & Koekemoer 4059, 4060.

Malieveld specimens

Perold & Koekemoer 3573, 4053, 4082.

F. porphyrorhiza

Wilms 024671 (G), incorrectly referred here.

F. pulvinata

Mildbraed 22182 (G).

F. renateae

Doidge CH3564. LUbenau-Nestle SA264 p.p., SA265 (Herbarium

Liibenau). Perold & Koekemoer 4073 (holotype), 4074, 4076, 4078.

ACKNOWLEDGEMENTS

I express my deep gratitude to Dr H. Stieperaere,

Meise, for refereeing this article and for his helpful sug-

gestions. I also sincerely thank Dr Theo Arts, St.-Job in’t

Goor, Belgium, for his generous gift of specimens to

PRE; also Dr P. Geissler, Geneva, Mrs P. Lorber (BOL),

as well as Dr M.S. Chuah-Petiot, University of Nairobi

and last, but by no means least. Dr R. Liibenau-Nestle,

Kempten, Germany, for the loan of specimens. Ms M.

Koekemoer, curator at PRE, is kindly thanked for her

unstinting help with fieldwork, likewise my thanks to Ms

G. Condy for the drawings, Mrs A. Romanowski for

developing and printing the photographs and to Ms D.

Maree for typing the manuscript.

REFERENCES

ARNELL, S.W. 1956. Hepaticae collected by O. Hedberg et al. on the

East African mountains. Arkiv for Botanik 3: 517-562.

CHUAH-PETIOT, M.S. 1995. Data to the bryoflora of Mount Kenya,

Kenya. Tropical Bryology 10: 41-53.

JONES, E.W. 1990. An artificial key to the genera of African hepatics.

Journal of Bryology 16: 9-40.

JOVET-AST, S. 1946. Hepatiques des iles du Cap Vert essai sur leur

repartition. Memoires de la Societe de Biogeographie 8;

363-367.

KRAUSS, F. (ed.) 1846. Pflanzen des Cap-und Natal-Landes, gesam-

melt und zusammengestellt von Dr Ferdinand Krauss. Flora 29:

135, 136.

82

Bothalia29,l (1999)

PEROLD, S.M. 1997a. Studies in the liverwort genus Fossombwnia

(Metzgeriales) from southern Africa. 2. An amendment to three

species from Western Cape, described by S.W. Amell. Bothalia

27: 29-38.

PEROLD, S.M. 1997b. Studies in the liverwort genus Fossombwnia

(Metzgeriales) from southern Africa. 4. A re-examination of

F. crispa, F. leucoxantha and F. tumidci. Bothalia 27: 105-

116.

PEROLD, S.M. 1997c. The lectotypification of Fossombwnia crispa.

Bothalia 27: 137, 138.

PROSKAUER, J. 1955. Notes on hepaticae. III. The Bryologist 58:

192-200.

SCOTT, G.A.M. & PIKE, D.C. 1988. Revisionary notes on Fossom-

bwnia. The Bryologist 9\ \ 193-201.

SMITH, A.J.E. 1990. The liverworts of Britain and Ireland. Cambridge

University Press.

STEPHANI, F. 1900. Species hepaticarum. Bulletin de T Herbier Bois-

sier 1: 383.

STEPHANI. F. 1911. Hepaticae. In G.W.J. Mildbraed, Wissetuchaf-

liche Ergebnisse der Deutschen Zentral-Afrika-Expedition

;907-790S, 2 (Botany): 113.

STEPHANI, F. 1917. Species hepaticarum. Bulletin de T Flerbier Bois-

sier 6: 75.

Bothalia 29,1: 83-93 (1999)

Studies in the liverwort genus Fossombronia (Metzgeriales) from

southern Africa. 10. Three new species from Northern and Western

Cape

S.M. PEROLD*

Keywords: Fossombronia Raddi, F. hyalorhiza Perold, F marindae Perold, F. monticola Perold, Hepaticae, Metzgeriales, Northern Cape, southern

Africa, Western Cape

ABSTRACT

Three new species of Fossombronia from the winter rainfall area of the Northern and Western Cape are described: F.

hyalorhiza, which has colourless rhizoids and is fairly widespread, but not common; F. marindae, which has purple rhi-

zoids, very attractive, highly ornamented spores and is so far known from a single locality only and F. monticola, which

also has purple rhizoids, with spores irregularly ridged, often partly areolate, and is quite common but fairly variable.

1. Fossombronia hyalorhiza Perold, sp. nov.

Plantae repentes, caulibus viridibus, nucleo centrali

purpureo. Rhizoidea hyalina. Folia undulata, saepe incisa

vel lobata, interdum forma valde variabili. Dioicae.

Antheridia in plantis masculis bracteis obtecta. Pseudo-

perianthium cupulatum; ore undulato, cum ± 11 promi-

nentiis irregularibus. Sporae 40.0-57.5 pm diametro,

interdum 55.0-67.5 pm; superficie distali cum 6-8(-9)

cristis irregularibus, rectis vel curvatis, brevibus vel lon-

gibus. Elateres 100.0-227.5 x 7.5-10.0 pm, omnino ter

spirales vel solum medio, extremitatibus bis spirales,

rarissime quater vel quinquiens spirales.

TYPE. — Northern Cape, 3219 (Wuppertal): Skurwe-

berge, on road from Ceres to Citrusdal, N of Op-die-

Berg, along turnoff to Boplaas; slightly beyond hiking

trail, near Sandrivier; below seep, at rocks with mosses,

(-AB), S.M. Perold 3867 (PRE, holo.).

Plants medium-sized to fairly large, gregarious, most-

ly growing together with mosses, green, but leaves soon

dying and turning translucent, sometimes marginally

purple, occasionally with 3 or 4 rows of purple cells at

base or sparsely blotched with purple throughout, shoots

often simple or once-furcate, rarely with short lateral

branches or buds near base, frequently arising from

tuberous apex of old stem, which is up to 500 x 600 pm,

flanked by small, often purple leaves (Figure IK), taper-

ing proximally and 3-5 mm long; male plants 5-8 x ±

3.2 mm, ± 1.6 mm high; female plants larger, 10-15 x ±

3.7 mm, ± 2 mm high. Stems prostrate, sometimes arch-

ing, distally fleshy, proximally tapering, dorsally green,

ventrally purple, in cross section with outer layers of lat-

eral and ventral cells green, core ± central or ventrally

displaced, purple, persistent in older shoots, ‘tailing off’

in young ones, plano-convex to slightly biconvex, in

male plants at apex (Figure IN) up to 550 pm (14 cell

rows) high, ± 600 pm wide, at base (Figure 10) ± 300 x

360 pm; in female plants at apex (Figure IP) up to 530

pm (14 cell rows) high, ± 620 pm wide, at base (Figure

IQ) ± 450 X 570 pm. Rhizoids hyaline, 15-20 pm wide.

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

MS. received: 1999-01-18.

some with internal mycorrhizal strands. Leaves overlap-

ping, suberect, or incurved to widely spreading, upper

margin undulating, often notched or lobed, lobes angular

or rounded, sometimes very variable in shape, shorter or

longer than wide, slightly decurrent, succubously insert-

ed; in male plants leaves (Figure lA-E) 875-1500

(-2050) X 875-1875 pm, generally narrower below; in

female plants mostly larger (Figure IF-J), 1625-2000 x

1300-1625 pm, also narrower below, at pseudoperianth

often larger, up to 2250 x 3025 pm, but narrower below;

margins with 3 or 4 well-spaced papillae, generally ses-

sile but near leaf base raised on 1 or 2 basal cells. Leaf

cells thin-walled, not appreciably different in male and

female plants, at upper margins (Figure IL) rectangular

across, occasionally isodiametric, 32.5-42.5 x 40.0-67.5

pm, at lateral margins long-rectangular, 80-100 x

37.5- 42.5 pm; upper laminal cells 5- or 6-sided,

62.5- 75.0 X 50.0-62.5 pm; middle laminal cells

112.5- 162.5 X 50.0-67.5 pm; basal cells 125.0-162.5 x

55-80 pm. Oil bodies absent in the material examined;

chloroplasts ± 5 pm diam. (Figure IM).

Dioicous. Antheridia dorsal on stem, near apex very

young, spaced, toward base mature, in groups (Figure

2A-C), short-stalked, globose, ± 250 pm diam., subtend-

ed by perigonial bracts (Figure IR-V), at maturity fairly

large, 450-550 x 150-220 pm, when 2 adjacent ones

joined together, 320-430 pm wide, marginal cells

67.5- 95.0 X 35-40, cells in interior 4-6-sided, 100-125

X 25.0-32.5 pm, above with finger-like appendages,

150-250 pm long, tapering to apices. Archegonia in a

row dorsally along stem (Figure 2D), naked, sometimes

2 adjacent ones becoming fertilised. Pseudoperianth

(Figure IW, X) cupulate, about as tall as leaves (Figure

2E, F), up to 2125 pm long, flaring widely from narrow

base, ± 750 pm wide, across mouth ± 3325 pm wide,

undulating, with folds or pleats and ± 1 1 irregular pro-

jections, 400^50 pm long and ± 550 pm wide across

base, at margin with 19 or 20 sessile papillae, sometimes

laterally incised, rarely 2 adjacent pseudoperianths part-

ly joined; cells comparable in shape and size to those of

leaves. Capsules globose, ± 850 pm diam., walls bis-

tratose, cells of inner layer irregularly shaped (Figure

lY), roughly rectangular, trapezoidal or triangular,

22.5- 57.5 X 27.5^5.0 pm, each cell wall with (1)2 or 3

84

Bothalia29,l (1999)

FIGURE 1 . — Fossomhronia hyalorhiz.a. A-E, male leaves; F-J, female leaves; K, apical leaf of old female stem; L, detail of leaf margin; M, medi-

an leaf cells with chloroplasts (oil bodies disintegrated); N, c/s male stem apex; O, c/s male stem base; P, c/s female stem apex; Q, c/s

female stem base; R-V, perigonial bracts; W, pseudoperianth from side; X, opened pseudoperianth; Y, cells in inner capsule wall; Z, c/s

seta. A-J, N, O. R-V, Y, Z, S.M. Pewld 3867', K-M, W, X, S.M. Perold 343I\ P, Q. SM. Perold3393. Scale bars: A-K, W, X, 500 pm; L,

Z, 100 pm; M, Y, 50 pm; N-V, 250 pm. Artist: G. Condy.

Bothalia29,l (1999)

85

FIGURE 2. — Fossombronia hyalorhiza. A, male plant with small perigonial bracts near apex of shoot; B, close-up view of perigonial bracts and

very young antheridium (indicated by arrow); C, perigonial bracts and antheridia (indicated by arrows) near base of shoot; D, female plant

with archegonia (one indicated by arrow); E, furcate female shoot, each branch with a pseudoperianth; F, close-up of pseudoperianth and

surrounding leaves from above. A-D, SM. Perold 3409\ E, F, S.M. Perald 3902. A, x 8.3; B, x 38.8; C, x 29; D, E, x 10; F, x 20.

dark brown, nodular and sometimes semi-annular thick-

enings. Seta 1. 5-4.0 mm long, in cross section (Figure

IZ) 230-250 |im diam., 7 or 8 cells across. Spores brown

to deep brown, hemispherical, 40.0-57.5 pm diam. in

some populations, in others 55.0-67.5 pm, including

ridges (or spines) projecting at margins; distal face

(Figure 3 A-D) convex, with 6-8(9) irregular, straight or

curved, short or long ridges across, 5.0-7. 5 pm high,

sometimes furcate, branches then running in different

directions but often ± parallel down sides of spore, ± 7.5

pm apart, crests crenulate, between bases faint cross-

striations and occasionally low, scattered papillae, rarely

with complete or incomplete areolae ± 10 x 7.5 pm,

sometimes with a central inclusion; proximal face

FIGURE 3. — Fossombronia hyalorhiza. A-E, spores; F, elater. A, B, distal face; C, side view of distal face; D, detail of part of distal face; E, prox-

imal face. A, S.M. Perold 3903\ B, D. F, S.M. Perold 343P, C, S.M. Perold 3902; E, S.M. Perold 3904. A, x 658; B, x 734; C, E, x 734;

D, X 1193;F, X 1285.

86

Bothalia29,l (1999)

(Figure 3E) mostly flat, triradiate mark absent, but occa-

sionally elements of ornamentation lined up to form part

of 1 or 2 radii, inner area densely covered with coarse,

irregular, erect, platelike projections or even amorphous

‘blobs’, in between and around circumference dotted

with spinules and papillae, at spore periphery up to 25

‘spines’ (‘ends’ of ridges) projecting 5. 0-7. 5 pm and

incompletely joined by a membranous wing or perispore,

its margin crenulate or entire. Elaters (Figure 3F) light

brown, 100.0-227.5 x 7.5-10.0 pm, tapering to tips and

sometimes ending in a loop, 5 pm wide, 3-spiral entirely

or only in middle, ends 2-spiral, very rarely partly 4- or

5-spiral, often faintly papillose.

Fossombronia hyalorhiza was collected several times

(November 1995, October 1997) at four different localities

in the Northern and Western Cape (Figure 4), which is in the

winter rainfall area of southern Africa. This is referred to as

the Fynbos Biome (sclerophyllous, microphyllous vascular

plant vegetation) (Cowling etal. 1997; Rutherford 1997), or

else the area is called the Cape Region (phytogeographic

region based on the distribution of vascular plants)

(Cowling & Hilton-Taylor 1997). The four localities

referred to above, (listed from north to south), are as fol-

lows; Matsikama Mountains, on plateau, east of

Vanrhynsdorp; the type locality at Sandrivier, northeast of

Op-die-Berg; Uitkyk Pass in the Cederberg Reserve; at the

top of Gydo Pass, north of Prince Alfred Hamlet. According

to Low & Rebelo (1996) the vegetation type in these local-

ities is Mountain Fynbos. The plants grow at seepage areas,

near storm water furrows or at streams, on light brown or

greyish, gravelly soil, containing quartz crystals. Mosses

found with the type specimen are Pleuridium ecklonii

(C.Miill.) Snider and Campylopus sp. A Dwsera sp. was

also growing with this collection, indicating that the soil is

nitrogen-poor. The specific epithet hyalorhiza, meaning

‘having colourless rhizoids’, was chosen to distinguish this

species from a sterile, undescribed Fossombronia species

with purple rhizoids, which was mixed with some of the

specimens with colourless rhizoids. Epithets like leucorhiza

and porphyrorhiza have previously been used for Fossom-

bronia species. In order to minimise damage to the speci-

mens, it is, however, not possible to examine the rhizoids of

every plant in the collections.

F. hyalorhiza can be distinguished by its colourless

rhizoids; stems with a purple, ± central core, persisting in

older shoots, but ‘tailing’ off in young ones; undulating,

suberect to spreading leaves and spores with the distal

face bearing 6-8(9), ridges, which are sometimes paral-

lel, at other times furcate, the branches often running in

different directions. The spore ornamentation is fairly

constant in all the specimens that were studied by light

microscope as well as by scanning electron microscope.

From the northern, summer rainfall area of southern

Africa, another new species with colourless rhizoids, F.

straussiana (Perold 1997a), has been described. It differs

from F. hyalorhiza by being monoicous, its leaves are

densely imbricate and highly convoluted, with basal

appendages partly curved around the antheridia and its

spores are ornamented with incomplete areolae contain-

ing fine inclusions.

To compare the two southern African species that

have colourless rhizoids with F. husnotii Corb., a loan of

FIGURE 4. — The distribution of Fossombronia hyalorhiza, •; F.

marindae. A; and F. monticola, □, in Western Cape.

the type material was requested from the Societe

Nationale des Sciences Naturelles et Mathematiques de

Cherbourg, but it was not available. Samples of speci-

mens of F. husnotii (with colourless rhizoids) collected

in Madeira by Dr T. Arts were kindly sent by him.

Examination of the spores of these samples by SEM,

demonstrated once again that the ornamentation on the

distal face of F. husnotii spores is extremely variable,

‘ranging from discrete spines to ± parallel lamellae or

irregular areolae’ (Paton 1973). In F hyalorhiza spores

there are no discrete spines on the distal face, only the

‘ends’ of the ridges which project around the periphery

of the spore, with some papillae between them. The orna-

mentation on the proximal face is also much coarser and

denser. Jones & Harrington (1983) report ‘the constant

production of descending stolons and tubers (that) form

useful diagnostic characters’ in F. husnotii. These are

absent in F hyalorhiza, and new growth is often initiated

from old stem apices.

The two southern African species with colourless rhi-

zoids, F. straussiana and F hyalorhiza, are thus shown to

be distinct from F. husnotii.

2. Fossombronia marindae Perold, sp. nov.

Plantae repentes, arete conturbatae. Folia imbricata,

suberecta, forma irregulari, interdum lobata. Solum plan-

tae femineae visae. Pseudoperianthium campanulatum;

ore undulato, saepe leviter vel profunde lobato. Sporae

42.5-55.0 pm diametro, superficie distali exquisite orna-

ta, cum 7 vel 8 cristis trans diametrum, inter cristos cum

spinulis multis tenuibus. Elateres 112.5-230.0 x 7.5 pm,

multi bis spirales, alii longitudinem secus alterne bis et

ter spirales.

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

at northern outskirts of town, Rustig Chalets, against

moderate slope, behind camp sites nos. 1 and 2, on damp,

clayey soil, in partial shade, (-CB), M. Koekemoer 1415

(PRE, holo.).

Since female plants were the only ones seen, the fol-

lowing description pertains to them only. Plants medium-

sized, creeping,' in closely crowded stands, clear green.

Bothalia 29,1 (1999)

87

turning brown and translucent with age; shoots simple

(Figure 6A), up to 8 mm long, ± 1.75 mm high, often

gradually becoming lower toward base, ± 1.5 mm wide,

occasionally once furcate, apical segments mostly short,

0.8-1. 5 mm long, moderately divergent, rarely with lat-

eral shoots apparently arising from buds. Stems prostrate,

green, occasionally arched (Figure 6D), some growing

from apices of old shoots, apically (Figure 5Q) fleshy, in

cross section ± 350 pm (10 cell rows) high, up to 600 pm

wide, tapering proximally and basally (Figure 5R) ± 350

X 300 pm, plano-convex. Rhizoids purple, 10-15 pm

wide. Leaves overlapping, suberect, undulating, suc-

cubously inserted on stem, often decurrent, those lateral

to pseudoperianth near stem apex larger, widely obovate,

1575-2625 x 1575-2375 pm, below 1000-1125 pm

wide (Figure 5N), shape of other leaves (Figure 5A-L)

irregular, sometimes lobed, shorter than wide above,

1150-1500 X 1225-2250 pm, narrower below, 800-1250

pm, rarely 2 leaves joined (Figure 5M) for some distance

above their bases and produced at branching off of later-

al shoot between it and main axis; margins with 2-4

widely spaced, sessile papillae. Leaf cells thin-walled, at

upper margins (Figure 50) subquadrate to rectangular

across, 25^0 x 37.5-52.5 pm, at lateral margins long-

rectangular, 40.0-92.5 X 25-30 pm; upper laminal cells

5- or 6-sided, 40.0-52.5 x 30-40 pm; middle laminal

cells 72.5-90.0 x 42.5-65.0 pm; basal cells 100.0-112.5

X 50.0-87.5 pm. Oil bodies (Figure 5P) smooth, globose

and shiny, 2-15 per cell, 5. 0-7. 5 pm diam.; chloroplasts

mostly spherical, 2. 5-5.0 diam.

Dioicous presumably, but no plants with antheridia

seen. Archegonia in a row dorsally along stem (Figure

6A), naked, often obscured by suberect leaves.

Pseudoperianth campanulate (Figure 5T), generally very

close to stem apices (Figure 6C), which are often raised,

sometimes 2 alongside each other or else 2 in acropetal

sequence (Figure 6E), 2250-2475 pm long, projecting ±

750 pm above leaves, widely flaring above, from narrow,

slightly stipitate base, up to 6090 pm wide across mouth,

margin undulating, often shallowly to deeply lobed

(Figure 5S), with ± 6 widely spaced papillae, mostly at

angulations, sessile, rarely raised on basal cells, some-

times along one side deeply incised (Figure 6F); cells

comparable in shape and size to those of leaves.

Capsules globose, up to 1375 pm diam., wall bistratose,

cells of inner layer polygonal, (35-)55-70 x

(20.0-)30.0-62.5 pm, each cell wall (Figure 5V) with

1-3, rarely 4, dark brown, nodular and occasionally

semi-annular thickenings. Seta 1. 8-8.0 mm long,

300-350 pm or 10 cell rows diam. (Figure 5U). Spores

light brown, hemispherical, 42.5-55.0 pm diam., includ-

ing ridges projecting at periphery; distal face (Figure

7A-D) convex, elaborately ornamented with 7 or 8

ridges across face, partly parallel, otherwise branched,

rather thin, curved or winding somewhat, slightly undu-

lating to lightly pleated, up to 7.5 pm high and 5-10 pm

apart, some ridges anastomosing and forming closed

areas, others open-ended, crests crenulate or eroded,

space between ridges containing numerous slender spin-

ules in 1 or more rows, crowned with fine granules, and

some basally connected to sides of ridges or to one

another by faint buttressing. lines; proximal face (Figure

7E) without triradiate mark, toward centre with irregular,

prominent, as well as smaller protrusions and a few short

ridges, surrounded by numerous fine spinules in 4 or 5

irregular rows, some coalesced into small ridges, around

spore periphery a variable number of projecting spines,

mostly joined by a 5. 0-7. 5 pm wide, incomplete mem-

branous wing with crenulate margin. Elaters (Figure 7F)

pale straw-coloured, 112.5-230.0 x ± 7.5 pm, tapering to

tips, 5 pm wide, finely papillose, many entirely 2-spiral,

others alternately 3-spiral, 2-spiral, 3-spiral and 2-spiral

along length or else 2-spiral, 3-spiral and 2-spiral, some

2-spiral in one half and 3-spiral in the other.

Fossombronia marindae was collected in August

1998 on the outskirts of Worcester in the Western Cape

(Figure 4) by Ms Marinda Koekemoer, after whom I

have named it, in grateful recognition of much help and

support, especially with fieldwork. It is so far only

known from the type locality, where it was growing

together with F. densilamellata S.W.Arnell and F. spin-

osa (Perold 1999: 32) on light brown, clayey soil, con-

taining some small quartz crystals. This locality falls in

the winter rainfall Fynbos Biome (Cowling et al. 1997;

Rutherford 1997), and more specifically, in the Little

Succulent Karoo vegetation type (Low & Rebelo 1996)

which is adapted to rather dry conditions. The plants

were sporulating profusely and the species must be more

widespread than the single locality in Worcester sug-

gests.

Schuster (1992) remarked that, ‘one interesting fea-

ture, which apparently has not been noted for the genus’,

is as follows, ‘leaves produced on either side (inner

sides) at the base of a dichotomy are coalesced with each

other for some distance above their bases’. He compared

this with ‘similar paired (or bilobed) leaflike structures

(that) also occur in Pliyllothallia, at the fork of a dichoto-

my’. In F. marindae this feature was noticed at the

branching off of a lateral shoot from the main axis, but it

has also occasionally been observed at an apical dichoto-

my in another, undescribed southern African species of

Fossombronia. It is not thought to be of any taxonomic

significance in discriminating between species.

F. marindae can be recognised by usually split-open

pseudoperianths with undulating margins at the mouth;

they are often situated very close to the stem apices and

sometimes on short shoots which are apically slightly

raised. The most important character, however, is the

very elaborate spore ornamentation with 7 or 8 ridges

across the distal face and numerous spinules between. It

is surely one of the most beautiful in the entire genus and

not to be confused with that of any other species.

3. Fossombronia monticola Perold, sp. nov.

Plantae repentes, sublaxae. Folia imbricata, suberecta

vel patentia, undulata, saepe 2- vel 3-lobata, margine

cum processibus nonnullis brevibus spinosis. Dioicae.

Antheridia in plantis masculis bracteis obtecta. Pseudo-

perianthium campanulatum; ore undulato, in 11 vel 12

lobis diviso, marginibus cum usque ad 35 prominentiis

spinosis. Sporae 35-50 pm diametro; superficie distali

cum 9-11 cristis, interdum subparallelis, ceteris cursibus

variis. Elateres 92.5-147.5 longi, plerumque bis spirales,

rarissime ter spirales.

88

Bothalia 29,1 (1999)

FIGURE 5. — Fossomhronia marindae. A-L, female leaves; M, partly joined leaves produced at side shoot branching off main axis; N, female leaf

adjacent to pseudoperianth; O, detail of leaf margin; P, median leaf cells with oil bodies (solid lines) and chloroplasts (dotted lines), Q, c/s

female stem apex; R, c/s female stem base; S, opened pseudoperianth; T, pseudoperianth from side; U, c/s seta; V, cells in inner capsule

wall. A-V, M. Koekemoer 1415. Scale bars: A-M, S, T, 5(X) pm; O, U, 100 pm; P, V, 50 pm; Q, R, 250 pm. Artist: G. Condy.

Bothalia 29,1 (1999)

89

FIGURE 6. — Fossombronia marindae. A, female shoot with row of archegonia; B, young pseudoperianth near apex of branch; C, close-up view

of young pseudoperianth; D, shoot seen from side, showing arched stem with leaves; E, two pseudoperianths in acropetal sequence, distal

one hidden (indicated by arrows); F, pseudoperianth slit at side, showing seta emerging from tom calyptra, capsule dehisced. A-F, M.

Koekemoer 1415. A, C, x 18.4; B, F, x 13.8; D, x 11; E, x 13.4.

TYPE, — Western Cape, 3219 (Wuppertal): Ceder-

berg, between Elizabethfontein turnoff and Pakhuis Pass,

at seepage area, on moderate mountain slope on northern

side of Pakhuis Pass, under Stoebe plumosa and Restio

bushes, (-AA), M. Koekemoer 1397 (PRE, holo.).

Plants smallish to medium-sized, creeping, in rather

loose stands, light green, becoming translucent with age;

shoots simple or once/twice shortly furcate, occasionally

with 1 or 2 lateral branches, frequently arising basally (or

sometimes laterally) from tuberous apices of old stems,

the latter ± 350 x 500 pm (Figure 8R), flanked by small-

er reddish leaves, rounded, ± 750 x 875 pm (Figure 80),

proximally naked and tapering, ± 5 mm long; male plants

up to 5.5 X 2.4-2. 5 mm, ± 1.0 mm high; female plants

larger, up to 9 x ± 2.8 mm, ± 1.6 mm high. Stems pros-

FIGURE 7. — Fossombronia marindae. A-E, spores; F, elater. A, B, distal face; C, side view of distal face; D, detail of part of distal face; E, prox-

imal face. A-F, M. Koekemoer 1415. A, x 879; B, x 910; C, x 876; D, x 1338; E, x 780; F, x 1506.

90

Bothalia29,l (1999)

FIGURE 8. — Fossomhronia monticola. A-F, male leaves; G-N, female leaves; O, small leaf from old, tuberous stem; P, detail of leaf margin; Q,

median leaf cells with oil bodies (solid lines) and chloroplasts (dotted lines); R, c/s old tuberous stem; S, c/s male stem apex; T, c/s female

stem apex; U, c/s female stem base; Vi-Vj, perigonial bracts; W, opened pseudoperianth; X, pseudoperianth from side, showing outline of

capsule within; Y, c/s seta; Z, cells in inner capsule wall. A-Z, M. Koekemoer 1397. Scale bars: A-O, W, X, 500 pm; P, Y, 100 pm; Q, Z,

50 pm; R-U, Vi-V.;, 250 pm. Artist: G. Condy.

Bothalia 29,1 (1999)

91

trate, occasionally arched or bent, green, in cross section

plano-convex, in male plants at apex (Figure 8S)

180-200 pm (± 8 cell rows) high, 340-370 pm wide, at

base ± 200 x 380 pm; in female plants at apex ± 210 x

400 pm (Figure 8T), tapering to base, ± 200 x 250 pm

(Figure 8U). Rhizoids deep reddish pink, 12.5-20.0 pm

wide. Leaves overlapping, suberect to spreading, upper

margin undulating, often 2- or 3-lobed and with several

short spinous projections, variable in size but mostly

larger proximally than distally, somewhat fan-shaped,

generally wider above and gradually narrowing below,

sometimes decurrent, succubously inserted; in male

plants (Figure 8A-F) (725-) 1000-1 350 x 725-1350 pm,

below 550-750 pm wide; in female plants (Figure

8G-N) mostly larger, 1500-2000 x 1700-2125 pm,

below 625-1125 pm wide, margins with 11-16 papillae,

rounded or conical, 15.0-32.5 x 17.5-22.5 pm, some-

times collapsed, sessile or raised on a single basal cell

(Figure 8Ph otherwise on tiers of up to 3 cell rows,

respectively consisting of 2, 3 and 4 cells alongside each

other and arranged in a triangle, ± 160 pm long and 150

pm wide at base. Leaf cells thin- walled, generally some-

what larger in female plants, which are more common,

hence their cell dimensions are given: at upper margins

(Figure 8P), rectangular across or occasionally isodia-

metric, 32.5-37.5 x 45-65 pm, at lateral margins long-

rectangular, 45-90 X 25.0-37.5 pm; upper laminal cells

5- or 6-sided, 30-85 x 40-50 pm; middle laminal cells

100-125 X 55-65 pm; basal cells 132.5-150.0 x 75-100

pm. Oil bodies (Figure 8Q) generally faintly granular,

not quite globose, shiny, 8-17 per cell, 3-5 pm diam.;

chloroplasts mostly spherical, ± 5 pm diam.

Dioicous. Antheridia dorsally along stem (Figure

9A-C), sometimes confined to proximal part only, in

groups, short-stalked, globose, ± 170 pm diam., subtend-

ed by perigonial bracts (Figure 8V1-V5), size and shape

rather variable, body 240-480 x 120-350 pm, with up to

4 finger-like appendages, 70-140 pm long and topped

with a papilla, marginal cells 30.0-47.5 x 50.0-77.5 pm,

cells in interior 4-6-sided, 75-80 x 30.0-42.5 pm.

Archegonia in a row dorsally along stem, sometimes

even distal to a pseudoperianth, occasionally 2 in

acropetal sequence becoming fertilised. Pseudoperianth

(Figures 8W, X; 9D-F) campanulate, about as tall as

adjacent leaves, up to 1 875 pm long, flaring widely from

narrow base, ± 750 pm wide, across mouth 3300-3750

pm wide, undulating, divided into 11 or 12 lobes, some

up to 375 X 625 pm, at margin ± 35 spinous projections,

1-5 cells high and topped with a papilla; cells compara-

ble in shape and size to those of leaves; sometimes later-

ally with loose outgrowths, ± 1250 pm long and 3 cells

wide. Capsules globose, 800-1250 pm diam., walls bis-

tratose, cells of inner layer (Figure 8Z) irregularly

shaped, roughly quadrangular, rectangular or trape-

zoidal, 32.5^5.0 X 15-50 pm, each cell wall with 1^

dark brown, nodular thickenings, semi-annular thicken-

ings rare. Seta up to 2 mm long, in cross section (Figure

8Y) ± 330 pm diam., 8 cells across. Spores light brown

to deeper brown, hemispherical, 35-45(-50) pm diam.,

including ridges projecting at margins; distal face

(Figure lOA-D) convex, covered with 9-11 branched

and rejoined or discontinuous ridges, straight or curved,

short or long, sometimes a few partly parallel to one

another, others running in different directions, up to 5

pm apart and ± 3 pm high, crests crenulate, between the

bases of adjacent ridges, thin buttressing lines, occasion-

ally joined across and forming areolae, their transverse

walls fainter than the others, rarely a partly open ring

formed around a central, star-shaped, spinous process

(Figure IOC); proximal face (Figure lOE) mostly flat, tri-

radiate mark absent, with fine granules surrounding inner

coarse ones, sometimes joined up to form short, radiating

ridges or larger ‘blobs’, around periphery ± 26 spines, ±

FIGURE 9. — Fossombronia monticola. A, furcate male shoot with perigonial bracts; B, simple male shoot with bracts and one antheridium; C,

close-up of bracts and antheridium; D, female shoot with pseudoperianth; E, close-up of pseudoperianth; F, pseudoperianth with capsule

from above. A-F, M. Koekemoer 1397. A, x 17; B, x 13.3; C, x 26; D, x 10; E, x 13.8; F, x 16.5.

92

Bothalia29,l (1999)

FIGURE 10. — Fossornbroniu monticola. A-E, spores; F, elater. A-C, distal face; D, detail of part of distal face; E, proximal face. A-F, M.

Koekemoer 1397. A, x 759; B, x 794; C, x 940; D, x 1804; E, 840; F, x 1526.

2.5 |jm high and mostly connected by a thin membranous

wing or perispore. Elaters (Figure lOF) light brown,

92.5-147.5 X 7.5 pm and tapering to looped tips, mostly

2-spiral, very rarely 3-spiral, smooth.

Fossombronia monticola has fairly frequently been

collected in the Western Cape (Figure 4). The localities,

which also fall in the winter rainfall Mountain Fynbos

vegetation type, are from north to south: Matsikama

Mountains, on plateau east of Vanrhynsdorp; the type

locality in the Cederberg; Uitkyk Pass, Cederberg; Gydo

Pass, north of Prince Alfred Hamlet; Farm Merino, at

foot of Waboomberg; Bain’s Kloof; south of Tulbagh;

Kogman’s Kloof; Farm Zandrivier, near Seweweeks-

poort. The plants grow at seepage areas, often on soil

pockets on vertical rock walls of mountain passes or on

streambanks. Several of these localities are mountainous,

hence the specific epithet, monticola. The type specimen

was growing together with Riccia crozalsii Lev. and R.

purpurascens Lehm., as well as a Fissidens sp. and a

Jungermannia sp., on greyish to pale yellow soil, con-

taining fine to rather large quartz crystals. Part of this

collection had to be kept in cultivation for some weeks,

to allow the spores to ripen. Repeated samplings of the

spores were then taken to make sure that the specimen

was not mixed with other Fossombronia species.

In one of my collections of F. monticola, Perold 3293,

from the Farm Zandrivier near Seweweekspoort, I

noticed some leaves with few to many cells deeply

stained with red and containing a dark, round, internally

granular body, as I mentioned in F. tumida (Perold

1997c). Scattered cells containing spherical bodies, were

also remarked on and illustrated by Sim (1926) and

Arnell (1963) in their respective treatments of F. tumida.

Perhaps it is possible that these cells had been parasitised

by some organism.

Vegetatively F. monticola is quite a variable species,

but it can generally be distinguished by somewhat fan-

shaped leaves that are 2- or 3-lobed and mostly wider

above than below; the upper margin is undulating, and

has several short spinous projections topped with a papil-

la; the pseudoperianth is campanulate, its mouth is divid-

ed into 11 or 12 lobes and the margin has up to 35 spin-

ous projections; the distal face of the spores is covered

with 9-1 1 irregular ridges, up to 5 pm apart and some-

times joined by fainter transverse walls to form areolae.

The ornamentation on the distal face of the spores in F.

monticola resembles that of F. montaguensis S.W. Arnell,

but it differs from the latter by having lobed, undulate, not

frilly leaves; each of the cells in the inner layer of the cap-

sule wall in F. montaguensis (Perold 1997b). has several

semi-annular thickenings, which are often ring-formed,

whereas F. monticola capsule wall cells almost exclusive-

ly have nodular and only rarely any semi-annular thick-

enings; the new species is also more common.

The distal spore face ornamentation of F. monticola also

bears some resemblance to that of F. rudis Scott & Pike

(1988) nom. inval. typ. non cit., from Stoke’s Creek,

Northern Territory, Australia, in that they depict and

describe it as ‘irregularly covered with thin scattered lamel-

lae and spines’; vegetatively, however, the species differ

greatly, F. rudis having interalia a pseudoperianth raised on

a ‘stalk into which the foot projects as a perigynium’.

SPECIMENS EXAMINED

Held at PRE. Bracketed numbers after collectors’ name

and number refer to the species in the text in alphabeti-

cal order, namely; F. hyalorhiza (1), F marindae (2) and

F. monticola (3).

Bothalia29,l (1999)

93

Koekemoer 1397 (type), 1408 p.p. (3), 1415 p.p. (type) (2).

S.M. Perold 3293, 3296, 3356 (3), 3393, 3409, 3431 (1), 3443, 3447,

3448, 3818, 3844, 3851, 3864 (3), 3867 (type) (1), 3871, 3883, 3884,

3901 (3), 3902-3904 (1).

ACKNOWLEDGEMENTS

My deep thanks to Dr H. Stieperaere, Meise, for kind-

ly refereeing this article and for his helpful suggestions.

I also sincerely thank my colleagues at NBI, i.e. the cura-

tor of PRE, Ms M. Koekemoer, and also Mr J. Meyer, for

their generous help with fieldwork. Dr Theo Arts,

Belgium, is heartily thanked for his gift of Fossombronia

specimens from Madeira (and elsewhere). Dr H.F. Glen

is also thanked for translating the diagnoses into Latin. In

addition, I extend my gratitude to Ms G. Condy for the

drawings, Mrs A. Romanowski for developing and print-

ing many photographs and to Ms D. Maree for typing the

manuscript.

REFERENCES

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

Science Council. Stockholm.

COWLING. R.M. & HILTON-TAYLOR, C. 1997. Phytogeography,

flora and endemism. In R.M. Cowling, D.M. Richardson &

S.M. Pierce, Vegetation of southern Africa. Cambridge Univer-

sity Press, Cambridge.

COWLING, R.M., RICHARDSON, D M. & MUSTART, RJ. 1997.

Fynbos. In R.M. Cowling, D.M. Richardson & S.M. Pierce,

Vegetation of southern Africa. Cambndge University Press,

Cambridge.

JONES, E.W. & HARRINGTON, A.J. 1983. The hepatics of Sierra

Leone and Ghana. Bulletin of the British Museum (Natural

History), Botany 11: 215-289.

LOW, A.B. & REBELO, A.G. (eds) 1996. Vegetation of South Africa.

Lesotho and Swaziland. Department of Environmental Affairs

& Tourism, Pretoria.

PATON, J A. 1973. Taxonomic studies in the genus Fossombronia

Raddi. Journal of Bryology 7: 243-252.

PEROLD, S.M. 1997a. Studies in the liverwort genus Fossombronia

(Metzgeriales) from southern Africa. I . Three new species from

Northern Province, Gauteng and Mpumalanga. Bothalia 27:

17-27.

PEROLD, S.M. 1997b. Studies in the liverwort genus Fossombronia

(Metzgeriales) from southern Africa. 2. An amendment to three

species from Western Cape, described by S.W. Amell. Bothalia

27: 29-38.

PEROLD, S.M. 1997c. Studies in the liverwort genus Fossombronia

(Metzgeriales) from southern Africa. 4. A re-examination of

F. crispa, F. leucoxantha and F. tumida. Bothalia 27: 105-115.

PEROLD, S.M. 1999. Studies in the liverwort genus Fossombronia

(Metzgeriales) from southern Africa. 8. F. elsieae and F. spinosa,

two new Western Cape species with spinose spores. Bothalia 29:

25-33.

RUTHERFORD, M.C. 1997. Categorization of biomes. In R.M. Cowling,

D.M. Richardson & S.M. Pierce, Vegetation of southern Africa.

Cambridge University Press, Cambridge.

SCHUSTER, R.M. 1992. The Hepaticae and Anthocerotae of North

America 5: 416. Field Museum of Natural History, Chicago.

SCOTT, G.A.M. & PIKE, D C. 1988. A new species of Fossombronia

from Australia. Beiheft zur Nova Hedwigia 90: 109-1 12.

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

Royal Society of South Africa 20: 35.

Bothalia29,l: 95-117 (1999)

Notes on African plants

VARIOUS AUTHORS

ERICACEAE

TWO NEW SPECIES OF ERICA FROM WESTERN CAPE, SOUTH AFRICA

Erica hansfordii E.G.H.Oliv., sp. nov., habitu gra-

cili repenti, florescentibus floris singularis, floribus

parvis pendulis, corolla flava laevi, sepalis latis ovatis,

antheris muticis, ovulo singulari in quoque loculo

dignoscenda.

TYPE. — Western Cape, 3319 (Worcester): Slanghoek

Valley, Witelsrivier Farm, 900 ft [274 m], (-CC), 16

December 1997, Oliver & Hansford 10999 (NBG, holo.;

BM, BOL, E, G, K, MO, NY, P, PRE, S, W). Figure 1.

Shrublet diffuse, straggling amongst restiads, some-

times stouter up to 300 mm high in open areas, single-

stemmed reseeder. Branches', a few thin intertwining

main branches up to 300 mm long with internodes 4-10

mm long, numerous secondary branches 2-15 mm long.

FIGURE 1. — Erica hansfordii. A, flowering branch; B, stem; C, flowering branchlet; D, leaf; E, flower; F, bract; G, sepal; H, anther, side, back

and front views; I, stamens and gynoecium with ovary opened laterally; J, gynoecium; K, capsule; L, seed; M, testa cells, with jigsawed

inner anticlinal portion. All drawn from type, Oliver & Hansford 10999. A, x 1 . Scale bars: B, H, 0.5 mm; D-G, 1 mm; C, 2 mm. Drawings

by Inge Oliver.

96

Bothalia 29,1 (1999)

reflexed, 1 (2) per node of main branch, bearing 1 apical

inflorescence; all branches reddish brown with fine

short spreading hairs. Leaves 4-nate, ± 3.5 x 0.3 mm,

subspreading to spreading, slightly incurved to slightly

recurved, acute, adaxially flattened, very sparsely and

shortly hairy, abaxially rounded, glabrous and narrowly

sulcate, margins rounded, sparsely ciliolate, sometimes

with a few small sessile glands; petiole appressed, ± 0.6

mm long, sparsely ciliolate, yellowish. Inflorescence: a

1 -flowered unit, terminal on secondary branches only,

mostly pendulous; pedicel very short ± 0.6 mm long,

glabrous to slightly hairy, yellowish green; bract basal,

minute, 0.4 x 0.3 mm, oblong, rounded to subacute,

lacking a sulcus, ciliolate, pale yellow to white; bracte-

oles 2, partially recaulescent, in middle position, other-

wise like the bract. Calyx 4-partite; lobes slightly imbri-

cate, 0.8 X 0.7 mm, ovate, subacute, glabrous, yellow,

margins edged with a few short hairs and non-sticky

gland-tipped hairs, very narrowly sulcate in upper ‘A to

‘A. Corolla 4-lobed, 2.0-2. 3 x 2. 5-2. 8 mm, broadly

campanulate to cyathiform, glabrous, smooth, yellow

with a tinge of green; lobes 1.0 x 1.5 mm, broadly

rounded or with slight acute apex, entire. Stamens 8,

free, included; filaments straight, ± 0.8 mm long,

glabrous, yellow to golden brown apically; anthers

bilobed, ovate, muticous or with remnant of minute

decurrent spur; thecae 0.5 x 0.25 mm, narrowly ovate,

aculeate, golden brown, pore '/a length of theca; pollen

in tetrads. Ovary 4-locular, 1. 5-2.0 x 2.5 mm, obovoid to

oblate, emarginate, glabrous but pustulate, semitrans-

parent when fresh, yellow with reduced basal nectaries;

ovules 1(2) per locule, spreading from a central position

on columella, placenta not swollen; style included, 1.2

mm long, straight, yellow with slightly wider brown

apex; stigma simple truncate. Capsule 2. 0-0.8 mm,

broadly cyathiform; valves splitting only half their

length with slightly incurved apices, splitting complete-

ly from columella to 135° angle, outside reticulate-fove-

olate; septa only on valves. Seeds 0.3 x 0.25 mm, ellip-

soid, golden brown and shiny, testa deeply reticulate,

anticlinal walls thick and jigsawed towards base becom-

ing semi-straight at outer limits.

E. hansfordii is distinguished by its delicate creeping

habit, single-flowered florescences, by the flowers being

small and pendulous with a clear yellow, smooth corolla,

small broad ovate sepals, muticous anthers and a single

spreading ovule per locule. This species is probably relat-

ed to another localised species, E. rehmii Dulfer, which

occurs nearby in the mountains of the northeastern end of

Bain’s Kloof and is also confined to marshy seeps. The

latter also has yellow flowers, similar sepals,

bract/bracteoles, anthers and ovary, but differs in being

much larger in stature (woody up to 1 m high), with

prominent larger flowers which are erect and arranged in

4-flowered florescences.

There are several Erica species that look very similar

to E. hansfordii with the same distinctive habit, which is

associated with marshy habitats dominated by short tuft-

ed restiads, but none occurs in the same region. E. myri-

ocodon Guthrie & Bolus occurs in the Franschhoek area

and has pale pink flowers, which are hairy with linear

sepals, a longer pedicel, appendiculate anthers and a

hairy ovary. The two very closely related species, E.

18° 19° 22! 2f

FIGURE 2. — The known distribution of Erica hansfordii, O; and E.

feminarum, •.

limosa L. Bolus and E. salteri L. Bolus, both endemic in

the Cape Peninsula also have similar muticous anthers

and a glabrous ovary, but the flowers are hairy and are a

dirty pinkish cream colour on a longer pedicel and with

linear sepals.

This species is a highly restricted one occurring in

only two seepage areas on the sandy flats in the

Slanghoek Valley between Worcester and Wolseley, flats

which are covered with short fynbos (Figure 2). These

seepage areas are rather far from the source of their

water, namely the very dominant Cossacks Peaks, and

thus it is surprising that this special habitat is situated

there. There is much encroachment of farming (mainly

vineyards) in the valley and therefore the species is clear-

ly endangered, but fortunately the owner of the ground is

aware of its importance.

The plants of Erica hansfordii are very inconspicuous

in their habitat being very delicate and intertwining

among the dominant matted restiad, Restio zwartber-

gense. Also they are mostly below the upper levels of the

restiad and have their flowers hanging downwards. The

habitat remains wet for the whole year so the plants tend

to have a continuous flowering period throughout the

year (G. Hansford pers. comm.).

With only very reduced nectaries and inconspicuous

pendulous pale yellow-coloured flowers, it is difficult to

postulate what the pollinating agent could be for this

species. None was evident when the type material was

collected. Elowering time: flowers can be found in most

months of the year, but the peak flowering period is in

early summer.

The species is named after Gerard Hansford, an agri-

cultural consultant from Tulbagh, who discovered it very

recently during one of his many botanical forays in the

district. He has provided the National Botanical Institute

with many useful and interesting records of plants.

FIGURE 3. — Erica feminarum. A, flowering branch; B, flowering branchlet; C, stem; D, leaf, abaxial and lateral views; E, flower; F, bracteole

and bract; G, sepal; H, half flower with ovary opened laterally; I, gynoecium; J, anther, side, front and back views. All drawn from type,

Oliver 10844. A, x 1. Scale bars: B, 4 mm; C-I, 3 mm; J, 1 mm. Drawings by Inge Oliver.

WESTERN CAPE. — 3319 (Worcester): Slanghoek area,

Witelsrivier Farm, 280 m, (-CC), 15-02-1997, Hansford 96 (K, NBG,

PRE, para.); ibid., 1-06-1997, Hansford 99 (MO, NBG, PRE, para );

ibid., 24-08-1997, Hansford 100 (BOL, NBG, para.).

Erica feminarum E.G.H.Oliv., sp. nov., floribus

atrorubentibus subtiliter pilosis, saepe in synflorescenti-

bus pseudospicatis, sepalis latis ovatis et antheris muticis

notabilis.

TYPE. — Western Cape, 3319 (Worcester): Stettyns-

berg, slopes east of Beacon 68 above Elandskloof, 3860 ft

[1 176 m], (-CD), 9 August 1997, E.G.H. & I.M. Oliver

10840 (NBG, holo.; BM, BOL, E, K, MO, NY, P, PRE, S).

Erect, slender shrublet 200-300 mm high, single-

stemmed reseeder. Branches: 1-3 erect main branches,

50-130 mm long, continuing vegetative growth, rarely

ending in a florescence, numerous erect secondary

branchlets, (2)3 at every node, 2-5 mm long, with a ter-

minal florescence; all branches with simple short spread-

ing hairs; internodes on main branches ± 10 mm long, on

secondary branchlets 0.5 mm long. Leaves 3-nate, erect,

± 4.0 X 0.5 mm, narrowly oblong, incurved, adaxially

flattened and minutely hairy, abaxially rounded and nar-

rowly sulcate, with subacute margins, minutely hairy

towards margins, ciliolate; petiole appressed, 0.8 mm

long, ciliolate (Figure 3). Inflorescence: 1 -flowered ter-

minal on short secondary branchlets, pendent, often

98

Bothalia29,l (1999)

aggregated into spike-like synflorescences towards ends

of main branches; pedicel 1.5-1. 8 mm long, curved,

minutely hairy, red; bract partially recaulescent in basal

position, appressed, 0.9 x 0.3 mm, lanceolate, without a

sulcus, minutely hairy, red; bracteoles 2, near middle of

pedicel, like bract but slightly smaller. Calyx 4-parti te,

appressed to corolla, minutely hairy; lobes imbricate at

base, 1.1 X 1.1 mm, broadly ovate, ciliolate, dark red.

Corolla 4-lobed, 4x3 mm, cyathiform, dark red, buds

almost black, covered with minute white unicellular

hairs; lobes 1.1 x 1.8 mm, broadly triangular, erect,

acute, ciliolate, entire. Stamens 8, free, included; fila-

ments 2 mm long, linear, erect, straight, glabrous, red;

anthers bilobed elliptic to ovate in outline, erect, light

brown, muticous; thecae appressed, 0.5 x 0.3 mm ellip-

tic to ovate, smooth, pore V3 length of theca; pollen in

tetrads. Ovary’ 4-locular, 1.0 x 0.9 mm, depressed glo-

bose, emarginate, glabrous, green, with basal nectaries;

ovules 4-6 per locule, spreading from placenta covering

entire columella; style included, 2 mm long, narrowly

cylindrical but broadening towards base, glabrous red;

stigma capitate dark red. Fruit not seen.

E. feminarum is a very distinct species with its dark,

wine-red flowers that appear almost black in direct sun-

light, are very finely hairy, occurring singly on very short

lateral branchlets, these often arranged in a pseudospike

on the main branches, and have muticous anthers.

There are indications that a relationship exists with

the E. xanthina/cremea complex despite the difference in

size of the corollas, the latter having long tubular flow-

ers. All have a finely hairy corolla with non-spreading

lobes, small broad sepals with similar appressed bract

and bracteoles and similar anthers. They also have the

same habitat preferences, namely seeps or moist gullies

at high altitude. E. cremea Dulfer with cream-coloured

flowers is a restricted endemic in the mountains of Bain’s

Kloof and E. xantluna Guthrie & Bolus occurs along the

western part of the Riviersonderend Range. It has pale

cream to pinkish flowers. There is also an alliance, but

more remote, with the long-tubed E. kogelbergensis

E.G.H.Oliv. with orange and yellow or plain yellow

flowers from the Kogelberg area and E. colorans Andr.

with white to pink flowers from the lower hills and flats

in the Bredasdorp District.

The specific epithet, feminarum = of women, was

chosen because the species was first collected by a

woman, Elsie Esterhuysen, 50 years ago and found again

by us on National Women’s Day, 1997.

The species is highly restricted in its distribution,

occurring, as far as is known, in only one small seepage

area on the eastern slopes of Stettynsberg in the

Villiersdorp area (Figure 2). These slopes are well pro-

vided with seeps, but a search of the area located only a

single population of plants that were very inconspicuous

due to the dark colour of the flowers. With the very

mountainous tenain in this region of the Western Cape

there could well be other habitats on the nearby moun-

tains of the under-collected Goudini Range. The plants

occur at sufficiently high altitude to be covered by snow

for short periods in winter.

The pollinating agent of the species is unknown. The

species is obviously entomophilous with its small,

included stigma and nectaries around the base of the

ovary. The flowers had a very slight musty odour in the

field. With the dark colour of the flowers that hang

downwards, there appears to be no clear attraction for a

foraging insect flying overhead. However, when viewed

from below with the sun shining through the flowers, the

colour turns to a glowing red that could well be an attrac-

tion for an insect below the plant. This feature coupled

with the odour may indicate that flies could be the polli-

nating agent of the species. Flowering time: July and

August.

WESTERN CAPE.— 3319 (Worcester): Stettynsberg, E side, 4000

ft [1 219 m], (-CD), 31-07-1949, Esterliuvseii 15587 (BM, BOL, K,

MO, NBG, NY, PRE, para.).

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

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

7735 Claremont, Cape Town.

MS. received: 1998-03-23.

SOLANACEAE

THREE NEW RECORDS OF SOLANUM SECTION OLIGANTHES IN SOUTHERN AFRICA

During the investigation of the specimens of Solamim

L. in PRE for a revision of the genus in southern Africa,

it became clear that three species of subgenus Lepto-

stemonum (Dunal) Bitter (1919) growing in the eastern

parts of southern Africa, were not recorded for the region

by Welman (1993). These are S. litoraneum A.E.G0115.,

S. torreanum A.E.Gon^. and S. usaramense Dammer of

section Oliganthes (Dunal) Bitter (1923), all described

from tropical Africa.

Solaniim litoraneum A.E.Gon^. in Kew Bulletin

52; 703, fig. 1 (1997). Type: Mozambique, Maputo Pro-

vince, Inhaca Island, Mogg 27597 (K, holo.; J!, PRE!,

SRGH, iso.).

Distribution and ecology

S. litoraneum seems to be restricted to the coastal

areas of southern Mozambique and northern KwaZulu-

Natal; from the Massinga District south to Black Rock

between Kosi Bay and Lake Sibayi, from where only one

specimen is represented in PRE. It is near-endemic to the

Maputaland Centre of Endemism of Van Wyk ( 1996).

It grows from sea level to ± 50 m in the littoral vege-

tation of dunes, in thickets or the margins of dense bush-

land or dune scrub forests, always on sandy soils. From

the few specimens available, it seems to flower from

September to April and fruit from September to July.

BothaIia29, 1 (1999)

99

Taxonomy and diagnostic characters

S. litoraneum is closely related to S. riibetorum Dunal

and may eventually prove to be part of it. In the southern

African context, S. litoraneum can be distinguished by

the large inflorescences with up to 14 flowers, the fairly

large flowers with corollas 8-15 mm diam., the mostly

recurved spines and the orange-red fruit as well as the

coastal habitat.

Specimen examined

KWAZULU-NATAL.— 2632 (Bela Vista); Black Rock, (-DD),

Venler 6264 (PRE),

Solanum torreanum A.f.Gonp. in Kew Bulletin 52;

706, fig. 2 (1997). Type: Mozambique, Maputo Province,

Boane, Matola District, Balsinhas 1466 (DISC, holo.;

LMA, iso.).

Distribution and ecology

S. torreaniim occurs in the Maputo province in south-

ern Mozambique, in Swaziland and also in the southern

Mpumalanga and northern KwaZulu-Natal provinces of

South Africa. It is endemic to the Maputaland Centre of

Endemism of Van Wyk (1996), and is mainly associated

with the Lebombo Range and its foothills.

S. torreaniim grows from ± 15-300 m altitude on flats

and on slopes, on black turf or sandy to sandy-clay and

stony soils, also soils rich in lime. It is found in dry

Acacia bushveld, in mixed scrub with Combretum/

Sclerocarya and Themeda, also in scrub forest margins

and in ruderal places. Flowering and fruiting times:

October to July.

Taxonomy and diagnostic characters

Gonsalves (1997) regards this species as close to S.

capense L. It can be distinguished from that species by

the 2-8-flowered inflorescences with corollas 15-25 mm

diam., and also the tendency of S. torreaniim to dry dark

brownish. This is the species which Ross (1972) listed as

Solanum sp. no. 29 from Zululand with the cited speci-

men Strey 4611 .

Specimens examined

MPUMALANGA. — 2531 (Komatipoort): Figtree, (-DD), Nel 68 (PRE).

SWAZILAND, — 2531 (Komatipoort): Tshaneni, (-DC), Barrett 246

(PRE). 2631 (Mbabane): Lubulla Ranch, (-AD), Corbett s.n. (PRE);

Lubuku Ranch, (-BD), Phillips s.n. (PRE). 2731 (Louwsburg); Ingwa-

vuma Poort, (-BB), Compton 28584 (PRE).

KWAZULU-NATAL.— 2632 (Bela Vista): Nkonjane/Aberkom Drift,

(-CC), Moll & Pooley 4224 (NH, PRE); Ndumu Hill, (-CD). Pooley

1672 (NU, PRE). 2731 (Louwsburg): Pongola Poort, (-BC), Strey 4611

(NH, PRE); Nkonkoma, (-DC), Wells 2155 (PRE). 2732 (Ubombo):

Ingwavuma foothills, (-AA), Strey 10258 (NH, PRE); False Bay Park,

(-CD), Taylor 730 (NH, PRE). 2831 (Nkandla); Zondela, Mahlabatini,

(-AD), Gerstner 4309 (PRE).

Solanum usaramense Dammer in Engl., Die

Pflanzenwelt Ost-Afrikas und der Nachbargebiete, Theil

C: 353 (1895). Type: Tanzania, Usaramo, Stuhlmann 7066

(B, holo.f).

Distribution and ecology

Recorded from coastal Tanzania and Mozambique

southwards to the northern KwaZulu-Natal coast in the

Ingwavuma, Ubombo and Hlabisa Districts.

S. usaramense grows just above sea level in shady

areas along the margins of dune forests; also found in

coastal swamp forests. Flowering time: November to

March. Fruiting time: Eebruary to April.

Taxonomy and diagnostic characters

S. usaramense is closely related to S.filicaiile Dammer

and may be part of S. cordatiim Eorssk. This is the species

which Ross (1972) and Vahrmeijer (1966) listed as 5.

zanzibarense Vatke var. vagans (C.H. Wright) Bitter from

Maputaland (Tongaland), without a cited specimen. Van

Wyk ( 1996) listed this as Solanum sp. nov. A [Vahrmeijer

739). Material of this species has also been filed under 5.

cf monotanthiim Dammer in South African herbaria.

In South Africa it can be distinguished by its climbing

habit, greyish green stems and lower leaf surfaces, its fair-

ly large flowers with blue-purple corollas which are 15

mm diam., its recurved spines and also its coastal habitat.

Specimens examined

KWAZULU-NATAL.— 2732 (Ubombo): Lake Sibayi, (-BC),

Vahrmeijer 739 (PRE); Lalanek Dune Forest, (-BD), Venter 4377

(PRE); Sordwana Bay, (-DA), Vahrmeijer 613 (PRE). 2832

(Mtubatuba): Futululu Forest, (-AC), Strey 10411 (NH, PRE).

ACKNOWLEDGEMENTS

The author is grateful for valuable comments by Dr

R.N. Lester and Prof. A.E. van Wyk.

REFERENCES

BITTER, G. 1919. Die papuasischen Arten von Solanum. Botanische

Jahrbiicher 55: 59-1 1 3.

BITTER, G. 1923. Solana africana IV. Repertorium specierum

novarum regni vegetabilis. Beiheft 16: 1-320.

DAMMER, U. 1895. Solanaceae. In A. Engler, Die Pflanzenwelt Ost-

Afrikas und der Nachbargebiete, Theil C: 351-356. Dietrich

Reimer, Berlin.

GONSALVES, A.E. 1997. Two new species of Solanum (Solanaceae)

from Mozambique. Kew Bulletin 52: 703-709.

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

of South Africa No. 39.

VAHRMEIJER, J. 1966. Notes on the vegetation of northern Zululand.

African Wild Life 20: 151-161.

VAN WYK, A.E. 1996. Biodiversity of the Maputaland Centre. In

L.J.G. van der Maesen, X.M. van der Burgt & J.M. van

Medenbach de Rooy, The biodiversity of African plants:

198-207. Kluwer Academic Publishers, Dordrecht.

WELMAN, W.G. 1993. Solanaceae. In T.H. Arnold & B.C. de Wet,

Plants of southern Africa: names and distribution. Memoirs of

the Botanical Suivey of South Africa No. 62: 617-621.

W.G. WELMAN*

* National Botanical Institute, Private Bag XI 01, 0001 Pretoria.

MS. received; 1998-05-21.

100

Bothalia29,l (1999)

AGAVACEAE

AGAVE VIVIPARA: THE CORRECT NAME FOR AGAVE ANGUSTIFOLIA

At least three taxa of Agave L. (Agavaceae) are natu-

ralised in southern Africa. Two of these, A. americana L.

var. americana and A. sisalana Perrine have been treated

by Smith & Mossmer (1996) in the series FSA contribu-

tions. A catalogue of the naturalised flora of any country

is by definition dynamic: new species names are added to

the list as and when they are reported. It is therefore not

surprising that the naturalisation of a third species, A.

decipiens Baker, only came to light recently (Smith &

Steyn in prep.). Some species of Agave are aggressive

growers and it is likely that further taxa will in due

course be added to the ever-expanding list of naturalised

aliens. A. vivipara L. (Figure 4), a native of Mexico

(Gentry 1982), is one such species. This agave is often

offered for sale at nurseries and is frequently grown as a

garden ornamental, as a hedge plant or as a barrier plant.

However, in South Africa this species is consistently,

but incorrectly known under a later homotypic synonym,

A. angustifolia Haw. Wijnands (1983) and Forster (1992)

have shown conclusively that the correct name for this

species is A. vivipara. The following nomenclature

should therefore be adopted for the two taxa of this

species, namely A. vivipara var. vivipara and A. vivipara

cv. Marginata, which are found in South Africa;

Agave vivipara L. var. vivipara. Species plan-

tarum 1; 323 (1753). Type: Commelin, Praeludia botani-

ca 65, t. 15 (1703) [lectotypified by Wijnands (1983)].

Agave angustifolia Haw: 72 (1812).

Agave vivipara cv. Marginata (Hort. ex Gentry)

PForster in Brittonia 44: 74 (1992). Type: not designated.

Agave angustifolia var. marginata Hort. ex Gentry: 564 (1982).

The leaves of this horticultural cultivar have distinct

yellow edges. The cultivar Variegata with leaves that are

variously striped with yellow or white has yet to be

recorded for southern Africa.

REFERENCES

FORSTER, PI. 1992. New varietal combinations in Agave vivipara

(Agavaceae). Brittonia 44: 74, 75.

FIGURE 4. — Agave vivipara growing on an empty stand in urban

Pretoria, South Africa. Photograph: A. Romanowski.

GENTRY, H.S. 1982. Agaves of continental North America. University

of Arizona Press, Tucson.

HAWORTH, A.H. 1812. Synopsis plantarum succulentarurn, edn 1.

Richard Taylor, London.

LINNAEUS, C. 1753. Species plantarum, edn 1. Salvius, Stockholm.

SMITH. G.F. & MOSSMER, M. 1996. FSA contributions 4:

Agavaceae. Bothalia 26: 31-35.

SMITH, G.F. & STEYN, E.M.A. in prep. A first record of Agave decip-

iens naturalised in South Africa. South African Journal of Botany.

WIJNANDS, O. 1983. The botany of the Commelins. Balkema, Rotter-

dam.

G.F. SMITH* and E.M.A. STEYN*

* National Botanical Institute, Private Bag XI 01, 0001 Pretoria.

MS. received: 1998-10-26.

LAMIACEAE

.SALVIA THERMARA, A NEW SPECIES FROM THE WESTERN CAPE, SOUTH AFRICA

Salvia thermara Van Jaar.sv., sp. nov., a S. granit-

ica Hochst. foliis oblanceolatis dimidio distali serrulatis,

racemis brevibus densis 3-4 verticillis, corolla 40-50

mm longa differ!. Figure 5.

Erect branched shrub, up to 1 m tall from a stolonifer-

ous base. Stolons horizontal with erect aerial branches.

Branches square, green, up to 3 mm diam., purplish when

dry; the young vegetative parts glandular hairy, becoming

Bothalia29,l (1999)

101

FIGURE 5. — Salvia thermara. A,

part of plant with inflores-

cence; B, calyx; C, leaf; D,

stamen, showing connective

branch with sterile theca; E,

part of calyx showing pistil,

ovary and young developing

nutlets. A-C, x 1; D, E, x 2.

glabrescent with age. Leaves simple, glabrous, sessile to

subsessile, oblanceolate to linear-oblanceolate, 20-35 x

4—7 mm, green (green group 138B, R.H.S. Colour Chart),

with prominent veins below, margin serrate in upper half,

rarely entire (teeth up to 2 mm long, with up to 6 pairs of

teeth); apex acute to acuminate; upper leaves gradually

smaller and similar to bracts. Inflorescence a short dense

terminal raceme 80-100 mm long (not branched); verti-

cillasters 3 or 4, 2-flowered, up to 7 mm apart. Bracts

oblanceolate-acuminate, glandular hairy, 7-8 x 2.5 mm.

Pedicels ascending, 5-6 mm long. Calyx tubular-campan-

ulate 23-25 mm long, up to 8-veined and covered with

short eglandular hairs, not expanding in fruit, two-lipped,

reddish purple (red-purple group 59B, R.H.S. Colour

Chart); upper lip oblong, up to 10 mm long, more or less

truncate, 3 subequally toothed; teeth 2 mm long, acumi-

nate; lower lip oblong, 10 mm long, 2-toothed; teeth

acuminate, 4 mm long. Corolla, 40-50 mm long, reddish

102

Bothalia29,l (1999)

FIGURE 6. — The inflorescence of Salvia thermara.

(red group 37A, R.H.S. Colour Chart); tube oblong, up to

22 mm long, 2-3 mm wide at base and expanding to 12

mm; hood falcate to subfalcate, oblong, sometimes

straight, 15-16 mm long, 5 mm deep; lower lip drooping,

two-lobed, up to 7 X 9 mm (Figure 6). Staminal connec-

tives 19-20 mm long, with sterile lower theca (4.0 x 1.5

mm) pointing inwards; filaments 4 mm long; fertile the-

cae oblong, 3 mm long; pollen white. Ovary broadly glo-

bose-ovate, 3 mm diam.; style slender, 50-63 mm long.

Nutlets dark brown, smooth, up to 3.5 x 3 mm. Flowering

time: December- January.

TYPE. — Western Cape, 3319 (Worcester): Badsberg,

Goudini Spa, near Worcester on the Slanghoek trail, 15

Dec. 1997, (-CB), Van Jaarsveld & Vorster 15533 (NBG,

holo!).

Salvia thermara occurs on the lower N slope of the

Badsberg (adjacent to Slanghoek Mountains) in Fynbos,

among quartzitic sandstone rock and soil (Figure 7). The

specific epithet thermara pertains to the Goudini warm

springs where the plant was collected. It occurs locally,

scattered in full sun forming small groups. Associated

species include Protea nitida, P. laurifolia, Diospyros

glabra, Leucadendron salignum, Montinia caryophyl-

lacea, Othonna parviflora, Oldenburgia paradoxa and

Maytenus oleoides.

It is interesting to note the galls on the stems as a

result of perhaps a parasitic wasp. These appear as small

growths of about 10 mm diam. bearing ovate, densely

hairy, imbricately arranged leaves in a rosette.

S. thermara is related to S. granitica Hochst., a

species confined to an area south of Citrusdal and in the

Caledon District. It is at once distinguished from S.

granitica by its large reddish corolla, 40-50 mm long,

and short dense racemes consisting of 3 or 4 verticil-

lasters. S. granitica is a smaller perennial up to 0.6 m

high with a mauve-pink corolla, 20 mm long and elon-

gated racemes. Among the South African species, it

shares with 5. thermara the two-flowered verticillasters,

the upper lip of the corolla distinctly falcate, the truncate

upper calyx lobe, its stoloniferous habit and also an

absence of aroma when the leaves are crushed. Among

the other Western Cape species it superficially resembles

FIGURE 7. — Distribution of Salvia thermara.

S. lanceolata Lam. and S. africana-lutea L., both species

bearing large reddish brown to rose corollas in 2-4-flow-

ered verticillasters and with a strong aroma. Both are

much-branched shrubs up to 2 m tall (not stoloniferous)

bearing grey velvety leaves.

The genus Salvia in Africa was treated by Hedge

(1974) and is represented by ± 60 species of which 27

have been recorded for South Africa and also treated by

Codd for the Flora of southern Africa (1985). Two cen-

tres are recognised: 1, along the Mediterranean coast in N

Africa; and 2, a centre in South Africa confined to the

southern parts of the Western and Eastern Cape. The

Western Cape species are mostly large shrubby species

bearing attractive flowers, whereas many of the Eastern

Cape species are herbaceous with weedy characteristics.

S. thermara shows promise as a garden subject together

with other South African species such as S. africana-lutea

L., S. muirii L.Bolus, S. lanceolata Lam., S. africana-

caerulea L., S. dentata Aiton, S. chamelaeagnea Berg, S.

albicaulis Benth., S. radula Benth. and S. dolomitica

Codd. The reddish colour of the corolla of S. thermara is

unique among the South African salvias.

ACKNOWLEDGEMENTS

Dr O.A. Leistner is thanked for the Latin diagnosis,

G. Germishuizen for editing the manuscript, and Vicky

Thomas for the illustration.

REFERENCES

CODD, L.E. 1985. Salvia. Flora of southern Africa 28: 79-101.

HEDGE, I.C. 1974, A revision of Salvia in Africa. Notes from the Royal

Botanic Garden Edinburgh 33: 81.

E.J. VAN JAARSVELD*

*National Botanical Institute, Private Bag X7, 7735 Claremont, Cape

Town,

MS. received: 1998-01-13.

Bothalia 29,1 (1999)

103

POLYPODIACEAE

POLYPODIUM ENSIFORME, THE CORRECT NAME FOR MICROSORUM ENSIFORME (POLYPODIOIDEAE)

INTRODUCTION

In southern Africa four fern species are generally

ascribed to the tribe Microsoreae of the family

Polypodiaceae. Sim (1895, 1915) included them all in

the genus Polypodium L. but Schelpe (1969a & b, 1970)

placed P. punctatum L. and P. pappei Mett. ex Kuhn in

Microsorum L. (as Microsorium) and P. ensiforme

Thunb. and P. scolopendria Burm.f. in Phymatodes

C.Presl. Phymatodes, however, is an illegitimate name

and was replaced by Phymatosorus Pic.Serm. who

placed both these species in that genus (Pichi Sermolli

1973). In an unpublished checklist of southern African

pteridophytes, pending the Flora of southern Africa,

Schelpe followed Copeland (1947) by including

Phymatosorus (as Phymatodes) in Microsorum. This

view was followed by Jacobsen (1983), Schelpe &

Anthony (1986) and Burrows (1990). In her revision of

the genus Microsorum (Bosman 1991), only M. puncta-

tum (L.) Copel. was retained in the genus. The other

species were either placed in Phymatosorus or suggested

as belonging to the genus Neocheiropteris Christ.

Although recognising their close affinity, all three of

these genera are widely accepted (Ching 1933; De Vol &

Kuo 1980; Tryon & Lugardon 1990; Hennipman et al.

1990; Bosman 1991). Nooteboom (1997), however,

placed the last-named genera in synonymy under

Microsorum. He also, quite erroneously, placed M. ensi-

forme (Thunb.) Schelpe in synonymy under M. scolpen-

dria (Burm.f.) Copel.

In view of these varied opinions it was thought neces-

sary to review the southern African taxa ascribed to this

group of ferns.

MATERIAL AND METHODS

Sections of rhizomes and fronds were obtained from

plants cultivated at Kirstenbosch National Botanic

Garden. Samples were also taken from specimens

housed in the Compton Herbarium from both the NBG

and SAM collections. Leaf clearings were prepared

using household bleach. Illustrations were prepared

using a drawing tube fitted to an Olympus CH-2 light

microscope.

DISCUSSION AND CONCLUSION

Microsorum was placed in the subfamily Micro-

sorioideae of the Polypodiaceae (Nayar 1970) but this

generic assemblage was amended and the redefined

group reduced to tribal level (Tu 1981). The group is cur-

rently defined as terrestrial, epilithic or epiphytic plants

with a creeping, dorsoventrally flattened rhizome that

bears basally attached or peltate, clathrate paleae. The

lamina is simple or pinnately dissected with entire or sin-

uate margins. The venation forms a complex reticulum

with free, simple or branched, recurrent and excurrent

veinlets in the areoles. The veinlets mostly terminate in a

hydathode. Sori are irregularly scattered or they are vari-

ously arranged but always occur at a plexus of connec-

tives or at a plexus of distinct soral veins and connectives

(Figure 8A). Paraphyses are mostly simple and uniseriate

but peltate paraphyses occur in a few species. The spores

have plain, irregularly rugate or tuberculate surfaces often

with small papillae and globules. The ultrastructure of the

exospore is described by Hennipman & Roos (1983).

Many of the characters that define the Microsoreae

are not unique to the group but also occur in other groups

within the Polypodioideae. As a result, the definition and

circumscription of monophyletic groups within the sub-

family is varied and still much debated. The microsorioid

ferns, however, are widely accepted as a group of ferns

with a common descent (Copeland 1947; Nayar 1970;

Hetterscheid & Hennipman 1984; Bosman 1991;

Nooteboom 1997). Perhaps the most diagnostic feature

of the Microsoreae is its venation pattern and soral inner-

vation that has been studied in great detail by

Hetterscheid & Hennipman (1984) and Bosman (1991).

The venation pattern and soral innervation of M.

pappei, M. punctatum and M. scolopendrium are typical-

ly that of microsorioid ferns. In M. ensiforme, however,

the veins form costal areoles and to a lesser degree small-

er areoles near the margin (Figure 8E, F). The costal are-

oles are very prominent and contain an excurrent, simple

or branched, free veinlet. In some instances this anadro-

mous vein anastomoses with a tertiary vein to form a nar-

row rectangular secondary areole. Apart from the soral

veins, free veinlets are rarely formed in any of the are-

oles, but excurrent ones do form along the margin. Some

veins may end in an inconspicuous enlarged vein ending,

while others do not. The sori are superficial, adaxially

pustulate, circular, and up to 5 mm diam. They are posi-

tioned in a single row on either side of the primary vein

and/or costa at a plexus of a distinct branched excurrent

soral veinlet. A single sorus occurs within each costal

areole (Figure 8F). This venation pattern and type of

soral innervation does not occur in any of the known

Microsorum species nor does the pattern correspond with

that of the Microsoreae as currently defined. It is there-

fore suggested that M. ensiforme be removed from the

Microsoreae and placed in the Polypodeae.

Generic delimitation within the Polypodeae, especial-

ly the Polypodium-Microgramma-Pleopeltis complex is

problematical (De la Sota 1973; Mitsuta 1984). The last-

mentioned two genera are often treated as subgenera of

Polypodium (Stolze 1981; Proctor 1989) whereas others,

although realising their close affinity, treat them as dis-

tinct (Copeland 1947; Holttum 1954; Tryon & Tryon

1982). Hennipman et al. (1990) treat these genera as ten-

tative groups of Polypodium. The close affinity of these

genera is emphasised by the occurrence of natural

hybrids between Polypodium and Pleopeltis (Anthony &

Schelpe 1985; Mickel & Beitel 1987) and Polypodium

and Microgramma (Gomez 1975). Ribulose-1, 5 buphos-

phate carboxylase (RbcL) data, however, support the

104

Bothalia29,l (1999)

FIGURE 8. — Indumentum, venation, soral innervation and paraphyses in A, Microsorum punctatum\ E, F, H, Hh, Polypodium ensi-

forme', G, L, LI, M, N, P. vulgurc, C, D, I, li, J, Jj, K, Pleopeltis macwcarpa and B, O, Oo, P, Q, Micrognimma lycopodioides.

A-G, venation and soral innervation; H, Hh, I, li, L, LI, O, Oo, rhizome paleae; M, N, P, K, lamina indumentum; J, Jj, Q, para-

physes. A, Roux 603 (NBG); B, Van Jaarsveld 2158 (NBG); C, D, Roux 2770 (NBG); E, F, H, Hh, Roux 1997 (NBG); G, Roux

2336 (NBG); I, li, J, JJ, Roux 439 (NBG); K, Roux 2274 (NBG); L, LI, M, N, Matthews 1032 (NBG); O, Oo, P, Roux 665

(NBG); Q, Buitendag 533 (NBG). Scale bars: A-G, 5 mm; H, I, L, O, 0,25 mm; Hh-Jj, LI, Oo, 0.2 mm; J, K, N, 0.1 mm; M,

0.25 mm; P, 0.25 mm; Q, 0.2 mm.

TABLE 1. — Comparative morphology of Polypodiiim ensiforme, Polypodium, Pleopeltis and Microgyamma

Bothalia29,l (1999)

O'

a;

S CQ

u oo

5 ^

o

E

105

-o

'5

'•B

o

Cl.

(Li

S o

C

U C

g ^ OJ

O Cl, ^

Q

£ ^

Paraphyses simple uniseriate hairs absent/simple uniseriate hairs/ absent/peltate clathrate paleae uniseriate (often glandular)

peltate clathrate paleae hairs/narrow paleae

106

Bothalia29,l (1999)

boundary between Polypodium and Pleopeltis (Haufler

& Ranker 1995).

In the Polypodeae, as in the Microsoreae, the charac-

ters that define generic boundaries are mostly shared.

The position of M. ensiforme, henceforth referred to as

Polypodium ensiforme, in the Polypodium-Pleopeltis-

Microgramma assemblage is therefore not clear. Table 1

summarises some of the most important morphological

characters of P. ensiforme and that of the Polypodium-

Pleopellis-Microgramme assemblage.

Based on the data provided in Table 1, it is evident that

P. ensiforme indeed belongs to Polypodium rather than

Pleopeltis or Microgramma, as their characteristics are all

identical. The inclusion of P. ensiforme in Goniophlebium

C.Presl as suggested by Fee (1852) and defined by Rodl-

Linder (1990) is not supported by the venation pattern,

spore morphology and distribution of that genus.

Tribus Polypodeae

Polypodium ensiforme Thunb., Prodromus plan-

tarum capensium: 172 (1800); Sim: 272 (1915). Mar-

ginaria ensiformis (Thunb.) C.Presl: 188 (1836). Phlebo-

dium ensifonnis (Thunb.) J.Sm.: 59 (1841). Goniophle-

bium ensiforme (Thunb.) Fee: 255 (1852). Phymatodes

ensifomns (Thunb.) Schelpe: 135 (1969b); Jacobsen: 316

(1983). Phymatosorus ensiformis (Thunb.) Pic.Serm.: 459

(1973). Microsorum ensiforme (Thunb.) Schelpe: 151

(1982); Schelpe: 165, t. 51, fig. 2, 2a (1986); Burrows:

202, t. 46, fig. 205 (1990), as 'Microsorium'. Type:

Grootvadersbosch, Thunberg s.n. {UPS-THUNB24492,

holo.).

Tribus Microsoreae

Microsorum pappei (Mett. ex Kuhn) Ching in

Bulletin of the Fan Memorial Institute of Biology,

Botany 4: 295 (1933); Jacobsen: 313 (1983); Schelpe:

163, t. 51, fig. 1, la (1986); Burrows: 199, t. 46, fig. 202

(1990). Polypodium pappei Mett. ex Kuhn: 150 (1868);

Sim: 277, t. 141, fig. 1 0915). Types: Prom. bon. spei.

Caffraria, Rawson s.n. (BM, ?isosyn.); Natalia, Sander-

son s.n. (K, ?isosyn.).

Microsorum punctatum (L.) Copel. in University

of California Publications in Botany 16: 111 (1929);

Jacobsen: 312 (1983); Schelpe; 163 (1986); Burrows:

199, t. 46, fig. 203 (1990), as ‘Microsorium' . Acro-

stichum punctatum L.: 1524 (1763). Polypodium puncta-

tum (L.) Sw.: 21 (1802), non Thunb. (1784); Sim: 165, t.

145, fig. 2 (1915). Pleopeltis punctata (L.) Bedd.: 22

(1876). Type: China, Fothergill s.n. (Lost).

Microsorum scolopendria (Burm.f.) Copel. in

University of California Publications in Botany 16: 112

(1929); Jacobsen: 314 (1983); Schelpe: 165 (1986); Bur-

rows: 200, t. 46, fig. 204 (1990), as 'Microsorium

scolopendrium' . Polypodium scolopendrium Burm.f: 232

(1768). Phymatodes scolopendria (Burm.f) Ching; 63

(1933). Phymatosorus scolopendria (Burm.f.) Pie.Serm.;

460 (1973). Type: India, Herb. Bumiann (G, holo.).

Polypodium phymatodes L.: 306, 307 (1771), nom.

illeg.; Sim: 273, t. 138 (1915). Type: Habitat in India ori-

entali, sine coll. s.n. (LINN 1251.6, holo.).

ACKNOWLEDGEMENTS

Ms M. Koekemoer is thanked for obtaining literature

whilst Botanical Liasion Officer at Kew.

REFERENCES

ANTHONY, N.C. & SCHELPE, E.A.C.L.E. 1985. X Pleopodiwn—a

putative intergeneric fern hybrid from Africa. Bothalia 15:

555-559.

BEDDOME, R.H. 1876. Supplement to the ferns of southern India and

British India. Madras.

BOSMAN, M.T.M. 1991. A monograph of the fern genus Microsorum

(Polypodiacette). Leiden Botanical Series 14: 1-161.

BURMAN, N.L. 1768. Flora indica. prodromus. Cornelius Haak, Leiden.

BURROWS, J.E. 1990. Southern African ferns and fern allies.

Frandsen Publishers, Sandton.

CHING, R.C. 1933. The studies of Chinese ferns. IX. Bulletin of the

Fan Memorial Institute of Biology, Botany 4: 47-106.

COPELAND, E.B. 1929. The oriental genera of Polypodiaceae.

University of California Publications in Botany 16: 44—128.

COPELAND, E.B. 1947. Genera fdicum. Waltham, Massachusetts.

DE LA SOTA, E.R. 1973. On the classification and phylogeny of the

Polypodiaceae. In A.C. Jermy, J.A. Crabbe & B.A. Thomas,

The phylogeny and classification of the ferns. Botanical Journal

of the Linneun Society 67, Suppl. 1: 229-244. Academic Press,

London.

DE VOL, C.E. & KUO, C. 1980. Flora ofTaiwan. Vol. 1, edn 2. Taiwan.

FEE, A. L. A. 1852. Memoires sur les families des fougeres. Genera fili-

cum, Polypodiacees. Berger-Levrault, Strasbourg.

GOMEZ, L.D. 1975. Contribuciones a la pteridologi'a costarricense.

VIII. La hibridacion en el tropica: Microgramma x Polypodium

y P. aspidiolepis Baker. Brenesia 6: 49-57.

HAUFLER, C.H. & RANKER, T.A. 1995. RbcL sequence provide

phylogenetic insight among sister species of the genus

Polypodium. American Fern Journal 85: 361-374.

HENNIPMAN, E. & ROOS, M.C. 1983. Phylogenetic systematics of

the Polypodiaceae (Filicales). Verlmndlungen des Naturwissen-

schaftlichen Vereins in Hamburg 26: 321-342.

HENNIPMAN, E., VELDHOEN, P. & KRAMER, K.U. 1990, Polypo-

diaceae. In K. Kubitzki, The families and genera of va.uular

plants: 203-230, Springer- Verlag, Berlin.

HETTERSCHEID, W.L.A. & HENNIPMAN, E. 1984. Venation pat-

terns, soral characteristics, and shape of the fronds of the micro-

sorioid Polypodiaceae. Botanische Jahrbiicher fiir Systematik

105: 11^7.

HOLTTUM, R.E. 1954. A revised Flora of Malaya. Vol. 2. Ferns of

Malaya. Government Printing Office, Singapore.

JACOBSEN, W.B.G. 1983. The ferns and fern allies of .southern

Africa. Butterworths, Durban, Pretoria.

KUHN, F.A.M. 1868. Filices africanae. Leipzig.

LINNAEUS, C. 1763. Species plantarum 2, edn 2. Laurentius Salvius,

Stockholm,

LINNAEUS, C. 1771. Mantis.sa plantarum altera. Laurentius Salvius,

Stockholm.

MICKEL, J.T. & BEITEL, J.M. 1987. Notes on X Pleopodium and

Pleopeltis in tropical America. American Fern Journal 11:

16-27.

MITSUTA, S. 1984. Studies in the venation and systematics of

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J.P. ROUX*

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

7735 Claremont, Cape Town.

MS. received: 1997-12-22.

ASTERACEAE

A NOTE ON THE GENUS PHILYROPHYLLUM

Anderberg (1991) in his generic revision of the tribe

Gnaphalieae (Asteraceae), described the capitula of the

genus Philyrophyllum O.Hoffm. as epaleate. However,

Hoffmann (1890) and Merxmiiller (1967) described

them as being paleate. Material from herbarium speci-

mens housed in PRE was dissected to investigate this

matter and it became clear that the capitula are in fact

paleate. Anderberg agreed on these results (pers. comm.).

The key to the genera presented by Anderberg (1991) is

still a problem, as the genus Philyrophyllum is keyed out

under the group containing epaleate capitula. The key

and the description of the genus will be corrected in the

Seed plants of southern Africa (in prep.).

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Gnaphalieae (Asteraceae). Opera Botanica 104: 43.

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lien 4,5: 206, 208.

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von Sudwestafrika 139: 146-147.

P.P.J. HERMAN*

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

MS. received: 1998-08-13.

RUBIACEAE

A NEW SPECIES OF PAVETTA FROM THE SOUTPANSBERG, SOUTH AFRICA

Pavetta tshikondeni N.Hahn, sp. nov., P. cato-

phyllae K.Schum. affinis sed lobis calycis ensiformibus

non triangularibus et floribus fructibusque in cymis lax-

ioribus dispositis differt; fortasse P. gracillimae S. Moore

affinis, species sylvarum oriente meridieque regionis

Zimbabwe dictae habitu tenniore lobisque calycis

anguste triangularibus non ensiformibus.

TYPE. — Northern Province, 2231 (Messina): Makhuya

Park, World’s View, 22° 30' 24.1" S and 31° 01' 59.6" E

(WGS 84 mapping datum), (-CA), 300 m, 27-01-1997, (in

flower), N. Hahn 1367 (K, holo.; PRE, Herb. Sout., iso.).

Multistemmed shrub up to 2 m tall. Bark pale to dark

grey, smooth. Branches when young, covered with a

white indumentum, becoming glabrous with age. Leaves

opposite, obovate to spatulate, up to 78 x 22 mm, apex

obtuse to rounded, base attenuate to obtuse, upper sur-

face pale green and sparsely hairy, lower surface grey-

green and hairy, bacterial nodules randomly scattered on

leaf lamina, domatia absent, principal lateral veins 5-8

pairs; petiole up to 2 mm long (Figure 9). Inflorescences

of terminal cymes, borne on slender lateral branches up

to 700 mm long; peduncle hairy; pedicels 0-3 mm long,

hairy (Figure 10). Flowers white, 4-merous. Calyx lobes

ensiform, fused at base, up to 9.0 x 0.5 mm, hairy.

Corolla sparsely hairy on both surfaces; tube up to 15

mm long; lobes 6x2 mm, apices obtuse to acute.

Anthers exserted, 4—5 mm long. Style 25-30 mm long.

Disc glabrous. Hypanthium (part fused with inferior

ovary) up to 1 .5 mm long. Fruit a fleshy berry, spherical,

up to 5 mm diam., turning black when ripe; calyx lobes

persistent, forming a crown at apex.

Distribution and habitat

Pavetta tshikondeni is associated with Androstachys

woodland on soils derived from Karoo Supergroup

108

Bothalia29,l (1999)

FIGURE 9. — Leaves of Pavetta tshikondeni. A, upper surface, x 0.6;

B, lower surface, x 0,6.

rocks. This particular type of habitat is of restricted

occurrence in the Soutpansberg region (Figure 11) and

contains many rare and endemic plant taxa. Andro-

stachys woodland consists of almost homogeneous

stands of the tree Androstachys johnsonii (Euphorbi-

aceae) and covers some of the sandstone ridges in the

eastern Soutpansberg area of the Northern Province.

Many plants endemic to the Soutpansberg are associated

FIGURE 10. — Pavetta tshikondeni. Inflorescence, x 2, with flower

buds and a few open flowers; note the long, ensiform, calyx

lobes.

FIGURE 11. — Distribution of Pavetta tshikondeni in South Africa.

with this vegetation type, at least three of which are

restricted to the northeastern sector of the mountain

range, namely Ceratotheca saxicola, Duvalia procum-

bens and Euphorbia rowlandii.

Within the boundaries of the Tshikondeni Mining lease

the new species has been found on Fripp Sandstone

Formation along the escarpment west of the Luvuvhu River,

as well as on Clarens Sandstone Formation ridges north of

the Mutale River. A specimen in the National Herbarium,

Pretoria, was collected in the extreme north of the Kruger

National Park, apparently from a similar habitat. Pavetta

tshikondeni is one of at least 36 plant species endemic to the

Soutpansberg range (including the Blouberg).

Taxonomic affinities

Pavetta tshikondeni is easily distinguished from all the

South African members of Pavetta. Its closest ally among

local members of the genus appears to be Pavetta cato-

phylla. However, it differs from Pavetta catophylla in its

calyx lobes being ensiform, forming a distinct fused tube

surrounding the fruit. Its flowers and fruit are also not

borne in as densely packed axillary cymes as those of P

catophylla. It may also be related to P. gracillima, a

species from eastern and southern Zimbabwe. P. gracilli-

ma is less robust in habit and its calyx lobes are narrowly

triangular in outline. Selected morphological characters

that distinguish P. gracillima from P. tshikondeni are

summarised in Table 2. The following key should aid in

TABLE 2. — Summary of characters distinguishing P. gracillima from

P. tshikondeni. Measurements in mm

Bothalia29,l (1999)

109

the identification of all known species of Pavetta occur-

ring within the Soutpansberg and its surroundings.

Key to Pavetta species in the Soutpansberg

la Calyx lobes less than 1 mm long:

2a Leaves large, up to 160 x 90 mm, circular in outline

Pavetta eylesii S. Moore

2b Leaves not circular in outline:

3a Leaves spatulate:

4a Leaves glabrous

Pavetta gardeniifolia A. Rich. var. gardeniifolia

4b Leaves hairy

Pavetta gardeniifolia A. Rich. var. subtomentosa K.Schum.

3b Leaves elliptic to lanceolate:

5a Petiole 5-15 mm long; forest trees

Pavetta inandensis Bremek.

5b Petiole 2-6 mm long; occurring in savanna or on forest

margins Pavetta lanceolata Eckl.

lb Calyx lobes more than 1 mm long;

6a Calyx lobes acute:

7a Leaves sessile Pavetta harborii S. Moore

7b Leaves stalked . . . Pavetta schumanniana F.Hoffm. ex K.Schum.

6b Calyx lobes ensiform:

8a Leaf lamina obovate to spatulate; calyx lobes up to 9 mm

long; Androstachys woodland

Pavetta tshikondeni N.Hahn

8b Leaf lamina ovate; calyx lobes up to 5 mm long; occur-

ring in high mountain mistbelt vegetation

Pavetta trichardtensis Bremek. sensu lato

Etymology

The new species was discovered on 4-05-1996 in the

course of a vegetation survey for the Tshikondeni Mining

Company. The specific epithet is derived from the name

of this company, in recognition of their responsible min-

ing practises and conservation efforts in the region.