Bothalia
i
A JOURNAL OF BOTANICAL RESEARCH
Vol. 32,2 Oct. 2002
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
devoted to the furtherance of botanical science. The main fields covered are taxonomy, ecology,
anatomy and cytology. Two parts of the journal and an index to contents, authors and subjects are
published annually.
Three booklets of the contents (a) to Vols 1-20, (b) to Vols 21-25 and (c) to Vols 26-30, are available.
STRELITZIA
A series of occasional publications on southern African flora and vegetation, replacing Memoirs of
the Botanical Surx’ey of South Africa and Annals of Kirstenbosch Botanic Gardens.
MEMOIRS OE THE BOTANICAL SURVEY OF SOUTH AFRICA
The memoirs are individual treatises usually of an ecological nature, but sometimes dealing with
taxonomy or economic botany. Published: Nos 1-63 (many out of print). Discontinued after No. 63.
ANNALS OF KIRSTENBOSCH BOTANIC GARDENS
A series devoted to the publication of monographs and major works on southern African flora.
Published: Vols 14-19 (earlier volumes published as Supplementary volumes to the Journal of
South African Botany). Discontinued after Vol. 19.
FLOWERING PLANTS OF AFRICA (FPA)
This serial presents colour plates of African plants with accompanying text. The plates are prepared
mainly by the artists at the National Botanical Institute. Many botanical artists have contributed to
the series, such as Fay Anderson, Peter Bally, Auriol Batten, Gillian Condy, Betty Connell, Stella
Gower, Rosemary Holcroft, Kathleen Lansdell, Cythna Letty (over 700 plates), Claire Linder-
Smith and Ellaphie Ward-Hilhorst. The Editor is pleased to receive living plants of general interest
or of economic value for illustration.
Erom Vol. 55, twenty plates are published at irregular intervals.
An index to Vols 1—49 is available.
FLORA OF SOUTHERN AFRICA (ESA)
A taxonomic treatise on the flora of the Republic of South Africa, Lesotho, Swaziland, Namibia
and Botswana. The FSA contains descriptions of families, genera, species, infraspecific taxa, keys
to genera and species, synonymy, literature and limited specimen citations, as well as taxonomic
and ecological notes.
Contributions to the FSA also appear in Bothalia.
PALAEOFLORA OF SOUTHERN AFRICA
A palaeoflora on a pattern comparable to that of the Flora of southern Africa. Much of the informa-
tion is presented in the form of tables and photographic plates depicting fossil populations. Now
available:
Molteno Formation (Triassic) Vol. 1. Introduction. Dicroidium, 1983, by J.M. & H.M.
Anderson.
Molteno Formation (Triassic) Vol. 2. Gymno.sperms (excluding Dicroidium), 1989, by J.M.
& H.M. Anderson.
Prodromus of South African Megalloras. Devonian to Lower Cretaceous, 1985, by J.M. &
H.M. Anderson. Obtainable from: A.A. Balkema Marketing, Box 317, Claremont 7735,
RSA.
Towards Gondwana Alive. Promoting biodiversity and stemming the Sixth Extinction, 1999,
by J.M. Anderson (ed.)
BOTHALIA
A JOURNAL OF BOTANICAL RESEARCH
Volume 32,2
Scientific Editor: G. Germishuizen
Technical Editor: B.A. Momberg
NATIONAL
.Botanical
INSTITUTE
2 Cussonia Avenue, Briimmeria, Pretoria
Private Bag XIOl, Pretoria 0001
ISSN 0006 8241
Oct. 2002
Editorial Board
D.F. Cutler
B.J. Huntley
P.H. Raven
J.P. Rourke
M.J.A. 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, The Netherlands
Acknowledgements to referees
Archer, Dr R. National Botanical Institute, Pretoria, RSA.
Archer, Mrs C. National Botanical Institute, Pretoria, RSA.
Barker, Dr N.P Rhodes University, Grahamstown, RSA.
Bouman, Dr F. Hugo de Vries Laboratory, Amsterdam, The Netherlands.
Braggins, Dr J.E. University of Auckland, New Zealand.
Bremer, Prof. K. University of Uppsala, Sweden.
Brummitt, Dr R.K. Royal Botanical Gardens, Kew, UK.
Burgoyne, Ms P. National Botanical Institute, Pretoria, RSA.
Ellis, Dr R.P Grassland Research Centre, Agricultural Research Council, Pretoria, RSA.
Goldblatt, Dr P. Missouri Botanical Garden, St Louis, USA.
Hanna, Dr W. Coastal Plain Experimental Station, Tifton, USA.
Herman, P.P.J. National Botanical Institute, Pretoria, RSA.
Ihlenfeldt, Prof. H.D. Ostseestr. 40, D-24369 Waabs, Germany.
Kallersjb, Dr M. Swedish Museum of Natural History, Stockholm, Sweden.
Killick, Dr D.J.B. 465 Sappers Contour, Lynnwood, 0081 Pretoria, RSA.
Leistner, Dr O.A. National Botanical Institute, Pretoria, RSA.
Marais, Dr E.M. University of Stellenbosch, RSA.
Miller, Dr J.S. Missouri Botanical Garden, St Louis, USA.
Mucina, Prof. L. University of the North, Qwa-Qwa Campus, Phuthaditjhaba, RSA.
Nelson, Dr C. Outwell, Wisbech, UK.
Oliver, Dr E.G.H. National Botanical Institute, Cape Town, RSA.
Pichi-Sermolli, Dr R.E.G. Montagnana Val di Pesa, Elorence, Italy.
Poes, Prof. T. Eszterhazy College, Botany Department, Eger, Hungary.
Robbrecht, Prof. Dr E. National Botanic Garden, Meise, Belgium.
Ronse De Craene, Dr L.P Katholieke Universiteit Leuven, Belgium.
Roux, Dr J.P National Botanical Institute, Cape Town, RSA.
Werger, Prof. M.J.A. University of Utrecht, The Netherlands.
CONTENTS
Volume 32,2
1. Systematics of the genus Daubenya (Hyacinthaceae: Massonieae). J.C. MANNING and A.M. VAN
DERMERWE 133
2. The genus Trichodesma {Boraginaceae: Boraginoideae) in southern Africa. E. RETIEE and A.E. VAN
WYK 151
3. Six new species and one new subspecies of Erica (Ericaceae) from Western Cape, South Africa. E.G.H.
OLIVER and I.M. OLIVER 167
4. Studies in the liverwort family Aneuraceae (Metzgeriales) from southern Africa. 4. Riccardia ohtusa.
S.M. PEROLD 181
5. Notes on African plants:
Amaryllidaceae. The typification of Cyrtaiidius smithiae Watt ex Harv. J.P. ROURKE 197
Asteraceae. A distinctive new species of Felicia (Astereae) from Western Cape, South Africa.
J.C. MANNING and P. GOLDBLATT 193
Asteraceae. A new species of Arctotlieca from Northern Cape, South Africa. J.B.P. BEYERS. . 185
Asteraceae-Anthemideae. Reduction of Foveolina alhida to Foveoliua diciwtoma. J.B.P.
BEYERS 185
Hyacinthaceae-Massonieae. A new species of Fachenalia from Namaqualand, South Africa.
G. D. DUNCAN & T.J. EDWARDS 190
Mesembryanthemaceae. Mesembs with nut-like schizocarpic fruit and Ruschianthemiini Fried-
rich sunk under Stoeberia Dinter & Schwantes. P. CHESSELET and A.E. VAN WYK . . 187
Portulacaceae. The synonymy of Ceraria namaqiiensis (Sond.) H. Pearson & E.L.Stephens.
H. F. GLEN 196
Pteridophyta. A new fern record for the Flora of southern Africa region. J.E. BURROWS and
S.M.BURROWS 195
6. A study of ovule-to-seed development in Ceratiosicyos (Achariaceae) and the systematic position of
the genus. E.M.A. STEYN, A.E. VAN WYK and G.F. SMITH 201
7. The grasslands and wetlands of the Sekhukhuneland Centre of Plant Endemism, South Africa. S.J.
SIEBERT, A.E. VAN WYK. G.J. BREDENKAMP and F. DU PLESSIS 211
8. Miscellaneous notes:
Poaceae. Chromosome studies on African plants. 17. The subfamilies Arundinoideae and
Danthonioideae. R. ROODT, J.J. SPIES, A.E. MALAN, F. HOLDER and S.M.C. VAN
WYK 233
Poaceae. Chromosome studies on African plants. 18. The subfamily Chloridoideae. R. ROODT
and J.J. SPIES 240
9. Book reviews 251
10. National Botanical Institute South Africa: administration and research staff, 31 March 2002, publica-
tions 1 April 2001-31 March 2002. Compiler: B.A. Momberg 253
11. Guide for authors to 269
New species, subspecies and combinations in Bothalia 32,2 (2002)
Arctotheca marginata Beyers, sp. nov., 185
Daubenya comata (Burch, ex Baker) J.C.Mauuiiig & A.M.vau der Merwe, comb, nov., 139
Daubenya zeyheri (Kiinthj J.C. Manning & A.M.van der Menve, comb, nov., 143
Erica amalophylla E.G.H.Oliv. & I.M.Otiv., sp. nov., 168
Erica annalis E.G.H.Oliv. & I.M.Oliv., sp. nov., 178
Erica cavartica E.G.H.Oliv. & I.M.Oliv., sp. nov., 167
Erica cymosa E.Mey. ex Bendi. siibsp. grandiflora E.G.H.Oliv. & I.M.Oliv., subsp. nov., 171
Erica lithophila E.G.H.Oliv. & I.M.Oliv., sp. nov., 174
Erica schelpeorum E.G.H.Oliv. & I.M.Oliv., sp. nov., 172
Erica umbratica E.G.H.Oliv. & I.M.Oliv., sp. nov., 176
Felicia josephinae & Goldblatt, sp. nov., 193
Lachenalia valeriae G.D. Duncan, sp. nov., 190
Trichodesma angustifolium Harv. subsp. argenteum Retief & A.E.van Wyk, subsp. nov., 161
iv
Bothalia 32,2: 133-150 (2002)
Systematics of the genus Daubenya (Hyacinthaceae: Massonieae)
J.C. MANNING* ** and A.M. VAN DER MERWE*’
Keywords: Ainphisiphon W.F.Barker, Androsiplion Schltr., Daubenya Lindl., Hyacinthaceae, Massonieae Baker, Neobakeria Schltr., South Africa, taxonomy
ABSTRACT
Daubenya Lindl. was until recently thought to comprise the single species D. aurea Lindl. but is now considered to
include the monotypic genera Androsiphon Schltr. and Amphisiphon W.F.Barker, as well as the species previously referred
to the genus Neobakeria Schltr. Eight species are now recognized in the genus, including the new combinations Daubenya
comata (Burch, ex Baker) J.C. Manning & A.M. van der Merwe and D. zeyheri (Kunth) J.C. Manning & A.M. van der Merwe.
Each species is fully described and illustrated in black-and-white and in colour. A key to the species, and distribution maps
are provided.
CONTENTS
Abstract 133
Introduction 133
Materials and methods 135
Morphological characters of taxonomic significance 135
Distribution and ecology 136
Pollination and seed dispersal 136
Evolution 137
Taxonomic history 138
Systematic treatment 138
Daubenya Lindl 138
Key to species 139
1. D. comata (Burch, ex Baker) J.C. Manning &
A.M. van der Merwe 139
2. D. namaquensis (Schltr.) J.C.Manning & Gold-
blatt 140
3. D. marginata (Willd. ex Kunth) J.C.Manning &
A.M. van der Merwe 142
4. D. zeyheri (Kunth) J.C.Manning & A.M. van der
Merwe 143
5. D. alba A.M. van der Merwe 144
6. D. capensis (Schltr.) A.M. van der Merwe &
J.C.Manning 145
7. D. stylosa (Barker) A.M. van der Merwe &
J.C.Manning 146
8. D. aurea Lindl 148
Specimens examined 149
Acknowledgements 149
References 150
INTRODUCTION
The Hyacinthaceae (excluding the North American
chlorogaloid genera) are now well established as a
monophyletic lineage within the order Asparagales (Eay
& Chase 1996; Pfosser & Speta 1999; Fay et al. 2000).
The family is distributed through Africa, across most of
Europe and central Asia to India, and in Andean South
America, with centres of diversity in southern Africa and
the Mediterranean (Speta 1998). The plants prefer open.
* Compton Herbarium. National Botanical Institute, Private Bag XV,
7735 Claremont, Cape Town.
** Department of Botany, University of Stellenbosch, Private Bax XI,
7602 Matieland, South Africa.
MS. received: 2002-01-09.
sunny habitats in seasonal climates and are correspond-
ingly rare in heavily wooded, tropical regions. Although
it has long proved difficult to devise a natural infrafamil-
ial classification of the Hyacinthaceae, recent molecular
analysis of the chloroplast DNA region trnL-F provides
strong support for the division of the family into four
subfamilies, Oziroeoideae, Urgineoideae, Omithogaloi-
deae and Hyacinthoideae (Pfosser & Speta 1999). Within
the subfamily Hyacinthoideae two clades are retrieved.
The first combines the Indian and African genera south
of the Sahara, and largely coincides with the delimitation
of the tribe Massonieae Baker (with the significant inclu-
sion of the sub-Saharan species previously placed in the
genus Scilla L.). The second clade includes the Medi-
terranean and Asian genera, corresponding to the tribe
Hyacintheae Dumort (Pfosser & Speta 1999). Despite
the strong molecular support for the recognition of these
two tribes, few corroborative morphological characters
are available. Further division of tribe Massonieae into
the subtribes Ledebouriinae and Massoniinae (Miiller-
Doblies & Miiller-Doblies 1997) is not supported by the
molecular data. The poor congruence between morpho-
logical and other characters within Hyacinthaceae has
also made generic circumscriptions very difficult. One of
the consequences of this has been the recognition of a
large number of genera that are poorly defined morpho-
logically (Speta 1998).
Within the tribe Massonieae the relationships between
the species traditionally placed in the genera Amphisiphon
W.F.Barker (1 sp.), Androsiphon Schltr. (1 sp.), Daubenya
Lindl. ( 1 sp.), Neobakeria Schltr. (1-3 spp.) and Massonia
Houtt. (± 6 spp.) have been especially controversial
(Jessop 1976; Miiller-Doblies & Miiller-Doblies 1997;
Goldblatt & Manning 2000). These genera share some
morphological characters, in particular two prostrate or
spreading leaves and a condensed inflorescence of more or
less tubular flowers but differ greatly from one another in
floral details. Traditional morphological methods have not
been useful in resolving the relationships between the
species but molecular analysis has proven extremely in-
formative. Analysis of chloroplast DNA (Van der Merwe
et al. in prep.) includes Amphisiphon, Androsiphon, Dau-
benya, Neobakeria and one species of Massonia in a
strongly supported monophyletic clade. This clade is resis-
tant to further segregation at the generic level and the cir-
134
Bothalia 32,2 (2002)
PLATE 1. — A. Daitbenya comata: B, D. namaquensis', C, D. margwata: D, D. zeyheri\ E, D. alba\ E, D. capensis\ G, D. stylosa', H, D. aurea. Photographer: John Manning.
Bothalia 32,2 (2002)
135
cumscription of the genus Daubenya is therefore broad-
ened to circumscribe the whole clade. The genus
Daubenya in this broader sense is characterized by a tuni-
cated bulb with the older tunics leathery and dark brown
and extending as a short papery neck, two ± glossy leaves
with impressed longitudinal striations on the upper sur-
face, and a racemose, corymbose or subspicate inflores-
cence of white, red or yellow flowers with a slender, cylin-
drical perianth tube. Species of Massonia s. str. are char-
acterized by their soft, fleshy leaves, corymbose inflores-
cences with large, leaflike bracts, white to pink flowers
with the tepals typically sharply recurved from the base
and then incurved in a characteristic sigmoid fold, and
tumbleweed-like infructescences.
Daubenya is centred along the western edge of the
southern African central plateau and all of the species are
winter- growing. Most are highly localized endemics
restricted to pockets of doleritic clays. The vegetative
similarity between the species contrasts sharply with the
unusual diversity of floral form evident in the genus.
This floral radiation represents adaptations to a variety of
pollinating agents, including insects such as bees, butter-
flies, moths and monkey beetles and sunbirds. Most of
the species exhibit adaptations to anemochory, especial-
ly in the development of large, winged capsules. These
winged, dehiscent capsules contrast markedly with the
indehiscent, ovoid capsules present in two of the species.
Eight species are recognized within this expanded con-
cept of Daubenya, some of them well established in hor-
ticulture but others very poorly known. This revision
provides full descriptions of all species as well as notes
on their history, ecology and distribution. All of the
species were studied in the field and each is fully illus-
trated in black-and-white and in colour.
MATERIALS AND METHODS
Herbarium specimens in BOL, K. NBG, PRE and
SAM were studied to gather data on morphology, flow-
ering time and distribution. Extensive fieldwork was also
undertaken and all species were visited in their natural
habitat. All observations on pollinators were made in the
field. Conservation status for each species was assessed
according to the new lUCN categories and criteria
(Victor 2000).
MORPHOLOGICAL CHARACTERS OE TAXONOMIC
SIGNIFICANCE
Bulb: the bulbs in Daubenya are turbinate to globose
or ovoid in shape and between 20-35 mm diam. The
outer tunics become leathery and dark brown on drying,
and split at the top into narrow, flat, papery segments that
form a characteristic neck. This papery neck is particu-
larly conspicuous in species or individuals with deeply
buried bulbs. A similar neck is found in some species of
Lachenalia but not in any species of Massonia 5. str.
Leaves: all species have paired leaves that are pros-
trate or spreading. The leaf bases are subterranean and
long or short depending on the depth of the bulb. The
blades are glabrous and rather glossy above with
impressed longitudinal striations along the main veins.
They are lanceolate to ovate in shape and measure
40-150x25-50 mm.
Leaf anatomy: leaves in Daubenya are amphistomatic
with a thick cuticle. The marginal epidermal cells are
columnar with slightly thickened walls and a unicellular
hypodermis is present at the leaf margin. The mesophyll
typically comprises three rows of palisade parenchyma
adaxially and three rows of spongy parenchyma abaxial-
ly. The vascular bundles are surrounded by parenchyma-
tous bundle sheaths. Slight indentations correlating with
the impressed longitudinal striations on the upper leaf
surface occur above the main veins. In these indentations
the epidermal cells are smaller than elsewhere with
thicker outer periclinal walls and they overlie two rows
of palisade parenchyma instead of three.
Inflorescence: is borne at ground level between the
leaves on a subterranean peduncle. It is essentially race-
mose but varies in shape from conical to capitate.
Shortening of the rachis in several species results in a
corymbose, rather capitate inflorescence, whereas sup-
pression of the pedicels results in a subspicate inflores-
cence, accompanied in some species by shortening of the
axis. Well-developed pedicels and a corymbose inflores-
cence are found in Daubenya aurea, D. alba, D. capen-
sis and D. zeyheri but the remaining species are charac-
terized by a subspicate inflorescence. This is usually con-
ical, protruding for up to 90 mm above the leaves in D.
marginata and D. namaquensis, but is ± capitate in D.
comata and D. stylosa. The inflorescence bracts are usu-
ally inconspicuous and pale in colour except in D. aurea,
which is characterized by large, green bracts. The bracts
typically increase in size acropetally but the degree of
this increase varies. In D. comata, D. stylosa and D. zey-
heri the upper bracts are only slightly larger than the
lower ones and form an inconspicuous greenish coma at
the top of the inflorescence. In D. namaquensis and D.
marginata, however, the increase in bract size is marked
and the inflorescences are topped by a more conspicuous
coma of greenish or pinkish to orange bracts. The inflo-
rescence remains at ground level until the fruits are
mature, at which stage the peduncle elongates rapidly
from the base, pushing the infructescence well above the
ground as a prelude to dispersal of the seeds.
Flowers: there is great variation in the morphology of
the flowers in Daubenya but all species are characterized
by a slender, cylindrical perianth tube. The tepals are
usually free and spreading from the level of the stamen
insertion but in D. stylosa they are fused well beyond this
level into a narrow tube around the staminal column.
Tepal colour ranges from white or pinkish in D. alba, D.
comata, D. marginata and D. zeyheri, to bright yellow in
D. capensis, D. namaquensis and D. stylosa, or brilliant
red in most populations of D. aurea. The flowers in most
species are actinomorphic but some degree of zygomor-
phy is evident in the lowermost flowers in D. comata and
D. namaquensis, and reaches an extreme degree in D.
aurea. In D. comata and D. namaquensis the outer flow-
ers are at most weakly bilabiate through a slight asym-
metry in the perianth tube, with the adaxial side slightly
longer than the abaxial side. An analagous situation is
also sometimes evident in the filament column in the
lowermost flowers of D. alba. In D. aurea, however, the
136
Bothalia 32,2 (2002)
zygomorphy is extremely marked and results from the
great enlargement of the lowermost tepals of the lower
flowers, accompanied by an elongation of the perianth
tube on this side rather than on the adaxial side. The sta-
mens in most species of Daubenya are ± fused at the base
into a staminal column or tube. This is most marked in D.
alba, D. aurea, D. capensis and D. stylosa. In the bilabi-
ate, lower flowers of D. aurea, the abaxial tepals are
actually fused to the staminal column and as a conse-
quence of this the abaxial filaments are inserted much
higher up the perianth than the adaxial and appear to be
free from one another. In D. capensis a thick, convex disc
occludes the top of the staminal tube with the style pro-
truding through a small pore in the disc. The staminal
column is very short in D. marginata and D. zeyheri, and
is lacking or only slightly evident in D. comata and D.
namaquensis .
Fruit and seeds: the fruit in Daubenya is a papery
capsule, typically dehiscing loculicidally in the upper
portion. In D. comata the capsule is obovoid and round-
ed but in most of the other species it is + inflated and
three-angled. These angles are particularly well devel-
oped in the upper part of the capsules in D. alba and D.
capensis, and the capsules in these two species are decid-
edly cuneate in shape. The species D. marginata, D.
namaquensis and D. zeyheri are distinctive in their deci-
duous capsules that fall free of the pedicels at maturity
and are dispersed individually. In other species of
Daubenya the capsules remain attached to the pedicel
and rachis and the entire infructescence functions as the
dispersal unit. Unique capsules with a persistent, horn-
like style characterize D. stylosa. The capsules in this
species are essentially indehiscent although the locules
eventually separate along the septa. The seeds in all
species are globose and black in colour with a smooth
testa. They range in size from 2-3 mm diam.
DISTRIBUTION AND ECOLOGY
Daubenya is a small genus of eight species that is
endemic to South Africa. It is largely restricted to the
fringes of the winter rainfall region along the Bokkeveld
and Roggeveld Escarpments, which mark the western
margin of the South African central plateau. Five of the
eight species occur here, at altitudes of 1 000-1 500 m.
Most of the species of Daubenya are highly local
endemics and only D. comata is more widespread, occur-
ring over much of the South African central plateau. All
of the species are winter-growing, despite the fact that
not all are restricted to the winter rainfall region. Plants
typically leaf and flower in the autumn or winter,
between April and July, with only one species, D. aurea,
flowering in the spring, in early September. Five of the
species, D. alba, D. aurea, D. capensis, D. stylosa and D.
zeyheri are restricted to the winter rainfall region and
respond largely to frontal rain that falls between April
and August. Apart from D. zeyheri, which is a coastal
species confined to calcareous sands near Saldanha, the
remaining four winter rainfall species occur along the
edge of the Bokkeveld and Roggeveld E.scarpments from
Nieuwoudtville and Calvinia in the north southwards to
Sutherland. The remaining three species occur partially
or wholly within the summer rainfall region. Daubenya
marginata is distributed along the Roggeveld Escarp-
ment but extends inland to Fraserburg, where it enters the
summer rainfall region; D. namaquensis occurs at the
edge of the summer rainfall region in Bushmanland, east
of Springbok; and D. comata is widespread across the
central parts of the summer rainfall region. Despite their
distribution, these three species respond to autumn thun-
dershowers and begin to grow slightly before the winter
rainfall species.
Most species of Daubenya are highly localized
endemics known only from a few populations. They typ-
ically occur in small colonies, sometimes in large num-
bers, in heavy clay soils, and are invariably restricted to
low-lying washes or drainage lines where the soil be-
comes seasonally waterlogged. The majority of the species
occur on the South African central plateau, where they
are largely restricted to clays derived from doled te. The
particular nature of doleritic clays ensures that these soils
retain moisture for longer than the surrounding clays
derived from shales of the Karoo series. The two remain-
ing species occur on more sandy substrates, D. namaquen-
sis in deep red sands and D. zeyheri in calcareous coastal
sands.
All of the species of Daubenya are vulnerable to dis-
turbance or transformation of their habitat, particularly
through agriculture. Although predation of the bulbs by
porcupines is common among the species growing on the
Roggeveld Escarpment, its impact on the populations is
unknown. The numbers of these rodents can be expected
to have increased as a result of the reduction in their
natural predators, particularly leopard. Most species of
Daubenya still appear to set large quantities of seed in
the wild to replace the adult plants consumed by herbi-
vores. An additional pressure on populations comes from
sheep and baboon, which eat the leaves and inflores-
cences, resulting in a drastic reduction in seed set in
some instances. A recent initiative aims at the protection
of a population of the yellow-flowered form of D. aurea
from overgrazing through co-operation of the landowner
and the local bulb growers’ association. There is, how-
ever, no formal conservation of this population and its
survival thus remains uncertain. In another encouraging
development, cultivated bulbs of D. aurea have become
available in commercial nurseries. All of the species of
Daubenya are worth cultivating for their compact habit
and brilliantly coloured, often fragrant flowers.
POLLINATION AND SEED DISPERSAL
The floral diversity evident in the genus Daubenya is
exceptional in the family Hyacinthaceae and reflects a
coiTesponding diversity in pollination strategies. These
include several strategies that, although well developed
in the Western Cape, are rare or absent elsewhere. Most
of the species produce fragrant, white or yellow flowers
that are visited for nectar or pollen by a variety of diur-
nal and nocturnal insects, including pollen- and nectar-
collecting bees, butterflies and moths in the family
Noctuidae. The species with generalist flowers include
D. namaquensis, D. comata, D. alba and D. stylosa. In
these species the nectar collects either in the perianth
tube only or also in the staminal tube above this. Both the
Bothalia 32.2 (2002)
137
perianth and staminal tubes are narrow and the nectar is
thus accessible only to long-tongued insects.
Two species, D. marginata and D. zeyheri, are adapt-
ed to pollination by sunbirds. The adaptations to
omithophily in these species include a reduced and
widened staminal tube that forms a shallow reservoir in
which significant quantities of nectar accumulates,
orange or red filaments (with purple bases in D. zeyheri)
and a lack of fragrance. Omithophily occurs in several
other genera of Hyacinthaceae in the Western Cape,
especially Lxichenalia and Veltheimia. Daubenya aurea
is adapted to pollination by monkey beetles (Scarabidae-
Rutellinae), a pollination strategy that appears to be
unique among the Hyacinthaceae, although well repre-
sented among Western Cape Iridaceae. These active,
hairy beetles utilize various brightly coloured, open or
bowl-shaped flowers as sites for reproduction and are
often encountered on various species of Asteraceae and
Iridaceae (Goldblatt et al. 1998). Flowers adapted to
these beetles are typically ornamented with dark mark-
ings that act as beetle mimics, decoying the insects
through their resemblance to potential mates. Adapta-
tions for this type of beetle pollination in D. aurea are the
asymmetric enlargement of the lower or peripheral flow-
ers in the inflorescence to imitate the ray florets of
Asteraceae like Arctotis and Gazania, the lack of nectar,
with associated vestigial staminal tube, and the absence
of floral fragrance. Although the pollination biology of
the remaining species, D. capensis, has not been studied,
the peculiar stmcture of the flowers and their yeasty
odour suggest that the species might be adapted to polli-
nation by rodents. This strategy is also evident in one or
two species of Massonia (Hyacinthaceae) (Johnson et al.
2001) as well as several species of Androcymbium Willd.
(Colchicaceae) among the geophytes in Western and
Northern Cape.
In all species of Daubenya the fruiting peduncle elon-
gates at maturity, raising the infructescence above the
ground, and then abscises to release it. Thereafter, the
species differ markedly in seed dispersal strategies. In D.
comata and D. stylosa no particular adaptations for seed
dispersal appear to be developed and the relatively small
capsules remain attached to the axis, shedding the seeds
immediately around the plants. These autochorous
species have short or vestigial pedicels that do not elon-
gate at maturity and ± ovoid capsules that are indehis-
cent, although the locules do eventually separate along
the septa to release the seeds over time as they disinte-
grate. The remaining species exhibit a suite of adapta-
tions to anemogeochory or wind dispersal through tum-
bling (Van der Fiji 1982; Snijman & Linder 1996). The
most prominent of these adaptations are the large, three-
angled or -winged, loculicidally dehiscent capsule and
the frequent detachment of the entire inflorescence as a
single unit before seed release. In D. marginata, D.
namaquensis and D. zeyheri the capsules abscise readily
from the pedicels at maturity and are dispersed individu-
ally but in the remaining species they remain attached to
the peduncle and the infructescence is dispersed as an
entire unit. The surface area of the dispersal unit in these
species is further increased by the prominent wings on
the capsules in D. alba and D. capensis, giving them a
kite-like appearance and by the large papery bracts in D.
aurea, which act as sails. The conversion of the in-
fructescence into a tumbleweed is enhanced in D. alba
and D. capensis through the elongation of the pedicels,
increasing the size of the infructescence and giving it a
rounded, balloon-like form. There is a remarkable simi-
larity in the adaptations to wind dispersal of the fruits
developed in these latter species and in the genus
Massonia, particularly the obtriangular, winged capsules
borne on relatively long pedicels to form a rounded, bal-
loon-like structure that is readily dispersed by the wind.
In Massonia, however, the pedicels are invariably sub-
tended by large, sail-like bracts resembling those found
in D. aurea. The smooth, globose seeds in the anemo-
chorous species are easily shaken out of the capsules but
do not appear to disperse significantly thereafter.
Anemogeochory is well known in the family Ama-
ryllidaceae; subtribe Amaryllideae (Snijman & Linder
1996) but has not been studied in the Hyacinthaceae. In
both of these families, however, this mode of seed dis-
persal is more common in South Africa than elsewhere in
the continent, and is best developed in the semi-arid,
winter rainfall region.
EVOLUTION
Sequence analysis of the tniL-tniF region of the
chloroplast genome (Van der Merwe et al. in prep.)
offers little resolution within the genus apart from indi-
cating a close relationship between the species D. alba
and D. capensis. This species pair is well defined mor-
phologically by the cuneate, winged capsules borne on
long pedicels. Their relationship to the remaining species
as well as the relationships between these species remain
unresolved. Although the similarity in floral form and
capsules in D. marginata and D. zeyheri suggests that
these two species are closely allied, further morphologi-
cal assessment of relationships within the genus is con-
founded by the high degree of autapomorphies displayed
by each species. Until further genetic analysis is com-
plete, little more can be said about possible relationships
between the species.
The genus Daubenya is essentially restricted to sea-
sonally moist, usually clay depressions and all of the
species respond to autumn rains, whether they occur in
the winter or summer rainfall regions. There is thus little
ecological diversification within the genus in response to
climate or soils, which is probably not surprising given
that all species are vegetatively indistinguishable. Apart
from D. comata and D. marginata, species of Daubenya
are local endemics that are rarely sympatric. Exceptions
are the species pairs D. aurea and D. marginata in one or
two localities, and D. stylosa and D. capensis. It is per-
haps significant that the members of these pairs flower at
different times, suggesting that flowering time operates
as a prezygotic isolating mechanism between them.
While there may be little diversification within the genus
in response to climate or soil, Daubenya is unparalled in
the Hyacinthaceae in its floral radiation. This extreme
variation can be linked to differences in pollination strat-
egy and is accompanied by some variation in capsule
morphology and seed dispersal mechanisms. Most
species are anemochorous to some degree and adapta-
138
Bothalia 32,2 (2002)
tions to wind dispersal of the seeds include both elabora-
tion of the capsules themselves as well as enlargement of
the inflorescence bracts. Exceptions are D. stylosa and
possibly D. comata, which have indehiscent capsules
with no apparent adaptations for seed dispersal.
Although the anemochorous species are more diverse
than those lacking obvious seed dispersal mechanisms,
the individual species adapted to anemochory are not
more widespread than the autochorous species. This sug-
gests that anemochory in Daubenya has facilitated salta-
tional speciation, whereby the occasional colonization of
suitable new habitats through seed dispersal has
favoured speciation through founder-effects and subse-
quent genetic drift.
TAXONOMIC HISTORY
The taxonomic history of the genus Daubenya reflects
the significance that has traditionally been attached to
floral differences among the Hyacinthaceae, resulting in
the establishment of several monotypic genera between
1835 and 1936 for newly discovered species with dis-
tinctive flowers. The first of these genera, Daubenya,
was established in 1835 by Lindley to accommodate the
species D. aurea, which is characterized by extremely
zygomorphic lower flowers. Floral zygomorphy is
uncommon in the family Hyacinthaceae and this degree
of zygomorphy does not occur elsewhere in the family.
In his treatment of the sympetalous genera of the family
Hyacinthaceae, Baker (1871) allied Daubenya with the
genus Massonia in his tribe Massonieae on the basis of
the reduced peduncle. Bentham (1883), however, includ-
ed these two genera in his tribe Allieae because their con-
gested inflorescences in which the flowers are subtended
by large bracts suggested the umbel-like inflorescences
of Alliaceae. Species related to D. aurea but with less
well-developed bracts were placed by Baker (1871) first
in section Astenuna of Massonia and later in subgenus
Astemma of Polyxena (Baker 1897). The species recog-
nized by Baker (1871) in subgenus Astemma are those
that are currently recognized as D. comata, D. marginata
and D. zeyheri and he remarked at the time on the simi-
larity between them and certain species of Massonia,
with the suggestion that the two groups were best com-
bined. This opinion was followed in recent treatments of
the group (Jessop 1976; MUller-Doblies & Miiller-
Doblies 1997) but was not the view of Rudolf Schlechter
(1924), who segregated the taxa placed by Baker in
Polyxena subgenus Astemma as a distinct genus,
Neobukeria, nor of Phillips (195 1 ) or Dyer (1976). At the
same time that Schlechter described Neobakeria, he also
established another genus, Androsiphon, for a florally
unusual species discovered a few years previously by his
brother. Following this, yet another monotypic genus
was established by Barker (1936), for the species
Amphisiphon stylosa. These two genera were combined
by Phillips (1951) but have been retained as distinct by
subsequent workers, starting with Dyer (1976) and
Jessop (1976). Recent studies on the molecular system-
atics of the group reject the recognition of these genera.
The characters on which these various genera were based
are best seen as extreme adaptations to a variety of polli-
nation and seed dispersal strategies (Goldblatt &
Manning 2()()0). Those species with more conventional
flowers that were previously placed in Neobakeria are
now also confirmed to be closely allied to these florally
more unusual species. Although all of these genera were
included in an expanded concept of the genus Daubenya
(Goldblatt & Manning 2000), not all of the species were
transferred to that genus and this is accordingly done
here.
SYSTEMATIC TREATMENT
Daubenya Lindl. in Botanical Register 21.' t. 1813
(1835); Baker: 394 (1871); Baker: 417 (1897); E.Phil-
lips: 193 (1951); R.A.Dyer: 940 (1976); Jessop: 431
(1976); U.Miill.-Doblies & D.Miill.-Doblies: 91 (1997).
Type: D. aurea Lindl.
Polyxena Kunth subgenus Astemma sensu Baker: 419
(1897) non Massonia Thunb. section Astemma Endl.:
145 (1836).
Androsiphon Schltr: 148 (1924); Jessop: 432 (1976);
U.Miill.-Doblies & D.Miill.-Doblies: 86 (1997). Type: A.
c opens e Schltr.
Neobakeria Schltr.: 150 (1924); U.Miill.-Doblies &
D.Miill.-Doblies: 86 (1997). Type: N. namaquensis Schltr.
Amphisiphon W.F.Barker: 19 (1936); Jessop: 432
(1976); U.Miill.-Doblies & D.Miill.-Doblies: 86 (1997).
Type: A. stylosa W.F.Barker.
Deciduous geophytes. Bulb turbinate to globose,
sometimes deeply buried; outermost tunics leathery, dark
brown, extending in a neck as narrow, flat, papery seg-
ments. Leaves 2, spreading to prostrate, lanceolate to
ovate, longitudinally striate, glossy green, bases clasping
peduncle for some distance. Inflorescence corymbose or
racemose to subspicate, congested and capitate to coni-
cal; peduncle subterranean in flower but elongating from
base in fruit and then well-exserted; bracts usually small,
rarely large, usually increasing in size acropetally, the
uppermost sterile, often forming a coma; pedicels
suberect, vestigial to well developed in lower flowers but
decreasing in length acropetally, sometimes elongating
slightly in fruit. Flowers white to lilac, greenish yellow
or red, strongly scented or unscented, actinomorphic or
sometimes dimorphic with lower flowers slightly or
strongly bilabiate, sympetalous; perianth tube cylindrical
or somewhat dorsoventrally compressed, shorter in
upper flowers, sometimes either adaxial or abaxial part
of tube longer than opposing part in lower flowers; tepals
spreading to suberect or recurved, rarely erect and con-
nate, linear to oblanceolate, usually subsimilar but some-
times lower three much enlarged. Stamens erect or slightly
spreading; filaments subequal or adaxial ones slightly
longer, free or fused below into short or long tube, inserted
on base of tepals but sometimes tepals fused above fila-
ment insertion and then filaments apparently arising
within perianth tube; anthers dorsifixed. Ovary ovoid;
style usually between two-thirds and as long as fila-
ments; stigma penicillate; ovules 6-8 in two series per
locule. Infructescence toppling over at maturity through
elongation of lower part of peduncle. Capsule papery,
obovoid, and 3-angled or sometimes somewhat inflated
and 3-winged, acute or retuse at apex, usually dehiscing
loculicidally from top, rarely indehiscent, style rarely
persistent as a beak. Seeds globose, smooth and glossy,
black.
Bothalia 32,2 (2002)
139
Species eight. South Africa, mainly the more arid
winter rainfall parts.
Key to species
la Lower flowers strongly zygomorphic with lower three tepals
much enlarged, oblanceolate and several times longer
than upper tepals; perianth red or yellow; lower bracts
large, 25-35 mm long 8. D. aurea
lb Lower flowers not or weakly zygomorphic with tepals sub-
similar; perianth yellow to white or pinkish; lower
bracts small or large, 1-30 mm long;
2a Filaments united into narrow tube 8-12 mm long;
3a Pedicels 1-3 mm long; tepals ± connate into a narrow
tube above stamen insertion 1. D. stylosa
3b Pedicels 12-25 mm long; tepals spreading and free
above stamen insertion;
4a Flowers white; perianth tube 12-27 X 1. 2-2.0 mm;
tepals linear-oblanceolate 5. D. alba
4b Flowers yellow; perianth tube 6-10 X 3^ mm; tepals
oblong to ovate 6. D. capensis
2b Filaments free or shortly united into wide-mouthed tube up
to 3 mm long;
5a Perianth tube of lower flowers 25-45 mm long; tepals
recurved; filaments free, white or flushed lilac . . .
\. D. comata
5b Perianth tube of lower flowers 10-25 mm long; tepals
spreading or suberect; filaments free or shortly unit-
ed into tube, yellow to red;
6a Lowermost flowers zygomorphic through basal fusion
of upper tepals; tepals linear; flowers fragrant . . .
2. D. namaquensis
6b Lowermost flowers actinomorphic; tepals ovate to
lanceolate; flowers unscented;
7a Inflorescence conical, topped with conspicuous coma
of coloured bracts; perianth tube compressed-
cylindrical, 2-3 mm diam.; stamens uniformly
yellow to red 3. D. marginata
7b Inflorescence capitate, sometimes with inconspi-
cuous coma of green bracts; perianth tube cylin-
drical, 1-2 mm diam.; stamens orange to red
with basal collar flushed deep purple 4. D. zeyheri
1. Daubenya comata (Burch, ex Baker) J.C. Man-
ning & A.M.van der Merwe, comb. nov.
Massonia comata Burch, ex Baker in Journal of the Linnean
Society, Botany 11; 392 (1871); Jessop; 421 (1976); U.Mull.-Doblies
& D.Miill.-Doblies; 68 (1997). Polyxena comata (Burch, ex Baker)
Baker; 419 (1897). Neobakeria comata (Burch, ex Baker) Schltr.: 150
(1924). Type; Northern Cape, Noupoort Dist., Carolus Poort. 19 March
1813, Burchell 2751 (K, holo.!).
Bulb subglobose, 20-35 mm diam., deeply buried;
outer tunics leathery, dark brown, extending in a papery
neck up to 30 mm long. Leaves 2, spreading to prostrate,
ovate to lanceolate, 60-150 x 30-90 mm, bases clasping
peduncle for up to 80 mm, dark green or flushed maroon.
Inflorescence subspicate, capitate, exserted up to 20 mm
above leaves; bracts increasing in size acropetally, lower-
most linear or awl-shaped, white, 7-12 mm long, those
above gradually becoming broader, ovate to lanceolate,
up to 10 X 2 mm, uppermost sterile, forming a coma,
green or flushed pinkish, up to 8 x 3 mm; pedicels vesti-
gial, lowermost up to 1 mm long, flowers subsessile.
Flowers white or flushed pink, strongly scented during
the day and night, fragrance sweet and spicy; weakly
dimorphic, lower flowers slightly bilabiate, upper flowers
actinomorphic; perianth tube cylindrical or abaxial sur-
face flattened, lightly curved outwards, (25-)30-35(^5)
mm long in lower flowers, ± 15 mm long in upper flow-
ers, 2-3 mm diam.; in lower flowers adaxial half of tube
± 1 mm longer than abaxial half, with short sinus between
upper three and lower three tepals; tepals spreading from
base and then recurved or coiling downwards, linear-
oblong to narrowly oblanceolate, up to 7 x 1. 5-2.0 mm.
Stamens erect at anthesis, later spreading slightly above;
filaments subequal or adaxial ones slightly longer, free,
inserted on or up to 1 mm above base of tepals, (7-)10-15
mm long; anthers pink, 2. 5-3.0 mm long before dehis-
cence. Ovary ovoid, ± 3 mm long; style reaching to just
below or slightly above filaments, 20-40 mm long.
Capsule obovoid, 3-angled but rounded on angles, 10-12
X 7-8 mm. Seeds globose, ± 2 mm diam. Flowering time:
mid April to mid May. Figure 1; Plate lA.
Distribution and ecology: Daubenya comata is scat-
tered across the western portion of the South African
FIGURE 1 . — Daubenya comata. A,
whole plant; B, lower flower
and bract; C, capsule. Scale
bars; A-C, 10 mm. Artist;
John Manning.
140
Bothalia 32,2 (2002)
central plateau where it is largely restricted to altitudes
between 1 200-1 500 m, bordering the western rim of the
drainage basin of the Orange/Gariep and Vaal Rivers
(Figure 2). It has been recorded from Beaufort West near
the northern border of the Western Cape, eastwards to
Molteno in the Eastern Cape, through the Free State and
eastern parts of the North-West as far north as
Barberspan near Delareyville. Plants grow in colonies in
seasonally waterlogged loam or clay, particularly the red,
glutinous clays derived from the doleritic sills that are
frequently exposed across the interior plateau. These
soils retain moisture and remain damp for longer than the
surrounding karoo shales. Like D. namaquensis, another
early flowering species, D. comata responds to late sum-
mer showers, flowering before the soils dry up and the
temperatures drop too low for active growth. It shares
this strategy and habitat with several other sympatric
dwarf geophytes with a similar growth habit, particular-
ly Andwcymhium astewides, Moraea falcifolia and
Polyxena ensifolia.
The white or pink flowers are strongly and sweetly scent-
ed throughout the day and night. They secrete small amounts
of nectar and are pollinated by pollen- and nectar-collecting
bees during the day (Apis meltifera and Anthophora sp.) and
by small noctuid moths during the night.
Diagnosis and relationships: D. comata is readily rec-
ognized by the capitate inflorescence of strongly scented,
white flowers with narrow, recurved or coiled tepals and
free fdaments. The lowermost Bowers are very slightly
bilabiate through a slight asymmetry in the length of the
perianth tube, in the same way but to much less a degree
than in D. namaquensis. The two species also resemble
FIGURE 2. — Distribution of Daube-
nya comata, •; D. namaquen-
sis, ■; D. marginata. A; D.
zeyheri, ♦.
one another in their free filaments but they differ in inflo-
rescence shape, flower colour and in the form and size of
the capsules. The small, subsessile capsules in D. coma-
ta are rounded on the sides and quite unlike the inflated
and angled or winged capsules that characterize the other
species in the genus.
History: D. comata was first collected on the Farm
Caroluspoort, just south of the Free State border near
Noupoort, on 19 March 1813 (McKay 1943) by the natu-
ralist and traveller William John Burchell. Fifty years were
to elapse before the species was formally published by the
Kew botanist J.G. Baker in 1871, with due acknowledge-
ment to Burchell, who had sketched and named the
species in his field notebook at the time of its original col-
lection. It is now known to be widespread, although scat-
tered, across the southern African central plateau.
Conservation status: Not endangered.
2. Daubenya namaquensis (Schltr) J.C. Manning
& Goldblatt [‘D. namaquana'], in Goldblatt & Manning
in Strelitzia 9: 713 (2000).
Neobakeria namaquensis Schltr.: 150 (1924); U.Miill.-Doblies &
D.Miill.-Doblies: 86 (1997). Type: South Africa, Northern Cape,
Springbok Dist., Zabies (= Sabies), 4 June 1896, M. Schlechter 90 (B,
holo.; BOL!, GRA, PRE, SAM!, Z).
Massonia angustifolia sensu Je.ssop: 419 (1976) in part.
Bulb turbinate, 20-35 mm diam., deeply buried; outer
tunics leathery, dark brown, extending in a papery neck
up to 30 mm long. Leaves 2, spreading to prostrate,
lanceolate, 100-150 x 25-70 mm, bases clasping pedun-
Bothalia 32,2 (2002)
141
cle for up to 100 mm, dark green. Inflorescence subspi-
cate, conical, exserted up to 90 mm above leaves; bracts
increasing in size acropetally, lowermost awl-shaped,
white, 1-2 mm long, those above gradually becoming
longer and oblanceolate, up to 6 x 3 mm, uppermost ster-
ile, forming a coma, spathulate, green or flushed pinkish,
up to 8 X 3 mm; pedicels suberect, decreasing in length
acropetally, lowermost up to 5 mm long, uppermost up to
0.5 mm long, flowers subsessile. Flowers greenish yel-
low, strongly scented during the day and night, fragrance
sweet and spicy, camation-like; dimorphic, lower flow-
ers slightly bilabiate, upper flowers actinomorphic; peri-
anth tube cylindrical or abaxial surface flattened, lightly
curved outwards, 13-15 mm long in lower flowers, 9-10
mm long in upper flowers, ± 2 mum diam.; in lower flowers
adaxial half of tube ± 2 mm longer than abaxial half with
pronounced sinus between upper three and lower three
tepals; tepals spreading from base and curving upwards
in outer half, linear-oblanceolate, 9-10 x 1-1.5 mm,
apices weakly cucullate. Stamens erect at anthesis, later
spreading slightly above; filaments subequal or adaxial
ones slightly longer, 13-15 mm long, free, inserted on
base of tepals; anthers yellow, up to 2.5 mm long before
dehiscence. Ovary ovoid, ± 3 mm long; style reaching to
between two-thirds and as long as filaments, 15-25 mm
long. Capsule abscising at maturity, obovoid, 20-23 x 15
mm, somewhat inflated, 3-angled and introrse at base but
3-winged and obtuse at apex, dehiscing loculicidally
from top. Seeds globose, ± 2 mm diam. Flowering time:
mid May to mid June. Figure 3; Plate IB.
Distribution and ecology: Daubenya namaquensis is
known from a few collections made in the semi-arid flats
east of Okiep and Springbok at an altitude of 1 000 m
(Figure 2). This region, just east of Namaqualand proper
and on the extreme western edge of Bushmanland, lies on
the boundary between winter and summer rainfall regions
and in consequence receives both erratic late summer
showers as well as some rain from the occasional winter
frontal system that extends inland from Namaqualand.
The species responds rapidly to autumn rains and flowers
in early winter before temperatures drop too low for
active growth. Plants of D. namaquensis occur in small
colonies in deep red sands in a vegetation dominated by
the willowy shrub, Sisyndite spartea (Zygophyllaceae)
and the tussock grass, Stipagrostis namaquensis. The
colonies are restricted to lower-lying drainage areas in
which the water table remains nearer the surface. The
very deeply seated bulbs are an obvious adaptation
enabling the plants to make use of this soil moisture.
The greenish yellow flowers are strongly and sweetly
clove-scented throughout the day and night. They secrete
small amounts of nectar and are probably pollinated by a
variety of diurnal and nocturnal insects, including bees
and moths.
Diagnosis and relationships: D. namaquensis is distin-
guished by the well-exserted conical inflorescence of
strongly scented, yellow flowers with linear tepals and
free filaments. Although confused by Jessop (1976) with
D. marginata, the resemblance between the two species is
very superficial, encompassing nothing more than the
prominent inflorescence topped by a coma of sterile
bracts. The flowers of D. marginata are unscented, with
ovate, whitish or pale yellow tepals and golden yellow or
orange filaments that are fused at the base into a short col-
lar. A distinct feature of D. namaquensis is the bilabiate
lower flowers in which the upper side of the perianth tube
is prolonged for ± 2 mm beyond the lower side, resulting
in a distinct sinus between the three upper and three lower
tepals. This differential in the tube length rapidly
becomes less evident higher up the inflorescence and the
upper flowers are quite actinomorphic. This asymmetry
FIGURE 3. — Daubenya namaquen-
sis. A, whole plant; B, lower
flower and bract. C, D, upper
flower: D, L/s. E, capsule.
Scale bars: A-E, 10 mm.
Artist: John Manning.
142
Bothalia 32,2 (2002)
in the lower flowers is not very evident without careful
examination, especially in pressed material. It occurs
also, although to a lesser degree, in some other species of
Daubenya, particularly D. comata and D. stylosa.
History. D. namaquensis was first collected by Max
Schlechter during a trip to Namaqualand with his broth-
er, the botanist Friedrich Richard Rudolf Schlechter,
soon after his arrival in South Africa in May 1896. Two
decades were to pass before it was described, when it
formed the basis of the genus Neobakeria, established by
Rudolf Schlechter to accommodate the group of species
until then treated as Polyxena subgenus Astemma. These
seven species were distinguished from Polyxena s. str. by
their uniseriate stamens, and from Massonia by their
inconspicuous bracts. Although struck by the well-
exserted, foxtail-like inflorescence of D. namaquensis,
Schlechter ironically overlooked the even more diagnos-
tic zygomorphy of the lower flowers through dissecting
only the more accessible upper flowers, which are per-
fectly actinomorphic. This characteristic was also over-
looked by Jessop ( 1976), who confused the species with
D. marginata. The curious zygomorphy of the lower
flowers of D. namaquensis was only recently highlight-
ed by Muller-Doblies & Muller-Doblies (1997). They
used it as the basis for restricting the circumscription of
the genus Neobakeria to this species alone, while in turn
overlooking the fact that floral zygomorphy of the same
type occurs, albeit to a lesser extent, in both D. comata
and D. stylosa. Over a century was to pass before the
species was collected for the second time and it is still
known from only a handful of specimens.
Conservation status: Endangered (Bl+2c, D).
3. Daubenya marginata (Willd. ex Kunth) J.C.
Manning & A.M.van der Merwe in Bothalia 32: 65 (2002).
Massonia marginata Willd. ex Kunth; 299 (1843). Polyxena mar-
ginata (Willd. ex Kunth) Benth. & Hook, ex T.Durand & Schinz: 366
(1895). Neobakeria marginata (Willd. ex Kunth) Schltr.; 150 (1924).
Type: South Africa, Caput Bonae Spei (B-WILLD 6373, holo.-NBG,
photo.!).
Massonia rugulosa Licht. ex Kunth: 299 (1843). Polyxena rugu-
losa (Licht.. ex Kunth) Baker: 420 (1897). Neobakeria rugulosa (Licht.
ex Kunth) Schltr.: 150 (1924). Type: Caput Bonae Spei, Lichtenstein
224 (B, holo., ?destroyed).
Polyxena haemanthoides Baker: t, 1727 (1888); E. Phillips; t. 56
(1922a). Neobakeria haemanthoides (Baker) Schltr.: 150 (1924). Type:
South Africa, Nuweveld Mountains near Fraserburg, April 1886, Bolus
5493 (BOL, lecto.l, designated by J.C. Manning & A.M.van der
Merwe: 65 (2002); G, SAM!].
Mas.sonia angustifolia auct. non M. angustifolia L.f. (= M. echina-
ta L.).
Bulb subglobose, 20-30 mm diam., shallowly buried;
outer tunics leathery, dark brown, extending as a papery
neck up to 40 mm long. Leaves 2, spreading, ovate to
elliptic, 50-100 x 25-50 mm, base clasping peduncle for
20-30 mm, apiculate, dark green or flushed maroon.
Inflorescence subspicate, capitate or conical, exserted
20-60 mm above leaves at flowering; bracts erect, low-
ermost triangular, 2-5 mm long, becoming oblanceolate-
spathulate acropetally, up to 8 x 3 mm, uppermost ster-
ile, imbricate, forming a short or elongate coma flushed
orange, up to 10 x 5 mm; pedicels 1.0-2. 5 mm long.
Flowers greenish yellow with yellow or orange stamens,
unscented; perianth tube compressed-cylindrical,
10-15(-20) X 2-3 mm; tepals suberect, inner three con-
nate for 0. 5-1.0 mm beyond outer three, lanceolate,
conduplicate, 6-8 x 2.5 mm. Stamens connate for
1.5-2. 5 mm in a wide-mouthed, slightly flaring staminal
collar; filaments suberect and lightly incurved,
10-15(-22) mm long; anthers yellow, 2. 0-2. 5 mm long
before anthesis. Ovary ovoid, ± 6 mm long; style lightly
deflexed, 1 1-20 mm long. Capsule abscising at maturity,
ellipsoid to obovoid, 12-15 x 15-20 mm, somewhat
inflated, introrse at base, obtuse at apex, 3-angled below
but 3-winged at apex. Seeds globose, ± 2.5 mm diam.,
glossy black. Flowering time: May to July, rarely to
August. Figure 4; Plate 1C.
Distribution and ecology: Daubenya marginata is
widespread across the Roggeveld Escarpment and west-
ern karoo at altitudes of 1 000-1 500 m, from near
Calvinia and Williston southwards to Sutherland, thence
eastwards along the Nuweveld scarp as far inland as
Fraserburg and Loxton (Figure 2). A few isolated popu-
lations occur to the west, below the Bokkeveld Es-
carpment, at an altitude of 200 m in the Knersvlakte
north of Vanrhynsdorp. Populations occur on silt or grit-
ty clay, more rarely red clays derived from dolerite, in
seasonally moist depressions or washlines. D. marginata
grows sympatrically with D. aurea in some colonies but
flowers several months earlier.
Diagnosis and relationships: D. marginata is mor-
phologically very close to D. zeyheri and the two were
regarded as conspecific by Jessop (1976). There are,
however, several differences between them, some rather
subtle. In D. marginata the inflorescence is typically
conical and protrudes above the leaves, and is invariably
surmounted by a coma of conspicuous yellow or orange,
often spathulate bracts. The globular inflorescence of D.
zeyheri, in contrast, does not protrude much above the
leaves and is at most topped by an inconspicuous coma
of narrow, green bracts. The two species differ also in
flower form and colour, although in both the perianth
tube varies greatly in length. In D. marginata the peri-
anth tube is always conspicuously flattened and 2-3 mm
in diameter, the tepals tend to be flushed yellow and
firm-textured and the filaments are uniformly pigmented
yellow to orange or red. The flowers of D. zeyheri, in
contrast, are characterized by a much more slender peri-
anth tube 1.0-1. 5 mm in diameter, white tepals that tend
to be almost papery in texture and, most conspicuously,
reddish filaments that are flushed deep purple at the base
where they are fused into a shallow collar.
The plants with their orange to reddish stamens and
conical inflorescence topped with enlarged orange or
yellow bracts are conspicuous when in flower and are
visited by Malachite sunbirds which probe the flowers
for nectar. The relatively broad tubes contain abundant
nectar that accumulates in the wide-mouthed staminal
collar where it is readily accessible to the birds, which
feed from the ground. The lack of floral scent is also con-
sistent with bird pollination.
Bothalia 32,2 (2002)
143
FIGURE 4. — Daubenya marginala.
A, whole plant. B, C, flower:
B, with bract; C, 1/s. D, cap-
sule. Scale bars: A-D, 10
mm. Artist: John Manning.
History: the true identity of this species has been mis-
understood ever since the Kew botanist J.G. Baker con-
fused it with Massonia angustifolia L.f., possibly as
early as 1871 but certainly by 1897. This mistake was
perpetuated by subsequent authors until recently, when it
was shown that M. angustifolia is actually conspecific
with Massonia echinata L. (Manning & Van der Merwe
2002). The species known by all authors after Baker as
M. angustifolia is in fact properly known as Daubenya
marginata, first described as Massonia marginata by the
Berlin Botanist Carl Kunth. Although known to his pre-
decessor, Carl Willdenow, it was not formally named
until Kunth (1843) published the name that had earlier
been suggested for it by Willdenow. The collector and
place of collection of the type specimen are unknown.
The species was later described as Polyxena haeman-
thoides by Baker in 1888, based on a plant collected by
the Cape Town stockbroker and botanist, Harry Bolus,
on the Nuweveld Mountains near Fraserburg in April
1886. It seems that Bolus intended describing the species
himself, as Haemantholirion capense, but Baker must
have dissuaded him from this step. Baker did, however,
retain the reference to the genus Haemanthus (Ama-
ryllidaceae) that the appearance of the plant had suggest-
ed to Bolus. Baker, although suspecting that his species
was similar to Kunth’s, had not seen material of M. mar-
ginata and thus described it as new. The species was re-
cognized until Jessop (1976) revised the genus. Un-
convinced that the apparent differences between it and D.
zeyheri were significant, he combined the two taxa under
the misapplied name M. angustifolia.
Conservation status: Not endangered.
4. Daubenya zeyheri (Kunth) J.C. Manning &
A.M.van der Merwe, comb. nov.
Massonia zeyheri Kunth in Enumeratio plantarum 4: 298 (1843);
U.Miill.-Doblies & D.Miill.-Doblies: 77 (1997). Polyxena zeyheri
(Kunth) Benth. & Hook, ex T.Durand & Schinz: 367 (1895). Type:
without locality or date, Zeyher 298 [fB, holo. destroyed; K, lecto.!,
designated by Jessop: 419 (1976)].
Massonia pedunculata Baker: 8 (1892). Type: South Africa,
Malmesbury Dist., near Hopefield, Schaapplaatsfontein, June 1887,
Bachmann 2043 (K. lecto.!, here designated; B,-BOL, drawing!).
Massonia burchellii Baker: 393 (1871). Polyxena burchellii
(Baker) Baker: 420 (1897). Neobakeria burchellii (Baker) Schltr.: 150
(1924). Type: South Africa, Caput Bonae Spei, Burchell s.n. (K,
holo.!,-BOL, drawing!).
Massonia angustifolia auct. non M. angustifolia L.f. (= M. echina-
ta L.).
Bulb subglobose, 20-30 mm diam., outer tunics leath-
ery, dark brown, extending as a papery neck up to 10 mm
long. Leaves 2, spreading, elliptic to lanceolate, 60-150
X (10-)30-80 mm, bases clasping peduncle for 10-100
mm, apiculate, dark green. Inflorescence corymbose,
capitate, exserted 20-40 mm above leaves; bracts erect,
lowermost triangular, 2-3 mm long, becoming lanceolate
acropetally and increasing in size, up to 8 x 2 mm,
uppermost sometimes sterile and forming inconspicuous
coma; pedicels 2-10 mm long, lowermost sometimes
longer than upper. Flowers translucent, whitish tinged
pink on tube, stamens reddish tinged purple at the base,
papery in texture, unscented; perianth tube cylindrical,
lightly constricted at mouth, 12-20 mm long in lower
flowers but rarely more than 13 mm in upper flowers,
1. 5-2.0 mm diam, inner tepals connate for 0. 5-1.0 mm
beyond outer ones; tepals spreading at base then
suberect, lanceolate, conduplicate, 6-10 x 2. 5-3.0 mm.
Stamens reddish but staminal collar purple, connate for
1.0-1. 5 mm in a flaring staminal collar; filaments
suberect, 12-19 mm long; anthers yellow or reddish, 2-3
mm long before anthesis. Ovary ovoid, ± 4 mm long;
144
Bothalia 32,2 (2002)
FIGURE 5. — Daiibenya zeyheri. A,
whole plant. B, C. flower: C,
1/s. D, capsule. Scale bars:
A-D, 10 mm. Artist: John
Manning.
Style 15-30 mm long. Capsule abscising at maturity,
broadly ovoid, 10-15 x 9-13 mm, somewhat inflated, 3-
angled, introrse at base, obtuse at apex. Seeds globose, ±
2.5 mm diam., glossy black. Flowering time: late May
and June to early July. Figure 5; Plate ID;
Distribution and ecology: Daubenya zeyheri is a coastal
species occuring near sea level, unlike the other species in
the genus, which occur at higher altitudes inland. It is
restricted to the west coast of Western Cape, where it is
now known from only three locations, one at Paternoster
and Cape Columbine and the others less than 20 km to the
south, near Saldanha (Figure 2). The material that formed
the type of M. pedunculata was collected 1 0 km southeast
of Hopefield on the Farm Schaapplaatsfontein but no fur-
ther collections have been made from here and this popu-
lation has presumably disappeared under wheat. D. zeyheri
grows on sandy calcareous soils overlying limestone,
sometimes in dense colonies.
The flowers, with their reddish filaments and con-
trasting glistening purple centres attract the attention of
Lesser double-collared sunbirds, which probe the flow-
ers for the copious nectar that is held in the wells formed
by the staminal collars. The lack of floral fragrance is
typical of bird-pollinated flowers.
Diagnosis and relationships: although closely related
to D. marginata and sometimes difficult to separate from
it in the dry state, living plants of D. zeyheri are readily
recognized by the conspicuous purple centres to the
flowers. The pigmentation is restricted to the staminal
collar and when this is filled with nectar the flowers
appear to have a glistening black centre that is quite lack-
ing in D. marginata, in which the stamens are uniformly
yellow or orange to reddish. Other more subtle differ-
ences between the two species are discussed under D.
marginata. The two were treated as conspecific by
Jessop (1976).
History: D. zeyheri has been described several times
under different names but the earliest name dates from
1843, when the species was described by the Berlin
botanist, Carl Kunth, from a collection made by the pro-
fessional plant collector, Carl Zeyher. Zeyher’s collec-
tion was made sometime between 1829 and 1834, prob-
ably near Saldanha Bay, which is still one of only two
sites where the species is known to occur. The species
was redescribed twice, both times by the Kew botanist
J.G. Baker. In 1892 he described Massonia pedunculata
from a specimen collected near Hopefield by the German
naturalist Frans Bachmann, who practised medicine in
the village from 1886-1887. A few years later he named
Massonia burchellii from a collection made by the natu-
ralist and traveller William John Burchell. Baker was
clearly unsure about the exact identity of several of his
species because he later transferred M. burchellii to
Polyxena while leaving M. pedunculata in Massonia,
although noting that it lacked the large bracts that are so
characteristic of the genus.
Conservation status: Endangered (Bl+2c, C2).
5. Daubenya alba A.M.van der Merwe, in
A.M.van der Merwe & Marais in South African Journal
of Botany 68: 312 (2002). Type: South Africa, Northern
Cape, 25 km SW Middelpos, Farm Botuin, June 2001,
Van der Merwe 195 (NBG, holo.!).
Bulb subglobose to turbinate, 10-15 mm diam., usu-
ally shallowly buried; outer tunics leathery, brown,
extending in a short, papery neck up to 5 mm long.
Leaves 2, suberect to spreading, ovate to lanceolate,
40-100 X 15-50 mm, bases flushed red and clasping
peduncle for up to 30 mm, dark green. Inflorescence
corymbose, capitate, exserted up to 20 mm above
leaves; bracts ovate-lanceolate, 3-4 mm long; pedicels
well developed, lowermost 12-19 mm long, elongating
slightly in fruit and ultimately 20-25 mm long. Flowers
mauve or white with tips of tepals, filaments and
exposed portion of style flushed pale lilac or mauve,
strongly scented during the day and night, fragrance
sweet and spicy, actinomorphic; perianth tube cylindri-
Bothalia 32,2(2002)
145
FIGURE 6. — Daubenya alba. A,
whole plant. B-D, flower: B,
with bract; C, 1/s; D, detail of
mouth of staininal tube. E.
capsules. Scale bars: A-C, E,
10 mm; D, 5 mm. Artist:
John Manning.
cal, ( 10-)12-25(-27) mm long, 1. 5-2.0 mm diam.;
tepals spreading from base, linear-oblong to narrowly
oblanceolate, (8-) 10-13 x 1. 5-2.0 mm. Stamens con-
nate below into cylindrical tube 8-10(-20) mm long,
free parts suberect or lightly incurved at tips, (6-)8-15
mm long; anthers brown to purple, ± 2 mm long before
dehiscence. Ovary' ovoid, ± 4 mm long; style reaching
up to 2 mm beyond anthers, 28-30(-50) mm long.
Capsule ovoid-cuneate to obtriangular, (12-)20-25 x
10-12 mm, tapering below, deeply retuse above, three-
winged. Seeds globose, 2. 0-2. 5 mm diam., glossy
black. FloweriiiQ time: mid May to mid June. Figure 6;
Plate IE.
Distribution and ecology: Daubenya alba is known
from a few scattered localities along the edge of the
Roggeveld Escarpment between Calvinia and Middelpos
at an altitude of 1 000-1 500 m (Figure 7). The plants
FIGURE 7. — Distribution of Daubenya alba, •; D. capensis, ■.
grow in colonies in seasonally waterlogged doleritic
clay, on low hills or at the foot of rocky dolerite outcrops.
Diagnosis and relationships: D. alba is readily recog-
nized by its white to lilac flowers with the stamens fused
below into a narrow tube 6-10 mm long. The slight
zygomorphy of the lowermost flowers, resulting from a
slight asymmetry in the staminal tube, that is mentioned
in the original description, is not always evident. The
relationships of D. alba appear to lie with D. capensis,
which it resembles in several respects, particularly its
relatively long pedicels, cylindrical staminal tube and
large, obtriangular capsules that are deeply retuse at the
apex. The two species are readily distinguished by their
flowers. In D. capensis the firm-textured flowers are
bright yellow with a shorter, broader tube, 6-10 mm long
and ovate tepals. The flowers of D. alba are altogether
more delicate, with a slender perianth tube, 12-20 mm
long and narrowly oblanceolate tepals. In addition D.
alba lacks the conspicuous disc occluding the mouth of
the staminal tube that is characteristic of D. capensis.
History: the most recently discovered species in the
genus, D. alba was in cultivation in a few specialist col-
lections by the early 1990s, although the type material
was only collected from the wild in June 2001.
Conservation status: Vulnerable (Bl+2c, D).
6. Daubenya capensis (Schltr.) A.M.van der
MerH’e & J.C. Manning, in Goldblatt & J.C. Manning in
Strelitzia 9: 713 (2000).
Androsiphon capense Schltr.: 148 (1924); Brandham: 124 (1990);
U.Miill.-Doblies & D.Miill.-Doblies: 86 (1997). Type: South Africa,
Northern Cape. Calvinia Dist., Onder-Bokkeveld, Oorlogskloof, 2500
ft, July 1897, R. Schlechter 10969 (B. holo.; BOL!, G, GRA, PRE, Z).
Bulb subglobose, 20-30 mm diam., outer tunics leath-
ery, dark brown, extending as a very short, papery neck
up to 5 mm long. Leaves 2, spreading, ovate to elliptic.
146
Bothalia 32,2 (2002)
FIGURE 8. — Dauhenya capensis. A,
whole plant. B-D, flower: B,
with bract; C, 1/s; D, detail of
mouth of staminal tube. E,
capsule. Scale bars: A-C, E,
10 mm; D, 5 mm. Artist: John
Manning.
50-150 X 25-90 mm, bases clasping peduncle for 20-80
mm, apiculate, dark green. Inflorescence corymbose,
capitate, 30 x 40 mm; bracts erect, linear-oblanceolate,
lowermost 25-30 x 2-3 mm, becoming shorter acro-
petally, uppermost oblanceolate-spathulate; pedicels
15-25 mm long, elongating slightly in fruit and ulti-
mately 20-30 mm long. Flowers firm-textured, golden
yellow with staminal column and operculum flushed red-
dish orange and style streaked with red, yeast-scented;
perianth tube subcylindrical, widening slightly upwards,
6-10 X 3^ mm; tepals suberect, inner three sometimes
connate for up to 1 mm beyond outer three, oblong to
oblanceolate, 11-15 x 3^ mm. Stamens connate in a
stiff, thick-walled cylinder for 9-12 mm, top of tube
occluded by convex disc or operculum through which
style protrudes, upper portion of filaments suberect and
lightly incurved, 10-14 mm long; anthers yellow,
2.0-2. 5 mm long before anthesis. Ovary ovoid, ± 6 mm
long; style erect, 20-25 mm long. Capsule ovoid-cuneate
to obtriangular, 20-30 x 10-12 mm, tapering below,
three-winged with apex deeply retuse. Seeds globose, ±
2.5 mm diam., glossy black. Flowering time', late June
and July, rarely early August. Figure 8; Plate IF.
Distribution and ecology: Dauhenya capensis is
endemic to the immediate vicinity of Nieuwoudtville,
where it is known from several populations around the
village at an altitude of 800 m (Figure 7). It is restricted
to seasonally moist dolerite Hats in red clay. The large,
prominently three-winged capsules borne on relatively
long pedicels form a rounded, balloon-like infructes-
cence that is readily dispersed by the wind.
The curious disc that occludes the mouth of the tube
is glossy and has the appearance of being wet even when
dry, although nectar does in fact ooze out of the pore in
the staminal di.sc and accumulate on it. Nothing is known
about the pollination biology of the species but the firm
texture of the flowers and their yeasty odour suggest
rodent pollination.
Diagnosis and relationships: D. capensis is readily
distinguished from other species of Dauhenya in which
the filaments are fused into a long, narrow column by the
peculiar disc that occludes the mouth of the column and
through which the style protrudes via a narrow pore. D.
capensis is most similar in floral form to D. alba, which
it also resembles in its distinctly pedicellate flowers and
large, obtriangular capsules that are deeply retuse at the
apex. The fruits in these two species are among the
largest in the genus. D. capensis is readily distinguished
from D. alba by its more robust, yellow flowers with
shorter perianth tube, 6-10 mm long and by the conspicu-
ous disc that occludes the mouth of the staminal tube.
History: D. capensis was first collected in July 1897 by
the German botanist, Friedrich Richard Rudolf Schlechter
and was described by him some years later, when it
formed the basis of his monotypic genus Androsiphon.
Here it remained until recently, when that genus was
included in Dauhenya (Goldblatt & Manning 2000).
Conservation status: Vunerable (Bl-i-2c, D).
7. Dauhenya stylosa (W.FBarker) A.M.van der
Mem’e & J.C. Manning, in Goldblatt & J.C. Manning in
Strelitzia9: 713 (2000).
Amphisiplwn stylo.fiim ['.stylosa' j W.F.Barker: 19 (1936);
Brandham: 58 (1989); U.MUll.-Doblies & D.Mull.-Doblies; 86 (1997).
Type; South Africa, Northern Cape, Calvinia Dist., 3 miles north of
Nieuwoudtville, 21 June 1934, Salter 4552 (BOL, holo.!).
Bulb subglobose to turbinate, 20-35 mm diam., usu-
ally shallowly buried; outer tunics leathery, dark brown,
extending in a papery neck up to 5 mm long. Leaves 2,
suberect to spreading, ovate to lanceolate, (20-)50-100 x
15-50 mm, bases clasping peduncle for up to 30(-60)
mm, dark green. Inflorescence subspieate, capitate or
rarely conical, exserted up to 50 mm above leaves; bracts
increasing in size acropetally, lowemiost lanceolate,
white, 3-5 mm long, uppermost oblanceolate, up to 8 x
3 mm; pedicels suberect, 1-3 mm long, flowers subses-
sile. Flowers greenish yellow with free part of filaments
and style golden yellow, honey-scented; perianth tube
cylindrical, 8-14 mm long in lower flowers with portion
Bothalia 32,2 (2002)
147
FIGURE 9. — Daubenya stytosa. A,
whole plant. B-E, flower: B,
with bract; C, 1/s; D, tepal
detail of lower part; E, tepal
detail of upper part. F, cap-
sule. Scale bars: A-C. F, 10
mm; D, E. 5 mm. Artist; John
Manning.
below level of filament insertion 4-7 mm long but 6-8
mm long in upper flowers with portion below level of fil-
ament insertion 2-3 mm long, 2.0-2. 5 mm diam.; lower
flowers lightly bilabiate with adaxial portion of tube ± 3
mm longer than abaxial half through a greater degree of
fusion of upper three tepals; tepals connate into tube
above level of filament insertion, free parts oblong to
ovate, 2-5 mm x 2.0-2. 5 mm. Stamens fused below into
tube 10-15 mm long, free parts of filaments suberect,
9-10 mm long; anthers yellow, up to 2.5 mm long before
dehiscence. Ovary ovoid, ± 3 mm long; style reaching to
top or just beyond filaments, 17-28 mm long. Capsule
depressed-ovoid, 10-12 x 7-10 mm, lightly retuse at
base and tapering above into peristent style which forms
slender beak ± 20 mm long, indehiscent but eventually
separating along septa. Seeds globose, ± 2 mm diam.,
glossy black. Flowering tune: mid May to mid June.
Figure 9; Plate IG.
Distribution and ecology. Daubenya stylosa is a high-
ly local endemic around Nieuwoudtville where it is
known from a few populations in the immediate vicinity
of the town, especially along the trekpath that passes the
Farm Glenlyon, at an altitude of 800 m (Figure 10). The
populations here are extremely dense. The plants typi-
cally grow in lower-lying drainage areas in red doleritic
clays. Scattered plants occur along the foot of dolerite
outcrops on the Wildflower Reserve and on Glenlyon
itself. The species is more or less sympatric with D.
capensis but the two cannot be confused, either in flower
or fruit. The compact infructescence in D. stylosa lacks
the adaptations for wind dispersal typically found in
most other species in the genus and the seeds are mainly
shed immediately around the parent plants.
The sweetly scented, greenish yellow flowers secrete
small amounts of nectar that ooze out of the mouth of the
staminal column. They are visited avidly for the nectar
by honeybees (Apis mellifera) and the Painted lady but-
terfly (Cynthia cardui) during the day and by several
species of noctuid moth at dusk.
Diagnosis and relationships: the species is readily rec-
ognized by its almost completely fused tepals, which form
a tube around the staminal column, and by the persistent
styles that remain attached to the capsule. These stiff,
prong-like beaks are a conspicuous feature of the subsessile
fruits. In addition, unlike most other species in the genus,
which have loculicidally dehiscent capsules, the fruits in D.
stylosa are essentially indehiscent although the locules
eventually separate along the septa to release the seeds.
History. D. stylosa was first collected relatively
recently, in June 1934, by retired Royal Navy Paymaster,
T.M. Salter, best known for his work on the genus
Oxalis. It was described soon thereafter in the monotyp-
ic genus Amphisiphon by the Cape Town botanist W.F
Barker. Included in Amphisiphon by Phillips (1951), the
genus was recently transferred to Daubenya (Goldblatt &
Manning 2000).
FIGURE 10. — Distribution of Daubenya stylosa, •; D. aurea, ■.
148
Bothalia 32,2 (2002)
Conservation status: Vulnerable (Bl+2c, D).
8. Daubenya aurea Lindl. in Botanical Register
21: t. 1813 (1835); Kunth: 301 (1843); E.Phillips: t. 71
( 1922b); Jessop; 43 1 ( 1976); U.Mull.-Doblies & D.Mull.-
Doblies: 91-95 (1997). Type: South Africa, Cape, with-
out locality or collector, cultivated by Messrs Young of
Epsom (icono.!).
Daubenya fulva Lindl.: t. 53 (1839); Kunth: 300 (1843). Type:
received from the Cape of Good Hope, with other bulbs, but [mistak-
enly] believed to have been collected somewhere on the East Coast of
Africa or in Madagascar, cultivated by Robert Barchard, Wandsworth
(CGE, holo.).
Daubenya coccinea Harv. ex Baker: 395 (1871). D. aurea Lindl.
var. coccinea (Harv. ex Baker) Marloth in E.Phillips (1922b). Type:
Caput Bonei Spei, Harvey s.n. (TCD, holo.).
Bulb subglobose to turbinate, 20-35 mm diam., usu-
ally shallowly buried; outer tunics leathery, dark brown,
extending in a short, papery neck up to 10 mm long.
Leaves 2, suberect to spreading, ovate to lanceolate,
50-100(-140) X 20-75 mm, bases clasping peduncle for
5-50 mm, dark green. Inflorescence corymbose, capitate,
exserted up to 50 mm above leaves; bracts decreasing in
size acropetally, greenish, lowermost obovate to oblance-
olate, 25-35 x 10-15(-20) mm long, uppermost up to 20
X 10 mm; pedicels suberect, lowermost 5-10 long,
uppermost up to 3 mm long. Flowers brilliant red or
canary yellow, rarely flushed reddish at tips of tepals, at
most lightly scented, ± zygomorphic and bilabiate; lower
flowers strongly bilabiate with abaxial or lower half of
tube much prolonged and adnate to filament column and
lower three tepals forming conspicuous lip separated
from upper tepals by pronounced sinus, sometimes dor-
sal or uppermost tepal arising very much below remain-
ing tepals, perianth tube obliquely cylindrical, 15-25
mm long on upper side but 20-40 mm long on lower
side, 3^ mm diam., upper tepals oblong-ovate, laterals
obliquely so, 2-A x 1. 5-2.0 mm, lower tepals oblanceo-
late to obovate, subequal or median larger, 20-40 x 5-15
mm; upper flowers weakly zygomorphic or actinomor-
phic, perianth tube 13-20 x 2. 5-3. 5 mm, tepals oblong.
2-A X 1 .5-2 mm. Stamens fused basally into an oblique
tube, 5-7 mm long on adaxial side but much longer on
abaxial side, especially in lower flowers, and there
adnate to lower tepals such that lower stamens appear to
be free, free parts of filaments suberect, 4—8 mm long;
anthers yellow, 2. 5-3.0 mm long before dehiscence.
Ovary ovoid, 7-8 mm long; style reaching to top or just
beyond all or only adaxial filaments, 20-25 mm long.
Capsule oblong-ovoid, 3-angled, 15-20 x 7-10 mm,
introrse at base but acute at apex. Seeds globose, ± 3 mm
diam., glossy black. Flowering time: late August to mid
September. Figure 11; Plate IH.
Distribution and ecology: D. aurea is a highly local-
ized endemic of the Roggeveld Escarpment known from
three or four localities midway between Sutherland and
Middelpos, at an altitude of ± 1 500 m (Figure 10). The
known populations lie along the eastern foot of a mean-
dering dolerite ridge, with the total distribution little
more than 10 km in extent. The plants grow in colonies
in low-lying drainage lines in seasonally damp red clay
derived from dolerite. Populations are sympatric with D.
marginata in several of the localities but flower much
later than that species. D. aurea occurs in two colour
forms, the more common one a brilliant red and the other
canary yellow. Populations are typically uniformly
coloured, although red populations will often contain
occasional yellow morphs, whereas yellow populations
may include a few plants in which the tepals are flushed
reddish at the tips. The yellow populations consistently
come into flower one or two weeks before the red but the
fruits appear to take much longer to ripen. The prominent
papery bracts that are characteristic of the species are
particularly conspicuous in fruit, serving as sails that
assist in the wind dispersal of the infructescences.
The inflorescence resembles a large daisy capitulum
in the genus Arctotis or Gazania due to the enlargement
of the lower florets, and the species is adapted to polli-
nation by monkey beetles (Scarabidae: Holpiinae) that
congregate on the large flower heads to mate. The flow-
ers, unlike those of the other species in the genus, do not
produce nectar and are also unscented or at most weakly
FIGURE 1 1 . — Daubenya aurea. A,
whole plant; B, lower flower
and bract. C, D, upper
flower: D, 1/s. E, capsule.
Scale bars: A-E, 10 mm.
Artist: John Manning.
Bothalia 32,2 (2002)
149
scented. Both of these characteristics are typical adapta-
tions to pollination by monkey beetles.
Diagnosis and relationships: D. aurea is a highly dis-
tinctive species that cannot be confused with any other
species in the genus. The highly zygomoiphic lower
flowers in which the outer tepals are greatly exaggerated
in size make this species instantly recognizable. It is also
characterized by large floral bracts that are persistent and
papery in fruit.
History: it is ironic that the source of the original
material of the best known species of Daiihenxa
remained obscure for so long. D. aurea was described in
1835 by John Lindley, Professor of Botany at the
University College. London and head of the Chelsea
Physic Garden, from a yellow-flowered plant cultivated
by the nurserymen, Messrs Young of Epsom. Nothing is
known of its origin beyond that the Youngs obtained
their plant from the Cape of Good Hope under the name
Massonia liitea. Three years later a reddish-flowered
variant of the species came to Lindley’s attention through
the agency of John Royle, then Professor of Materia
Medica at King’s College, London, from the collection
of Robert Barchard of East Hill, Wandsworth. Lindley
was struck by the apparently more extreme zygomorphy
evident in the flowers of the red form and he recognized
this collection as a different species, D. fulva. on this
basis. The name is an allusion to the tawny colour of the
plant, but this, along with the rather etiolated appearance
of the plant, was probably the result of cultivation in less
than ideal light. From the illustration it is clear that this
plant was in fresher flower than the first one that Lindley
had seen and he presumably examined only the lowest,
most zygomorphic flowers in the inflorescence. This
plant had been received from the Cape of Good Hope
with other bulbs but was believed to have been collected
somewhere along the East Coast of Africa or in
Madagascar. This was, of course, not the case but it is
possible that Barchard received his plant from the same
source as did the Youngs. A dried plant of D. fulva was
later seen by Baker (1897) in the herbarium of William
Wilson Saunders, who had a notable garden at East Hill
from 1857 to 1874 and was a neighbour of Barchard. It
is thus probable that his specimen came from Barchard
himself and is possibly even the one illustrated in the
protologue. At the same time that Lindley was describing
his two species of Daubenya. a specimen appears to have
been in the possession of William Henry Harvey in Cape
Town. Harvey, although nominally Colonial Treasurer,
occupied most of the four years that he spent in Cape
Town between 1835 and 1842 on botany. Harvey's speci-
men formed the basis of D. coccinea, described by Baker
(1871), which was distinguished from D. fulva by its
compact inflorescence and more robust flowers.
Although the provenance of the plant is unknown, its
appearance is typical of wild collected material or that
grown under good light, such as would have been expe-
rienced in Cape Town. It is impossible to be certain on
this point but the more or less coincidental appearance of
material in three different hands strongly suggests that it
all originated from the same source. Circumstantial evi-
dence favours Capt Walter Synnot as this source. Synnot
spent four years in Clanwilliam as deputy landdrost,
returning to Ireland in 1825 with numerous bulbs and
seeds. These were disposed of in England to various
nurseries and included various species from the
Bokkeveld and Roggeveld Escaipments, some of which
only flowered in the 1830s. It is thus quite possible that
among them were seeds or plants of D. aurea.
Almost a century was to pass without further infor-
mation until, in 1920, Rudolph Marloth came upon
Daubenya aurea in the wild on the Roggeveld
Escarpment. Marloth, a retired chemist with a passionate
interest in South African botany, sent plants and exact
details of the locality and habitat to the Pretoria botanist,
E.P. Phillips, who published the story of the rediscovery
in 1922. At the same time it became evident that no sig-
nificant structural differences existed between the three
species hitherto known and they were consequently
reduced to one. Although further plants were sent to
Kirstenbosch gardens by a Mr Metelerkamp in 1936
(Hall 1970), they soon languished and it was left to
HaiTy Hall, then Senior Horticulturalist at the Gardens,
to locate Marloth’s collecting site in 1968 and bring the
species into cultivation once more. He has provided a
lively account of his search, illustrated with fine photo-
graphs of the plants in the wild (Hall 1970).
Consen'ation status: Vulnerable (Bl-i-2c, D).
Specimens examined
Acocks 16344. 22097 ( 1 ) PRE. Archibald 3066, 3118 ( 1 ) PRE.
Barker 9736, 10179, 10237, 10244 (4) NBG; 6703 (2) NBG; 9366,
9389 (6) NBG; 9390, 10545 (7) NBG. Boucher 5153 (3) NBG.
Cloete & Ha.felau 96 (8) NBG. Coetzer 67 (3) NBG. Compton 702/28,
7470 ( 3) NBG; NBG1531/26 (6) NBG.
Hall 225, 3242, 4253 (3) NBG; 3240. s.n. (8) NBG; 3269 (8) NBG.
PRE; 3276 (8) PRE. Narrower 305 (5) NBG. Henrici 3983 ( 1 ) PRE.
Herman 530 ( 1 ) PRE.
Leipoldt 796 (6) NBG. Leistner 1900 (1) PRE; 2508 (2) KMG. M.
PRE. Louw 1 828 (\)nOL. PRE.
Manning 2175 (4) NBG; 2250 (2) NBG; 2263. 2343. .t.n. (3) NBG;
2327. 2329 (\) NBG; 2345 (5) NBG; Marloth 10346 (8) PRE; 10415
(8) BOL; 10415 (8) NBG. PRE. McCarthy SAM23080 (1) SAM.
Meiring BOL2775/15 ( 8) BOL. F.W. Meterlekamp 355/37 (8) NBG.
Mo.stert 819 (\) PRE. Muller sub Marloth 9550 (8) PRE.
Oliver 4423. 9464 (3) NBG; 4412, 8932. 8972 (8). NBG.
Peny 3511 (8) NBG. Pern,' & Snijnian 2132, 2143 (7) NBG.
Ro.s.toitw .'i.n. (3) NBG.
Saaiman 125 (I) PRE. Salter 7558/34 (7) NBG; 4452 (3) BOL.
Schlechter 90 (2) SAM. BOL. Schlieben 9017 {2) BOL. BM. BR. M.
PRE. Z. Snijnuin 1228, 1705 (7) NBG. Stayner .s.n. (3) NBG. Steytler
.s.n. (4) NBG.
Thomp.son 2439, 3011, 3093 (3) NBG.
Van der Merwe 195 (5) NBG. Van W\'k 934 0) PRE; 2578 (6) BOL;
2579 (7) BOL. Vlok 633 (6) NBG.
Wilnian s.n. ( 1 ) BOL.
Zainbatis 202 ( 1 ) PRE. Zietsman & Zietsman 1700 ( 1 ) PRE.
ACKNOWLEDGEMENTS
We are grateful to A. Horstmann and G. Summerfield
for providing material for illustration and to the curators
and staff of the cited herbaria, particularly Clare Archer.
The colour illustrations were kindly sponsored by Rachel
and Rod Saunders of Silverhill Seeds.
150
Bothalia 32,2 (2002)
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Bothalia 32.2: 151-166 (2002)
The genus Trichodesma (Boraginaceae: Boraginoideae) in southern
Africa
E. RETIEF* and A.E. VAN WYK**
Keywords: Boraginaceae. Boraginoideae. morphology, pollen, southern Africa, taxonomy, Trichodesma R.Br., Trichodesmeae
ABSTRACT
Trichodesma R.Br. is a genus of about 45 species known from tropical and subtropical regions of Africa. Asia and
Australia. The group comprises predominantly perennial herbs, the flowers characterized by anthers with prolonged con-
nectives. often twisting above the thecae, and a prominent accrescent calyx. Five species and three subspecies of
Trichodesma are currently recognized in southern Africa, with T. angusufoiium Harv. subsp. argenteum Retief & A.E. van
Wyk newly described. These taxa are widespread in southern Africa, occurring in various vegetation types. A key to the
species, descriptions, distribution maps and illustrations of various micro- and macromorphological as well as palynologi-
cal features are provided.
CONTENTS
Abstract 151
Introduction 151
Materials and methods 151
Historical outline 151
Tribal delimitation 152
Morphological characters of taxonomic signifi-
cance 152
Phytogeography 156
Trichodesma R.Br 158
Key to species 159
1. T. zeylaniciim (Burm.f.) R.Br 159
2. T. africanum (L.) Lehm 160
3. T. angiistifolium Harv 160
4. T. physaloides (Ttnz\) A.DC 161
5. T. ambacense Welw 163
Specimens examined 163
Acknowledgements 164
References 165
INTRODUCTION
Brown described Trichodesma in 1810, a genus well
defined by flowers with a deeply divided and strongly
accrescent calyx, the absence of fomices, anthers usually
with fairly long, soft hairs on the back and conspicuous
long, linear, often twisted connectives produced above
the thecae. It belongs to the family Boraginaceae estab-
lished by Jussieu (1789), and subfamily Boraginoideae.
This family is characterized amongst others, by coiled,
cymose inflorescences, setae with multicellular bases and
in most genera, fruit that develop into four nutlets.
The genus Trichodesma R.Br. comprises ± 45 species,
known from tropical and subtropical regions of Africa,
Asia and Australia (Mabberley 1997). It is currently rep-
* National Herbarium. National Botanical Institute. Private Bag XIOI,
0001 Pretoria.
** H.G.W.J. Schweickerdt Herbarium, Department of Botany, University
of Pretoria, 0002 Pretoria.
MS. received: 2001-06-05.
resented by five species and three subspecies in southern
Africa; about 15 species occur in Africa south of the
Sahara (Lebrun & Stork 1997). Members of Tricho-
desma are widespread in southern Africa and occur in
various vegetation types, ranging from grassland and
savanna (bushveld) to succulent shrubland.
The purpose of this paper is to present a taxonomic
revision of the genus Trichodesma in southern Africa.
Diagnostic characters, two keys, one based mainly on
fruiting characters, the other on the leaf and flowering
characters, full descriptions of all species and infraspe-
cific taxa, illustrations and distribution maps are provid-
ed. The genus and species descriptions are based on
material from southern Africa and the adjacent Flora
zambesiaca (FZ) region. This paper forms part of a revi-
sion of the Boraginaceae currently in progress for the
Flora of southern Africa (FSA) region, henceforth
referred to as southern Africa.
MATERIALS AND METHODS
Herbarium specimens of Trichodesma in BFLU, BM,
BOL, COI, E, GRA, J, K. NBG, NH, NMB, NU, PRE.
PRU, SAM and WIND (acronyms as in Holmgren et al.
1981) were examined to gather data on morphological
characters, phenology and distribution. Specimens of
taxa occurring outside southern Africa that were exam-
ined, are not cited. This was supplemented by field work
which included observations on the effect of fire on
plants of T physaloides in the Pretoria National Botani-
cal Garden, where this species occurs naturally. Pollen
and various plant parts were studied with an ISI-SX-25
scanning electron microscope. Measurements of pollen
grains were done from acetolysed grains mounted in
glycerine jelly. Acetolysis followed the standard method
of Erdtman ( 1960).
HISTORICAL OUTLINE
Brown (1810) based his concept and description of
Trichodesma on Borago zeylanica Burm.f. and B.
152
Bothalia 32,2(2002)
africanum L. When De Candolle (1846) revised the
genus, he recognized eight species. In the description of
T. africanum, De Candolle mentioned that it was collect-
ed by Thunberg and by Drege in the ‘Cape of Good
Hope’, the first southern African record for the genus. De
Candolle also included Friedrichsthalia physaloides
Fenzl, a species previously described from the Sudan.
Wright (1904) noted that T. physaloides (Fenzl) A. DC.
also occurred in the Transvaal, South Africa. He added a
third southern African species, T. angustifolium,
described by Harvey (1859), but was not aware of T. z.ey-
lanicum (Burnt. f.) R.Br., today also known from southern
Africa, although originally described from Sri Lanka
(Ceylon), and most probably introduced to the region by
man. Two species, T. africanum and T. angustifolium,
were recognized in the flora of South West Africa (now
Namibia) by Friedrich-Holzhammer (1967). Herman
( 1993) listed five species and a subspecies for southern
Africa, namely T. africanum, T. angustifolium, T areni-
cola Giirke, T. physaloides, T. zeylanicum and T. amha-
cense Welw. subsp. hockii (De Wild.) Brummitt. How-
ever, specimens from southern Africa previously identi-
fied as T. arenicola, proved to be T. amhacense Welw.
subsp. hockii. In the present paper we recognize five
species and three subspecies of Trichodesma in southern
Africa, namely T. africanum (L.) Lehm., T. amhacense
Welw. subsp. hockii (De Wild.) Brummitt, T. angustifoli-
um Harv. subsp. angustifolium, T. angustifolium Harv.
subsp. argenteum Retief & A.E.van Wyk subsp. nov., T.
physaloides (Fenzl) A. DC. and T. zeylanicum (Burm.f.)
R.Br. Brummitt (1990), in his account on Trichodesma
for the Flora zamhesiaca (FZ) region, recognized the
same species for the FZ region except for T. africanum,
a species with a disjunct distribution between the FSA
region, Angola and North Africa, and which is not known
in the region. T. zeylanicum, T. physaloides and T. amha-
cense extend further north into Africa, supplemented by
other species (Verdcourt 1991).
TRIBAL DELIMITATION
De Candolle ( 1846) assigned Trichodesma to the tribe
‘Borrageae’ and subtribe Cynoglosseae, a classification
followed by most subsequent authors, for example
Hooker (1885), Giirke (1897), Baker & Wright (1905),
Brand (1921), Melchior (1964) and Hilger (1985). In
1941 Zakirov placed Trichodesma in a tribe of its own,
Trichodesmeae, recognized by RiedI (1967) and is also
accepted in the present contribution. Trichodesnia is well
defined by a combination of floral and palynological
characters: deeply divided and strongly accrescent calyx;
the absence of fornices in the corolla tube; anthers that
usually have fairly long, soft hairs on the back; and con-
spicuous long, linear, often twisted connectives above
the thecae. Pollen of the genus shows essentially no
interspecific differences, is isopolar, tricolporate and
with a nodular tectum. This pollen type is strikingly dif-
ferent from that of most genera of the tribe Cynoglosseae
in which the grains are heterocolpate and the tectum psi-
latc (Retief & Van Wyk 1999a, b). Avetisian (1956)
agreed with Zakirov in removing Trichodesma from the
tribe Cynoglosseae, based on the structure of its flowers
and fruits, because the pollen grains of the genus have
nothing in common with those of the Cynoglosseae.
Pollen morphology thus strongly supports the recogni-
tion of the separate tribe, Trichodesmeae.
Members of Trichodesma differ significantly from
each other in nutlet morphology (see note on fruit mor-
phology). However, the presence of glochidia on the
outer surface of the nutlets of T. africanum and T angus-
tifolium could be used to argue that the placement of
Trichodesma in its own tribe is not Justified, because
glochidia also occur on the outer nutlet surface of
Cynoglossum and Afrotysonia. The types of glochidia
found in Trichodesma, however, differ from those of
Cynoglossum and Afrotysonia. Hilger ( 1985) studied the
development and morphology of flowers and fruits of 23
species of the Cynoglosseae and Eritrichieae, and com-
mented on the implications of his findings on the tax-
onomy of these groups. He found that in most
Cynoglosseae the nutlets are initially positioned with
their disc (the flat base) parallel to the basal area (the
nectary disc) of the gynoecium. Subsequent growth in a
vertical direction brings the nutlets to their final oblique
position. In both species of Trichodesma studied by
Hilger, he observed that the nutlets are, from the begin-
ning, in an oblique position. This can also be used to sup-
port the recognition of a separate tribe for this genus.
Two other genera are assigned to the tribe Tricho-
desmeae, namely Caccinia Savi and Suchtelenia Kar. ex
Meisn. (Riedl 1967). Giirke ( 1897) separated five genera
using the accrescent calyx, either enclosing the fruit or
expanded, as the key character; Trichodesma, Such-
telenia, Caccinia, Brachyhotrys Maxim, ex Oliv. and
Heliocarya Bunge. Heliocatya is regarded as congeneric
with Caccinia (Mabberley 1997) and Brachyhotrys
belongs to the tribe Trigonotideae (Riedl 1997). Such-
telenia has 6-heterocolpate pollen grains, showing simi-
larity to pollen of members of Heliotropioideae,
Cynoglosseae, Eritrichieae and Myosotideae. The tria-
perturate pollen grains of Trichodesma and Caccinia, on
the other hand, show similarity to pollen of the subfami-
lies Wellstedioideae and Ehretioideae. The tribe
Trichodesmeae can be regarded as ‘primitive’ within the
subfamily Boraginoideae, showing similarities to various
other genera within the Boraginaceae s.l. The family is
sometimes treated as two separate entities, Boraginaceae
s. str. and Ehretiaceae, but various characteristics sup-
port the recognition of one family divided into several
subfamilies (Retief & Van Wyk 1999c).
MORPHOLOGICAL CHARACTERS OF
TAXONOMIC SIGNIFICANCE
Hahit
Trichodesma is a genus of perennial and annual herbs
or subshrubs. T. africanum is the only southern African
species that is usually annual or occasionally biennial.
This growth form can be correlated with its distribution
in arid and desert regions where fast growth under opti-
mal conditions is essential.
The other species are perennial herbs or subshrubs
occurring in summer rainfall regions, mainly in grassland
and savanna. They are subjected to winter drought and
Bothalia 32,2 (2002)
153
also to regular natural fires and frost. Grassland species
are usually quick in responding to the effects of fire
(smoke, change in temperature and the release of nutri-
ents), and plants may sprout a number of inflorescences.
With sturdy, often very old. fire-resistant rootstocks and
mass seed production by fire-stimulated flowering, these
species are well-adapted to survive unfavourable condi-
tions. Most examples of fire-stimulated flowering plants
do flower in the absence of fire, but not as profusely as
when subjected to fire. It was, however, observed that
unbumed plants of T. physaloides did not flower at all or
produced only a few inflorescences, as opposed to burned
plants of the species in the same grassland that sprouted
and produced inflorescences abundantly after a natural fire
(Figure lA, B). Burning of this particular piece of grass-
land under controlled circumstances to remove moribund
and/or unacceptable grass material had no effect on the
number of flowers produced, because the intensity of the
fire was too low. A cool or low-intensity fire of less than
1 000 kJ/s/m is usually applied (Trollope 1992).
Leaf
Infraspecific taxa of Trichodesma differ remarkably
in the indumentum of the leaf blade. The leaf trichome
complement of the southern African members of
Trichodesma consists of setae with multicellular bases,
and simple, unbranched hairs. T. physaloides and T.
ambacense subsp. hockii have flat, large-based setae, but
differ in the structure of the setae (Figure 2A, B, D. E).
T ambacense subsp. hockii (Figure 2D) has slender
setae, orientated in different directions, whereas T.
physaloides (Figure 2E) has shorter, attenuate setae, ori-
entated more or less in a direction parallel to the midrib.
Upper leaf surfaces of the taxa concerned are relatively
densely hairy (Figure 2A, C, H), whereas the lower leaf
surfaces of T. physaloides (Figure 2B), T. ambacense
subsp. hockii, T. angustifoliiim subsp. angustifoliwn and
T. africanum (Figure 21) are sparsely hairy, with setae
scattered on the midribs and along the veins. T. angiisti-
foliurn subsp. angustifoliiim is characterized by an indu-
mentum of appressed setae, orientated in a direction par-
allel to the midrib and varying in length and in size at the
base, with distinct spaces between the trichomes. The
leaf surfaces of T. angustifolium subsp. argenteum
(Figure 2C, F) are silver-grey, covered with a dense layer
of setae on both surfaces, a feature distinguishing this
taxon from all other southern African taxa of Tricho-
desma. The outline of the multicellular bases of the setae
is not circular, but slightly asymmetrical.
Leaves of T zeylanicum (Eigure 2G) are characterized
by a lower surface that is densely hairy, with mainly sim-
ple, small hairs, but also with scattered setae along the
midrib and some veins. The upper surface is covered by
setae of which the large, 2-layered multicellular bases are
± circular in outline, with much smaller setae in between
(Figure 2H). T africanum has a spinose indumentum
with stiff setae on the lower surface (Figure 21). Cells of
the multicellular base of the setae are narrower compared
to those of T zeylanicum. The upper surface of T.
africanum is characterized by well-spaced setae with a
prominent row of swollen cells at the point where the
seta and multicellular base join. Cells of the multicellu-
lar bases of the older leaves are impregnated with silica
and some calcium, giving them a spotted appearance.
Flower
Various flower characters in Trichodesma are taxo-
nomically significant. Anthers with connectives lengthe-
ning into usually twisted appendages above the thecae,
distinguish the genus from all other southern African
members of the family (Figure 3A, B). Long, shaggy
hairs (Figure 3A, C, F) are present on the dorsal surfaces
of the connectives, whereas the inside of the thecae are
glabrous (Figure 3F). The subglobose stigma displays
two types of receptive surfaces (Figure 3D, E, G, H). T.
angustifolium has papillae with prominent contiguous
caps (distal swellings) and crenulate margins (Eigure 3D,
E). Papillae without distinct caps occur in the other taxa
studied (Eigure 3G, H). Corolla lobes are often twisted
(Figure 31) and acuminate to long-acuminate. The colour
of the lobes is usually shades of blue except for T. physa-
loides which has white lobes with a brownish rim.
A prominent feature of the flower in Trichodesma is
the calyx which is strongly accrescent when in fruit
FIGURE 1. — Trichodesma physaloides. A. plant in unbumed grassland; B, plant flowering abundantly after a natural veld fire.
154
Bothalia 32,2 (2002)
FIGURE 2. — Trichodesma leaf surfaces. A, Trichodesnui amhacense subsp. hockii, Schoenfelder 159. upper surface; B, T. physaloides, Jacobsen
1813, lower surface; C, T. anguslifoliiim subsp. argentewn. Relief 1404.01, upper surface; D, T. amhacense subsp. hockii, Schoenfelder
159. setae, slightly hooked, on upper surface; E, T. physaloides, Onderstall 1248, stiff setae on upper surface; F. T. angustifoliinn subsp,
argenteum. Relief 1404.01, slender setae on upper surface. G, H, T. zeylanicuni, Stahnans 1076: G, lower surface; H, upper surface. I, T.
africannm, Sinook 7695, stiff setae on midrib of lower surface. Scale bars; A, G 18 um; B, 1 1 pm; C, 8 pm; D, 49 pm; E, 26 pm; F, 51
pm; H, 2 pm; I, 37 pm.
(Figure 4). T. phy,saloides and T. amhacense subsp.
hockii are characterized by a single mature nutlet with a
persistent style. The style is initially gynobasic, but due
to abortion of three ovules, it becomes orientated side-
ways. These single nutlets, together with the calyx and
style are shed from the plant. This type of nutlet disper-
sal also occurs in Cryptantha flava (A.Nels.) Payson,
another member of the Boraginaceae. According to
Casper & Wiens (1981), the abortion of three of the four
ovules may be an adaptation for dispersal by wind, the
entire floral structure serving as a relatively light disper-
sal unit. The attached, accrescent, papery calyx may aug-
ment the buoyancy of nutlets in air currents and increase
the dispersal distance. In the case of T. africanum it can
be speculated that the spinose indumentum of the calyx
protects the nutlets, allowing them to mature before any
damage can be done by herbivores.
Pollen
Pollen studies revealed no significant differences
between the species of Trichodesma included in this
Bothalia 32,2 (2002)
155
FIGURE 3. — Trichodesma flower parts. A-E, Trichodesma angiistifoliwn subsp. angustifolium. Peelers, Gericke & Burelli 163: A, style, stigma
and upper parts of anthers; B, twisted connectives; C. hairs on back of anthers; D. stigma; E, papillae. F, T. physaloides. Fourie 2804, sec-
tion of two anthers. G-I. T. africanum, Gennishuizen 4575: G, stigma; H, papillae; I, twisted apices of corolla lobes. Scale bars; A, 375
pm; B. 178 pm; C, 264 pm; D, 51 pm; E, 8 pm; F, 750 pm; G, 40 pm; H. 18 pm; I, 259 pm.
Study, a finding that is in agreement with Brummitt
(1982). Pollen grains of the genus are isopolar, radially
symmetrical, tricolporate, subspheroidal, with P =
7.0-11.5 pm, E = 6.0-9. 5 pm, P/E = 1.1-1. 2 (Eigure
5A-F). The shape of the pollen grains in polar view is ±
triangular, with convex mesocolpia and sunken apertures
(Eigure 5B). The equatorial view is elliptic to rounded
with protruding ora; grains are angulaperturate (Figure
5A, C). Long apertures, extending over about Vs of the
length of the polar axis, are characteristic of the grains.
They are comparatively narrow with acute ends and the
margins are not conspicuously different. Ora are coarse-
ly granular, with endo-apertures lalongate (Figure 5D).
The tectum is nodular with nodules ± of the same size
(Figure 5E), sometimes absent in the vicinity of the ora
(Figure 5C, D). As noted in the discussion on the tribal
delimitation of the genus, some authors prefer to classify
Trichodesma in the tribe Cynoglosseae but the tricolpo-
rate pollen of the genus supports its placement in the sep-
arate tribe Trichodesmeae. Pollen grains of members of
the tribe Cynoglosseae are usually heterocolpate, quite
different from those of Trichodesma. The similarity in
pollen morphology between Trichodesma and Cordia,
does not support the separation of the Boraginaceae into
two separate families (Retief & Van Wyk 1999a).
Fruit
In Boraginaceae fruit characters are often used to dis-
tinguish species. Brand (1921) recognized six sections in
Trichodesma, based on the outer surface morphology of
the nutlets. Members of four sections are present in
southern Africa, similar to the diversity reported by
Verdcourt (1991) for East Africa. T. physaloides and T.
156
Bothalia 32.2 (2002)
FIGURE 4. — Trichodesma physaloides. Mature fruit and accrescent
calyx, X 1. Relief 2223 (PRE).
shaped, silken-haired nutlet (Figures 4; 6A; lOE).
Nutlets of T. physaloides are covered with thin attenuate
hairs (Figure 6B), whereas those of T. ambacense var.
hockii have an indumentum of hooked hairs (Figure 6C).
The other species have a fruit of four nutlets and are
divided into two groups. Nutlets of T. zeylanicum (sect.
Trichodesma) are emarginate and smooth on the outer
face (Figure 6F), but rugose on the inner face. T. angus-
tifoliwn (sect. Acanthocaryum Brand) and T. africanum
(sect. Serraticaryum Verde.) are characterized by the
presence of glochidia (Figure 6D, E, G). The glochidia
differ, however, in structure (Figure 6E, G). T. angusti-
folium is characterized by much smaller barbs than T.
africanum. Nutlets of 71 africanum are ovoid-cup-shaped
with distinct, serrate, raised margins and a concave,
glochidiate inner face (Figure 6G, H). The family
Boraginaceae is characterized by flat, conical or pyrami-
dal gynobases as e.g. in 71 africanum where the gynobase
is pyramidal (Figure 61).
PHYTOGEOGRAPHY
ambacense subsp. hockii (sect. Friedrichsthalia (Fenzl) Trichodesma is widespread in southern Africa (Figure
A.DC.) are characterized by a single, + circular, cushion- 7) where members occur in various vegetation types.
FIGURE 5. — Trichodcswci pollen, A-E: unacclolysed; E, ucelolysed. A, B, T. africaiuint: A, Genni.shuizen 4575. side view; B, Nicholas 2576,
polar view. C, I), T. zeyUmicunr, Strey 5656: C, group; D, equatorial view of grain. E. T. ungustifoUum subsp. aiigiislifoliiim, Hanekom
I4W. teetuin; F', T. ambacense subsp. hockii, Codd 7416, group. Seale bars; A, B, 2.8 pm; C, 6.5 pin; D, 3.2 pm, E, 1.0 pm; E, 4.8 pm.
Bothalia 32.2 (2002)
157
FIGURE 6. — Trichodesma nutlet surfaces. A. T. physaloides. JBG6150. margin; B. hairs; C. T. amhacense subsp. hockii. Schoenfelder 159.
hooked hairs. D. E. T. angustifolium subsp. angustifoliunv. D, Gerstner 5133. glochidiate surface; E. Hanekom 1498. glochidia. E, T. zey-
laniciim. Netshiungani 915. smooth nutlet. G-I. T. africanum: G. Bengis 463. margin with glochidia; H. Bengis 463. setae on dorsal sur-
face; I. Van Jaarsveld 8825. pyramidal gynobase, one nutlet removed. Scale bars: A. 838 pm; B. C, 84 pm; D, 749 pm; E. 131 pm; F. 713
pm; G. 79 pm; H. 49 pm; 1. 419 pm.
ranging from grassland and savanna to succulent shrub-
land, mainly in the Savanna and Grassland Biomes. All
southern African species extend further north into Africa.
Trichodesma angustifolium subsp. argenteum, occur-
ring in the northern parts of Namibia, is the only taxon
endemic to the FSA region. Brummitt (1985) described a
new Trichodesma species from the volcanic regions of
Kenya, T. marsabiticum Brummitt, which is very similar
in facies to T. angustifolium, and can be regarded as a
vicariant of the southern African species, which differs in
calyx and corolla characters. T. angustifolium subsp.
angustifolium itself is disjunct in distribution in southern
Africa (Figure 9), and its current occurrences may repre-
sent relicts of a once much wider distribution. Brummitt
(1985) considered that an early record of the species
from Bulawayo in Zimbabwe should be disregarded
unless modem collections can substantiate it.
The floristic connection between the dry areas of
southern Africa, especially Namibia, and northeastern
tropical Africa is well known, and is ascribed to a so-
called arid corridor which connected the two regions at
various times in the past (Verdcourt 1969; De Winter
1971; Thulin & Johansson 1996). T. africanum and T.
angustifolium are examples of species confined to arid
158
Bothalia 32,2 (2002)
FIGURE 7. — Distribution of Trichodesma in southern Africa.
climatic conditions: in Africa it is known from arid parts
of southern Africa, Angola and North Africa, thus
belonging to the disjunct Afro-arid element.
The genus Trichodesma occurs in all six phytogeo-
graphical regions of southern Africa recognized by White
(1983): Zambesian region, Kalahari-Highveld Transition
Zone, Karoo-Namib Region, Tongaland-Pondoland
Region, Afromontane Region and Cape Region. T physa-
loides occurs mainly in the Afromontane floristic region
which extends from the northern parts of Africa to south-
ern Africa along the eastern mountain ranges. The Afro-
montane region comprises a series of isolated highland
areas and is well represented in eastern Africa from Yemen
to South Africa, but is also present in the Cameroon
(Denys 1980). The region in southern Africa is character-
ized by numerous small forest patches in a grassland or
fynbos matrix. The edges, or ecotones between forests and
grasslands are usually sharp (typically just over a few
metres) and are mainly maintained by both natural and
more anthropogenic fires. T ambacense subsp. hockii,
closely related to T. physaloides, is found in some regions
where T. physaloides occurs but is also found in regions
such as Nigeria (see Brummitt 1985).
Trichodesma zeylarucurn is an ancient Old World
weed of human cultivation, introduced in southern
Africa, ± 1905. It is no longer possible to distinguish nat-
ural occurrences of this species from introductions,
hence its original native region is uncertain. T. zeylan-
icum occurs in grassland, woodland, in dry river beds, is
a common weed in old and new cultivated land, a pioneer
in disturbed areas, and is found along roadsides. The
species is known from Namibia (Caprivi), the eastern
parts of Northern Province, Mpumalanga, Swaziland and
northern KwaZulu-Natal.
TAXONOMIC TREATMENT
Trichodesma R.Br., Prodromus florae novae hol-
landiae: 496 (1810), nom. conserv.; Harv.: 241 (1838);
A.DC: 172 (1846); Benth.: 845 (1876); Hook.f.: 153
(1885); Gurke: 99 (1897); C.H.Wright: 10 (1904); Baker
& C.H.Wright: 44 (1905); Brand: 19 (1921); E.Phillips:
628 (1951); Friedr.-Holzh.: 2 (1967); Riedl: 219 (1967);
Kazmi: 515 (1971); Taton: 36 (1971); Kabuye & Agnew:
520 (1974); S.P.Banerjee & Pramanik: 109 (1975);
R.A.Dyer: 510 (1975); J.R.Edm.: 435 (1978); Qaiser: 31
(1979); Tolken: 1172 (1986); Brummitt: 94 (1990);
Nowicke & J.S.Mill.: 24 (1991); Verde.: 91 (1991);
E.S. Martins & Brummitt: 61 (1993); Riedl: 124 (1997);
Relief: 182 (2000). Type: Trichodesma zeylanicum
(Burm.f.) R.Br. (= Borago zeylanica Burm.f.)
Borraginoides Boehm.: 18 (1760). Pollichia Medik.:
247 (1783); non Pollichia Alton: 5 (1789). Boraginella
Kuntze: 435 (1891). Type: Borago indica L.
Friedrichsthalia Fenzl: 53 (1839). Type: Friedrichs-
thalia physaloides Fenzl.
Leiocarya Hochst.: 30 (1844). Type: Leiocarya kotschy-
ana Hochst.
Annual and perennial herbs, often with annual stems
from a woody rootstock, variously hairy. Leaves simple,
opposite, subopposite or alternate, basal leaves usually
opposite and petiolate, upper ones alternate and sessile or
all leaves sessile, diminishing in size towards inflor-
escences; blade variously shaped, entire; stipules absent.
Inflorescence a scorpioid, cymose panicle, bracteate,
usually terminal. Flowers bisexual, regular, pedicellate,
calyx, corolla and stamens usually 5-merous. Calyx
deeply divided to base or lobes loosely adherent along
lower margin, finely setulose on abaxial side, variously
hairy on adaxial side; lobes ovate or narrowly ovate,
sometimes winged, base rounded to cordate, apex acute,
acuminate or cuspidate, strongly accrescent and papery
in fruit. Corolla blue or white; tube campanulate or fun-
nel-shaped, naked in throat, but with gibbosities between
lobe sinuses; lobes ovate to broadly triangular, apex trun-
cate or acute to acuminate, sometimes cuspidate, often
spreading or reflexed. Stamens arising from base or from
throat of corolla tube, sessile or filaments shorter than 1
mm and broader than long; anthers linear-oblong or lin-
ear-lanceolate, with long hairs on back, connectives pro-
longed above anthers, often twisted together at apex,
usually exserted. Ovary 4-lobed, with a single ovule in
each loculus; style persistent, gynobasic, terete, narrow-
ing above; stigma subglobose, papillate, papillae with or
without distinct caps, caps with crenulate margins. Fruit
either 4 nutlets or by abortion 1; nutlets ovoid, planocon-
vex or biconvex, smooth or variously ornamented.
The name Trichodesma alludes to the twisted hairs or
awns that terminate the anthers; Greek thrix, trikhos = hair
and desme = a band or bundle (Tolken 1986).
Brand (1921) divided the genus into six sections but
his taxonomy and nomenclature have been queried since.
Some of the rather striking differences could be consid-
ered almost of generic importance (Verdcourt 1991), but
we found no supporting palynological differences to
justify this view in agreement with Brummitt (1982).
Five species and three subspesies are recognized in south-
ern Africa. Southern African species of Trichodesma rep-
resent the following four sections:
Bothalia 32.2 (2002)
159
Sect. Trichodesma; Riedl: 225 (1967).
Fruit of four emarginate, smooth, shining nutlets,
gynobase pyramidal with four strongly concave sides,
each with ± winged margin; nutlets compressed-ovoid,
outer face usually mottled with grey and brown, inner
face rugose, covered with indistinct papillae {T. zeylan-
icum).
Sect. Serraticaryum Verde/. 92 (1991) (sect
Friedrichsthalia Brand non (Fenzl) A. DC.).
Fruit of four nutlets, ovoid-cup-shaped with distinct
serrate margins and glochidiate hairs on outer face, i.e.
base of cup; outer face convex, inner face concave, ser-
rate margin also with glochidia; gynobase pyramidal
with four strongly concave sides, each with ± winged
margin, verrucose (T. africanum).
Sect. Acanthocaryum Brand: 27 (1921).
Fruit of four nutlets, glochidiate, marginal glochidia
usually coalescing at bases to form distinct rim; gynobase
pyramidal with four strong concave sides, each with ±
winged margin; nutlets compressed-ovoid {T. angustifoU-
um). Brummitt (1992) suggested that this section might
be combined with sect. Friedrich.sthalia Brand non
(Fenzl) A.DC., now sect. Serraticaryum Verde.
Sect. Friedrichsthalia {Fenzl) A.DC.: 173 (1846).
Fruit a single, ± circular, cushion-shaped, silken-
haired nutlet, concealed in accrescent calyx until
released, attached by most of its lower surface to recep-
tacle; gynobase flat; style persistent, lateral (T. amba-
cense subsp. hockii, T. physaloides).
Key to species
(based on fruiting characters)
la Fruit of four nutlets:
2a Nutlet with outer surface smooth and shiny; lower leaf blade surface with setae only along midrib and veins,
intercostal surfaces with dense cover of small bulbous-based hairs, orientated in different directions, upper
leaf blade surface covered with long and much smaller setae in between 1. 71 zeylanicum
2b Nutlet with outer surface glochidiate; indumentum of leaf blade not as above:
3a Glochidia thick, mainly along serrate, raised margin of nutlet, thick short setae on dorsal side of nutlet; leaf
blade ovate to narrowly ovate, lower surface with tubercle-based, stiff setae usually only along midrib and
veins, upper surface with well-spaced, discoid-based setae 2. 71 africanum
3b Glochidia slender, covering whole surface of nutlet; leaf blade linear to narrowly elliptic, lower and upper
surfaces with well- to densely spaced, discoid-based, slender setae 3. 71 angiL^tifolium
lb Fruit a single nutlet:
4a Corolla lobes white with a light brown rim, rounded to truncate with short cuspidate apices; lower leaf blade
surface with short, stiff, discoid-based setae scattered on midrib and veins, upper leaf blade covered with
similar setae well spaced 4. 71 physaloides
4b Corolla lobes blue or rarely whitish, ± triangular, with long acute apices; lower and upper leaf blade surfaces
with broad, discoid-based, slender setae, well spaced 5. 71 ambacense
Key to species
(based on leaf and flower characters)
la Leaf blade linear to narrowly elliptic 3. 71 angustifoUum
lb Leaf blade broadly ovate, ovate, narrowly ovate or elliptic:
2a Corolla lobes scarcely spreading when fully developed,
rounded to truncate, cuspidate white and with a
brownish rim; leaf blade sparsely hairy with stiff,
short setae 4. 71 physaloides
2b Corolla lobes reflexed over calyx when fully developed,
acute to acuminate at apices, blue or rarely whitish,
not with a brownish rim; leaf blade hairy, setae slen-
der:
3a Accrescent calyx lobes cordate at base, ± 22 x 15 mm in
fruit 5. 71 ambacense
3b Accrescent calyx lobes ± rounded at base:
4a Stems, inflorescence axes and calyx spinously hairy;
lower leaf surfaces sparsely hairy with setae scat-
tered along midribs and veins 2. 71 africanum
4b Stems, inflorescence axes and calyx patently hairy;
lower leaf surfaces with setae only along midribs
and veins, intercostal surfaces with a dense cover
of small, bulbous-based hairs, orientated in differ-
ent directions 1 . 71 zeylanicum
1. Trichodesma zeylanicum (Burm.f.) R.Br.,
Prodromus florae novae hollandieae; 496 (1810); A.DC.: 172
(1846); A.Rich.: 91 (1850); Hook.: t. 4820 (1854); Klotzsch:
255 (1861); Gurke: 99 (1897); Baker & C.H. Wright: 51
(1905); Brand: 40 (1921); F.W. Andrews: 90 (1956); Ivens: 83
(1967); Taton: 37 (1971); Kabuye & Agnew: 520 (1974);
S.P.Baneijee & Pramanik: 109 (1975); Tdlken: 1172(1986);
Bmmmitt: 95 (1990); Nowicke & J.S.Mill.: 25 (1991);
Verde.: 92 (1991); E.S.Martins & Bmmmitt: 62 (1993);
Retief & P.P.J.Herman: 354 (1997); Riedl: 125 (1997). Type:
Ceylon, Garcin in Herb. Bunn. (G, lecto.-K, photo.!).
Borago zeylanica Burm.f.: 41 (1768). PolUchia zeylanica (Burm.f.)
F.Muell.: 100 (1882). Boraginelia zeylanica (Burm.f.) Kuntze: 435 (1891).
Borraginoides zeylanica (Burm.f.) Hiem: 720 (1898).
Leiocarya kotschyana Hochst.: 30 (1844). Types: Ethiopia,
Djelajeranne, Schimper 625 (B, syn.t, BM. K! isosyn.) & Sudan,
Nubia, Camamil and Gebbel Kassan, Kotschy 542 (Bt, K!, W, isosyn.).
Erect, perennial herb with annual stems from a woody
rootstock, up to 1.5 m high. Stems, inflorescences and
petioles patently hairy. Leaves sessile or petiolate; blade
elliptic, 30-110(-120) x 10-30(-35) mm, base rounded
to subcordate, apex acute, discolorous, upper surface with
long, up to 0.75 mm, larged-based, 2-layered setae and
much smaller setae in between, orientated ± in same
direction, lower surface with long setae on midrib and
major veins only, intercostal area densely covered with
small, bulbous-based setae orientated in different direc-
tions; petiole up to 10 mm long. Flowers with pedicels up
to 30 mm long in fmit. Calyx lobes ovate, ±9x3 mm in
flower, 15-18 X 7-9 mm in fruit. Corolla very pale blue.
160
Bothalia 32,2 (2002)
darker at edges of lobes, with reddish purple marks at
junction of lobes or lobes with white along mid-area, blue
in outer areas; tube funnel-shaped, 5-7 mm long; lobes 5
or 6, broadly ovate, ± 5 x 1.5 mm, with abrupt, acuminate
apices, usually twisted. Fruit of four nutlets; nutlets grey-
ish marble brown, compressed ovoid, ± 4.0— 4.5 mm long,
external face ± convex, shiny, internal face 3-angled,
rugose. Flowering time: January to December.
Although different infraspecific taxa of T zeylanicum
have been described, the species does not warrant subdi-
vision in southern Africa. In Australia the species does
show differences and three forms are recognizable
(Randall 1993).
Distinguishing characters: stems, inflorescence axes
and petioles patently hairy; leaves discolorous because
upper and lower leaf blade surfaces differ in indumentum;
calyx lobes 15-18 x 7-9 mm in fruit; apices of corolla
lobes acuminate; nutlets smooth, shiny and grey-marbled.
Distribution: Namibia (Caprivi), eastern parts of the
Northern Province, Mpumalanga, Swaziland and north-
ern KwaZulu-Natal (Figure 8); also known from various
regions in eastern Africa, the Comoros, Madagascar, the
Mascarenes, India, Sri Lanka, Malaya, Java, the
Philippines and Australia (Verdcourt 1991), having
reached at least some of these areas presumably as a weed
of cultivation. Habitat: grassland, woodland, in dry river
beds, common weed in old and new cultivated land, pio-
neer in disturbed areas, along roadsides; well-drained
sandy, gravelly turf, poorly drained loam or in rocky well-
drained soil; full sun. Illustrations: Hooker: t. 4820
(1854); Ivens: 83 (1967); Moriarty: t. 103 (1975);
Verdcourt; 93 (1991); Bromilow: 157 (1995).
2. Trichodesma africanum (L.) Lehm., Plantae e
familia Asperifoliarum nuciferae: 195 (1818) non R.Br.;
C.H. Wright: 11 (1904); F.W. Andrews: 88 (1956); Riedl:
220 (1967); Friedr.-Holzh.: 3(1967); H.Heine: 323(1963);
Kazmi: 519 (1971); Qaiser: 31 (1979). Type; ‘habitat in
Aethiopia’, LINN 188.4 (LINN, syn.-PRE, microfiche!).
Borago africana L.: 138 (1753). Pollichia africana (L.) Medik.:
247 (1783). Borraginoides acideata (L.) Moench: 515 (1794).
Borraginella africana (L.) Kuntze: 435 (1891).
Borago verrucosa Forssk.: 41 (1775). Type: Egypt. Cairo. Forsskd!
288 (C. holo.).
Borraginoides africana (L.) Hiern: 721 (1898). Type: Angola.
Mossamedes. Welwitsch 5303 (PRE. syn.!).
Erect, annual or biennial herb, up to 1 m high, harsh-
ly scabrid. Stems branched, covered with stiff, tubercle-
based setae; setae up to 2 mm long. Leaves opposite, ses-
sile or petiolate; blade ovate to narrowly ovate, rarely
oblong, 50-105(-l20) x 5-30(-80) mm, base auriculate,
apex acute, upper surface with setae of different sizes;
setae on lower surface stiff, with tubercle-like multicel-
lular bases, mainly on veins; petiole up to 40 mm long.
Inflorescences terminal at ends of stems and also termi-
nal on lateral branches; Bowers with pedicels up to 15
mm long in fruit. Calyx: lobes narrowly ovate, ±6x2
mm in flower, ±12x6 mm in fruit, ba.se truncate to sub-
cordate, apex acuminate, midrib and margins densely
hairy with long, stiff setae, setae interspersed with small,
fine, simple, unbranched hairs. Corolla pale pink when
12 14 16 18 20 22 24 26 28 30 32
FIGURE 8. — Distribution of Trichodesma zeylanicum. A; and T.
africanum, ■. in southern Africa.
young, changing to blue, fading white with blue margin,
throat yellow with purple spots; tube funnel-shaped, 3-4
mm long; lobes ovate-triangular, ± 2. 5-3.0 mm long,
apices long-acuminate, often twisted. Fruit of 4 nutlets;
nutlets ovoid-cup-shaped, up to 4 mm long, with
glochidia along serrate, raised margins and single ones
along midrib, base of cup with thick, short setae and ver-
rucose epidermal protuberances. Flowering time:
January to December.
Trichodesma africanum is a rather plastic species with
small leaves and much-branched inflorescences under
arid climatic conditions, compared to large leaves and
less branched inflorescences when growing in the shade
of boulders or during times of above-average rainfall.
The different varieties recognized by Brand (1921) and
El-Hadidy & Boulos (2000) are not upheld here, but this
needs further attention.
Distinguishing characters: spinously, patently hairy;
leaves variable in size; calyx lobes 12 x 6 mm in fruit;
apices of corolla lobes long and acuminate; nutlets
ovoid-cup-shaped with distinct serrate, raised margins
and a concave, glochidiate inner face. Distribution:
Namibia, Northern Cape and Western Cape (Eigure 8);
also known from North and West Africa, Afghanistan,
Iran and Pakistan. Habitat: disturbed places, road verges,
granite or sandstone hills, mountain slopes, among rocks,
rocky ridges, dry, sandy river beds, calcareous sandy
flats and river banks. Utilization: grazed (Henrici 3393).
Illustrations: Heine: 323 (1963); Qaiser: 32 (1979).
3. Trichodesma angustifolium Harv., Thesaurus
capensis 1: 26 (1859); C.H.Wright: 11 (1904); Baker &
C.H.Wright: 47 (.1905); Brand; 27 (1921); Friedr.-
Holzh.: 3 (1967); J.H.Ross: 297 (1972); Brummitt: 852
(1985); Brummitt; 96 (1990); E.S. Martins & Brummitt:
64 (1993); Relief & P.P.J. Herman: 353 (1997); Pooley:
468 (1998). Syntypes: Magaliesberg, Burke 60, 313 (K,
iso.!); Zeyher 1250 (BM, iso.!); Zeyher 1251 (K, iso.!);
Rhenoster River, Free State, Sanderson 157 (K, iso.!).
Boraginella angustifolium (Harv.) Kuntze: 436 (1891).
Bothalia 32,2 (2002)
161
Perennial herb or subshrub, up to 1.2 m high, with a
woody rootstock, appressed hairy throughout, with setae
orientated parallel to midrib of leaves. Stems erect or
decumbent-ascending; setae variable in size or of ± equal
length; epidermis often flaking off in older plants.
Leaves shortly petiolate or blades decurrent; blade linear
to narrowly elliptic to linear-elliptic, 20-50 x ( 1 .5-)
3.0-6.0(-15.0) mm, base narrowly cuneate, apex acute,
surfaces with setae dense or with spaces in between; petiole
up to 3 mm long. Inflorescences terminal at ends of main
stems and also terminal on lateral branches, lowermost
flowers occasionally solitary in leaf axils. Calyx setu-
lose, winged, ±11x4 mm in flower, ± 25 x 20 mm in
fruit, base cordate, apex long-acute. Corolla pinkish in
young stage, changing to blue or mauve, fading white;
tube cylindric to slightly campanulate, ± 9-15 mm long;
lobes triangular, ± 5.5 x 4 mm, long-acuminate, often
twisted. Fruit of 4 nutlets; nutlets glochidiate, ± 5-6 mm
long; glochidia usually coalescing at base to form dis-
tinct rim.
Distinguishing characters: resprouting perennial,
stems, inflorescences and calyces appressed hairy; leaf
blades linear to narrowly elliptic; calyx winged, ± 25 x
20 mm in fruit; nutlets with several-barbed glochidia;
glochidia usually coalescing at their bases to form a dis-
tinct rim.
Key to subspecies
Indumentum setulose, greyish green; setae well spaced; usually
on red. sandy loam; widespread subsp. angiistifolium
Indumentum tomentose, silvery grey to greyish green; setae
densely packed; usually on lime-rich soils; only known
from northern Namibia subsp. argenteum
subsp. angustifolium
Trichodesma lanceolatwn Schinz: 269 (1888). Type: Namibia.
Scap River. Schinz 758 (COI!, GRA!, K!. NUI. syn.).
Flowering time: October to May. Distinguishing char-
acters: indumentum setulose, setae with distinct spaces in
between, green; usually on red, sandy loam. Distribution:
Namibia, Botswana, Northern Province, North-West,
Gauteng, Free State, KwaZulu-Natal, Northern Cape
(Figure 9); also in Mozambique. Habitat: open grassland,
open shrubland, mixed shrub savanna, mesophytic closed
bush, abandoned lands, disturbed places, mixed ruderal
vegetation; red sandy loam, stony soils; plants reach a con-
siderable age and large rootstocks may be more than a cen-
tury old. Common name: blue-bells, according to Smith
(1966); 'motlhagapele' (Moss BMP735). Medicinal use:
dried ground root powder mixed with cold water is used
for diarrhoea (Moss BMP735). Illustrations: Harvey: t. 40
(1859); Pooley: 469 (1998).
subsp. argenteum Retief & A.E.van \Vyk. subsp.
nov., subsp. angustifolio affinis sed folds dense pilosis (non
interv'allis conspicuis inter bases setarum), plerumque in
solo calcareo, non luto arenoso rubro habitat.
TYPE. — Namibia, 1916 (Gobaub): Etosha National
Park, between Gemsbokvlakte and Olifantsbad, (-AA),
Retief 1404.01 (PRE, holo.; WIND).
FIGURE 9. — Distribution of Trichodesma angustifolium subsp. angus-
tifolium. ■; and T. angustifolium subsp. argenteum. A, in
southern Africa.
The prominent silver-grey to greyish green indumen-
tum is reflected in the specific epithet: ‘argenteum’ = sil-
ver. The difference in the density of the setae on the leaf
surfaces, distinguishes the two subspecies. This distinc-
tion is further supported by habitat differences.
Flowering time: November to April. Distinguishing
characters: indumentum tomentose, silvery grey to grey-
ish green; setae densely packed; usually on lime soils.
Distribution: northern part of Namibia (Eigure 9).
Habitat: grassland, savanna, margins of pans, road verges;
usually in grey, lime-rich soils.
4. Trichodesma physaloides (Fenzl) A. DC.,
Prodromus 10: 173 (1846); Giirke: 99 (1897);
C.H. Wright: 11 ( 1904); Baker & C.H. Wright: 46(1905);
Brand: 22 (1921); Pole Evans: t. 351 (1929); Eetty: 280
( 1962); Taton: 39 ( 1971); Agnew: 521 (1974); Brummitt:
439 (1982); Brummitt: 96 (1990); Verde.: 100 (1991);
E.S. Martins & Brummitt: 65 (1993); Germish. in
A. Fabian & Germish.: 338 (1997). Retief & P.P.J. Her-
man: 353 (1997); A.E.van Wyk & C.E.Malan: 46 (1997);
Pooley: 178 (1998). Type: Sudan, Eazokl, Kotschy 577
(BM, K!, iso.).
Friedrichsthalia physaloides Fenzl: 54 (1839). Boraginella physa-
loides {Fenz\) Kuntze: 435 (1891).
Trichodesma droogmansianum De Wild. & T.Durand: 69 (1900). T.
droogmansianum De Wild. & T.Durand var. glahrescens (Giirke)
Brand: 24 ( 1921 ). Type: Zaire, Lualaba, Descamps s.n. (BR, holo.).
T. glahrescens Giirke: 389 (1901). Type: Tanzania, Kinga-Berge.
Ussangu, Goetze 1267 (BR, E, iso.-K, photo.!).
T. ringoetii De Wild.: 100 (1914). Type: Zaire, Katanga, Nieuw-
dorp. Ringoet 6 (BR, holo.).
Perennial herb with 1 to several annual stems from a
woody rootstock, up to 0.5 m tall. Stems ± glabrous, with
scattered setae; young stems fleshy, pinkish. Leaves ses-
sile, bluish green; blade broadly ovate to narrowly ovate,
30-50(-75) X 12-16(-32) mm, ba.se cuneate to rounded,
apex acute or obtuse, upper surface clothed with short.
162
Bothalia 32.2 (2002)
FIGURE 10. — Trichodesma physcdoides, Gennishuizen 2156 (PRE). A, setae on upper surface of leaf, x 23; B. flowering stem, x I ; C, rootstock
with young shoots, x 1 ; D. longitudinal section of corolla, x 1.5; E. nutlet, x 3; F, habit reduced, plant in Pretoria National Botanical
Garden. Artist: Gillian Condy.
Stiff setae; setae all pointing ± in same direction, multicel-
lular base discoid, 3-5-layered; lower surface with scat-
tered setae on midrib and veins only. Inflorescences termi-
nal, cymose panicles, cymules .scorpioid; pedicels up to 30
mm long, drooping. C«/yx brownish puiple; lobes 13-15
X 5-7 mm in flower, up to 30 x 18 mm in fruit. Corolla
white with brown gibbosities at sinuses between lobes,
apices of lobes with a light brown rim; tube funnel-shaped,
hairy inside at level where anthers arise; lobes broadly
ovate, apices cuspidate, glabrous or den.sely pilose down
middle of lobes on adaxial side. Frail one discoid nutlet;
nutlet with a prominent rim, den.sely hairy, with style and
stigma lateral due to abortion of three other nutlets.
Flowering;; time: August to November. Figure 10.
Distingaishing characters: mullistemmed greyish
green suffrutex; stems glabrous or with setae with promi-
nent discoid, multicellular bases; flowers drooping;
calyx brownish purple; corolla white with a light brown
rim at apices of lobes; fruit a hairy, discoid nutlet.
Disfrihation: Northern Province. North-West, Gauteng,
Mpumalanga, Swaziland, KwaZulu-Natal (Figure II);
also known from southern Sudan, western Ethiopia,
Kenya, Uganda, Burundi, Tanzania, Zaire, Zambia,
Malawi, Zimbabwe and Mozambique. Habitat: grass-
land, woodland, open mixed bushveld, hill slopes, dis-
turbed areas, roadsides, waste places; sandy loam, clay,
loam or rocky soils, gravel; large plants may have root-
stocks well over a century old. Common name: chocolate
bells. Illustrations: Pole Evans: t. 351 (1929); Letty: t.
139, 3 (1962); Moriarty: t. 30 (1975); Brummitt: 431
(1982); Onderstall: 167 (1984); Brummitt: 97 (1990);
Eabian & Gennishuizen: 339 ( 1997); Van Wyk & Malan:
47 (1997); Pooley: 179 (1998).
Bothalia 32.2 (2002)
163
12 14 16 18 20 22 24 26 28 30 32
FIGURE 11. — Distribution of Trichodesma physaloides. ■: and T.
ambacense subsp. hockii. A. in southern Africa.
5. Trichodesma ambacense Welw. in Annaes do
Conselho ultramarino 1: 589 (1859) as 'ambacensis';
Brummitt: 442 ( 1982); Brummitt: 98 ( 1990); Verde.; 101
(1991). Boraginella ambacensis (Welw.) Kuntze; 435
(1891). T. angolense Brand; 26 (1921), nom. superfl.
illegit. Type; as for T. ambacense above. Type: Angola,
Welwirsch 5450 (LlSC, holo.; BM, BR, COl!, K!). "
Currently T. ambacense is divided into two subspecies
(Brummitt 1982). T. ambacense subsp. ambacense is
endemic to Angola. It differs from subsp. hockii in the
leaves which are obovate to oblanceolate (not ovate to
elliptic) and calyx lobes which are 3-5 mm long in
flower (not 5-8(-9) mm long).
subsp. hockii (De Wild.) Brummitt in Kew Bulletin
37: 443 (1982); Roessler: 39 (1984); Brummitt: 98
( 1990); Verde.: 101 ( 1991); E.S. Martins & Brummitt: 67
tl993);E.S.Martins: 5 (1993); Bolnick: 32 ( 1995). Type;
Zaire, Shaba, Lubumbashi [Elisabethville], Hock (BR,
holo.,-K, photo.!).
Trichodesma hockii De Wild.: 546 ( 1913).
T. ledennannii Vaupel: 529 (1912). Type: Cameroon, near Laro.
Ledennann 3080 (B. holo.t).
T. tinctoriiim Brand: 393 (1914). Type: Zaire. Shaba. Lukafu.
Verdick 104 (BR. holo.).
T. verdickii Brand: 329 ( 1914). Type: Zaire. Shaba, Lukafu, Verdick
740 (BR. holo.).
Perennial herb with annual stems from a woody root-
stock, up to 0.5 m high. Stems hairy or setose. Leaves
sessile or occasionally shortly petiolate; blade ovate to
elliptic, 50-65 x 20-27 mm, base cuneate, apex acute;
petiole up to 1.5 mm long. Inflorescences terminal at
ends of stems, cymose panicles, cymules scorpioid;
flowers slightly pendent; pedicels up to 25 mm long in
fruit. Calyx ± 14 x 8 in flower, ± 22 x 15 mm in fruit.
Corolla deep blue, pale lavender to white with raised
amber-brown gibbosities at sinuses of petal lobes; tube
campanulate, lobes reflexed, apices acuminate. Fruit:
one, densely silken-haired nutlet; hairs usually hooked.
Flowering time: June to September.
Distinguishing characters: multistemmed suffrutex;
stems hairy; leaf blade elliptic; setae with prominent dis-
coid, multicellular bases, usually hooked at apices; flow-
ers slightly pendent; corolla deep blue, lobes reflexed;
fruit a silken-haired nutlet. Distribution: Namibia,
Botswana (Figure 11); also known from Sudan, Nigeria,
Cameroon, Uganda, Kenya, Tanzania, Zambia, Malawi,
Mozambique and Zimbabwe. Habitat: grassland, wood-
land, abandoned fields, roadsides; sandy soil. Common
name: bells of St Mary’s. Illustrations: Moriarty; t. 103
(1975); Martins & Brummitt: 69 (1993); Bolnick: t. 15
(1995).
SPECIMENS EXAMINED (southern Africa only)
Numbers in brackets signify the identity of the speci-
mens: (1) Trichodesma zeylanicum; (2) T. africanunr,
(3a) T. angustifolium subsp. atigustifolium: (3b) T.
angustifolium subsp. argenteum; (4) T. physaloides', (5)
T. ambacense subsp. hockii.
Acocks 1541 (3a) PRE; 1619. 18133 (3a) K, PRE; 2026, 2533 (2) BOL,
K. PRE; 2578 (3a) BOL, PRE; 2587 (3a) K. Acocks & Hafstrdm 1314.
1350 (2) PRE; 1315 (4) PRE. Adamson D147 (2) PRE. Anderson N18
(3a) PRE.
Balkwill 1499 ( 1 ) NU. PRE. Balkwill & Cadman 3501 (4) E, PRE.
Balsinhas 3014 (3a) K. PRE. Bal.sinhas & Kersberg 2006 (2) PRE,
WIND. Barber s.n. (3a) K; PRE13748 (3a) PRE. Barker 102 (4) J; 791
(3a) PRE. Barnard SAM36092 (2) SAM. Barrett 132 (1) K, PRE.
Bengis 463 (2). Bolus 641 (2) BM, BOL, K; 5713 (3a) K; 6443 (3a)
BOL. PRE; 9706 (4) BOL; 10399 (2) BOL. Bond 842 (2) NBG. Boshoff
& Mason 2541 (3a) PRE. Bo.ss 18. A80. TRV35550 (3a) PRE;
TRV36168 (2) PRE. Botha & Ubbink 1733 (3a) PRE. Boucher 5147 (2)
NBG. Bradfield 583 (2) K, PRE. Bremekamp TRV27519 (1) PRE.
Breyer PRE59361. TRV17580 (1) PRE; PRE59405, TRV20582 (3b)
PRE. Brown & Kolberg 277 (3b) WIND. Biyant J118 (2) PRE; 894B (2)
K. Buchanan sub Wood 7032 ( 4) PRE. Buitendag 626 (4) K, PRE.
Burchell 1264 (2) K, PRE. Burger & Louw 297 (2) NBG, PRE.
Burgovne 408 (4) PRU. Burke 60. 313 (3a) K; PRE13749 (4) PRE. Burtt
Davy 2051 (4) PRE; 7048 (3a) K; 11009, 11278, 14393 (3a) NBG.
Catell 314 (2) NBG. Chennells 151 (3a) BOL. PRE. Clarke 22/57 (I)
PRE; 1318 (4) PRE. Codd 627 (4) PRE; 2084, 6645 (3a) K, PRE. Cole
1388 (3a) PRE. Comins 663 (2) K, PRE; 1801 (2) PRE. Compton 2447.
11530. 20616 (2) NBG; 27100 (4) K. PRE; 28934 (I) NBG. PRE.
Craven 14. 184. 216. 1556. 1939. 3136 (2) WIND. Crook M37, 737 (4)
PRE. Crosby 863 (2) PRE. Cross 228 (4) PRE.
Davidse 6285 (2) PRE. Davies, Thompson & Miller 48, 104 (2) PRE,
WIND. De Kruifl231 (4) PRE. De Lange UN1N6718 (4) PRE, UNIN.
De Sousa 559 (4) PRE. De Winter 419 (5) BM, K, PRE; 24 73 ( 3a) K,
PRE, WIND. De Winter & Giess 6158 (2) K. PRE, WIND. De Winter
& LeLstner 5685 (2) K, PRE, WIND. Dean 325 (2) BOL, PRE. Dimer
415 (3a) K, SAM; 925 (3b) NBG; 1213, 1273 ( 2) SAM; 3503 (3a) K.
PRE; 4296 (3a) BM. SAM; 4799 (2) K; 8423 (3a) BM, K. Dlamini
PRE40761 (4) PRE. Drege s.n. (2) K. Driver 52 (2) PRE. Du Ple.ssis
814. 1046 (4) PRE. PRU; 3100 (4) K, PRE.
Edwards PRE40742 (4) PRE. Elan-Puttick 292 (4) PRE. Esterhuvsen
2712 (2) NBG. PRE; 4488 (2) BOL. Evrard 9034, 9243 (2) PRE. '
Fourie 2804 (4) PRE. Francois 45 (4) E. Fries, Norlindh & Weimarck
1966 (4) PRE. Fuller PRE10596 (4) BM.
Galpin 502M, 503M (3a) PRE; 504M, 6994 (4) PRE; 9223 (1) PRE;
9288 (4) K, PRE; 11380 ( 1 ) K, PRE; 13700 ( 3a) K. PRE; 14111 (2)
BOL, K. PRE; 15097 (5) E. Galpin s.n. (4) K. Galpin & Pearson 1561
(2) BOL; 7506 ( 2) K, PRE, SAM. Germishuizen 2156 (4) K, PRE;
3086 (4) PRE; 4575, 5377, 5431 (2) PRE. Gerstner 3435 (4) NH; 4101
(3a) NH; 5119 (4) K. PRE; 5133 (3a) PRE; 6616 (1) PRE. Gibbs
164
Bothalia 32,2 (2002)
Russell, Robinson & Herman 311 (2) PRE. Giess 3628 (2) WIND. 8033
(2) PRE, WIND; 8311 (3a) PRE, WIND; 9488 (5) WIND; 10700 (3a)
WIND; 1331 (5) PRE, WIND. Giess & Loutit 14101 (3b) K, PRE,
WIND. Giess & Miiller 11980 (2) K, PRE. WIND. Giess & Van Vuuren
947 (3a) K, PRE, WIND. Giess & Wiss 3295 (2) WIND. Giess. Volk &
Bleissner 6276 (2) WIND; 6512 (3b) WIND. Gillet 2599 (3a) K; 4625
(4) K. Goldblatt 1986 (2) PRE. WIND; 2349A (2) NBG, PRE; 7003 (2)
PRE. Goosens PRE40757 (4) PRE. Greenway 8113 (4) PRE. Greater
21604, 21802 (2) PRE. Gubb 32.92 (3a) PRU.
Hall 57, 74 (\) NBG; 400, 798 (2) NBG. Harris 127 (4) PRU.
Hanekom 1498 (3a) K. PRE; 1619 (3a) K; 2151 (2) K, PRE. Hansen
3277 (3a) PRE. Hardy 1950 (2) K, WIND; 2470 (2) K. PRE; 5656 (1)
K. PRE. Hardy & Bayliss 1239 (3a) K, PRE. Hardy & De Winter 1393
(2) PRE. Hardy & Venter 4565 (2) K, PRE, WIND. Henrici 74 (3a)
PRE; 3393 (2) PRE. Heymans 13 (4) J. Hill PRE40760 ( 3a) PRE.
Hilliard 4731 ( 1 ) NU. Hillary & Robertson 601 (3a) PRE. Hines 400
(3b) WIND. Hitchins 319 ( 1 ) PRE. Hojfmann 1233 (2) NBG. Holt 8 (4)
NH, PRE. Homann, Benseler & Mittendorf 22 (2) WIND. Howlett
PRE40752 (4) PRE. Hugo 505, 2533 (2) K, NBG, PRE. Hutchinson
2987 (3a) BM. BOL, K, PRE. Hutchinson & Gillett 3590 (5) COI.
Ihlenfeldt, De Winter & Hardy 3226 (2) PRE. Immelman 569 (3a) PRE.
WIND; 579 (2) PRE.
N. Jacobsen 1813 (4) PRE. W.J. Jacobsen 1059 (3a) PRE. JBG6154 (4)
PRE. Jejfers 414 (3a) PRE. Jenkins 7432, 9130, TRV9130 (4) PRE;
7538 (3a) PRE; TRV8185 ( 1 ) PRE. Jensen 49, 70, 195, 241, 281, 1395
(2) WIND. Joffe 332 (4) PRE. Joubert 100/1447 (3b) WIND. Junod
548 iX) K, PRE; 604 (4) K; 5022 ( 1 ) E; 604 (4) K, PRE.
Karsten PRE31340, PRE31341 (4) PRE. Kers 7062, W1ND26641 (2)
WIND. Killian 20 (1) K, PRE. Kinges 1851 (4) PRE; 2305, 2399 (2)
PRE. WIND. Kluge 113 (4) PRE, PRU. Koch A15 (2) PRE. Kok 265
PRE, PRU; 1046 (4) PRU. Kraeusel & Wiss 1913 (2) WIND. Krynauw
70 (3a) PRE; 1283 (4) PRE. Kubirske 186 (2) K. Kubirske, Strohbach
& Swart 186 (2) WIND.
La Croix 3339, 4874 (4) PRE. Laidler 662 (2) NBG, PRE. Lang
TRV32136 (1 ) PRE. Lavranos 11019 (2) PRE. Lavranos & Pheleniann
20148 (3a) WIND. Lea 43 (4) PRE. Leendertz 267. 360, 824 (4) PRE.
Le Roux 13 (4) PRE, 483 (3b) PRE, WIND; 3064 (2) NBG. Leach &
Bayliss 13067 (2) K, PRE, WIND. Leeman PRE40744 (4) PRE.
Leendertz 267, 300, 824, 7594 (4) PRE; 713A, 8519, TRV11437 (3a)
PRE. Leistner 1240 (3a) K, PRE; 2338 (2) K, PRE. Leistner & Joynt
2846 (2) PRE. Leistner, Oliver, Steenkamp & Vorster 139 (2) K, PRE.
Leuenberger, Raus & Seiners 3266 (2) WIND. Levvns 1742 (2) BOL.
Liebenberg 5008 (2) PRE, WIND; 5103 (3a) WIND; 5158 (2) K, PRE,
WIND; 5703 (3a) PRE. Lloyd 35, 36 (2) PRE. Louw 2029 (3a) PRE.
MacCallum 541, 1589 (4) PRE. MacDonald 97, 357, 490, 568 (2) BM;
383 (3a) BM. Maguire 365 (2) NBG. Malan 9(1) PRE. Mathibe 64 (4)
PRU. Matthews 230 (2) PRU. Methuen 166 (4) PRE. Miller 2489 (4)
PRE. Marloth 780 (3a) PRE; 1188 (2) NBG; 1411, 2034 (2) PRE;
10162 (4) PRE; 12380 (2) NBG, PRE. McMurtry 4100, 10773, 15325,
19179, 23918 (4) J. Meebold SAM51370 (1) SAM. Merxmiiller 739,
1669 (2) PRE, WIND; 1325 (3b) K, PRE. Merxmiiller & Giess 1325
(3b) WIND; 1669 (2) WIND; 2827, 28107 (2) PRE, WIND. Mogg
8176 (3a) PRE, SAM; 8363, PRE11704, SAM44701 (3a) PRE; 9450 (4)
SAM; 15325, PRE9449, PRE10773 (4) PRE; 19179, 23918 (4) J.
Moore 44 (3a) PRE. Moran PRE40756 ( 3a) PRE. Morris &
Engelbrecht 1135, 1155 (3a) K, PRE. Moss BMP735 ( 3a) PRE; 4736,
22227 (4) J; 8855 (4) BM, J. Moss & Jacobsen K218 (2) PRE. Muller
29 (3a) PRU, WIND; 124 (2) PRE, WIND; 781 (3a) PRE; 1290 (3a)
PRE, WIND. Miiller & Umtit 1198 (2) WIND. Murray 614 (3a) PRE;
641 (4) PRE. Myre 27 ( 1 ) PRE.
Nation 288 (3a) K. Nelson 236 (3a) PRE. Netshiungani 915 (1) PRE.
Nicholas 2576 (2) PRE, WIND,
Obenneyer 711 (\) PRE; NH27321 (4) NH; TRV34735 (4) BM, PRE.
Oliver & .Steenkamp 6291 (2) K. PRE, WIND. Oliver, Tolken & Venter'
89 (2) K, NBG, PRE. Onder.stall 443, 1248 (4) PRE.
Pager P35 (2) WIND. Papendorf 375 ( I ) PRE. Parry PRE8016 ( 1 )
PRE. Patterson 21 (1) PRE. Pearson 3665 (3a) BOL, K, PRE, SAM;
3674 (3a) K, PRE; 4961 (2) K, NBG, PRE; 4995, 6005 (2) K; 6106 (2)
BM, K; 8551, 9123 (2) BOL, K; 9110, 9531 (3a) BOL, K, Peeters,
Gericke & Burelli 163 (3a) PRE. Pegler 979 (3a) BOL, K, PRE, SAM.
Peyper 1333 (3a) PRE. E.Phillips 1117 (4) J; 3275 (4) J, PRE.
J. Phillips 1616 (4) PRE. Phillips & Liebenberg 916 (3a) PRE. Phipps
778 (4) PRE. Pienaar 1089 (2) BOL, PRE. Pienaar & Vahnneijer 478
(1) PRE. Pillans 5074 (2) BOL, K; 5856 (2) BOL. Plowes 7046 (3a)
PRE. Pole-Evans 2827 (5) PRE; s.n. (4) K; PRE40746 (4) PRE;
PRE12870, PRE13241, PRE40765 (3a) PRE. Pont 512 (3a) PRE. Potts
BLFU2676 (3a) BLEU. Pooley 435 (3a) NU; 524 (1) NU. Prosser
P1071 (4) K, PRE; J029438 (4) J; JBG1204 (4) PRE.
Range 1030 (3a) SAM. Rehmann 4522 (3a) BM, BOL; 4759 (4) BM,
BOL. Repton 283 (4) PRE; 3410 (3a) K, PRE; UNIN4540 (4) PRE.
Relief 1404.01 (3b) PRE, WIND; 2223, 2224 (4) PRE. Roberts
PRE15862 (4) PRE. Rodin 2862 (3a) BOL, K; 3657 (3a) BOL, K, PRE.
Rogers 5039, 22270 (3a) PRE; 8290 (4) BOL; 13453 (1) PRE; 15322
(2) K; 18123 ( 1 ) K; 21405 (4) SAM; 22072, TRV26665 (4) PRE. Rdsch
& Le Roux 109, 179 (2) PRU, WIND. Riisch 45 (2) WIND.
SAGP/SAAB 1/76 (4) K, PRE, PRU. Sanderson 157 (3a) K. Scheepers
549 (1) K, PRE, PRU, UNIN; 1166 (4) BM, K, PRE, PRU; 1616 (3a)
PRE. Schenck 49 (3a) PRE. Schinz 758 (3a) COI, GRA, K, NU.
Schlechter 3601 (4) BM, E, J, PRE; 3677 (3a) BM, BOL, K, PRE; 5565
(3a) PRE; 10871 (2) BM, BOL, COI, K, PRE. Schlieben 8996 (2) BM,
K. PRE. Schmidt 295 (2) PRE, WIND. Schoenfelder 227 (3a) PRE.
Schulze 8 (1) PRU. Schweickerdt 1102 (3a) PRE. Schwerdtfeger 4181
(3a) WIND; 4286 (2) WIND. Seely 2009 (2) WIND. Seely & Robinson
306 (2) WIND. Seely & Ward 8, 9 (2) WIND. Seydel 314 (2) K; 1845
(3a) WIND; 1847, 2074 (3a) K; 2076 (3b) WIND; 1049, 2470, 4049
(3a) K, PRE, WIND; 2971, 4334 (2) K. Shearing 115, 620, 1083 (2)
PRE. Sim 20638 ( 1 ) PRE. Smit 1759 (4) PRU. Smith 153, 875, 1082A,
1215, 2131, 3250 (3a) PRE; 821, 837 (4) PRE. Smook 7695, 8691 (2)
PRE. Smook & Harding 713 (2) K, PRE. Smuts & Gillett 3290 (4) PRE.
Snyman C (1) PRE. Stalmans 158, 1408 (4) PRE; 1076 (1) PRE; 1996
(1 ) J. Steiner 624 (2) NBG. Stewart TRV8883 (4) PRE; 8966 (3a) PRE;
8970 ( 1 ) PRE. Stohr 219 (4) BOL. Story 758 (3a) PRE; 1361 (4) PRE;
5745 (2) K, PRE. Strey 2020 (2) BOL, PRE; 2153 (3a) K, PRE; 6588
(1) K, NU, PRE; 7890 (1) PRE. Strohbach 29 (2) WIND. Sutton 228
(3a) PRE.
Taylor 3396 ( 1 ) NBG; 3462B, 11537 (2) NBG. Theiler PRE59972 (3a)
PRE. Theron 1517 (4) PRE, PRU; 3899 (2) PRU. Thode 4764, 4766
(3a) PRE; A1453 (3a) K, PRE. Thorncroft TRV4550 (4) PRE; NH11609
(4) NH. Thorne SAM51582 (2) SAM. Tinley 1108 (3b) PRE, WIND.
Tolken & Hardy 777 (3a) K, PRE, WIND. Trapnell 507 (4) K. Tuck 813
(3a) BOL, SAM; 2275 (3a) BM.
Ubbink 744, 1141 (3a) PRE. Ueckennann 7295 (2) PRE.
Van Breda 4009, 4331, 4416 (2) PRE. Van Dam PRE59662. TRV25002
(1) PRE. Van der Schijff 542 (1) KNP, PRE; 5752 (1) K, KNP, PRE;
5952 (1) PRU; 8195 (2) PRE, PRU. Van Hoepen 1952 (2) BOL. Van
Jaarsveld 1496, 2529, 8413, 11921 (2) NBG; 8825 (2) NBG, PRE. Van
Rooyen 2017 (4) K, PRE, PRU; 2268 (2) PRU. Van Wyk 707 (2) PRE;
734 (3a) PRE; 1748 (4) PRE; 5505 (1) PRE, PRU; 6582 (2) PRE, PRU;
8578A (2) PRE, PRU, WIND; 8595 (2) PRE, PRU; 8778 (2) PRE,
PRU, WIND; 12682 ( 1 ) PRU. Vinjevold CVll (2) WIND. Visser 210
(2) PRE, PRU. Vlok 1491 (2) PRE. Volk 867 (3a) PRE; 2465A (2) PRE.
Von Koenen 559, W1ND26672 (3a) WIND.
Wallich 1847, s.n. (4) K. Walter 981 (3b) WIND. Wanntrop 676 (3a)
PRE; 979 (2) K, PRE. Ward 2596 (1) PRE; 3216 (3a) K, NU, PRE;
12614 (1) NU. Wasserfall 1098 (2) K, PRE. Watmough 866 (2) PRE.
Way PRE31343 (4) PRE. Wells 2217 (3a) K, PRE. Wendt 25/4, 32. 85.
2616, C/20, sub Giess 13614, sub Giess 14762 (2) WIND. Werdermann
& Oberdieck 587 (2) PRE; 2557 (3b) K, NBG, PRE. Wild 3331 (1)
PRE. Williams 169 (4) E. Williamson 3958, 5562 (2) NBG. Wilnian
PRE40755 (3a) PRE. Wilms 943 (3a) BM, K. Wiss 1944 (2) WIND.
Wood 7002 (4) PRE. Worsdell s.n. (3a) K.
Young A190 (4) PRE.
Zeyher 98 (4) SAM; 1239 (2) BM, K, SAM; 1249 (4) K, PRE, SAM;
1250 (3a) BM; 7257 (3a) BM, K, SAM. Zietsman 1747 (2) WIND;
2700 (2) PRU. Zmn SAM63451 (2) SAM.
ACKNOWLEDGEMENTS
The assistance of Drs O.A. Leistner, J. Van Rooy,
Mmes G. Condy, E. du Plessis, J. Ready and Prof. W.
Bothalia 32.2(2002)
165
Trollope, in preparing the manuscript is gratefully
acknowledged. The directors, curators and staff of the
cited herbaria are thanked for allowing the authors to
study their material and for sending specimens on loan.
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Bothalia 32.2: 167-180(2002)
Six new species and one new subspecies of Erica (Ericaceae) from
Western Cape, South Africa
E.G.H. OLIVER* and I.M. OLIVER*
Keywords: Erica L., new species. South Africa, taxonomy. Western Cape
ABSTRACT
Two new species and one new subspecies belonging in %Ceramia of the genus Erica L. are described from the western
half of the province — E. cavartica known from a single collection growing in a cave in the Cederberg. E. amalophylla con-
fined to a shaded rocky overhang and known only from a single collection in the Porterville Mountains and E. cymosa
subsp. grandiflora occuiring in the mountains between Ceres and Worcester. Four new species are described from moun-
tains in the Little Karoo area — E. schelpeorum occuning widespread mostly in dry watercourses in arid vegetation at low
altitudes. E. lithophila found only in dry rock crevices at high altitude in the eastern Swartberg and Kammanassie
Mountains. E. umbratica confined to shaded rocky habitats and known only from a few. very localized small populations
at low altitude in Meiringspoort. and E. annalis restricted to the arid habitats on lower northern slopes of Kammanassie
Mountains. Each is provided with a detailed drawing and a distribution map.
SPECIES IN §CERAMIA
In §Ceramia of the genus Erica L. there are many
species which are associated with damp, shaded or wet
habitats. Most are soft, low shrublets, either erect and
compact or diffuse and sprawling, with long delicate
main branches and often with open-backed leaves. The
section contains a group of species (the E. planifolia
group) which are closely related due to the position of the
bract being on the main stem and leaf-like and not
recaulescent on the pedicel (bract = leaf, in our terminol-
ogy), and to a thickened midrib which is distinctly
enlarged towards the distal end of the lamina and/or
beyond the lamina apex. Within this group there were col-
lections of herbarium material made by Thomas Stokoe
and Elsie Esterhuysen that had not been placed with any
certainty within a species and remained unnamed.
The three new taxa have a very similar facies with
similar stems, leaves and bract but with differences noted
in the corolla size, the presence of anther appendages, the
ovary indumentum and the filament shape. They all
occur in shaded, damp habitats.
1. Erica cavartica E.G.H. Oliv. & I.M.Oliv.. sp.
nov., bractea non recaulescenti, bracteolis basalibus,
corolla ±9x4 mm, filamentis angustis rectis obspathu-
latis, antheris appendiculatis, ovario glabro, fructu irreg-
ulariter rugoso dignoscenda. Figura 1 .
TYPE. — Western Cape, 3219 (Wuppertal): Clanwil-
liam Dist., Cederberg, near Crystal Pool, 4000 ft [1 220
m] (-AC), March 1932, Stokoe 2600 (BOL, holo.); ibid.,
as Stokoe SAM50109 (SAM).
Shrublets soft, sparse, procumbent, single-stemmed
reseeders. Branches: few old main branches spreading
* Compton Herbarium. National Botanical In.stitute. Private Bag X7.
7735 Claremont. Cape Town.
MS. received: 2002-07-04.
up to 0.75 m long; numerous secondary branches 50-100
mm long, some bearing flowers; stems with mixture of
fine, short, simple hairs and slightly longer, gland-tipped
hairs, internodes 10-20 mm long on main branches, 3-10
mm long on secondary branches. Leaves 3-nate, spread-
ing, broadly oblong. 4. 0-6. 5 x 15 mm, completely open-
backed, flattened very thin, both surfaces and margins
with sparse, delicate, gland-tipped hairs with fewer on
abaxial surface and with a few, shorter, simple hairs
admixed, midrib becoming thickened towards apex and
beyond apex forming an apical, non-sticky, gland-tipped
seta, ± 0.7 mm long; petiole ± 0.7 mm long with sparse
gland-tipped hairs. Inflorescence: flowers 3-nate in 1-3
whorls raceme-like on main and secondary branches,
subapical to far removed from apex, continuing growth
apically; pedicel 10-20 mm long, covered with long and
short gland-tipped hairs and short, simple hairs; bract not
recaulescent, identical to a leaf on stem; bracteoles 2,
near base of pedicel, ± 4.0 x 1 . 1 mm, otherwise like bract
or leaf. Caly.x 4-partite, oblong-ovate, ± 3.5 x 1.5 mm,
green and leaf-like, idumentum as in bract and leaf.
Corolla 4-lobed, shortly and broadly tubular, ± 9.0 x 4.5
mm, sparsely and finely hairy with simple hairs, colour?;
lobes erect, ±1x2 mm, subacute, margins entire.
Stamens 8, free, just included; filamentr elongate nar-
row-linear with broad, ± elliptic ba.se, glabrous; anthers
dorsally attached near base, oblong, ± 1.2 x 0.5 mm in
adaxial view, bilobed, muticous; thecae erect adpressed,
elliptic-oblong, ± 1.2 x 0.4 mm in lateral view, smooth,
brown, pore ± Vz length of theca; pollen shed in tetrads.
Ovary 4-locular, 8-lobed, oblate-globose, ± 1.2 x 1.4
mm, emarginate, with distinct basal nectaries, glabrous;
ovules ± 90 per locule, spreading from placenta the full
length of columella; style exserted, ± 9.5 mm long,
glabrous; stigma simple truncate. Emit a dehiscent cap-
sule, ± 2.2 X 3 mm, valves splitting ± Vz way down and
opening 30°^0°, with large, irregular wrinkles/folds,
not hard and woody, septa mostly on valves. Seeds ellip-
soid, ± 0.4 X 0.3 mm, shiny, ± smooth, orange, testa cells
elongate, ± 75-125 x 25 pm, anticlinal walls thin jig-
sawed, inner periclinal walls with numerous small pits.
Elowering time: March. Figure 1 .
168
Bothalia 32,2 (2002)
1 mm
FIGURE I, — Erica cavarticu. A. Oowering branch, natural size; B, flower; C, stem; D, leaf; E, bract; F, sepal; G, stamen, full in abaxial view,
anther in adaxial and lateral views; H, gynoecium; I, ovary opened laterally; J, capsule; K, seed; L, testa cells. All drawn from the type,
Stokoe 2600. Scale bars: B-J. 2 mm; K, 1 mm; L, 50 pm.
Diagnostic features: bracteoles basal, leaf-like; corol-
la long, ± 9 X 4.5 mm; filaments narrow straight with
spoon-shaped ba.se; anthers muticous; ovary glabrous;
fruit with large, irregular wrinkles/folds.
The species is closest to the similar looking E. cymosa
E.Mey. ex Benth. For differences between this and the
other species in this group see Table 1.
Erica cavartica is known only from the type collec-
tion made by Thomas Stokoe in the Cederberg (Figure
2). He gave the habitat details to Neville Pillans in the
Bolus Herbarium as ‘in rock-crevices and on lloor of a
cave near C’rystal Pool’. The name is derived from this
habitat preference — cavarticiis = born or living in caves
(Fatin). Pillans noted at the time that the main branches
were up to I'/j ft |0.75 m] long and procumbent on large
plants but on small plants were erect.
2. Erica ainalophylla E.G.H.Oliv. & l.M.Oliv., sp.
nov., bractea non recaulescenti, bract^jolis basalibus,
corolla ± 3.5 x 3.5 mm, filamentis rectis vel ad apicem
sigmoideis, antheris appendiculatis, ovario lanato, valvis
fructus laevibus dignoscenda. Figura 3.
TYPF. — Western Cape, 3319 (Worcester); Piketberg
Div., rocky plateau on Twenty Four Rivers Mts above
Porterville, (-AA), 16 Dec. 1949, Esterinivsen 16612
(BOF, holo.; BM, K, NBG, NY, PRF).
Bothalia 32.2 (2002)
169
FIGURE 2. — Known distribution of Erica cavartica. A: E. amalo-
phylla, ■: E. cymosa subsp. cymosa. •; and E. cymosa subsp.
grandiflora. □.
Shrublets soft, semi-erect to procumbent. 70-12 x 6-36
mm. single-stemmed reseeders. Branches: few main
branches 100-300 mm long, thin, weak, secondary
branches few, ± 20-50 mm long; stems with scattered,
long, simple hairs and longer, gland-tipped hairs, intem-
odes (2-)5-10 mm long. Leaves 3-nate, spreading, elliptic,
4.5-7. 0 X 2 mm. acute, flat and very thin, almost com-
pletely open-backed, margins slightly recurved, midrib
thickening towards apex and tapering into external simple
or sometimes gland-tipped seta, sparsely covered on both
sides with long, thin, simple hairs and longer, gland-tipped
hairs, margins with a few of same hairs; petiole ±0.3 mm
long, glabrous. Inflorescence: flowers 3-nate in 1-3
whorls, raceme-like towards apex of main and secondary
branches; pedicel 6-15 mm long, with a few scattered,
long, gland-tipped hairs; bract not recaulescent, identical
to a leaf on stem; bracteoles 2, near base of pedicel. ± 3 x
0.9 mm, otherwise like a leaf. Calyx 4-partite; segments
adpressed to corolla, ovate-elliptic, ± 2.5 x 1 mm, other-
wise like a leaf. Corolla 4-lobed, cyathiform, ± 3.5 x 3.5
mm, thinly textured, white, sparsely hairy with thin, long
hairs; lobes erect, broadly rounded, deltoid, ± 0.8 x 1.8
mm, entire. Stamens 8, free, included; filaments linear-
oblong with slightly broader elliptic base, straight or with
slight sigmoid bend below anther, glabrous; anthers
bilobed, erect, dorsally attached ± V3 up the back, muti-
cous; thecae adpressed with slightly spreading apical por-
tion, yellow, smooth, pore ±^U-Bzof theca; pollen shed in
tetrads. Ovary 4-locular, broadly ovoid to broadly ellip-
soid. ± 0.8 xl mm, emarginate, covered with numerous
long, erect hairs, with nectaries around base; ovules ± 15
per locule, suberect from placenta the full length of col-
umella; style included or just beyond corolla mouth, ± 2
mm long, glabrous; stigma subsimple truncate. Fruit a
dehiscent capsule, valves splitting to base and opening up
to 45°, smooth, septa only on valve. Seeds ellipsoid, ± 0.4
X 0.2 mm. smooth, orange; testa cells elongate, 60-100 x
20-22 pm. inner anticlinal walls thin, evenly broadly jig-
sawed. inner periclinal walls with medium-sized pits.
Flowering time: December. Figure 3.
Diagnostic features: bracteoles basal, leaf-like; corol-
la short (± 3.5 X 3.5 mm); filaments broadish, straight or
with slight S bend subapically, with slight spoon-like
base; anthers muticous; ovary woolly; fruit smooth, not
wrinkled.
The plants within this alliance of three species are all
rather soft and delicate with the leaves thin, flat and
open-backed. The name of the species is derived from
this feature — amalophylla from amalos = soft, tender,
weak; phyllon = leaf (Greek). For species alliances and
differences see Table 1 .
Erica amalophylla is known only from the type col-
lection made by Elsie Esterhuysen in the mountains
southeast of Porterville which drain into the Twenty Eour
Rivers system (Eigure 2). She noted that the plants were
‘growing on shaded sandy floor of overhanging rock shel-
ter’— a similar habitat description to that of E. cavartica.
3. Erica cymosa E.Mey. ex Benth. in DC.,
Prodromus 7: 670 (1839). Type: ad Dutoitskloof prov.
Worcester, Drege 1185 (K!, lecto.; BM!, BOL!, NBG!,
P!, TCD!. W), lectotype selected here.
Shrublets soft, sparse, low, erect to spreading, single-
stemmed reseeders. Branches: main branches 50^00
mm long, occasional secondary branches 20-100 mm
long; stems delicate, thin with dense, short, gland-tipped
hairs to subglabrous, no infrafoliar ridges, intemodes
3-10 mm long. Leaves 3-nate, spreading, elliptic-ovate.
TABLE 1 . — Diagnostic characters in Erica cavartica, E. amalophylla and E. cymosa
170
Bothalia 32.2 (2002)
2 mm
FIGURE 3. — Erica amalophyUa. A, tlowering branch, natural .size; B. leaf, atiaxial surface; C, leal', abaxial surface; D, flower; E, sepal; F, sta-
men, abaxial view, and anther in lateral and adaxial views; G, gynoecium; H, ovary opened laterally; 1, capsule with one valve removed;
J, seed; K. testa cells. All drawn from the type, Esreriniysen 16612. Scale bars: B-E, I, 2 mm; F-Fl, J, I mm; K, 50 pm.
3,5-10.0 X 1-3 mm, open-backed and Hat, delicate,
adaxially covered with sparse, short, simple hairs and a
row of longer, gland-tipped hairs near margins, abaxially
with dense, very short, simple hairs and a few scattered,
longer, gland-tipped hairs, midrib thickened towards
apex and extended into a stout seta terminated by a small
gland and very short-haired, sparsely ciliate with gland-
tipped hairs; petiole adpressed to spreading, 0.4-1. 0 mm
long, glabrous to fine-haired. Inflorescence: flowers 3-
nate in 1-8 whorls, raceme-like towards ends of main
and secondary branches, continuing vegetative growth
on main branches but often terminating secondary
branches; pedicel 4—12 mm long, glabrous or with short,
gland-tipped hairs; bract not recaulescent, leaf-like;
bracteoles 2, from Vi way up pedicel to approximate to
calyx, seldom opposite, narrowly ovate to narrowly
elliptic, 0. 9-2.0 x 0.2-0. 4 mm, with indumentum and
midvein same as in leaf. Ccilyx 4-partite, ovate to broad-
ly ovate to broadly elliptic, 0.7-2. 5 x 0.4-1. 1 mm, indu-
mentum and structure like a leaf. Corolla 4-lobed,
cyathiform to broadly urceolate, 1.5-5. 5 x 2. 2-4.0 mm,
covered with sparse, short, simple hairs, white to very
pale pink or tinged with red; lobes 1-2 mm long, round-
ed to subacute, margins entire. Stamens 8, free, included;
filaments elongate, very narrowly ovate, 2. 0-2. 5 x 0.2
mm, with subapical S-bend, glabrous; anthers dorsal ly
attached near base, bilobed, narrowly ovate in adaxial
views, appendiculate; thecae erect, adpressed, ovate ±
0.7 x 0.4 mm in lateral view, slightly prognathous at
base, smooth, yellow, appendages at base of thecae or
partially decurrent along apex of filament, pendulous,
narrow, ± Vi as long as thecae, pore Vi-Vi length of
theca; pollen shed in tetrads. Ovary 4-locular, slightly
depressed obovoid, 0.6- 1.5 x 0.8- 1.9 mm, emarginate.
Bothalia 32.2(2002)
171
glabrous or covered in upper half with sparse, short, sim-
ple hairs, nectaries present around base; ovules 35-50
per locule, spreading from placenta almost full length of
columella; style manifest, 1.6^.0 mm long, glabrous;
stigma small, capitellate. Fruit a dehiscent capsule,
valves splitting open ± 40° for ± V2 their length, smooth,
septa only on valves. Seeds ellipsoid, ± 0.4 x 0.2 mm,
rounded to angled, orange, shallowly alveolate; testa
cells iiTegularly elongate. ± 50-75 x 15 pm, anticlinal
walls thin, iiregularly jigsawed, periclinal walls with
numerous fairly large pits.
Diagnostic features: bracteoles positioned 'A way up
pedicel to approximate to calyx, mainly much reduced
but still leaf-like; corolla short to long, 1.5-5. 5 x 2. 2-4.0
mm; filaments very elongate, naiTOwly ovate with sub-
apical S-bend, without a basal, spoon-like expansion;
anthers appendiculate; ovary glabrous or hairy; fruit
smooth not wrinkled.
This species forms a close alliance with the two new
species described above. It differs in having the bracte-
oles placed further away from the bract ('A way up pedi-
cel to approximate to the calyx and not basal) and the sta-
mens with appendiculate anthers and filaments not
spoon-like at the base (Table 1 ).
Variation in characters between the populations of E.
cymosa are present. The most noticeable is in the size
and shape of the corolla. There are however, two cleat-
size groupings which occur in two disjunct distribution
areas (Figure 2). These are also coupled with a difference
in indumentum of the ovary. We have therefore decided
to recognize these two regional variants at subspecific
level.
Bentham (1839) described the species based on a
Drege collection which had been provisionally named E.
cymosa by Meyer who worked on and distributed many
Drege collections. Bentham saw many collections from
Berlin and usually cited these as ‘v. s. sp. in herb. reg.
Berol." Since he did not cite this reference for E. cymosa
he did not see his material there and must have relied
solely on the rather scant material at Kew which he anno-
tated. He noted in the protologue that he saw no cymes
in the material, and unfortunately he proceeded to use
Meyer’s manuscript name ‘'cymosa' for the species.
3a. subsp. cymosa
Guthrie & Bolus: 141 (1905); Dulfer: 77 (1965).
Illustration: Schumann & Kirsten: 122, t.l (1992).
Diagnostic features: corolla 1.5-2. 8 x 2. 2-3.0 mm;
ovary glabrous.
The typical subspecies occurs in the mountains of the
Du ToitskloofAVellingtonAVemmershoek area where it
grows in moist places on cliffs (Figure 2). Where we
have seen it, the plants grow on a ledge under an over-
hang in deep shade in a river gorge. Elowering time:
October to March, one record in June.
3b. subsp. grandiflora E.G.H.Oliv. & l.M.Oliv.,
subsp. nov., a subspecie typica tJoribus majoribus et
ovario puberulo differt. Figura 4.
TYPE. — Western Cape, 3319 (Worcester): Worcester
Div.; Blaaskloof, W slopes of Keeromsberg, 5000 ft [1 525
m], (-DA), 23 February 1958, Esterhuvsen 27590 (BOL,
holo.; BM, E, K, MEL, MO, NBG, NY, P, PRE, S, W).
Diagnostic features: corolla larger, 3. 5-5. 5 x 3^
mm; ovary hairy (Figure 4; Table 1).
This subspecies occurs on the mountains in the Hex
River Range just southwest of Ceres to near Worcester
and on the Keeromsberg across the Hex River Valley
(Figure 2). These populations are cut off from those of
the typical subspecies by the much drier, lowland
Worcester-Tulbagh Valley.
The plants are all recorded growing in rock crevices
on overhanging rocks in shady, moist areas. One collec-
tion on Schurfteberg was described as on ‘sheltered
rather dry slightly earthy cliff face’.
Compton (1953) under the protologue of his E.
monantha, noted that several collections by Elsie
Esterhuysen ‘may also belong to this species’. He noted
that they possessed open-backed leaves and sepals which
could be ascribed to their habitat preferences. These
included two collections of this subspecies, Esterhuysen
14719 & 15202 and the type of E. amalophylla. We
regard Compton’s E. monantha as being conspecific with
E. flacca Benth. Flowering time: November to March.
Parats'pe material for subsp. grandiflora
WESTERN CAPE. — 3319 (Worcester): Ceres Div., Schurfteberg,
3500 ft [1 070 m], (-AD), 7-12-1948. Esterhuysen 14719 (BOL. K.
NBG. NY. PRE); Ceres Div., Mitchell Peak. 4000 ft [1 220 m]. (-AD),
27-03-1949. Esterhuysen 15202 (BOL. NBG. PRE); Ceres Div,,
Waaihoek Peak facing Tarantula Peak, 5800 ft [1 770 m], (-AD). 16-
01-1961. Esterhuysen 28737 (BOL. PRE); Worcester Div.. Milner
Peak, E side. 5000 ft [1 525 m]. (-AD), 18-12-1948. Esterhuysen
14938 (BOL, PRE); Worcester Div.. Audensberg Ridge Peak, 2()0() ft
[610 m], (-CB). 1-01-1950. Esterhuysen 16744 (BOL. NBG).
Specimens examined for subsp. cymosa
WESTERN CAPE.— 3319 (Worcester): Du Toitskloof. (-CA),
Drege 1185 (K. lecto.; BM. BOL. NBG, P, TCD); ibid., Drege
SAM10592 (SAM); ibid.. 2000-3000 ft [600-900 m], October, Drege
s.n. (BOL); Malbrokskloof, 2500-3000 ft [760-900 m], IIIAe. (-CA).
20-02-1828, Drege s.n. (P); Seven Sisters above Groen Kloof, 3500 ft
[1 070 m], (-CA), 14-01-1951. Esterhuysen 18322 (BOL); kloof
between Krom River Peak and Krom River Dome. 3000 ft [915 m],
(-CA). 30-12-1958. Esterhuysen 28062 (BOL, K); ibid.. 4000 ft [1 220
m], 26-10-1960. Esterhuysen 28687 (BOL, NBG); Haelhoek Sneeukop
overlooking Wemmershoek Valley, 3500 ft [1 070 m], (-CA). 14-03-
1959. Esterhuysen 28225 (BOL, PRE); Klein Drakenstein Mtns,
Donkerkloof. ± 3000 ft [915 m], (-CA), 31-06-1960. Esterhuysen
28435 (BOL, K. PRE); ridge between Witteberg and Molenaarsberg.
4500 ft [1 370 m], (-CA), 27-12-1960. Esterhuysen 28693 (BOL);
Du Toitskloof Mtns, upper Krom River SE of New Year's Peak, 3500 ft
[1 070 m], (-CA). 22-12-1959. Oliver 418 (NBG) [voucher for
Schumann & Kirsten: 122, t. 10. 1992]; ibid., 1 000 m, 7-01-1961,
Oliver 1144 (NBG); Seven Sisters, Wellington. (-CA), 10-1931, Stokoe
6695 (BOL).
172
Bothalia 32.2 (2002)
2 mm
FIGURE 4. — Erica cymosa subsp. grandiflora. A, flowering branch, with bracteoles mostly in halfway position, natural size; B, flower, with
bracteoles subapproximate; C. bracteole; D, sepal; E, stamen, lateral, back and front views; F, gynoecium; G, ovary opened laterally; H,
capsule; 1, seed; J, testa cells. All drawn from the type, Esterhuysen 27590. Scale bars: B-D, F-H, 2 mm; E, I, I mm; J, 20 pm.
LITTLE KAROO SPECIES
4. Erica schelpeorum E.G.H.OUv. & l.M.Oliv.,
sp. nov., foliis 4-natis, ramis, foliis, bractea bracteolisqoe
pilis glandulosis, bractea non recaulescenti, inflorescen-
tibus umbellati.s, pedicello longo (5-9 mm), calyce sine
glandibus, corolla glabra, antheris exsertis muticis vel
calcaribus minnlis dignoscenda. Figura 5.
TYPE. — Western Cape, 3321 (Ladismith): Swarlbcrg
Mtns, northern foothills due N of highest point of road
before descent into Die Hel, 4 ()()() ft 1 1 220 m|, 6-05-
1963, Oliver STE32I0H (NBG, holo.; BM, BOL, K, MO,
NY, PRE).
Shrub erect, much branched, twiggy, 0.5-1. 5 m tall,
grey, single-stemmed resecder. Branches: numerous,
main branches 30-120 mm long and secondary branch-
es 10-30 mm long; internodes 2-5 mm long; stem with
short, spreading, simple hairs and fewer, long, gland-
tipped hairs admixed. Leaves 4-nate, subspreading,
incurved but not imbricate, oblong, 4-5 x 0.8-1. 2 mm.
with rounded margins, obtuse, finely short-haired on
both surfaces, abaxially with scattered, long, gland-
tipped hairs admixed, sulcus narrow, open at base; peti-
ole ± 0.8 mm long, short-haired. Inflorescence: flowers
4-nate in 1-4 whorls umbel-like at apex of most main
and secondary branches; pedicel 5-9 mm long, finely
short-haired with a few, long, gland-tipped hairs
admixed; bract not recaulescent, 2-3 x 0.6 mm, other-
wise like leaf, green or pinkish red; bracteoles 2, just
above mid position on pedicel, 1 .5-2.2 x 0.3-0. 6 mm,
like miniature leaf with long petiole. Caly.x 4-partite;
segments broadly elliptic, ± 3 x 1.5 mm, adpressed to
corolla, subacute, margins entire at base, slightly
toothed/lobcd in uppei' half, glabrous with upper sulcate
region short-haired, purple-red often with green apical
portion, sulcus narrow, '/a length of .segment. Corolla 4-
lobed, globose-urceolate, 4. 0-4. 5 x 3-4 mm, glabrous,
whitish pink base to dark pink apex; lobes suberect to
erect, ± I x 1.5 mm, broadly rounded, entire to slightly
erose. Stamens 8, exserted; filaments free, linear-
oblong, ± 2.6 X 0.25 mm, slightly widened towards base,
straight or very slightly geniculate subapically, glabrous.
Bothalia 32,2 (2002)
173
FIGURE 5. — Erica schelpeorum. A, flowering branch, natural size; B, stem, with leaves removed; C, leaf; D, flower; E, bracteole; F, sepal; G,
stamen, side, back and front views; H, ovary; I, ovary opened laterally to show ovules and placentae; J, capsule with one valve removed;
K, seed; L, testa cells; M. anther variant with appendages. A-I drawn from the Oliver 11863 & 11879, M from Vlok 2454. Scale bars: B-J,
M, 2 mm; K. 1 mm; L, 50 pm.
white; anthers dorsally attached near base, lanceolate to
oblong in adaxial view, bipartite, minutely appendicu-
late or muticous; thecae lanceolate in lateral view, ±1.1
X 0.4 mm, adpressed to slightly spreading, smooth, dark
brown, spurs (when present) minute, pore ± ‘A length of
theca; pollen in tetrads. Ovary 4-locular, very broadly
obovoid, 1.5 X 1.7 mm, emarginate, 8-lobed, hairy with
hairs longer at apex of lobes, with large nectaries around
base; ovules 30^0 per locule spreading from placenta
in upper V4 of locule; style exserted, ± 5 mm long,
glabrous, white; stigma simple truncate. Fruit a dehis-
cent capsule, ± 1.8 X 2.8 mm, valves splitting V4 their
length to 45° angle, septa only on valves, placentae
large. Seed small, obovoid, rounded to sub-angular, ± 0.3
X 0.2 mm, orange; testa shallowly reticulate, not thick,
cells 50-70 X 25 (im, anticlinal walls slightly thickened,
jigsawed, inner periclinal walls sparsely pitted. Flower-
ing time: this would appear to be autumn to early winter
(May to July) but can be earlier when summer rains have
occurred. This is certainly the case with the Kykoe pop-
ulation which was in full flower for the Viviers & Vlok
specimen in July and the Schumann specimen in May
but just past peak flowering for us in early March.
However, in the latter case only a few of the shrubs had
flowered, with the others not having produced any flowers
at that stage. Figure 5.
174
Bothalia 32,2 (2002)
Diagnostic features-, leaves 4-nate; gland-tipped hairs
on stems, bract and bracteoles but not sepals; bract non
recaulescent (bract = leaf); pedicel long, 5-9 mm; corol-
la glabrous; anthers exserted with no, or only minute
appendages.
This new species is allied to several species in
§Ephebiis — E. hirta Thunb. (E. sphaeroidea Dulfer), E.
perlata Sinclair, E. tradouwensis Compton, and also to
E. arinata Klotzsch ex Benth. (^Myra), E. mitchellensis
Dulfer i^Oroplianes) and E. strigosa Sol. (^Ceramia).
These species are widely scattered through the current
poor, i.e. unnatural, system of sections in the genus.
It is most similar to Erica hirta, E. perlata and E.
tradouwensis in having the long pedicels and flowers
with slightly exserted anthers. E. hirta differs in the short
spike-like inflorescence (not umbel-like), the hairy
corolla and 3-nate leaves, E. perlata in the hairy corolla,
the subopen-backed, spreading leaves and the spurred
anthers, and E. tradouwensis in the 3-nate leaves which
are slightly open-backed, the hairy corolla and the eglan-
dular, long hairs on the stems and leaves.
Erica schelpeorum was stated above as being similar
to three other species in other sections. With E. arinata it
shares the bract on the main axis (non recaulescent), the
4-nate leaves, similar glands and umbel-like inflores-
cence but that species has larger, hairy flowers, pedicel
shorter than the corolla and longer anther appendages. E.
mitchellensis, which is similar to E. arinata, has a re-
caulescent bract and inflorescence of only a single whorl.
E. strigosa has the non recaulescent bract and glabrous
corolla, but differs in having a spike-like inflorescence
and short pedicel.
This species was first collected in 1954 by Mrs
Sybella Schelpe ( 1917-2001) while on a field trip with
her husband, Prof Ted Schelpe (1924-1985), who was
the lecturer in taxonomy for both of us during our student
years at the University of Cape Town. We thus take this
opportunity of honouring them both in naming this
species which was discussed with Sybella shortly before
her death.
Erica schelpeorum is widespread on mountains and
hills in and bordering the Little Karoo (Figure 6). In all
cases where we have seen the species, it occurs in dry,
arid areas usually associated with a dry watercourse.
Despite the high altitude of the type collection, the plants
there were growing in arid renosterveld vegetation on the
southern slopes of the northern foothills to the main
range of the Great Swartberg. In the Kykoe locality
which was shown to us by Jan Vlok and which is the
same population as Viviers & Vlok 184 and Schumann
684, the plants were growing in renosterveld vegetation
on shale/clay with a thin overlay of quartzite pebbles
washed down in a small dry watercourse.
All the species mentioned as being similar in some
respects, do not occur sympatrically with E. schelpeo-
rum, they all occur in the region to the southwest from
the Hex River Mountains to the Cape Peninsula east-
wards as far as the Langeberg, just east of Swellendam.
Erica hirta and E. tradouwensis are both low altitude
species with the former growing on clay/gravel slopes
sometimes on the border between dry fynbos and renos-
terveld vegetation in the Malmesbury, Stellenbosch and
Genadendal areas.
Paratype material
WESTERN CAPE. — 3321 (Ladismitli); Swartberg, in ravine
between Kliphuisvlei and Gamkaskloof, 1 200 m, (-BD), 7-05-1989.
Vlok 2112 (NBG); Calitzdorp area, Gainka Mountain Reserve, upper
Tierkloof. 500 m, (-DB), 7-05-1983, P. & J.Cattell 240 (BM, BOL, K.
MO, NBG, NY, PRE, S); ibid., N slopes of Dwarsberg, 900 in. (-DB).
8-09-1987, Allardice 1730 (NBG). 3322 (Oudtshoorn): Swartberg
Pass. N side, 3500 ft [1 060 m], (-AC). 12-02-1954, A.S.L.ScheIpe 98
(BOL); Great Swartberg. Meiringspoort, 2 100 ft [640 m], (-BC), 23-
06-1997 (fruiting), Oliver 10820 (NBG); Groot Swartberg, eastern end
at N base of Snyberg, 4000 ft [ 1 220 m], (-BD). 1 2-03- 1991, Vlok 2454
(BOL. NBG); Kamnianassie Mtns. Klein Plaas River Valley, 3600 ft
[1 100 m], (-DA), 10-08-1983, Mathews 1170 (K. NBG, P, PRE);
Ghwernaberg, above Kykoe, 600 m, (-DD), 11-01-2001, mainly in
bud. Oliver 11863 (NBG); ibid., 7-03-2001, Oliver 11879 (NBG);
Potjiesrivierhoogte Pass, southern foot, 550 m, (-DD), 25-05-1989,
Schumann 684 (NBG); ibid., next to national road on southern slope.
(-DD). 3-07-1987, Viviers Vlok 184 (BM. K, NBG, NY, PRE). 3323
(Willowmore); Antoniesberg. S side, ± 1 000 in, (-AD), 10-03-2001,
Van Wijk 2302 (NBG, SCHG),
5. Erica lithophila E.G.H.Oliv. & I.M.OUv., sp.
nov., Ericae kirstenii E.G.H.Oliv. proxima sed fruticulo
parvulo, ovulis 20-30 in quoque loculo non ± 60, ereclis
non effusis vel pendulis, antherarum calcaribus angus-
tioribus, testa parietibus anticlinalibus pertenuibus non
percrassis multo undulatis dignoscenda. Figura 7.
TYPE. — Western Cape. 3322 (Oudtshoorn): Swart-
berg at Meiring’s Poort. 5000 ft [I 524 m], (-BC), 16
October 1955, Esterhuysen 24865 (BOL, holo.; K!).
Small, brittle, delicate shrublets, 20 x 20-80 x 50 mm,
rarely 120 mm in diam., erect and compact to slightly
FIGURE 6, — Known distribution of
Erica schelpeorum.
Bothalia 32,2 (2002)
175
FIGURE 7. — Erica lithophila. A, flowering branch, natural size; B. stem with leaves removed; C. leaf; D. flower; E. pedicel; F, bract; G, bracte-
ole; H. sepal; I, stamen, side, back and front views; J, gynoecium; K, ovary partially opened laterally to show ovules; L, capsule with one
valve removed; M. seed; N, outer portion of anticlinal testa cell walls; O, inner portion of anticlinal testa cell walls. A-K, drawn from Vlok
2476', L-N. from Oliver 11552'. O from Oliver 11833. Scale bars; B, C. I. M, 1 mm; D, E-H. J-L, 2 mm; N. O, 100 pm.
spreading, single-stenimed reseeders. Branches', thick,
woody, basal stem with numerous main branches, 10-15
mm long, secondary branches few, ± 2 mm long; stems
with white, short, dense, simple hairs and a few plumose
hairs; intemodes very short. Leaves 2- or 3-nate. imbri-
cate, subspreading, elliptic, 2. 5-3.0 x 0.8 mm, abaxially
rounded, adaxially flattened, covered with dense, grey-
white, short, simple hairs, margins with short, plumose,
gland-tipped hairs, sulcus narrow open at base; petiole ±
0.4 mm long, short-haired to sparsely hairy, margins with
simple hairs and/or short, gland-tipped hairs. Inflores-
cence'. flowers 1-3 in 1 whorl at ends of main branches
and very rarely, secondary branchlets; pedicel ± 2 mm
long, red. covered with very short, fine hairs and in upper
half also with stouter, white, crisped, plumose hairs,
0.4— 1.0 mm long, these with or without an apical gland;
bract partially recaulescent about up pedicel, ovate, ±
2.2 X 1 .7 mm, subacute, pink, sparsely covered with very
short, fine, simple hairs, unevenly toothed, teeth gland-
tipped, sulcus naiTow, ± V3 length of bract; bracteoles 2,
approximate to calyx, slightly lopsided, otherwise same as
bract. Caly.x 4-partite, segments imbricate, broadly ovate,
± 2.8 X 2.8 mm. subacute, pink, otherwise like bract.
Corolla 4-lobed, urceolate, ± 7.5 x 4 mm. pink, glabrous;
lobes recurved, broadly rounded, ± 1.8 x 0.7 mm, erose
sometimes with very broad, plumose hairs at base.
Stamens 8, included, free; filaments linear, ± 3 x 0.4 mm,
slightly geniculate below theca, white, glabrous; anthers
bipartite, ovate in adaxial view, appendiculate; thecae
oblong, 0.9-1. 2 x 0.9 mm in lateral view, dark brown,
smooth, appendages elongate, naixow-lanceolate, ± 1 mm
long, strigulose, iiregulaiiy toothed, pore ± length of
theca; pollen shed in tetrads. Ovary 4-locular, sub-
sphaeroid, ± 1.8 x 1.5 mm, emarginate, completely cov-
ered with longish, spreading, simple hairs, with well-
developed nectaries around base; ovules 25-30 per locule.
176
Bothalia 32,2 (2002)
erect to suberect on a complete central placenta; style
included, ± 6 mm long, straight, glabrous; stigma manifest
capitellate. Fruit a dehiscent capsule, ± 3 x 4.2 mm, valves
splitting to base and 45°, septa mainly on columella, hard,
woody. Seeds ellipsoid, ± 0.7 x 0.4 mm, slightly flattened
laterally; testa colliculate, yellow-orange, cells ± oblong,
70-100 X 50 pm, outer part of anticlinal walls straight to
slight undulate, inner part irregularly undulate, inner peri-
clinal walls sparsely and finely pitted. Flowering time:
August to November. Figure 7.
Diagnostic features: shrublets very small with flowers
relatively large compared to the plant and leaves; leaves
grey; pedicel with long, plumose hairs in upper part;
anther appendages narrow; ovules 20-30 per locule,
erect; seeds with delicate testa having thin anticlinal
walls.
This new species is most closely related to E. kirstenii
E.G.H.Oliv. (Oliver & Oliver 2000) which is distin-
guished by the much larger stouter shrubs, leaves ± twice
as long, pedicel long and curved with short plumose
hairs all over, ovules ± 60 and not erect and testa cells
with thick anticlinal walls.
Erica lithophila occurs on the eastern parts of the
Great Swartberg Range in the Meiringspoort area and on
the Kammanassie Mountains just to the south where it is
confined to high altitudes (Figure 8). The species is easi-
ly overlooked due to the small size of the plants with very
old gnarled plants often only 30 x 30 mm and growing.
The habitat is rock crevices in large rocks usually with no
soil and north-facing. The largest plant we have seen was
growing in some humus in a vertical crack facing east and
was open and ± 120 mm in diameter. In these dry, rocky
habitats, the shrublets are often cryptic, melting in with
the lichen-covered rock. In younger plants the flowers at
± 7.5 mm long, are rather disproportionately large for a
plant only 20 mm in diameter. The habitat preference pro-
vides the specific epithet for the species — litlios = rock
(Greek), -philos = loving (Greek).
Erica kirstenii is confined to rocky places on the
Klein Swartberg in the region of Ladismith and
Seweweekspoort ± 100 km to the west of the distribution
range of E. lithophila.
There are several other species that grow in rocky
places and that have similar flowers with large, petaloid
bracts, bracteoles and sepals edged with teeth or stout
plumose hairs — E. taylorii E.G.H.Oliv. & I.M.Oliv.
(Oliver & Oliver 2001) and E. modesta Salisb. The for-
mer occurs in two very disjunct regions, the Cederberg
and the Klein and central Groot Swartberg and produces
large spreading mats on rocks at high-lying altitudes,
whereas the latter occurs along the summit ridges in the
Riviersonderend Mountains and forms a taller, erect
shrub up to 0.75 m.
Paratype material
WESTERN CAPE — 3322 (Oudtshoorn): Swartberg, Spitzkop neck
to the east, 1 600 m, (-AD), 15-04-2000 (fruiting), R. Taylor s.ii.
(NBG); ibid., 5000 ft [1 524 m], 9-03-1991 (fruiting), Vloks.n. (NBG);
Blesberg, 6000 ft [1 828 m], (-BC), 17-10-1955, Esterlmysen 24934
(BOL); ibid,, N & E slopes, 2 060 m, 13-07-2000 (flowers & fruiting),
Oliver 11552 (NBG); Swartberg Mtns, N slopes above Vrolikheid E of
Meiringspoort, 5200 ft [1 584 m], 16-08-1991, Vlok 2476 (NBG);
Kammanassie Mtns near Mannetjiesberg, 1 700 m, (-DB), 7-11-1995,
Hitchcock K73 (NBG); Mannetjiesberg Peak, N slopes, 6200 ft [1 890
m], (-DB), 2-10-1971, Oliver 3612 (NBG); Kammanassie Mtns due N
of Buffelsberg, 1 500 m, (-DB), 9-01-2001 (fruiting), Oliver 11807
(NBG); Kammanassie Mtns, Molenrivier 114, W of Mannetjiesberg, 1
840 m, (-DB), 9-01-2001 (fruiting), Oliver 11833 (NBG).
6. Erica umbratica E.G.H.Oliv. & I.M.Oliv., sp.
nov., fruticulo tenello fragili pendulo, foliis 3-natis effu-
sis, pilis brevibus glandulosis et glande magno apicali,
floribus viscidis, pedicello longo, sepalis glandibus ses-
silibus marginalibus, calcaribus antherarum longis undu-
latis, ovario pubenti dignoscenda. Figura 9.
TYPE. — Western Cape, 3322 (Oudtshoorn); Meiring’s
Poort, northern end near Wadrif, 600 m, (-BC), 5
January 2001, E.G.H. & I.M. Oliver 11757 (NBG, holo.;
K, PRE).
Shrublet delicate, brittle, pendulous, 200-500 mm
long, single-stemmed reseeder. Branches: main branches
200—400 mm long, with numerous secondary branches,
10-80 mm long, these not at each node, occasional,
short, tertiary branchlets; stems sparsely hairy with short,
spreading, simple hairs and stalked glands. Leaves 3-
nate, subspreading to spreading to 90°, 5-7 x 0.8 mm,
rounded on both sides, margins rounded to slightly acute,
sulcus narrow, sparsely hairy on both surfaces with
short-stalked glands on margins, apex tipped with soli-
tary large gland, open at base, with occasionally at base
of each lateral branch, l-3(-5) whorls of small brown
bract-like leaves; petiole ± 0.6 mm long, hairy on both
FIGURE 8. — Known distribution of
Erica lithophila.
Bothalia 32,2 (2002)
177
FIGURE 9. — Erica umbratica. A. flowering branch, natural size; B, stem; C, leaf; D. flower; E, flower opened laterally; E, bract; G, bracteole; H,
sepal; I, stamen, side, back and front views; J, gynoecium; K, ovary opened laterally to show ovules and placentae. All drawn from the
type collection, Oliver & Oliver 11757. Scale bars: B-H. J, K, 2 mm; I, 1 mm.
sides, glands on margins. Inflorescence: flowers 3-nate
in 1 or 2 whorls, umbel-like at ends of main and sec-
ondary branches; pedicel 6-7 mm long, pink, glabrous
with a few scattered short-stalked glands and simple
hairs at attachment of bract and bracteoles; bract partial-
ly recaulescent, ± 'A way up pedicel, ovate, ± 1.2-1. 4 x
0.6 mm, margins lined with sessile glands, apex with
larger, red gland, otherwise glabrous, greenish; bracteoles
2, V3-V2 way up pedicel, ± 1 .4 x 0.6 mm, otherwise like
bract. Calyx 4-partite, adpressed to corolla; segments
ovate, ±2x1 mm, margins lined with sessile very viscid
glands, the terminal one larger and dark red, otherwise
glabrous, green. Corolla 4-lobed, ovoid, ±4x3 mm,
glabrous, viscid, white; lobes recurved, ± 0.7 x 1.2 mm,
rounded, margins entire. Stamens 8, included, free; fila-
ments linear, ± 5 x 0.5 mm, slightly widened at base,
subapically geniculate, spiculate near base, otherwise
glabrous; anthers bipartite, dorsally attached near base,
lanceolate to oblong in adaxial view, appendiculate; the-
cae ovate in lateral view, ± 1.8 x 1.1 mm, finely spicu-
late, orange-yellow, appendages elongate, narrowly
lanceolate, ± 2.2. x 0.2 mm, slightly zigzagged, sparsely
toothed and with a few short, simple hairs, pore ± 'I2
length of thecae; pollen in tetrads. Ovary 4-loculate,
spherical, ± 1.5 x 1.8 mm, emarginate, short-haired, with
large basal nectaries; ovules 80-100 per locule, lateral
and spreading on large, bulbous placenta the full length
of locule; style ± 2.4 mm long, just exserted, glabrous;
stigma capitellate. Fruits not seen. Flowering time: this
is variable and possibly determined by rainfall, which is
mainly in winter but also as summer showers. Flowers
do not seem to be profuse on the plants. Figure 9.
Colour illustrations: Schumann & Kirsten: 251, t. 10
& t. 11 (1992).
Diagnostic features: delicate, brittle, pendulous shrub-
let; leaves 3-nate, spreading (up to 90°), delicate; pedicel
long (6-7 mm); flowers viscid; anther appendages long,
narrow, zigzagged and irregularly toothed; ovary short-
haired.
This new species shares similarities with a range of
viscid-flowered species such as E. carduifolia Salisb., E.
nubigena Bolus, E. ixanthera Benth., E. wittebergensis
Dulfer, and two undescribed species — one from the
Kouga Mountains (Esterhuysen 10673, 27097), the other
from the northern side of the Swartberg Pass (Schumann
798 and Vlok 2502). With these it shares long, 3-nate
leaves mostly with sticky glands, often spreading and
caducous, bract-like leaves at base of flowers and side
branches, long pedicel with few to many glands, similar
sized and positioned bract and bracteoles, large sessile
marginal glands on the sepals, long anther appendages,
and ovary with large placenta bearing numerous ovules
producing small seeds. It differs in its growth form, being
178
Bothalia 32.2 (2002)
pendulous with thin, very brittle branches, much longer
zigzagged and irregularly toothed anther appendages and
short-haired ovary. Each of the species has rather specif-
ic leaves.
Most of the other species grow in shaded, moist places
often on southern slopes, E. nuhigena and E. carduifolia
at high altitudes, E. ixanthera, E. sp. nov. (Kouga) and
E. umbratica at a much lower altitude. Erica sp. nov.
from the Swartberg Pass is the odd one out as it grows on
dry, rocky slopes in open veld on the northern side of the
Swartberg and forms erect, woody shrublets.
Erica umbractica has rather viscid flowers which
easily collect dirt on them in the very windy conditions
that prevail in Meiringspoort. This made Schumann in
frustration, refer to this new species as Erica “puer-sordi-
dus” — ^the dirty boy, when trying to produce perfectly
clean material for photographing (Schumann & Kirsten
1992).
Erica umbratica is known only from a few small
populations in Meiringspoort (Figure 10) where it grows
on very large, shaded rocks, just above the main stream
which runs through the poort. It could be postulated to
have evolved with the formation of the poort. The plants
we have studied grow in cracks and on ledges sheltered
under overhangs on a large rocky outcrop where they
receive very little direct sunlight. We have thus chosen
the name to reflect this habitat, umhraticus - of the shade
(Latin). Due to the inaccessible location, material for
study is very limited.
The discovery of this rare species was made by Jan
Vlok who has an eye for recognizing unusual and inter-
esting plants — the plants look so unlike an Erica species,
and more like some members of the family Asteraceae
such as Stoebe.
Paratype material
WESTERN CAPE. — 3322 (Oudtshoorn): Swartberg Mtns,
Meiringspoort, 550 m. 8-08-1987, (-BC), Schumann 548 (NBG); ibid.,
near Herriesklip, (-BC), 15-04-1986, Vlok 7-/60 (NBG; PRE); ibid., near
Waterfall picnic area, (-BC), 1 900 ft [580 m|, 26-06-1990, Vlok 2312
(NBG).
7. Erica annalis E.G.H.Oliv. & I.M.Oliv., sp. nov.,
a speciebus nobis notis bene distincta, foliis 4-natis pau-
cis spicLilis, intlorescentia non spicata, pedicello com-
parate longo (10-15 mm), bractea bracteolisque parva
remota, corolla perpubenti, filamentis styloque sparse
pilosis dignoscenda. Figura 1 1 .
TYPE. — Western Cape. 3322 (Oudtshoorn): Kamma-
nassie Mountains, Perdekloof, northern foothills of
Mannetjiesberg above Buffelsklip, 820 m, (-DB), 3
September 2001, E.G.H. & I.M. Oliver 11929 (NBG,
holo.; BOL, K, NY, PRE).
Sparsely branched, erect to spreading shrub, 0.2-1 .0 m
taU, often with gnarled old basal stems/branches, single-
stemmed. Branches: main branches 50-100(-200) mm
long; secondary branches vestigial; stems grey, very fine-
ly short-haired, sometimes with a few gland-tipped hairs
admixed; intemodes up to 5 mm long. Leaves 4-nate,
12-15 X 1 mm, subspreading to spreading, recurved,
glabrous or with a few, sparse, basal hairs and a few
spicules abaxially, ± circular in cross section, sulcus nar-
row, open at base; petiole 1 mm long adpressed, fine-
haired, yellow. Inflorescence: 1-3 flowers in a single
whorl at ends of main and secondary branches, sub-
spreading to spreading; pedicel 10-15 mm long, finely
short-haired with numerous longer, stalked, red, gland-
tipped hairs admixed, green to reddish; bract partially
recaulescent in basal position, oblong, ± 1.7 x 0.4 mm,
finely short-haired, margins with minute sessile glands,
green-brown, sulcus minute; bracteoles 2 in lower half of
pedicel, otherwise same as bract but sulcus often absent.
Calyx 4-partite; segments lanceolate, ± 5 x 0.4 mm, not
laterally imbricate, fine-haired with glandular margins,
green to reddish; sulcus ± 'A length of segment, narrow.
Corolla 4-lobed, ± 25 x 4 mm, tubular, funnel-shaped,
curved with slight constriction above ovary, with scat-
tered, short and a few longer, simple hairs, bright orange-
red; lobes ±3x3 mm, subacute, margins entire, spread-
ing to recurved. Stamens 8, free, manifest to exserted; fil-
aments linear, straight, very sparsely pilose, white with
pale red apex; anthers bilobed, dorsally attached ± 'A way
up, narrowly oblong-elliptic in adaxial view, muticous;
thecae narrowly oblong and ± 5 x 0.6 mm in side view,
smooth, golden brown to reddish brown, pore ± ‘A length
of theca; pollen in tetrads. Ovary 4-locular, cylindrical to
nan'owly ellipsoid, ± 3.3 x 2.2 mm, 4-lobed with round-
ed apex, covered with dense, long, erect hairs, nectaries
present around base; ± 35 ovules per locule, spreading
from full-length placenta; style ± 27 mm long with a few
hairs towards base, white, reddish towards apex; stigma
truncate simple, reddish. Fruit a dehiscent capsule, ± 5 x
4.5 mm, obovoid, hard and woody, valves splitting 35°
for Y4 their length, placenta whole length of capsule, sep-
tum mostly on valve. Seeds ± 1 x 0.7 mm, broadly ellip-
soid with one side slightly flattened, brown; testa slightly
alveolate, cells ± hexagonal ± 100 x 75 pm, anticlinal
FIGURE 10. — Known distribution of
Erica umbratica, •; and E.
annalis, ■.
Bothalia 32,2 (2002)
179
FIGURE II. — Erica amalis. A, flowering branch, natural size; B, stem with leaves removed; C, leaf; D, flower; E, pedicel enlargement; E, bract;
G, bracteole; H, sepal; I, anther, front, side and back views; J, gynoecium; K, ovary, opened laterally; L, capsule, with one valve removed;
M, seed; N, testa cells. All drawn from the type collection, Oliver & Oliver 11929. Scale bars: B,C, F-H, I-L, 2 mm; D, 4 mm; E, M, 1
mm; N, 100 pm.
walls thin straight inner periclinal wall with numerous
small pits. Flowering time: July to October, but probably
at other times depending on summer rains as evidenced
by old flowers on flowering material. Figure 11.
Diagnostic features: leaves 4-nate; corolla hairy with
a distinct swelling at the base; bract and bracteoles
small, remote; anthers well exerted, muticous, long and
narrow; sparse hairs on filaments and style; pedicel rela-
tively long, 10-15 mm, with small, sticky, red glands;
ovary densely hairy, cylindrical; leaves with very few
sclereids.
E. annalis could be related to several long-tubed species
that occur in the region — E. wendlandiana Klotzsch, E.
densifolia Willd. and E. abelii E.G.H.Oliv. and to a lesser
extent E. maximilianii Bolus. The first three species have
the narrow, elongated anthers which are included to slight-
ly manifest, but these are appendiculate, they have the nar-
row cylindrical ovary but all of them are glabrous, and the
hairy corolla. E. wendlandiana has the hairy filaments and
style which can be present in E. densifolia but these are
lacking in E. abelii. E. abelii differs in having a long bract
and bracteoles which are approximate to the calyx, not
small and remote. All three species differ from E. annalis
in having 3-nate leaves and no similar basal swelling of
the corolla. E. densifolia has spike-like synflorescences on
the main stems. E. wendlandiana is remarkable in the
genus for the large swelling at the base of the style and E.
mciximilianii differs in having a glabrous corolla and a
glabrous, broadly obovoid ovary.
This new species is known only from the type local-
ity where it was found during a survey of water catch-
ments in the Kammanassie Mountains by a local Nature
Conservation officer, Jan de Jaar. The plants are con-
fined to rock faces, either facing south or north in a sin-
gle side kloof running east-west. They grow in crevices
in the quartzitic rock. On the hot, dry, north-facing cliffs
they are the only Erica species present but on the cool-
er, south-facing slopes they grow on steep sheet-rock
with small plants of E. viridiflora subsp. primulina,
which is widespread in similar situations in the sur-
rounding mountains (Oliver & Oliver 2002). The sur-
rounding vegetation is arid scrub with no elements of
fynbos, which only starts appearing ± 300 m higher up
the slopes.
The epithet for this new species honours the dis-
coverer, the Nature Conservation field officer, Jan de
Jaar. We have used the word, jaar = year (Afrikaans),
annalis = relating to a year (Latin), even though his name
is probably derived from De Jager, the hunter (Afrikaans/
Dutch).
180
Bothalia 32,2 (2002)
Paratype material
WESTERN CAPE — 3322 (Oudtshoom): Uniondale, Buffelsklip,
in side kloof next to Buffelskliprivier. 775 m. (-DB), 30-08-2000. De
Jaar 508 (NBG).
REFERENCES
BENTHAM, G. 1839. Erica. In A.P. De Candolle. Prodromiis 7:
580-733. Treuttel & Wiirtz. Paris.
COMPTON, R.H. 1953. Plantae novae africanae. Journal of South
African Botany 19: 109-135.
DULFER, H. 1965 Revision der sudafrikanischen Arten der Gattung
Erica L. Annalen des Naturhistorisches Museums. Wien 668;
25-177.
GUTHRIE, F. & BOLUS, H. 1905. Erica. Flora capensis 4,1: 4-315.
Reeve, London.
OLIVER, E.G.H. & OLIVER, I.M. 2000. Erica kirstenii, a new rock-
loving species from South Africa. Yearbook of the Heather
Society 2000: 57-62.
OLIVER, E.G.H. & OLIVER, I.M. 2001. Five new species of Erica
(Ericaceae) from the Swartberg Range, Western Cape, South
Africa and a note on E. esterhuyseniae. Bothalia 31; 155-165.
OLIVER, E.G.H. & OLIVER, I.M. 2O02. The genus Erica (Ericaceae)
in southern Africa: taxonomic notes 1. Bothalia 32: 37-61.
SCHUMANN, D. & KIRSTEN, G. 1992. Ericas of South Africa. Fern-
wood Press, Cape Town.
Bothalia 32,2: 181-184 (2002)
Studies in the liverwort family Aneuraceae (Metzgeriales) from southern
Africa. 4. Riccardia obtusa
S.M. PEROLD*
Keywords: Aneuraceae, Riccardia Gray, R. obtusa S.W.Amell, southern Africa
ABSTRACT
For a clearer understanding of Riccardia obtusa S.W.Amell. it is here described and illustrated in greater detail than in
AmelTs (1952) initial publication of his new species. Subsequently, with minor alterations, Amell ( 1963) repeated his treat-
ment of R. obtusa in Hepaticae of South Africa. Wigginton & Grolle ( 1996) have, however, remarked that this species needs
to be clarified, along with the other two Amellian species of Riccardia. namely R. capensis and R. rhodesiae. which will be
treated in detail in future papers.
INTRODUCTION
Riccardia obtusa S.W.Amell appears to be quite
widespread in southern Africa, ranging from Karkloof in
KwaZulu-Natal to Joubertina in Eastern Cape and
Knysna in Western Cape, according to Arnell (1963).
The specimen from Pretoria that Arnell (1963) men-
tioned. had been identified earlier by Sim as R. com-
pacta, which is not correct either, as the thalli are much
too thin. According to my own observations, R. obtusa is
also known from Long Tom Pass and Buffelskloof
Nature Reserve, Mpumalanga, as well as Kirstenbosch,
Cape Town. Samples of fresh collections of it from Fern
Forest, Diepwalle (Figure 1), taken in October 2000,
were kept alive for as long as 17 months, in order to
observe the branching pattern of the thalli, as well as the
oil bodies (see Specimens examined). Although Arnell
did not remark upon the oil bodies of R. obtusa, they are
quite distinctive in being very dark and irregular in
shape. By the time Amell described his new species, the
oil bodies must have already disappeared.
Riccardia obtusa S.W.Amell in Botaniska Notiser
1952: 142; S.W.Amell: 96 (1963). Type: Cape Province,
Knysna, Deepwall (= Diepwalle) Forest Reserve, on a
wet sandstone slope, Arnell 1602 (BOL, holo.!; S!).
Thalli prostrate, in densely overlying patches, 2-A
layers thick and tightly adherent to substrate, as well as
to each other, often highly contorted and difficult to sep-
arate, bottle green and rather brittle; apices of distal
branches thickened and fleshy, variously shallowly
lobed, margins opaque and obtuse (Figure 2 A, B); pri-
mary and secondary branches with winged and translu-
cent margins (Figure 2A, C, D); when dry, dark green to
brown; smallish to medium-sized. Main axis mostly
10-15 mm long, rarely longer, clearly differentiated, dis-
tally trifurcate to subpalmately divided, the closely
adjoining or overlapping branches lobulate and short,
500-1550 pm wide, their combined width across up to
4.9 mm, apically each branch once, or 2 or 3 times shal-
* National Botanical Institute, Private Bag XlOl, 0001 Pretoria.
MS. received: 2002-06-27.
lowly notched, with continued growth elongating and
becoming fan-shaped and thinner; proximally, main axis
generally retaining its dominance, dorsally convex, up to
900 pm wide, narrowing somewhat toward base, branch-
ing somewhat irregularly pinnate on both sides, rarely
with a much elongated side branch up to 8.5 mm long,
similarly branched and in all probability, eventually
forming a new main axis. Primary branches/pinnae
opposite or subopposite, single, 350-2375 pm long, up
to 525 pm wide, obliquely spreading at angles of 40°-70°
with the main axis and separated by intervals of
825-2300 pm between them; sometimes with only 1
weaker, rarely with 2 (Ion each side) secondary branch-
es/pinnules, so that occasional trifurcate lateral branches
arise. Stolons (Figure 2C) quite rare, developing lateral-
ly from upper or lower part of main axis, or from tip of
primary pinna, seldom branched. Dorsal epidermal cells
in median part of apical segment of main axis from
above, 5-7-sided (Figure 2E), cell walls somewhat
thickened, 45.0-60.0(-87.5) x 25.0-32. 5(-37. 5) pm, sub-
dorsal cells larger, 112.5-212.5 x 50-85 pm, subventral
cells 105-200 x 52.5-77.5 pm, ventral epidermal cells
50-75 X 25^0 pm. Oil bodies very densely crowded at
meristematic apical notches of branches and then present
in up to 90% of dorsal and ventral epidermal cells, as
FIGURE 1 . — Distribution of Riccardia obtusa in southern Africa.
2 mm 2 mm
182
Bothalia 32,2 (2002)
FIGURE 2. — Riccardia ohtusa. A-C, thallus: A, with subpalmate distal branching and irregularly pinnate lateral branching; B, with several gynoe-
cial branches; C, with single male branch and a stolon. D, primary branch. E, median dorsal epidermal cells (solid lines) of ultimate seg-
ment of axis with smaller oil bodies, and large subdorsal cells (stippled lines) with larger oil bodies (oil globules indicated by stippling);
F, marginal cells of ultimate segment of axis, oil bodies stippled; G, H, c/s axis at ± middle of ultimate segment; I-K, c/s primary branches;
L, c/s stolon; M, mucilage papillae at ventral apical notch of branch; Ni, Ni, gemmae. 0-Q, antheridial branch: O, from above; P, from
side; Q, c/s. R, gynoecial branch from side; S, calyptra; T, c/s calyptra wall; U, c/s seta; V, c/s part of bistratose wall of capsule valve showing
different cell walls: abr, abaxial radial; adr, adaxial radial; it, inner tangential; mw, median; ot, outer tangential. W|, W2, spores; X, elater.
A, B, L, N-Q, S, Perold & Koekemoer 4456\ C, F, H-K, R, T, V-X, Perold & Koekemoer 4457\ D, Koekemoer 2246'. E, Perold. Burgoyne
& Smithies 4743', G, M, Perold & Koekemoer 4462', U, Perold & Koekemoer 4460.
Bothalia 32,2 (2002)
183
well as in marginal and internal cells, mostly 1 , but occa-
sionally 2 or 3 per cell, smaller in dorsal and ventral epi-
dermal cells, 12.5-20.0 x 12.5-17.5 pm, larger in inter-
nal cells, 25.0-32.5 x 15.0-27.5 pm, irregular or kidney-
shaped, occasionally subspherical, very dark, composed
of many tiny globules. Margins of ultimate segments of
main axis (Figure 2F) with outer cells generally some-
what smaller than intramarginal ones, from above
42.5-57.5 X 27.5-47.5 pm, rectangular to subquadrate,
free walls bulging slightly outward, most cells with a sin-
gle oil body, 12.5-20.0 x 10-20 pm; intramarginal cells
5- or 6-sided, 57.5-100.0 x 35.0-62.5 pm, with 1, occa-
sionally 2 oil bodies each. Cross section at ± middle of
ultimate segment of main axis biconvex (Figure 2G, H),
840-1040 pm wide and 5-8(-l 1 ) cell rows or up to 325
pm thick medianly, gradually tapering to obtuse margins,
thickness of dorsal cells 12.5-15.0 pm, subdorsal cells
25-30 pm, medullary cells (40-)60-100 pm, subventral
cells ± 30 pm and ventral cells ± 15 pm; base of main
axis generally narrower than younger part further along.
± 680 pm wide, 6 cell rows or ± 200 pm thick, margins
shortly acute; cross section of primary branch/pinna
(Figure 2I-K) 85-110 pm or 3 or 4 cell layers thick
medianly, unistratose margins 3 or 4 cells wide; cross
section of secondary branch/pinnule ± 55 pm or 3 cell
layers thick; cross section of stolons (Figure 2L), oval to
elliptical, ± 140 x 210 pm. Mucilage papillae (Figure
2M) ventral, crowded together at shallowly notched apex
of branch and then in 2 spaced rows, one on either side
of midline, 150-225 pm between successive ones, club-
shaped. 50-80 pm long, 17.5-37.5 pm wide above,
tapering to foot, 10.0-12.5 pm wide. Rhizoids ventral
along branches, 10.0-12.5 pm wide. Asexual reproduc-
tion by gemmae (Figure 2N) occasionally observed on
dorsal surface of upper branches, consisting of 2 cells
joined together, ± 65 x 40 pm.
Monoicous. Antheridial branches rather few in num-
ber, arising laterally on main axis (Figure 2C), immediate-
ly below base of primary branch or subopposite to base of
primciry branch, sometimes 2 together, sometimes shortly
stipitate, oblong-linear, 1125-1300 pm long, width
250-270 pm, in cross section ± 240 pm high, bearing up
to 1 1 pairs of antheridia, on same plant occasionally short-
er male branches, 460-830 pm long, width up to 220 pm,
with 4—9 pairs of antheridia, antheridial cavities from
above 50.0-62.5 x 32.5-47.5 pm, surrounding cells
37.5^2.5 X ± 30 pm, 1 or 2 adjoining cells between pairs
of cavities; margins (Figure 20) crenulate, with single
erect layer of swollen cells, 65.0-87.5 x 45-65 pm.
Gynoecial branches (Figure 2R) short, arising laterally on
same side along length of main axis, 2 or 3 in a row, some-
times in pairs on opposite sides of primary branch, ± 260
pm high. 450 pm wide, surrounding paraphyses up to 310
pm long, composed of 3 or 4 cells joined end to end.
archegonia in 2 rows. Calyptra (Figure 2S) clavate, length
2.0-3. 5 mm, ± 0.85 mm wide above and narrowing below
to ± 0.6 mm; cross section of wall (Figure 2T) 170-200
pm or up to 8 cell layers thick, many cells of outermost
row markedly protuberant. 115-135 x ± 25 pm, cells in
corona up to 175 pm long. Seta ±5.7 mm long, 220-280
pm wide, with 4 inner and 12 outer cell rows, i.e. 4 cells
diam. (Figure 2U). Capsule ellipsoidal, 1150-1675 pm
long, with 4 valves, ± 325 pm or 21-23 cell rows wide,
bistratose; cells of epidermal layer in external longitudinal
view (Figure 3A) 67.5-90.0 x 10.0-12.5 pm, end walls
straight or oblique, with vertical (nodular) thickenings; in
cross section (Figure 2V ) cells rectangular in shape, thick-
enings on adaxial radial and inner tangential walls, bands
on one side of median wall alternating in a minor image
with those on the other side; inner cells in internal longi-
tudinal view, 65-90 x 10.0 xl7.5 pm, without nodular
thickenings (Figure 3B); in cross section (Figure 2V) cells
faintly thickened on adaxial radial walls. Spores 12.5-15.0
pm diam., finely scabrate, pink with internal green areas.
Elaters 115-340 x ± 12.5 pm, reddish, with single spiral
band, 7.5-10.0 pm wide, the tapered tips at one or both
ends without spirals.
FIGURE 3. — Riccardia obtusa. A,
cells of epidermal layer of
wall of valve in external lon-
gitudinal view, X 389. B,
cells of inner layer of wall of
valve in internal longitudinal
view, X 389. A, B, Perold &
Koekemoer 4456.
184
Bothalia 32,2 (2002)
DISCUSSION
In his paper on the genus Riccardia in tropical Africa,
Jones (1956) remarked that he had not seen specimens
‘of R. campanuliflora S.Amell, R. capensis S.Arnell or
R. rhodesiae S.Amell (Botaniska Notiser 1952)’, but he
made no reference to R. obtusa S.W.Arnell, which was
published in the same paper as Arnell’s other three new
species. Later on, Engel (1975) identified a specimen
collected on Reunion as Riccardia cf. obtusa S.Arnell,
but added a question mark as follows: ‘Wet woods, Foret
de Bebour, 1300 m (?), lOllSg' (collecting number of
Harold E. Moore). It would appear that this altitude is
questionable.
Grolle (1995) refers to the above in his publication.
The Hepaticae and Anthocerotae of the East African
Islands. An annotated catalogue, excluding it from
Reunion. In Wigginton & Grolle (1996) it is observed
that R. obtusa needs to be clarified.
Bearing in mind Meenks’ (1987) admonition that
herbarium collections of Riccardia more than 20 years
old, are often useless, Arnell’s identification of three col-
lections of R. obtusa specimens is accepted, because he
had the advantage of studying them when fresh.
According to Arnell, this species resembles R. latifrons,
in that it has short, blunt branches. Schuster (1992)
described R. latifrons as closely adherent to the substrate
and subpalmately branched distally; the epidermal cells
of the thalli are, however, typically very large, always
thin-walled with colourless walls and the oil bodies are
absent near the meristematic tips and in all the epidermal
cells. It is also confined to the Northern Hemisphere.
There are, therefore, significant differences between
these two species.
R. obtusa specimens are distinguished by the follow-
ing characters: 1, the obtuse margins (hence the specific
name) of the distal branches; 2, the distal trifurcate to
subpalmate branching pattern; 3, the closely adjoining or
overlapping, lobulate apical branches; 4, the markedly
thickened ultimate segments of the main axes, 5-8(-ll)
cell rows thick; 5, the primary and secondary branches
with unistratose margins; 6, the prominent cell protru-
sions on the calyptra wall; 7, the distinct oil bodies,
which are very dark and irregular to kidney-shaped or
subspherical.
SPECIMENS EXAMINED
Arnell 1602 (holotype), Deepwall (= Diepwalle) Forest Reserve, on a
wet sandstone slope (BOL), (isotype) (S), 1620 Deepwall Forest, wet
slope (S), 1628, near Parkes Station, Knysna, wet sandstone slope
(BOL).
Esterhuysen 24255, N side of Zitzikamma Mts, near Joubertina, wet
shaded rock at side of stream on steep slope (BOL).
Koekemoer 2246, Buffelskloof Nature Reserve, south of Lydenburg, in
stream south of Research Cottage, on roots and stem of tree fern (PRE).
Perold, Burgoyne & Smithies 4742, 4743, Long Tom Pass, near
‘Staircase’, ± 25 km from Lydenburg, in gulley down steep slope
(PRE).
Perold & Koekemoer 4456, 4457, 4460, 4462, Valley of Ferns, on rot-
ting logs, soil or stone along footpaths, 4592, Kirstenbosch, in dark
gorge above ladders (PRE).
Scheipe 5154, ‘Braco’, Karkloof, Natal, on fallen logs in forest (BOL).
ACKNOWLEDGEMENTS
The curators of BOL, G and S are thanked for the loan
of specimens. I also express my sincere gratitude to Dr
M. Koekemoer, curator of PRE, and to the ladies P.
Burgoyne & S. Smithies, for all their assistance with
fieldwork, as well as to the referees for their helpful sug-
gestions and advice. The artist, Ms G. Condy, the pho-
tographer, Mrs A. Romanowski and the typist, Ms D.
Maree are thanked for their valued contributions.
REFERENCES
ARNELL, S.W. 1952. South African species of Riccardia. Botaniska
Notiser 1952: 138-156.
ARNELL, S.W. 1963. Hepaticae of South Africa. Swedish Natural
Science Council, Stockholm.
ENGEL. J.J. 1975. Hepaticae and Anthocerotae collected by Dr. Harold
E. Moore Jr in New Caledonia, Seychelles, Mauritius and Reunion
in 1972. The Biyologist 78: 361, 362.
GROLLE, R. 1995. The Hepaticae and Anthocerotae of the East
African Islands. An annotated catalogue. Bryophytonim Biblio-
theca 4^:
JONES, E.W. 1956. African hepatics XL The genus Riccardia in trop-
ical Africa. Transactions of the British Biyological Society 3:
74-84.
MEENKS, J.L.D. 1987. Studies on Colombian cryptogams XXVIII. A
guide to the tropical Andean species of Riccardia (Hepaticae).
Journal of the Hattori Botanical Laboratory 62: 161-182.
SCHUSTER, R.M. 1992. The Hepaticae and Anthocerotae of North
America 5: 1-854. Field Museum of Natural History. Chicago.
WIGGINTON, M.J. & GROLLE. R. Supplemented by GYARMATl.
A. 1996. Catalogue of the Hepaticae and Anthocerotae of sub-
Saharan Africa. Biyophytorum Bibliotheca 50. Cramer, Berlin,
Stuttgart.
Bothalia 32,2: 185-199 (2002)
Notes on African plants
ASTERACEAE-ANTHEMIDEAE
REDUCTION OF FOVEOLINA ALBIDA TO FOVEOLINA DICHOTOMA
In the past it has been difficult to distinguish between
Foveolina albida (DC.) Kallersjb and F. dichotoma
(DC.) Kallersjd, resulting in a rather confused collection
of herbarium specimens. These two species were consid-
ered species of Matricaria by Harvey in Flora capensis
(1865). Harvey (1865) regards Matricaria dichotoma
(DC.) Eenzl ex Harv. to be near M. albida (DC.) Fenzl ex
Harv., but the leaves are less compound. In M. dichotoma
the leaves are simple pinnatisect, the lobes flat and lin-
ear, whereas in M. albida the leaves are bipinnatisect.
and the lobes and lobules narrow. Kallersjd (1988) dis-
tinguishes between the two species on the shape of the
corolla of the disc florets. In F. dichotoma, the corolla
limb is broadly campanulate and the tube conspicuously
dilated at the base. In F. albida on the other hand, the
limb is not broadly campanulate and the tube is almost
cylindrical. After careful scrutiny of all the specimens of
these two species in NBG, no distinct difference between
the two species could be found. Sometimes in a single
specimen both simple pinnatisect and bipinnatisect
leaves occur. Similarly the limb of the corolla can be
broadly to narrowly campanulate and the tube may vary
from almost cylindrical to dilated at the base within one
specimen. In the distinguishing characters of Harvey
(1865) and Kallersjd (1988) there is a gradation from the
one form to the other. Since these two species are con-
specific, F. albida is placed into synonomy under F.
dichotoma. Both species were described in the same pub-
lication under the different genera (De Candolle 1838).
Since F. dichotoma is the type species of the genus
(Kallersjd 1988), it is therefore chosen here to represent
the species.
Foveolina dichotoma {DC.} Kdllersjd in Botanical
Journal of the Linnean Society 96: 319 (1988). Pentzia
dichotoma DC.: 138 (1838). Matricaria dichotoma (DC.)
Fenzl ex Harv.: 167 (1865). Type: ad Caput Bonae Spei
prope Olifantsrivier, Drege s.n. (G-DC, holo.; K!, S).
Tanacetwn alhidiim DC.: 132 ( 1838). Matricaria albida (DC.) Fenzi
ex Harv.: 166 (1865). Pentzia albida (DC.) Hutch.: 250 (1917).
Foveolina albida (DC.) Kallersjo: 319 (1988). Type: ad Caput Bonae
Spei in Klein Namaqualand. Drege s.n. (G-DC, holo.-PRE, microfiche!).
Pentzia annua DC.: 138 (1838). Type: ad Caput Bonae Spei in
regione Gariepina. Drege s.n. (G-DC, holo.-PRE, microfiche!).
Matricaria hirsulifoUa S. Moore: 1019 (1904). Type: Gross-
Namaland. Dinter 1221 (not seen).
REFERENCES
DE CANDOLLE, A.P. 1838. Compositae. Prodroimis 6. Treuttel &
Wiirtz. Paris.
HARVEY, W.H. 1865. Compositae. Flora capensis 3: 44-530. Hodges,
Smith, Dublin.
HUTCHINSON. J. 1917. Notes on African Compositae 3. Kew Bulletin
1916: 241-254.
KALLERSJO, M. 1988. A generic reclassification of Pentzia Thunb.
(Compositae-Anthemideae) from southern Africa. Botanical
Journal of the Linnean Society 96: 299-322. '
MOORE, S. 1904. In H. Schinz, Beitrage zur kenntnis der
Afrikanischen Flora. Bulletin de I'Herbier Boissier, ser. 2, 4:
995-1025.
J.B.P. BEYERS*
* Compton Herbarium, National Botanical Institute. Private Bag X7,
7735 Claremont. Cape Town.
MS. received: 2002-03-07.
ASTERACEAE
A NEW SPECIES OF ARCTOTHECA FROM NORTHERN CAPE, SOUTH AFRICA
Arctotheca marginata Beyers, sp. nov., habitu A.
prostratae (Salisb.) Britten similis sed foliis lineari-ellip-
ticis vel lineari-obovatis, adaxiale tomentosis, abaxiale
coactis, margine erosa revoluto distincte aculeato,
involucro abaxiale coacto, pappo flosculorum discorum
squamis 7-10 hyalinis differt.
TYPE. — Northern Cape, 3119 (Calvinia): Nieuwoudt-
ville, Farm Grootvlei, along edge of pan, ± 760 m, (-AC),
2M\\-2m,Bosenberg 1 (NBG, holo.; BOL, K, MO, PRE).
Perennial, tufted, stoloniferous herb. Stem prostrate,
rooting at nodes. Leaves alternate, petiolate; blade linear-
elliptic or linear-obovate in outline, 35-75 x 3-10 mm,
adaxially tomentose, abaxially felted, margin erose, revo-
lute and distantly aculeate; petiole 25-70 x 2-3 mm, up to
7 mm wide at base, felted. Capitula heterogamous, radiate,
solitary on long peduncles; peduncles 40-80 mm long, felt-
ed. Involucre widely cup-shaped, 10-13 mm diam.
Involucral bracts 5- or 6-seriate, imbricate; outer narrowly
ovate or narrowly oblong with acute apex, 2.8— 4.7 x
0.9-1. 0 mm, coriaceous, abaxially felted; inner gradually
larger, narrowly elliptic to elliptic with acute apex, coria-
ceous, abaxially felted; innermost elliptic to obovate, 6-9 x
2.2-2. 3 mm, with broad scarious margin, central coria-
ceous portion abaxially tomentose, apex rounded to obtuse.
Receptacle flat, honeycombed, fimbrilliferous. Ray florets
neuter, 13 or 14; tube cylindrical, 2.0-2.4 mm long; lami-
na spreading, narrowly elliptic, 8. 7-9. 2 x 1. 7-2.0 mm, yel-
low above, reddish brown below, 4-veined, apically
minutely 3-lobed; vestigial staminodes 2—4; ovary with
aborted style; pappus absent. Disc florets bisexual, numer-
ous; corolla yellow, narrowly cyathiform tapering into tube
below, 3.4-4.4 x 1.0-1. 2 mm; corolla lobes 5, ascending.
186
Bothalia 32.2 (2002)
FIGURE I, — Arctotheca marginata, Bosenberg I (NBG). A, portion of a plant, x 1; B. portion of leaf blade; C & D, involucral bracts (abaxial
view): C, outer; D, innermost. E, ray floret, E-I, disc floret: G, stamens; H, stamen and style; I, style. Scale bars: B, 4 mm; C-F, 2 mm;
G-l, 2 mm. Artist: Inge Oliver,
ovate-triangular, up to 1.3 x 0.5-0.8 mm; stamens 5;
anthers 2. 0-2. 2 mm long, linear, with sterile flat,
depressed-ovate apical appendage, base triangular; fila-
ment scabrous; style 4.9-5. 7 mm long, terete, thickened
above, bifid, style branches up to 0.2 mm long; cypsela
narrowly ellipsoid, 1.5-1. 7 mm long, densely pilose, hairs
tending to cohere; pappus scales hyaline, 7-10, narrowly
elliptic,! .0-1 .3 x 0.3-0.5 mm, apex acute. Figure 1.
Diagnostic characters'. Arctotheca marginata is mor-
phologically rather close to A. prostrata due to the
stoloniferous growth, but is distinguished from it by the
leaves which are linear-elliptic or linear-obovate in out-
line with an erose, revolute and distantly aculeate mar-
gin, tomentose above and felted beneath, and the involu-
cre which is abaxially felted. In A. prostrata the leaves
are lyrate-pinnatifid, green above, white-woolly beneath,
the involucre is glabrous or thinly tomentose with white-
woolly apices and the pappus is absent.
Distribution and ecology: Arctotheca marginata is
known only from the Farm Grootvlei in Nieuwoudtville.
It occurs in deep, waterlogged, sandy soil along the edge
of a pan. Figure 2.
Bothalia 32,2 (2002)
187
Etymology: the specific epithet, marginata, meaning
edge, was prompted by this species growing on the edge
of the pan.
Other specimen examined
NORTHERN CAPE. — 3119 (Calvinia): Nieuwoudtville, Farm
Grootvlei. along edge of pan, ± 760 m, (-AC), 7-1 1-2001, Roux 3168
(NBG).
ACKNOWLEDGEMENTS
I would like to thank Dr Ted Oliver for the Latin diag-
nosis, and Mrs Inge Oliver for the line drawings.
J.B.P. BEYERS*
* Compton Herbarium, National Botanical Institute. Private Bag X7,
7735 Claremont, Cape Town.
MS. received; 2002-02-04.
MESEMBRYANTHEMACEAE
MESEMBS WITH NUT-LIKE SCHIZOCARPIC FRUIT AND RUSCHIANTHEMUM FRIEDRICH SUNK
UNDER STOEBERIA DINTER & SCHWANTES
INTRODUCTION
Eruit of most Mesembryanthemaceae are hygrochas-
tic: they open, disperse seed and close again in response
to moisture. This moisture-mediated mode of seed dis-
persal is achieved through a sophisticated mechanism
based on the hygroscopic properties of expanding keels
that function to open and close the valves or lids of
loculicidal capsules (Hartmann 1988; Croizat 1993). In
the more complex hygrochastic fruit, the amount of seed
dispersed and their trajectories, are controlled through
various structures such as covering membranes and clos-
ing bodies (Parolin 2001). However, modes of dispersal
in Mesembryanthemaceae vary widely depending on the
construction of the fruit. The genus Carpohrotus N.E.Br.,
for example, has edible berry-like fleshy fruit (sour fig),
with seeds embedded in thick, sticky mucilage. A few
genera possess xerochastic fruit that dehisce when dry.
XEROCHASTIC FRUIT
Mesemb genera with fruit which open when dry are
found mostly in the tribe Apatesieae emend. Chesselet,
G.F.Sm. & A.E.van Wyk (Chesselet et al. 2001 ). Species
of Conicosia N.E.Br., for example, have capsules which
function as shakers, in a way similar to that of poppies.
In addition to having seeds in their locules, the fruit of
Conicosia and Skiatophytum L. Bolus have seed cham-
bers or pockets, the ‘Samentaschen’ of Schwantes (1949,
1957). in which a few seeds are entombed in woody tis-
sue. A few genera possess schizocarpic fruit which break
up into mericarps when dry. The terms schizocaip and
mericarp are not used in their strictest sense here,
because the mericarps of mesembs are formed by two
halves of neighbouring carpels (Leistner 1958). The
genus Hymenogyne Haw. has schizocarpic fruit which
break up into 8-12 one-seeded, flat, broadly winged, cir-
cular mericarps. Seeing that the fruit of certain species of
Conicosia may disintegrate in a comparable way,
Schwantes (1927) erected the genus Herrea which is no
longer considered distinct. The retention of seeds in seed
pockets outside the locules was mentioned above for
Conicosia and Skiatophytum. Among the Apatesieae this
syndrome has reached its highest degree of development
in Caryotophora skiatophytoides Leistner (Figure 3G,
H). Its fruit is a schizocarp which breaks up into 3 or 4
nut-like mericarps. Two genera outside the Apatesieae
also encompass species with seed enclo.sed in hard,
woody, nut-like mericarps: Brownanthiis and Ruschian-
themiim.
EVOLUTION OF NUT-LIKE MERICARPS
The evolutionary development of nut-like fruit from
hygrochastic capsules may be regarded as a convergent
adaptive feature, since this has occun'ed more than once
in the evolution of the Mesembryanthemaceae, with
Ruschianthemum gigas (Dinter) Friedrich as an example
from subfamily Ruschioideae (Figure 3A-D), and
Pseudobrownanthus niicifer Ihlenf. & Bittrich from sub-
family Mesembryanthemoideae (Figure 3E, F). We spec-
ulate that in these two species, the evolution of nuts may
be a seed-protecting mechanism that has evolved under
the extreme arid conditions that these species experience,
both taxa being restricted to southern Namibia and the
Richtersveld, South Africa. From a structural perspec-
tive, Hartmann ( 1988) ascribes the evolutionary deriva-
tion of nuts or nutlets in the mesembs to a process of
increased sclerenchymatization of all tissues in the fruit.
This phenomenon is supposedly associated with the evo-
lutionary replacement of raindrops by wind as the prime
dispersal agent in species exhibiting these characteristics
(Hartmann 1988, 2001). However, diaspores are not
accompanied by wings or plumes and the evolutionary
change to nutlets may rather be an adaptation to larger
seed size which may confer a competitive advantage to
the seedlings, especially in areas with unpredictable fol-
low-up rains. In Caiyotophora skiatophytoides Leistner
(Figure 3G, H) and in Skiatophytum tripolium (L.)
L. Bolus (Figure 31), the selective pressures are different
from those acting in the arid parts of Namibia and South
Africa. Both species occur in the fynbos vegetation of
Western Cape, South Africa, with C. skiatophytoides
from near Bredasdorp and S. tripolium from the Cape
Peninsula and sunounding areas. Ecological factors that
characterize this Mediterranean climate region include
summer aridity, mineral-poor soils, wind and fire. C. ski-
atophytoides is only found in post-fire vegetation. This
perennial plant is known to resprout from suckers fol-
lowing fire, whereas the annual S. tripolium reseeds after
fire. In both cases seeds germinate with difficulty
(Hickey & Van Jaarsveld 1995) and the nut-like fruit of
188
Bothalia 32,2 (2002)
FIGURE 3. — Mesembs with nut-iike schizocarpic fruit. A-D, Stoeheria gigcis', E, F. Brownanthiis luicifer, G. H, Caiyotophora skiatopliytoicles:
I, Skiatophytiim Iripolium. Scale bars: A-C. E-1, 5 mm; D. 1 mm. Artist: R Chesselet.
C. skiatophytoides decay slowly in habitat. Hickey &
Van Jaarsveld ( 1995) suggest that delayed seed germina-
tion is a survival strategy commonly found in fynbos
plants.
UROWNANTHUS. RUSCHIANTHEMUM AND STOEBERIA
The unusual nut-like fruit of Riiscliiaiitlieimim gigas
and of Pseudohrownanthus uucifer were thought to be
sufficient reason to establish the monotypic genera
Ruschianthemum and Pseiidohwwiumtluis (Friedrich
I960; Ihlenfeldt & Bittrich 1985). The widely accepted
importance of fruit structure in the taxonomic subdivi-
sion of the family provided justification for these gener-
ic separations e.g. Dehn (1992). however, increasingly,
fruit types are not regarded as superior characters for
generic delimitation and all characters need to be consid-
ered (Klak 2001). In a recent study of Brownandms,
Pierce & Gerbaulet ( 1997) argued that P. uucifer is but a
specialized member oi Browmmdnis which now contains
12 species, including the new combination Browmtuthus
niicifer (\h\enf. & Bittrich) S.M. Pierce & Gerbaulet. This
taxonomic decision is supported by molecular data, and
Pseudohrownanthus is deeply embedded within
Brownanthiis (C. Klak. BOL, University of Cape Town,
pers. comm.), suggesting that the nut-like fruit represents
an autapomorphy for the species rather than a character
providing resolution at genus level. The relative impor-
tance assigned to a character set, such as fruit structure,
Bothalia 32.2 (2002)
189
and especially its relevance at a particular rank in the
taxonomic hierarchy, needs to be carefully considered
when establishing new genera based on autapomorphies.
In an analysis of the genera of the Mesembryanthe-
maceae, Chesselet et al. ( 1995) showed a high number of
monotypic genera in this family. Many taxonomists have
misgivings about very large genera and genera with only
one species, yet both are integral to classification
schemes (Williams 1964). But, some monotypic genera
are founded on peculiarities, and under such circum-
stances genus and species concepts may converge, as in the
case of Pseudohrownanthiis. In this paper, we use simi-
lar argumentation to resurrect Stoeberia gigas (Dinter)
Dinter & Schwantes.
The type of Ruschianthemiim gigas (Dinter) Friedrich
was collected in the Klinghardt Mountains of southern
Namibia in September 1922 by Moritz Kurt Dinter, a
renowned botanist and botanical explorer of Namibia.
Prior to the establishment of Ruschianthemiim, Mesem-
bryanthemiim gigas Dinter (Dinter 1923) was placed in
the genus Stoeberia Dinter & Schwantes emend. Friedrich,
emend. Dehn (Schwantes 1927-1928; Friedrich 1960;
Dehn 1992), together with S. beetzii (Dinter) Dinter &
Schwantes and S. nipis-arcuatae (Dinter) Dinter &
Schwantes, a species that is now classified in the genus
Amphibolia L. Bolus. Ruschianthemiim was established
by Friedrich (1960) on account of its unusual fruit struc-
ture— the mature fruit of R. gigas is a five-locular schizo-
carp which breaks into nutlets consisting of parts of the
septum enclosing one seed and the remains of valve
wings and expanding keels adhering to each unit.
Following the release of the nutlets, persistent dorsal and
apical connecting vascular bundles present in the capsule
remain on plants as a fibrous, basket-like skeleton (see
Smith et al. 1998: 363; Burgoyne 2000: 8). Although
used as a diagnostic feature of Ruschianthemiim (Dehn
1992), similar basket-like fruit remains have now also
been noted in Stoeberia frutescens (L. Bolus) Van Jaarsv.
When not in fruit, R. gigas may be easily mistaken for a
Stoeberia which it closely resembles (Van Jaarsveld
1994).
Ruschianthemiim has been classified together with
Stoeberia, Amphibolia and Eberlanzia Schwantes in the
"Eberlanzia Group’ by Hartmann (1998). This group
shares the following characteristics: stems whitish; cap-
sules with valve wings and closing bodies which are
mostly small and often deep inside the locules so that
they appear absent. In this group, Ruschianthemiim is
closest to Stoeberia (Hartmann 2001), sharing with it the
nearly club-shaped leaves and the rich dichasial inflores-
cences. The flowers of Ruschianthemiim closely resem-
ble those in the genus Stoeberia — they are relatively
small and numerous, with pink-tipped filamentous sta-
minodes arranged in a cone; S. carpii Friedrich with its
large white flowers, is unique in the genus but fruit and
vegetative characters support its current placement in
Stoeberia. Both genera have the lophomorphic holonec-
tary characteristic of genera placed in Tribe Ruschieae
Schwantes by Chesselet et al. (2001, in press).
In Schwantes’ (Schwantes 1927-1928) key to
mesemb genera, Stoeberia is distinguished from other
genera by its capsule with rudimentary covering mem-
branes that are developed as a narrow rim, with valve
wings, and very large placental tubercles (closing bod-
ies) and stigmas that are short, dark and feathered. In the
present circumscription of Stoeberia the valves do not
close again completely once they have opened, and hard
(sclerified) valve wings and recurved valve rims charac-
terize the fruit (Chesselet et al. 2000). If we amend these
diagnostic characters to sclerification of fruit tissue in a
broader sense, we can accommodate R. gigas in
Stoeberia and reassign this species accordingly.
Stoeberia gigas (Dinter) Dinter & Schwantes in
Zeitschrift fur Sukkulentenkunde 3: 17 (1927).
Mesembryanthemum gigas Dinter in Feddes Repertorium 19; 153
(1923). Ruschianthemum gigas (Dinter) Friedrich in Mitteilungen der
Botanischen Staatssammlung. Miinchen 3: 564 (1960). Type: Namibia.
Klinghardt Mountains. September 1922. Dinter 3791 (B!. holo.).
REFERENCES
BURGOYNE. RM. 2000. Will the real Ruschianthemiim gigas please
stand up! Aloe 37: 8.
CHESSELET. R. MOSSMER. M. & SMITH. G.F. 1995. Research pri-
orities in the succulent plant family Mesembryanthemaceae
Fenzl. South African Journal of Science 91: 197-209.
CHESSELET, R, SMITH, G.F., BURGOYNE, R.M., KLAK, C„ HAM-
MER. S.A.. HARTMANN, H.E.K., KURZWEIL, H., VAN
JAARSVELD, E.J., VAN WYK, B-E., & LEISTNER, O.A.
2000. Mesembryanthemaceae. In O.A. Leistner, Seed plants of
southern Africa; families and genera. Strelitzia 10: 360-410.
CHESSELET, R, SMITH, G.F. & VAN WYK, A.E. 2001. A new trib-
al classification for the Mesembryanthemaceae Fenzl based on
characters of the floral nectary. Aloe 38: 25-28.
CHESSELET. R, SMITH. G.F. & VAN WYK, A.E, in press. A new
tribal classification of the Mesembryanthemaceae: evidence
from floral nectaries. Taxon.
CROIZAT, L, 1993, On the structural and developmental history of the
capsule of Mesembryanthemum s.l. Kirkia 14: 145-169.
DEHN, M. 1992. Untersuchungen zum Verwandtschaftskreis der
Ruschiinae (Mesembryanthemaceae Fenzl). Mitteilungen aus
dem Institut fiir AUgemeine Botanik. Hamburg 24: 91-198.
DINTER, K. 1923. Beitriige zur Flora von Siidwestafrika. Feddes
Repertorium 19: 153.
FRIEDRICH, H.-Chr. 1960. Mesembryanthemenstudien 1. Beitrag zur
Kenntnis der Gattungen Stoeberia Dtr. & Schw. und Ru.schi-
anthemum Friedr. gen nov. Mitteilungen der Botanischen
Staatssammlung, Miinchen 3: 554—567.
HARTMANN, H.E.K. 1988. Fruit types in Mesembryanthema.
Beitrdge zur Biologic der Pfianzen 63: 313-349.
HARTMANN, H.E.K. 1998. A letter to the MSG. Mesemb Study Group
Bulletin 2: 32-34.
HARTMANN, H.E.K. 2001. Illustrated handbook of .succulent plants:
Aizoaceae F-Z. Springer- Verlag, Berlin.
HICKEY, B. & VAN JAARSVELD, E. 1995. Rropagation of three rare
and endangered vygies. British Cactus and Succulent Journal
13: 141-146.
IHLENFELDT, H-D. & BITTRICH, V. 1985. Morphologie, Gliede-
rung und Abgrenzung der Gattung Psilocaulon N.E.Br. s.l.
(Mesembryanthemaceae). Botanische Jahrbiicher 105: 289-322.
KLAK, C. 2001. Phylogeny and character evolution in the
Lampranthus group (Aizoaceae-Ruschioideae). Rh.D. thesis.
University of Cape Town.
LEISTNER, O.A. 1958. A new monotypic genus of the Mesem-
bryanthemaceae. In H.M.L. Bolus, Notes on Mesembryanthe-
mum and allied genera 3; 289-291.
RAROLIN, R. 2001. Seed expulsion in fruits of Mesembryanthema
(Aizoaceae): a mechanistic approach to study the effect of fruit
morphological structures on seed dispersal. Flora 196:
313-322.
RIERCE, S.M. & GERBAULET, M. 1997. Brownanthus Schwantes
(Mesembryanthemoideae, Aizoaceae); two new species and a
new combination from the Richtersveld and southwestern
Namibia. Aloe 34: 42-44.
190
Bothalia 32,2 (2002)
SCHWANTES. G. 1927. Einige neue Mesembrianthemaceen. Mailers
Deutsche Gdrtner-Zeitung 42: 436.
SCHWANTES. G. 1927-1928. Zur Systematik der Mesembrian-
themen. Zeitschrift fur Sukkulentenkunde 3: 16-30.
SCHWANTES, G. 1949. Biology and systematics of Mesembryan-
themum. The National Cactus & Succulent Journal, December
1949: 73, 74.
SCHWANTES, G. 1957. Flowering stones and mid-day flowers. Ernest
Benn, London.
SMITH, G.F., CHESSELET, R, VAN JAARSVELD, E.J., HART-
MANN, H., HAMMER, S., VAN WYK, B.-E., BURGOYNE,
R. KLAK, C. & KURZWEIL, H. 1998. Mesembs of the world.
Briza Rublications, Rretoria.
VAN JAARSVELD, E. 1994. A synopsis of the genus Stoeberia. Aloe
31: 68-76.
WILLIAMS, C.B. 1964. Patterns in the balance of nature and related
problems in quantitative ecology. Academic Rress, London.
R. CHESSELET* and A.E. VAN WYK**
* Compton Herbarium, National Botanical Institute, Rrivate Bag X7,
7735 Claremont. Cape Town. E-mail: chesselet@nbict.nbi.ac.za;
http://www.nbi.ac.za
** H.G.W.J. Schweickerdt Herbarium, Department of Botany, University
of Rretoria, 0002 Rretoria. E-mail: avanwyk@scientia.up.ac.za
MS. received: 2001-10-31.
HYACINTHACEAE: MASSONIEAE
A NEW SRECIES OF LACHENALIA FROM NAMAQUALAND, SOUTH AFRICA
INTRODUCTION
Lachenalia Jacq.f. ex Murray is the largest endemic
genus within the southern African Hyacinthaceae and
comprises ±117 species. The distribution of the genus is
concentrated in the winter rainfall zone of the subconti-
nent, but also extends into areas of intermediate as well as
summer rainfall (Duncan 1998a). All its species follow a
strictly winter-growing, summer-dormant Ufe cycle, with
the exception of Lachenalia pearsonii (Glover)
W.F.Barker from southern Namibia, which is surmner-
growing and winter-dormant (Duncan 1999). The genus is
exceptionally diverse in flower shape, colour and orienta-
tion, as well as in leaf shape, surface sculpturing and
markings, and is cytologically also very varied. Lache-
nalia comprises several very widely distributed species,
but a much larger number are confined to narrow distribu-
tion ranges, such as the species described here. The genus
occurs across a wide variety of habitats and vegetation
types, including Succulent Karoo, Nama-Karoo, Fynbos
and Grassland Biomes. Many of the species have great
ornamental value (Duncan 1988, 1989a, b), and hybrids
developed by the Agricultural Research Council at
Roodeplaat near Pretoria, have recently been introduced
onto the international pot plant market. The new species
described here forms part of a series of papers towards a
revision of the genus (Duncan 1996, 1997, 1998b).
Lachenalia valeriae G.D. Duncan, sp. nov.
Planta 100-350 mm alta; bulbus subglobosus, 15-20
mm diam., albus, tunicis externis tenuibus brunneis; folia
2, late lanceolata ad anguste ovata, 120-170 x 20-30 mm,
patentia, pagina superiore venis longitudinaliter depressis
pustulis minutis dense tecta; flores oblongo-urceolati,
sessiles, patentes, pallide viridiflavi, tubo perianthii cre-
meo 2-3 mm longo, tepalis exterioribus ovatis 7-8 x 4-5
mm, tepalis interioribus obovatis, apicibus parum recur-
vatis, tepalis duobus superioribus imbricatis 9-10 x 5
mm, tepalo inferiore longiore angustioreque 10—11 x 3^
mm, in parte superiore vivide magenteo; stamina inclusa
declinata 7 mm longa.
TYPE.— Northern Cape, 2917 (Springbok): north-
western Namaqualand, Klein.see Nature Reserve, Kleinsee,
on west-facing granite slopes in brownish red sand,
(-CA), Duncan 444 (NBG, holo.).
Deciduous, wintes^growing geophyte, 100-350 mm
high. Bulb subglobose, 15-20 mm dianr, usually soli-
tary, occasionally duinp-forraing, white with thin, mem-
branous, pale to dark brown outer tunics, produced into a
very short neck; cataphyll subterranean, translucent
white with minute longitudinal veins, loosely clasping
leaf base. Leaves 2, broadly lanceolate to narrowly ovate,
120-170 X 20—30 mm, spreading or suberect, canalicu-
late, upper surface dark green with distinct depressed
longitudinal veins, densely covered with minute, dark
green pustules, lower surface plain or lightly to heavily
flushed with maroonish magenta; clasping leaf base
20-50 mm long, yellowish green or heavily flushed with
dark maroonish magenta above soil level, shading to
white below soil level. Inflorescence an erect, many-
flowered dense spike up to 120 mm long with short ster-
ile tip; peduncle erect, sturdy, up to 230 mm long, lower
half pale green with minute brownish purple speckles,
upper half heavily mottled with brownish purple; rachis
pale purplish brown in lower half, shading to electric-
blue in upper half and at tip; bracts much reduced, ovate
throughout inflorescence, 1-2 x \-A mm. Flowers ses-
sile, suberect in bud stage, spreading at flowering stage,
oblong-urceolate, pale greenish yellow, with conspicu-
ous, pale to bright magenta lower, inner tepal; perianth
tube cup-shaped, cream-coloured, with or without a very
pale blue tinge, 2-3 mm long; outer tepals ovate, 7-8 x
4-5 mm, cream-coloured at base, shading to yellowish
green above, with bright green keels and gibbosities;
inner tepals obovate, tips slightly recurved, protruding
well past outer tepals, upper inner tepals translucent dull
white, overlapping, 9-10 x 5 mm, with bright green
keels; lower inner tepal deeply canaliculate, 10-11 x 3^
mm, lower half translucent dull white, upper half pale to
bright magenta with pale greenish yellow keel. Stamens
included within perianth, declinate; filaments white, 7
mm long; anthers dull maroon prior to anthesis, yellow at
anthesis. Ovary ellipsoid, 3^ x 2 mm, pale green; style
6-7 mm long, white. Capsule ellipsoid, 8x5 mm, bright
green. Seed ovoid, E7 x 1.0 mm, shiny black with short,
ridged strophule, up to 0.4 mm long. Flowering time: late
July to mid-August. Figure 4.
Bothalia 32,2 (2002)
FIGURE 4. — Lachenalia valeriae. A, foliage and inflorescence; B,
bulb; C. single flower; D, L/s flower. Scale bars; A, B, 10 mm;
C, D, 5 mm. Artist; Vicki Thomas, drawn from type, Duncan
444 (NBG).
Etymology: Lachenalia valeriae is named for Valerie
Fay Anderson (Mrs R. Geary-Cooke), in recognition of
the wonderful contribution she has made to the knowl-
edge of the flora of southern Africa, through the medium
of watercolour paintings.
Diagnostic features and affinities: L. valeriae falls
into the small group of species with sessile, urceolate or
oblong-urceolate flowers with included, declinate sta-
mens, its closest relatives being L. framesii W.F.Barker
and L. camosa Baker. L. valeriae is recognized by its
moderately dense inflorescence of patent, oblong-urceo-
late, pale greenish yellow flowers with the deeply
canaliculate lower inner tepal distinctly longer than the
two upper lateral tepals, and conspicuously marked with
pale to bright magenta in the upper half (Figure 4). The
inner tepals are all slightly recurved at their tips, and the
outer tepals have bright green gibbosities and keels.
Mature bulbs always produce two opposite, spreading or
suberect, slightly canaliculate, broadly lanceolate or nar-
rowly ovate leaves with depressed longitudinal veins on
the upper surface, which is densely covered with minute,
rounded green pustules. The clasping leaf base is usual-
ly plain yellowish green but may also be heavily flushed
with dark maroonish magenta. The white, subglobose
bulb is surrounded by pale to dark brown, membranous
191
outer tunics and its relatively large, ovoid seeds have a
shiny black testa and a short, ridged strophule.
Lachenalia framesii resembles L. valeriae in the
shape of its yellow or greenish yellow, oblong-urceolate
flowers with the upper part of the inner tepals recurved,
and in the similar pale to bright blue upper portion of the
rachis, but its flowers nre mwch smaller and the upper
part of its inner tepals are all pale to bright magenta,
whereas in L. valeriae only the lower inner tepal has this
colouring. L. framesii also differs in being a dwarf
species with much shorter, suberect flowers, and it has
much shorter, suberect, canaliculate, narrow-lanceolate
leaves with strongly undulate margins, and the upper leaf
surface is always smooth. Its bulb is globose and much
smaller than that of L. valeriae, and its globose seeds are
minute in comparison, and have a reticulate, matt black
testa. L. framesii is a common species throughout central
Namaqualand and the Knersvlakte, and its distribution
extends close to that of L. valeriae in the Komaggas
Flower Reserve, but does not overlap that of L. valeriae,
which occurs west of this area. It usually occurs on flats
in large colonies in quartzitic sand.
The similar urceolate flowers of L. carnosa are short-
er and wider than those of L. valeriae and have dull white
perianth segments, with all the inner tepals having broad,
pale to dark mauve tips, and the gibbosities dark purplish
brown. Its two spreading or suberect leaves differ in
being broadly ovate and flat, with a distinct purplish
maroon, cartilaginous margin, whereas those of L. vale-
riae are broadly lanceolate to narrowly ovate, and slight-
ly concave. The upper surface of L. carnosa is almost
always smooth but may occasionally have irregularly
scattered, large flattened brown pustules, whereas those
of L. valeriae are always covered with numerous small
green, rounded pustules. The bulb tissue of L. carnosa is
usually pale to dark yellow, surrounded by strong, dark
brown tunics, whereas that of L. valeriae is always
white. The similarly shaped, ovoid seeds of L. camosa
are less than half the size of those of L. valeriae, and like
this species, also have a short, ridged strophule. L. car-
nosa is a very common species in central and western
Namaqualand, and the Kamiesberg, where it usually
grows in cracks and depressions of granite outcrops in
sandy soil, or less frequently in sandy gravel on open
flats. Its distribution does not overlap that of L. valeriae,
but comes close to it in the Komaggas area.
Distribution and habitat: L. valeriae is currently
known from five populations in the sandy coastal plain
of northwestern Namaqualand in the Succulent Karoo
Biome, where it occurs in shallow or deep brownish red
sand on east- and west-facing slopes of granite outcrops
(Figure 5). At its type locality near the mouth of the
Buffels River in the Kleinsee Nature Reserve, it occurs
only on west-facing slopes, near to or amongst low suc-
culent vegetation including Aloe framesii. Pelargonium
fulgidum and several Crassula and Euphorbia species,
but at a locality just north of Kleinsee it is found on both
east- and west-facing slopes. Individuals grow singly or
in colonies on rock depressions and in between rock
cracks (Figure 6), either in full sun or in partial shade of
the surrounding vegetation. The distribution of L. valeri-
ae needs to be further investigated north of Kleinsee, as
192
Bothalia 32,2 (2002)
FIGURE 5. — Distribution of Luchenatia valeriae.
it is likely that it extends all the way up the coast to the
mouth of the Holgat River, which is currently its north-
ernmost limit.
Materia! examined
NORTHERN CAPE. — 2816 (Oranjemund): Holgat River mouth,
1.6 km east of road, in sand dunes, (-DC), Wisiira 1610 (NBG). 2917
(Springbok): Kleinsee Nature Reserve, (-CA), Duncan 444 (NBG); 2
km N of Klein.see, (-CA), Duncan 448 (NBG); 8 km W of Komaggas,
(-CD), Lavranos 28585 (NBG).
ACKNOWLEDGEMENTS
We extend our thanks to Paul Kruger, environmental
officer for Namaqualand Mines at Kleinsee, as well as to
Ernst van Jaarsveld, Max Michael and Adam Harrower
for their assistance at various stages of this study, and to
Dr Ted Oliver for translating the diagnosis into Latin.
REFERENCES
DUNCAN, G.D. 1988. The Lachenalia handbook. Annals of Kirstenhasch
Hotanic Gardens 17. National Botanical Institute, Cape Town,
FIGURE 6. — Lachenalia valeriae, Kleinzee Nature Reserve. A, group
of flowering plants; B, natural habitat on west-facing granite
slopes. Scale bar: 10 mm.
DUNCAN, G.D. 1989a. Lachenalia. In B.J. Jeppe, Spring and winter
flowering hulhs of the Cape. Oxford University Press, Cape Town,
DUNCAN, G.D. 1989b, Lachenalia. In N.M. du Plessis & G.D. Duncan,
Bulbous plants of southern Africa. Tafelberg, Cape Town.
DUNCAN, G.D. 1996. Four new species and one new subspecies of
Lachenalia (Hyacinthaceae) from arid areas of South Africa.
Bothalia 26: 1-9.
DUNCAN, G.D. 1997. Five new species of Lachenalia (Hyacinth-
aceae) from arid areas of South Africa. Bothalia 27: 7-15.
DUNCAN, G.D. 1998a. Notes on the genus Lachenalia. Herhertia 53:
40-48.
DUNCAN, G.D. 1998b. Five new species of Lachenalia (Hyacinth-
aceae) from arid areas of Namibia and South Africa. Bothalia
28: 131-139.
DUNCAN. G.D. 1999, Lachenalia violacea. Curtis’s Botanical Maga-
zine 16: 252-255.
G.D. DUNCAN* & T.J. EDWARDS**
* National Botanical Institute, Kirstenbo.sch. Private Bag X7, 7735
Claremont. Cape Town.
** Botany Department, University of Natal, Private Bag XOl, 3209
Scottsville, Pietermaritzburg.
MS. received: 2002-01-24.
Bothalia 32,2 (2002)
193
ASTERACEAE
A DISTINCTIVE NEW SPECIES OF FELICIA (ASTEREAE) FROM WESTERN CAPE, SOUTH AFRICA
INTRODUCTION
Felicia Cass, is a genus of 83 species of annuals,
perennials and shrubs of tribe Astereae of the Asteraceae
(Grau 1973). The genus is centred in southern Africa but
a few species extend into tropical Africa as far north as
Nigeria, Ethiopia and the Arabian Peninsula (Bremer
1994; Herman et al. 2000). Felicia is distinguished from
related genera of the Astereae by an epaleate receptacle,
uniseriate pappus of numerous barbellate bristles, ecau-
date anthers and strongly compressed cypselas with
swollen margins. Most species have blue or mauve to
pink rays (rarely white or yellow) and yellow disc florets
(Grau 1973). The species described here, F. josephinae,
is remarkable in the genus for its broad white or cream-
coloured ray florets and deep purple disc florets.
Similarly coloured disc florets are otherwise known in
the genus only in F heterophylla (Cass.) Grau. Felicia is
well represented by 54 species in the Cape Floral Region
and F. josephinae was listed as Felicia sp. 1 in the
account prepared for the recent conspectus of the Cape
flora (Goldblatt & Manning 2000). Additional collec-
tions made in the spring of 2001 have enabled us to
describe it formally.
Felicia josephinae J.C. Manning & Goldblatt, sp.
nov.
Herba annua usque ad 200 mm alta hispida, folds
inferioribus oppositis oblanceolatis, capitulis het-
erogamis radiatis solitariis pedunculis usque ad 50 mm
longis trichomatibus glanduliferis vestitis insidentibus,
involucro anguste ovoideo ± 5 mm alto et 4 mm in
diametro, bracteis involucralibus biseriatis supra medi-
umtrichomatibus glanduliferis et setis sparsis omatis
marginibus membranaceis apicibus ciliatis, flosculis
radii 8 vel 9 ligula elliptico-oblanceolata, alba ad cremea
epapposa, flosculis disci 14 vel 15 atropurpureis cylin-
dricis supra dilatatis setis pappi ± 25 barbellatis et prope
medio subplumosis, cypselis obovatis compressis mar-
ginibus incrassatis trichomatibus adpressis vestitis pal-
lide brunneis.
TYPE. — Western Cape: 3218 (Clanwilliam), hills be-
tween Elands Bay and Leipoldtville, deep sands, (-BC),
15 Sept. 2001, Goldblatt & Porter 11887 (NBG, holo.;
K, MO, PRE, S, iso.).
Annual herb 150-200 mm high, branching near base;
stems sparsely leafy, hispid with short and long multicel-
lular hairs and with numerous shortly stipitate glands in
upper parts. Leaves decreasing in size up stem; lower
leaves opposite, soon withering, oblanceolate, 30-70 x
6-13 mm, narrowed to petiole-like base, thinly hispid
with long multicellular hairs; upper leaves mostly alter-
nate, becoming progressively lanceolate to linear, hispid
with long and short multicellular hairs. Capitula heteroga-
mous, radiate, solitary on terminal and axillary peduncles
up to 50 mm long, with few, scattered, subulate bracts.
Involucre narrowly ovoid, ± 5 mm high and 4 mm diam.;
involucral bracts biseriate, 11-13, glandular-hairy and
thinly setose in upper half, 4-5 mm long, with scarious
margins and ciliate tips, outer bracts linear with narrow
margins, ± 1 mm wide, inner bracts oblanceolate with
broad margins, ± 1.5 mm wide. Receptacle flat, epaleate,
alveolate. Ray florets female, 8 or 9; tube cylindrical,
glandular-hairy, 2. 5-3.0 mm long; lamina spreading,
elliptic-oblanceolate, 10-12 x 4-6 mm, 4- veined, white
to cream-coloured. Ovary narrowly elliptic, adpressed-
hairy; style terete with linear branches, not or shortly
exserted, branches ± 1 mm long, acute, margins stigmat-
ic. Pappus absent. Cypselas obovate, 2. 5-3.0 x ± 1.5 mm,
flattened with thickened margins, faces and margins
covered with adpressed hairs, pale brown. Disc florets bi-
sexual, 14 or 15, deep purple, inner florets functionally
male; tube cylindrical but widening slightly in upper part,
glandular-hairy, 2. 5-3.0 mm long, limb obliquely obcon-
ical, 5-lobed; lobes triangular, recurved, ± 1 x 0.8 mm,
with thickened margins. Anthers ± 2 mm long including
ovate, somewhat keeled apical appendage, deep blue with
white pollen; anther base obtuse, ecaudate. Ovary nar-
rowly elliptic, adpressed-hairy; style terete, branches ± 1
mm long, purple, incurved, linear, flattened with stigmatic
margins and sterile, triangular, papillate apical
appendages. Pappus bristles uniseriate, ± 25, spreading in
fruit, caducous, 3-4 mm long, barbellate but subplumose
near middle, connate at base into short collar, white.
Cypselas as in ray florets. Flowering time: September and
October. Figure 7.
Distribution and biology: Felicia josephinae has only
been collected from a small area along the Western Cape
coast between Elands Bay and Lambert’s Bay, extending
inland to Sandberg, east of Leipoldtville (Figure 8). It
occurs on hills and dunes, where it is restricted to
Strandveld Succulent Karoo (Low & Rebelo 1996). This
strand vegetation occurs on deep, calcareous sands in a
belt along the coastal plain and is dominated by scattered
shrubs, especially Salvia lanceolata (Lamiaceae),
Nylandtia spinosa (Polygalaceae) and succulent species
of Euphorbia (Euphorbiaceae). Numerous annual species
flourish seasonally in the open spaces between the
shrubs, including Alonosoa unilabiata, Hemimeris race-
mosa, Lyperia tristis, Nemesia affinis and N. bicornis
(Scrophulariaceae), and Arctotis hirsuta, Dimorphotheca
pluvialis and Trichogyne verticillata (Asteraceae).
Although Strandveld Succulent Karoo extends along the
coast for some 500 km, F josephinae appears to be
restricted to the extreme southern part where the sands
are relatively shallower and overlie more solid sand-
stone-derived substrates.
History: F josephinae appears to have been first col-
lected in September 1933 by C. Louis Leipoldt, poet,
author, medical practitioner and plant collector, from
between Elands Bay (incorrectly transcribed on the
herbarium labels as Elandskloof) and Clanwilliam, on
the Western Cape coast. During the ensuing two decades
a handful of further collections were made by various
Cape botanists, all from the same small stretch of coun-
Bothalia 32,2 (2002)
FIGURE 7, — Felicia josephinae. A, whole plant. B, C, involucral bracts: B, inner; C, outer. D, E, ray floret: D, floret; E, cypsela. F-1, disc floret:
F, floret with pappus bristles removed from front; G, two stamens; H, stigmatic branches; I, cypsela (rear pappus bristles not shown). Scale
bars: A-C, 10 mm; D-1, 1 mm. Artist: John Manning.
try between Elands Bay and Lambert’s Bay. These early
collections were all identified either as a new species or,
rather tentatively, as a variant of Chareis heterophylla
Cass, [now Felicia heterophylla (Cass.) Grau]. None of
these collections was apparently seen by Grau (1973) for
his revision of the genus. F. josephinae first came to our
attention in 1995 when we found it on the Farm
Sandberg, east of Elands Bay, possibly at or very near the
spot at which Leipoldt had originally located it. It was
clear to us then that it was not a form of F. heterophylla.
We encountered it again in the spring of 2001, at which
time we took the opportunity of collecting material for
description and illustration. It gives us great pleasure to
name it for our colleague, Josephine Beyers, in recogni-
tion of her long interest in the Asteraceae.
Diagnosis and relationships: this distinctive, highly
floriferous species is readily distinguished from all other
species of Felicia by its broad, white or cream-coloured
rays and deep purple disc. The rounded plants become
Bothalia 32,2 (2002)
195
FIGURE 8. — Distribution of Felicia josephinae in Western Cape.
covered with dainty white flower heads that in their
broad rays and small, convex, purple discs are quite
unlike any other species of Felicia. The central disc
florets, which are functionally male, have somewhat
longer ovaries than the surrounding florets, with the
result that the disc is conspicuously convex and projects
prominently above the rays. F. josephinae is one of a
handful of annual species in section Neodetris Grau that
have a strictly biseriate involucre and that lack pappus
bristles on the ray florets. Among these species it appears
to be most closely allied to F. heterophylla, the only other
species in the genus known to have blue or purple disc
florets. The two species also resemble one another in
their involucral bracts, which bear both glandular and
setose hairs, in their cypselas, which are rather scantily
clad with long hairs, and in their subplumose pappus
bristles. F. josephinae is distinguished from F. hetero-
phylla by its relatively broader (10-12 x 4-6 mm vs 15
X 4 mm) rays that are white to cream-coloured rather
than deep blue, by the shorter peduncles (30-50 mm long
vs 50-150 mm long), resulting in a more compact
appearance to the plants, and by the narrower involucres
(4 mm vs 6-8 mm diam.) that contain relatively few disc
florets in which the petal lobes are conspicuously
recurved rather than suberect or slightly recurving.
Herbarium material of the two species in which the
colour of the rays has faded or is not noted, can always
be separated by the characteristically short peduncles and
compact habit of F. josephinae. The species appears to
have significant horticutural potential.
Additional material examined
WESTERN CAPE. — 3218 (Clanwilliam): Lambert’s Bay, Otter-
dam, 6-09-1953, (-AB), Compton 24170 (NBG); Elands Bay, base of
Rooikransberg, 18-10-1935, (-AD), Pilians 8035 (BOL); between
Elands Kloof [sic] and Clanwilliam, (-BC), 09-1933, Leipoldt 21268
(BOL); between Elands Bay and Leipoldtville, (-BC), 09-1947, Zinn
s.n. (SAM); Farm Sandberg, east of Leipoldtville, sandy slopes of
Sandberg, (-BC), 27-09-1995, Goldblatt & Manning 10325 (MO,
NBG).
ACKNOWLEDGEMENTS
Our thanks to Roy Gereau for his incomparable Latin
diagnosis. Material for study was collected on a permit
provided by Western Cape Department of Nature Con-
servation.
REFERENCES
BREMER, K. 1994. Asteraceae. Ciadistics and classification. Timber
Press, Portland, Oregon.
GOLDBLATT, P. & MANNING, J. 2000. Cape plants. A conspectus of
the Cape flora of South Africa. Strelitzia 9. National Botanical
Institute and Missouri Botanical Garden, Cape Town.
GRAU, J. 1973. Revision der Gattung Felicia (Asteraceae).
Mitteilungen der Botanischen Staatssammlung, Munchen 9:
1-195.
HERMAN, P.P.J., RETIEF, E., KOEKEMOER, M. & WELMAN,
W.G. 2000. Asteraceae. In O.A. Leistner, Seed plants of south-
ern Africa: families and genera, Strelitzia 10: 101-170. National
Botanical Institute, Pretoria.
LOW. A.B. & REBELO, A.G. (eds). 1996. Vegetation of South Africa,
Lesotho and Swaziland. Department of Environmental Affairs
and Tourism, Pretoria.
J.C. MANNING* and P. GOLDBLATT**
* Compton Herbarium, National Botanical Institute, Private Bag X7,
7735 Claremont, Cape Town.
** B.A. Krukoff Curator of African Botany, Missouri Botanical
Garden, P. O. Box 299, St. Louis, Missouri 63166, USA.
MS. received: 2001-10-15.
PTERIDOPHYTA
A NEW FERN RECORD FOR THE FLORA OF SOUTHERN AFRICA REGION
Asplenium unilaterale (Figure 9) is a widespread fern
of the Old World tropics; on continental Africa it is fair-
ly common in West Africa, becoming less frequent south
of the equator. In the Flora zambesiaca region it is con-
fined to three known localities: in Malawi from the
Misuku Hills and Thyolo Mtn, and in Zimbabwe from a
single locality in Chirinda Forest. Its recently discovered
occurrence in the northern mountains of Swaziland
therefore represents an intriguing southward extension of
its distribution. It is also interesting to note that the val-
ley in which this fern occurs is also occupied by an iso-
lated population of the tree Trilepisium madagascarien-
sis DC. (Moraceae) (Burrows & Burrows in press), a
species with which it is always associated where it is
found in Zimbabwe and Malawi, suggesting that these
two taxa are possibly the last surviving remnants of a
relict vegetation type which persists in this hidden valley.
The plants were found growing on the wet margins of a
rivulet flowing over rocks, in deep shade of evergreen for-
est. Although only one small population was seen, it is
probable that other groups occur on the several other suit-
able sites in this valley. Since the locality is only ± 1 km
from the South African border, it is also possible that A.
unilaterale occurs on the South African side. This ribbon of
forest is severely threatened by several species of invading
196
Bothalia 32,2 (2002)
FIGURE 9. — Aspleniiun unilaterale, J.E. & S.M. Burrows 7693, A,
frond and rhizome; B, median pinnae. Scale bars: 15 mm.
Artist: S.M. Burrows.
alien plants, particularly Chromolaena odorata (Aster-
aceae) and Solarium mauritianum (Solanaceae). Somewhat
lesser threats at present include the uncontrolled harvesting
of medicinal plants, and the clearing of the forest for the
cultivation of Cannabis sativa (Cannabaceae).
Asplenium unilaterale iMm., Encyclopedic metho-
dique. Botanique 2,1: 305 (1786); Sim: 152 (1915);
Tardieu-Blot: 182, t. 37 (1953); Tardieu-Blot: 189 (1958);
Alston: 56 (1959); Tardieu-Blot: 195, t. 29 (1964);
Schelpe: 174 (1970); W.B.G.Jacobsen: 340, t. 250 (1983);
J.E.Burrows: 222, t. 50 (1990); J.E.Burrows & S.M.Bur-
rows: 92 (1993). Type: Mauritius, Commerson s.n. (P-LA).
Asplenium resectum Sm.: t. 72 (1791). Type: Reunion, Commerson
(Thouin) s.n. (P).
SWAZILAND. — 253 1 (Komatipoort): Ntfonjeni Dist., Wyldsdale
Farm, Ugutugulu River, 25° 48.36’S, 31° 17.01’E, 600 m, (-CD), 11
May 2002, J.E. & S.M. Burrows 7693 (K, LYD, NBG, NU, PRE,
SDNH, Buffelskloof Herb.).
ACKNOWLEDGEMENTS
Bob Johns of the Royal Botanic Gardens, Kew, is
thanked for tracing types.
REFERENCES
ALSTON, A.H.G. 1959. The ferns and fern allies of West tropical
Africa. Suppl. to Flora of West tropical Africa, edn 2. Crown
Agents, London.
BURROWS, J.E. 1990. Southern African ferns and fern allies. Frand-
sen Publishers, Sandton, Johannesburg.
BURROWS, J.E. & BURROWS, S.M. 1993. An annotated check-list
of the pteridophytes of Malawi. Kirkia 14: 78-99.
BURROWS, J.E. & BURROWS, S.M. in press. Trilepisium madagas-
cariensis (Moraceae) in Swaziland. PlantLife 27.
JACOBSEN, W.B.G. 1983. The ferns and fern allies of southern Africa.
Butterworths, Durban & Pretoria.
LAMARCK, J.B.A.P.M. DE 1786. Encyclopedic methodique. Bota-
nique 2,1: 1^00. Pancouke, Paris.
SCHELPE, E.A.C.L.E. 1970. Pteridophyta. In A.W. Exell & E. Launert,
Flora zambesiaca. Crown Agents, London.
SIM, T.R. 1915. The ferns of South Africa, edn 2. Cambridge Univer-
sity Press, Cambridge.
SMITH, J.E. 1791. Plantarum icones hactenus ineditae, plerumque ad
plantas in herbario Linneano conservatas delineatae, 3. Londini.
TARDIEU-BLOT, M.L. 1953. Les pteridophytes de I’Afrique inter-
tropicale franqaise. Memoires de ITnstitut Franqais d'Afrique
Noire 2^-. 1-241.
TARDIEU-BLOT, M.L. 1958. Polypodiacees (sensu lato). Dennstaed-
tiacees-Aspidiacees. In H. Humbert, Flore de Madagascar et
des Comores 5. Paris.
TARDIEU-BLOT. M.L. 1964. Pteridophytes. In A. Aubreville, Flore
du Cameroun 3. Museum National d'Histoire Naturelle, Paris.
J.E. BURROWS* and S.M. BURROWS*
* Buffelskloof Herbarium, P.O. Box 710, 1120 Lydenburg, South Africa.
MS. received: 2002-05-16.
PORTULACACEAE
THE SYNONYMY OF CERARIA NAMAQUENSIS (SOND.) H.PEARSON & E.L.STEPHENS
In his description of Ceraria namaquensis (Sond.)
H. Pearson & E.L. Stephens, Dyer (1984) suggests that
C gariepina H. Pearson & E.L. Stephens should proba-
bly be regarded as a synonym of the former name.
Pearson & Stephens (1912) had separated the two taxa
as follows:
‘Living stems not readily inflammable. Surface of stem dull
grey. Leaves 3-4.5 mm long C. namaquensis
Living stems readily inflammable. Surface of stem pale yellow.
Leaves less than 2 mm long C. gariepina'
However, Dyer (1984) reports that cultivated material
at the Pretoria National Botanical Garden that was other-
Bothalia 32,2 (2002)
197
wise identified as C. gariepina, only burned when dry.
Leaves of specimens in PRE show a continuous range
from 2.0 to 4.5 mm long, often on the same specimen.
Many plants, indeed nearly all the specimens in PRE, have
both grey and yellowish branches. In fact, the PRE dupli-
cate of Pearson s.n. in Percy Sladen Memorial Expedition
3789 (an isosyntype of C. gariepina) has dark coppery
brown stems, a colour matched by the long-leaved speci-
mens Pole Evans 2250 (PRE) and Tolken 5308 (PRE).
It may be concluded that the distinguishing characters
given by Pearson & Stephens (1912) are not constant in
practice. Examination of the specimens in PRE has failed
to yield any other characters which can be used to distin-
guish between these two taxa, and they have been
regarded informally as one for many years.
With the production of a major new work on the
southern African flora at PRE, it has become necessary
to regularise the position of informal decisions such as
this, and the purpose of this note is to place on record the
reasons for the following synonymy.
C. namaquensis (Sond.) H. Pearson & E.L.Stephens
in Annals of the South African Museum 9: 33 (1912);
Podlech: 8 (1967); Dyer (1984).
Portulacaria namaquensis Sond. 2: 386 ( 1862).
C. gariepina H. Pearson & E.L.Stephens 9: 33 (1912), synon. nov.
The names are typified by two syntypes each. As it
has not yet been possible to examine all four gather-
ings with type status, no lectotypes are chosen at this
stage.
REFERENCES
DYER, R.A. 1984. Ceraria namaquensis. Flowering Plants of Africa
45: t. 1892.
PEARSON, H.H.W. & STEPHENS, E.P. 1912. List of the plants col-
lected in the Percy Sladen Memorial Expeditions, 1908-1909,
1910-1911. Portulacaceae. Annals of the South African
Museum 9: 30-35.
PODLECH, D. 1967. Portulacaceae, In H, Merxmuller, Prodromus
einer Flora von Siidwestafrika 29: 1-12. Cramer, Lehre.
SONDER, O.W. 1862. Portulacaceae. In W. Harvey & O. Sender, Flora
capensis 2: 381-386. Hodges Smith, Dublin.
H.F. GLEN*
* National Herbarium, National Botanical Institute, Private Bag
XlOl. 0001 Pretoria.
MS. received: 2002-05-14.
AMARYLLIDACEAE
THE TYPIFICATION OF CYRTANTHUS SMITHIAE WATT EX HARV.
INTRODUCTION
While researching a collection of original water colour
paintings of South African flora by John and Margaret
Herschel, executed at the Cape between 1834 and 1838, a
painting was found in the collection which appears to
have an important bearing on the typification of the name
Cyrtanthus smithiae Watt ex Harv. The original plate is in
the Brenthurst Library, Johannesburg — no. 53 in the
Herschel collection (Warner & Rourke 1998).
Cyrtanthus smithiae is similar to C. helictus Lehm.
and, as there has been some confusion between the two
(Dyer 1939), it is important to typify these names as
accurately as possible.
TYPE OF CYRTANTHUS SMITHIAE
Harvey’s name Cyrtanthus smithiae is based on the
following brief description which appeared under the
generic account of Cyrtanthus in his Genera of South
African plants (1838): ‘There are several species, one of
the most elegant of which, C. Smithiae, Watt MSS. was
brought from Cafferland by Mrs Col. Smith, in 1836, and
blossomed in Mr Watt’s garden at Rondebosch last year.
It has large white flowers, each segment marked with a
rose coloured band, and spiral strap-shaped leaves’.
The Mr Watt, to whom Harvey attributes the name, is
James Duff Watt, Deputy Assistant Commissary Gene-
ral, who lived at Mowbray Place, Rondebosch. No mate-
rial seems to have been preserved or, if it was, it appears
to have been lost, as Dyer (1939), who was the first to
recognize this name, did not cite a type specimen in his
account of this species. The curator of the herbarium at
Trinity College Dublin (TCD) confirms that there is no
material of Cyrtanthus smithiae in Harvey’s herbarium
(M. Eoody pers. comm.). Dyer’s interpretation of the
name C. smithiae was evidently based on the above brief
but sufficient description.
Two years after- Harvey’s description appeared,
Herbert (1840) published Cyrtanthus smithianus Herbert,
based on the same collection — a bulb from Caffraria col-
lected by Mrs Smith and sent to him by Watt & Harvey.
Apparently he was unaware of Harvey’s earlier name. It
subsequently flowered in Herbert’s garden in England
from which Herbert’s description was prepared. Again,
no type material seems to have been preserved and no
illustration was published. Some of Herbert’s types are at
Kew but both Baker (1896) and Dyer (1939) failed to
trace any material used by Herbert in his description of
Cyrtanthus smithianus. As the Herschels’ watercolour
was also based on the same collection — collected by Mrs
Juana Smith (wife of the then Col. Harry Smith) — it has
particular relevance to the typification of the name Cyr-
tanthus smithiae as well as the later name, C. smithianus.
The Herschels’ painting is annotated 'Cyrtanthus
smithiae' in Sir John Herschel’s hand and is dated ‘Dec.
27 1837’, the same year in which Harvey had described
a flowering specimen from Mr Watt’s garden. However,
a day before the painting was made. Col. and Mrs Smith
198
Bothalia 32.2 (2002)
/S' 3
V-
.2^
FIGURE 10. — Lectotype of Cyrtanthus
smithiae Watt ex Harv., outlined
in pencil using a camera lucida
by Sir John Herschel and painted
in water colour by Margaret
Herschel on 27th December
1837. Mrs Juana Smith brought
the bulbs from ‘Cafferland' in
1836. Plate 53. Herschel collec-
tion. Brenthurst Library. Johannes-
burg. Photo: Clive Hassall.
dined with the Herschels at Feldhausen, on Dec. 26th
1837, according to Sir John Herschel’s diary [Evans et
al.\ 333 (1969)]. Knowing the Herschels’ interest in
botanical matters, it was clearly the Smiths themselves
who brought John and Margaret Herschel the flowering
specimen of the Cyrtanthus which Mrs Smith had also
given to Mr J.D. Watt where Harvey had originally
examined it. It is therefore reasonable to assume that the
Her.schels fine colour plate is the only visual image we
have of part of the type collection of Cyrtanthus smithi-
ae (Figure 10). This plate can therefore be regarded as
having the status of a lectotype and is here designated as
such. (See Warner & Rourke 1998, plate 1 10 for a colour
reproduction of this illustration.) In following this line of
argument one must assume that Mrs Juana Smith made a
single collection of this particular Cyrtanthus, that all the
material was of the same species, and that there were suf-
ficient bulbs to distribute to Mr J.D. Watt, in Ronde-
bosch. Cape Town, William Herbert in England and also
the Herschels in Cape Town.
SYNONYMY
Cyrtanthus smithiae Watt ex Harv., Genera of
South African plants: 338 (1838); Dyer: 100 (1939);
Reid & Dyer: 38 (1984). Type: Cafferland, Mrs Smith,
watercolour painting by J.E.W. Herschel and M.B.
Herschel, plate 53 labelled Cyrtanthus Smithiae in J.E.W.
Herschel’s hand, in Brenthurst Library (lecto.!, here de-
signated).
Cyrtanthus smithianus Herbert in Curtis’s Botanical Magazine 66
sub. tab. 3779 (1840); Baker: 227 (1896). Type: Caffraria, Mrs Smith,
plate 53 labelled Cyrtanthus Smithiae by J.F.W. Herschel and M.B.
Herschel in Brenthurst Library (lecto.! here designated).
ACKNOWLEDGEMENTS
I am grateful to the Brenthurst Library, Johannesburg,
for having the original Herschel painting of Cyrtanthus
smithiae photographed for this article and for permission
Bothalia 32,2 (2002)
199
to reproduce it here. Miss Mary Foody, Dept, of Botany,
Trinity College, Dublin, kindly searched the Harvey
herbarium at Trinity College, for material of Cyrtanthus
smithiae.
REFERENCES
BAKER. J.G. 1896. Amaryllidaceae. Flora capensis 6: 221 .
DYER. R.A. 1939. A review of the genus Cyrtanthus. Description,
classification and phylogeny. Herbertia 6: 100.
EVANS. D.S.. DEEMING, T.J., EVANS, B.H. & GOLDFARB, S. 1969.
Herschel at the Cape. Balkema, Cape Town.
HARVEY, W.H. 1838. The genera of South African plants. Robertson,
Cape Town.
HERBERT, W. 1840. Cyrtanthus smithianus. Curtis's Botanical Maga-
zine 66: sub. t. 3779.
REID, C. & DYER. R.A. 1984. A review of the southern African spe-
cies of Cyrtanthus. American Plant Life Society, La Jolla, Cali-
fornia.
WARNER, B. & ROURKE, J.P. 1998. Flora Herscheliana. Brenthurst
Press, Johannesburg.
J.P. ROURKE*
* Compton Herbarium, National Botanical Institute, Kirstenbosch, Private
Bag X7, 7735 Claremont, Cape Town.
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Pi v' ■ *?‘**j^*iy'iv**
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■ t'. ", '‘^rM'i, " ■!'
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■llV.. -UV llrf .‘4 ••";,.JPH'^ 'il‘' l '- .’(*?- I
^ItJplWr'v • m^ ;■
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i>-l * **, V '.i'iji',); W
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. ,® , ■
r>>'i
.•,4fc»~.»i»l-*.itti. '- t>»,. •■ >.-•^;'«w^
' .^<*1 ’ -Ai/f»|kiikilli: '*■' '
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e^’
Bothalia 32,2: 201-210 (2002)
A study of ovule-to-seed development in Ceratiosicyos (Achariaceae)
and the systematic position of the genus
E.M.A. STEYN* \ A.E. VAN WYK** and G.F. SMITH*
Keywords: Achariaceae, Ceratiosicyos Nees, embryology, fringe layer, sarcotesta, seed coat, transfer cells
ABSTRACT
A light microscope study of developing ovules and seeds of Ceratiosicyos laevis (Thunb.) A.Meeuse was undertaken to aug-
ment an investigation of ovule and seed structure in Achariaceae, a tri-generic family comprising three species of herbaceous
perennials endemic to southern Africa. Tests for myrmecochory suggest that seed of Ceratiosicyos Nees is not dispersed by ants
like those of Acharia Thunb. and Guthriea.Bolus. Structural differences include the absence of a raphal ridge and imbibition lid
and the presence of long funicles and medium-sized embryos in Ceratiosicyos.
INTRODUCTION
Ceratiosicyos laevis (Thunb.) A.Meeuse is one of
three species of herbaceous, dicotyledonous perennials
that make up an entire family of southern African en-
demics, the Achariaceae. The family is regarded as high-
ly modified and its relationships have been much debat-
ed (see e.g. Bernhard 1999; Steyn et al. 2001 and refer-
ences therein). Traditionally, Achariaceae were placed
among families belonging to Violales (Dahlgren 1980).
Based on evidence from phylogenetic analyses of mole-
cular data (Savolainen et al. 2000) the family is placed in
Malpighiales, where it has been linked with Kigge-
lariaceae which consists of woody perennials from
southern and East tropical Africa (Kiggelaria L.), Assam
and Burma {Gynocardia R.Br.) and Sri Lanka and
Malaysia {Trichadenia Thwaites).
In niche preferences and vegetative morphology, the
three herbaceous species of Achariaceae are so diverse
(Dahlgren & Van Wyk 1988) that they were placed in
separate genera. Yet, in breeding habit and floral struc-
ture Ceratiosicyos laevis shares many characters with
Acharia tragodes Thunb. and Guthriea capensis Bolus.
Notable similarities include the presence of few-flow-
ered inflorescences containing both male and female
flowers, absence of rudimentary organs of the opposite
sex in the unisexual flowers, sympetaly (petals loosely
coherent in Ceratiosicyos Nees), conspicuous (yellow-
ish), antipetalous floral glands, and anthers with broad
connectives and unusual, swollen trichomes (Dahlgren &
Van Wyk 1988; Bernhard 1999). In addition, recent
reproductive biological studies in Achariaceae have
shown that ovules and seed characters in Acharia Thunb.
and Guthriea Bolus are remarkably similar (Steyn et al.
in press). Furthermore, both genera are myrmecochorous
and their seeds have the same unusual adaptations for
seed germination and dispersal, namely an imbibition lid
and a pronounced raphal ridge to serve as a handle for
* National Botanical Institute, Private Bag XlOl, 0001 Pretoria.
** H.G.W.J. Schweickerdt Herbarium, Department of Botany, University
of Pretoria, 0002 Pretoria.
t Corresponding author, e-mail: elsie@nbipre.nbi.ac.za
MS. received: 2002-02-22.
carrying the smooth seed.
For Ceratiosicyos, very little information is available
on ovule and seed structure or seed dispersal. A brief
report on ovule structure by Bernhard (1999) and a refer-
ence to seed coat structure (exotegmy, according to
Dahlgren & Van Wyk 1988) suggest important embryo-
logical differences between Ceratiosicyos and Acharia
or Guthriea. It is not known whether Ceratiosicyos also
forms part of the herbaceous myrmecochorous flora of
southern Africa — it does not bear its fruit near the ground
like Acharia and Guthriea for easy collection by ants,
but is a vigorously growing, nontendrilliferous twiner
that reaches considerable heights along streams at the
edge of Afromontane forest, particularly along the east-
ern escarpment of southern Africa.
For the present study we investigated embryo sac for-
mation, mature ovule characters, ovule-to-seed develop-
ment and mature seed and seed coat structure in
Ceratiosicyos. We also tested the seed for possible dis-
persal by ants. Results are compared with those recently
obtained on Acharia and Guthriea (Steyn et al. 2001;
Steyn et al. in press) to determine the embryological char-
acters of the family and to evaluate our findings in the
light of available embryological data on Kiggelaria L.
MATERIAL AND METHODS
Floral buds, mature female flowers and developing
fruit of Ceratiosicyos were collected in Eastern Cape
from a population growing on the banks of the Maitland
River, in the Maitland River Forest Reserve (voucher
specimen: Van Wyk 13555 PRU). Additional material
that included seeds at dispersal stage was gathered at
Kowyns Pass near Graskop in Mpumalanga (voucher
specimen: Steyn 24 PRE). All flowering and fruiting
stages were immediately immersed and stored in a 0.1
M cacodylate-buffered solution (pH 7.4) containing 4%
formaldehyde and 2.5% gluteraldehyde. Flowers and
fruits were later dissected, ovules and developing seeds
removed, sorted according to size and rinsed in the
buffer. Dehydration and impregnation with glycol
methacrylate (GMA) followed the methods of Feder &
202
Bothalia 32,2 (2002)
FIGURE 1 . — Ovule and embryo sac of Cenitiosicyos as seen in sagittal section. A, mature ovule; B, early stage in development of bisporic embryo
sac; C, D, consecutive sections of same ovule as in A to show details of mature embryo sac. a, egg cell; c. central cell nucleus; curved
arrows indicate megaspore nuclei in chalazal dyad cell; d, disintegrating micropylar dyad cell; n, derivatives of nucellus epidermis; r, outer
epidermis of inner integument; v. vascular bundles in chulaza. Scale bars; A, 100 pm; B, 10 pm; C, D, 50 pm.
O’Brien (1968). A .selection of impregnated structures
was individually imbedded in GMA, hardened in the
oven at about 5S°C and sectioned transversely and sagit-
tally. Selected sections were stained with the periodic
acid/Schiff reaction and counterstained with toluidine
blue O by using the protocols of O’Brien & McCully
(1981).
Tests for myrmecochory: at the collection site on the
Kowyns Pass, dehiscing capsules were carefully removed
Bothalia 32,2 (2002)
203
from the plants, the seeds collected and immediately
strewn onto the trails or near the nests of ants of varying
sizes found at the collection site. Mature capsules were
stored in air-tight bags overnight and taken to Pretoria
where seeds, removed from capsules that had in the mean-
time split open, were again offered to ants. Dried-out seeds
with tuberculate surfaces were also offered to ants.
RESULTS
Placentation and orientation of ovules
The hypogynous female flowers of Ceratiosicyos
contain elongated, pentagonal ovaries borne on gyno-
phores. The ovaries are unilocular and usually contain
7-15 anatropous ovules that are borne singly on five
parietal placentae. The latter regions are not ridged and
run longitudinally along the inner surface of the ovary
wall, opposite the five median carpel traces. Ovules from
five placentae are so arranged that they form a single row
in the narrow locule and this alignment is maintained
throughout ovule-to-seed development. The alignment is
achieved by the ability to vary the orientation of the
ovules and the lengths of the funicles — some ovules
point upwards, others downwards and the funicles may
be short or more than twice the length of the ovular body
to place the ovules neatly in a single row.
Structure of mature ovule
Mature ovules are anatropous, bitegmic, crassinucel-
late structures (Figure lA) with an ovoid shape and ±
520 pm long (funicle excluded). The integuments are
multilayered, the outer consists of four to five layers in
its central part, while the inner is up to seven layers thick
in this region. The outer epidermis of the inner integu-
ment is very conspicuous — the cells are four to six times
longer than any other cell in the ovule with the exception
of the embryo sac. On the antiraphal side of the ovule,
the tip of the outer integument increases in thickness by
tangential divisions in its inner epidermal layer. In pre-
fertilization stages of ovules, the outer integument is as
long as the inner integument so that the micropyle canal
is formed by the inner integument only. The raphe is not
ridged. The vascular bundle of the raphe branches as
soon as it enters the chalaza, but the integuments are not
vascularized (Figure lA).
The mature embryo sac is about one-third the length
of the nucellar cylinder and lies in the centre of massive
nucellar tissue. About six layers of nucellus cells cover
the embryo sac on all sides. Below the micropyle, at least
three of these layers result from periclinal divisions of
the nucellus epidermis (Figure 1C, D). The nucellus apex
is slightly attenuate, but does not protrude into the
micropyle.
The embryo sac develops from the chalazal dyad cell
while the micropylar dyad cell degenerates (Figure IB).
During the second meiotic division the chalazal dyad is
not partitioned by a transverse cell wall so that both
megaspore nuclei are included in the same cell (Figure
IB). After two mitotic divisions an eight-nucleate, bis-
poric embryo sac of the Allium Type is formed. The
mature embryo sac contains many starch grains. A small
egg apparatus occurs below the parietal nucellar tissue
(Figure 1C). The short neck regions of the synergids con-
tain a filiform apparatus. A large central cell nucleus lies
in about the central part of the embryo sac, while three
ephemeral antipodal cells (not shown) develop in the
elongated and narrow chalazal base (Figure lA) of the
embryo sac.
Early development of endosperm and embryo
Fertilization is porogamous in Ceratiosicyos and
endosperm formation is nuclear. After entering the
micropyle, the tip of the pollen tube swells and stains
darkly with PAS and toluidine blue (Figure 2A). During
the initial stages of embryo sac enlargement (Figure 2A),
free endosperm nuclei become arranged in a single layer
alongside the embryo sac wall. When the growing seed
has reached a size of ± 5 x 2.5 mm, the first cell walls are
laid down between adjacent endosperm nuclei, and the
embryo sac then gradually becomes filled, layer upon
layer, with thin-walled endosperm cells.
The zygote remains inactive during the nuclear stage
of endosperm formation. The first division of the zygote
was not seen, but pro-embryos in the tetrad stage of develop-
ment (Figure 2B) were found before the endosperm start-
ed to become cellular. These four-celled pro-embryos are
T-shaped which shows that the apical cell (ca) has divid-
ed in a vertical plane and the basal cell (cb) transversely
(Figure 2B). The two daughter cells of ca then both
divide obliquely (Figure 2C) so that a bicellular, wedge-
shaped epiphysis (e) is formed in the apical tier during
the quadrant stage of the pro-embryo (i.e. when the deriv-
atives of ca comprise four cells). The epiphysis later
forms the shoot apex, whereas the remaining cells of the
quadrant form the cotyledons (Natesh & Rau 1984: 390).
The pro-embryo is globular in shape and has no suspen-
sor. Below the cells of the quadrant, the uppermost deriva-
tive of cb is a discoidal cell (h), designated ‘hypophysis’
by Hanstein (1870). The hypophysis later forms the ini-
tials of the root cortex and the root cap (Crete 1963).
In Johansen’s (1950) classification of embryogenic
types, T-shaped pro-embryos are characteristic of both
the Onagrad Type and Asterad Type, but it is only in the
latter type that the basal cell (cb) contributes significant-
ly to the formation of the embryo proper. Ceratiosicyos
embryos have no suspensors, all derivatives of the basal
cell are incorporated into the embryo proper which indi-
cates an Asterad Type embryo. The presence of the epi-
physis also points towards the Asterad Type — it is in taxa
conforming to this type that an epiphysis is formed dur-
ing the quadrant stage (Natesh & Rau 1984: 386, fig.
8.6). The lack of a suspensor places the embryo of
Ceratiosicyos in the Penaea variation of the Asterad
Type (Natesh & Rau 1984: 390, 414).
Development and structure of the seed coat
The seed coat of Ceratiosicyos is mainly derived from
the outer integument. This integument forms the sar-
cotestal layers and the outer, wavy layer of sclereids that
protrude peak-like into the sarcotesta (Figures 3D; 5D).
204
Bothalia 32,2 (2002)
FIGURE 2. — Early development of endosperm and embryo in Ceratiosicyos. A, developing seed during resting stage of zygote; B, T-shaped pro-
embryo; C, suspensorless pro-embryo during quadrant stage, ca, apical cell after vertical division; cb, basal cell after transverse division;
e, epiphysis; h, hypophysis; i, inner integument; I, bulge of inner integument; o, outer integument participating in formation of micropyle
canal; t, pollen tube in elongating embryo sac. Scale bars: A, 200 pm; B-C, 50 pm.
The inner integument contributes a few layers of pericli-
nally elongated fibres to the seed coat. During seed coat
development the cuticle between the outer and inner
integument gradually disappears so that a study of the
mature seed coat alone does not show which part each
integument plays in the formation of the seed coat.
Contribution of the outer integument (testa) to the seed coat
In pre-fertilization stages, the outer integument con-
sists of about five cell layers (Figure lA), except at its
rim where the number of layers increases through tangen-
tial divisions of the inner epidermis. The cells in the rim
remain meristematic and in early post-fertilization stages
the distal part of the outer integument grows beyond the
inner integument to take part in the formation of the
micropyle (Figure 2A). In developing seeds a layer of
actively dividing endotestal cells (s) can be .seen inside
the developing sarcotesta (Figure 3A, B, D). These cells
are the derivatives of the inner epidermis of the outer
integument. At first, the derivatives lie in radial rows and
the layer is of even thickness (Figure 3 A, B). When the
seed has reached its final length of ± 6 mm, the
endotestal layer becomes wavy and starts forming pro-
jections into the sarcotesta (Figure 3D). The endotestal
cells later develop into closely packed, thick-walled scle-
reids with starch grains and single, large crystals of cal-
cium oxalate in some of the cells, but the crystal-con-
taining cells do not form a continuous layer. At seed dis-
persal stage, the contents of the endotestal sclereids stain
intensely with toluidine blue, indicating the presence of
phenolic sub.stances (Figures 3C; 4A, B).
The epidermis and mesophyll of the outer integument
form a sarcotesta that envelops the whole seed, including
the chalaza and raphe. The raphal region is not pro-
nounced, i.e. a raphal ridge is not formed (Figures 4A;
5D). The raphal bundle lies imbedded in thin-walled.
Bothalia 32,2 (2002)
205
FIGURE 3. — Development and structure of seed coat in Ceratiosicyos. A, t/s young seed coat just after fertilization; B, 1/s seed coat during first
stages of pro-embryo formation; C, 1/s mature seed coat of dispersed seed; D, 1/s seed coat during maturation of fibrous exotegmen. Curved
arrows indicate position of stomata; n, nucellus cell remains; p, inner epidermis of tegmen; r, outer epidermis of tegmen with anticlinal
divisions; s, derivatives of inner epidermis of testa; u, sarcotesta; y, mesophyll of tegmen. Scale bars; A, 50 pm; B, C, 100 pm; D, 200 pm.
parenchymatous, sarcotestal tissue that, especially in this
area, contains large numbers of starch grains (Figure 4A).
Stomata, not seen in the outer epidermis of the ovule,
were found at regular intervals in the developing (Figure
3A, B) and mature (Figure 3C) epidermis of the sarco-
testa.
When the seeds are dispersed, the cells of the inner-
most layer of the sarcotesta have developed small, fibril-
lar protuberances on their inner tangential walls (Figures
3C; 4A, B). The fringe-like wall ingrowths are strongly
PAS-positive and also stain dark blue with toluidine blue.
The fringe layer possibly represents a layer of transfer
206
Bothalia 32,2 (2002)
FIGURE 4. — Development and structure of seed coat of Ceratiosicyos (continued): A, raphe and underlying seed coat layers in a median t/s of
seed; B, t/s fringe layer and adjacent cell layers seen at higher magnification than in Figure 3C. f, fringe layer; r, fibres of exotegmen; s,
derivatives of inner epidermis of testa; u, sarcotesta. Scale bars: A, 200 pm; B, 50 pm.
cells (Gunning & Pate 1969), often found in reproductive
structures for the short-distance transport of solutes
(Johri & Ambegaokar 1984: 29, fig. 1.13A-F).
Contribution of the inner integument (tegmen) to the seed
coat
After fertilization the conspicuously elongated outer
epidermal cells of the inner integument (Figure lA) ini-
tially divide anticlinally to form a single layer of dense-
ly packed, radially elongated meristematic cells (Figure
3A). While the first divisions of the pro-embryo are tak-
ing place, these meristematic cells of the tegmen divide
periclinally once or twice to form three to four layers of
cells that are stretched in a direction parallel to the lon-
gitudinal axis of the seed (Figure 3Bk At this stage the
cuticle between the developing endotesta and exotegmen
starts to disappear, but in longitudinal sections of imma-
ture seeds the boundary between the two layers is clear
because of the difference in the orientation of the cells
(Figure 3B, D). When the seed reaches its final size, the
FIGURE 5. — Fruit and seed of
Ceratiosicyos. A, mature, 5-
valved capsule, x 0.8; B, dis-
persed seed, X 0.8; C, de-
hisced fruit, showing funicles
of varying lengths left on the
placentae, x 0.8; D, 1/s seed,
f, fringe layer; m, mechanical
layers; u, sarcotesta; v, vas-
cular bundle branching in
chalaza. Scale bar: 1 mm.
Artist: G. Condy.
Bothalia 32,2 (2002)
207
exotegmic cells contain many starch grains and the outer
elements have started maturing into fibres (Figure 3D).
At seed dispersal stage the fibres have thick, lignified
walls with simple pits and contain no starch (Figures 3D;
4A, B).
The mesophyll and inner epidermis of the tegmen do
not play a significant role in the structure of the mature
seed coat. The thin-walled mesophyll tissue initially
shows divisions in various planes so that the layers
increase in number (Figure 3B). The innermost layer of
mesophyll cells becomes conspicuous by their large size
and darkly staining properties (Figure 3B, D). At first,
the inner epidermis of the tegmen keeps pace with the
growth of the seed by dividing anticlinally so that a layer
of small, densely packed cells is formed (Figure 3B).
Eventually, when the seeds are dispersed, all layers
inside the exotegmic fibres are obliterated and a struc-
tureless pellicle remains between the fibres and the flat-
tened cells of the nucellar tissue (Figures 3C; 4A).
Macromorphology of the fruit and seed and tests for myr-
mecochory
Mature fruit consists of 50-90 mm long, 5-valved cap-
sules (Figure 5A, C) that are thin-walled and light green
at seed dispersal stage. Five to twelve stalked seeds are
arranged in a single row, sometimes packed close to one
another in the locule. When the valves split open (not
very forcefully), the seeds break off, leaving their funicles
of various lengths attached to the centre of the valves
(Figure 5C). Seeds are ovoid to short-cylindrical, 6. 0-6. 5
mm long (Figure 5B), dark green to brown and covered
with a thin, succulent and translucent sarcotesta. The seed
surface becomes tuberculate when the sarcotesta dries
out, reflecting the projections formed by the underlying
mechanical layers (Figure 5D). At seed dispersal, the
axile embryo (sensu Martin 1 946: 520) is of medium size
(i.e. it occupies about three-quarters of the length of the
endosperm). It lies straight in the seed, has thin, spatulate
cotyledons and a well-formed radicle (Figure 5D).
Tests for myrmecochory were negative. If ant-seed
interactions are not species specific as claimed by
Slingsby & Bond (1981) and found by Steyn et al. (in
press), our results suggest that Ceratiosicyos seeds are
not dispersed by ants.
DISCUSSION
Differences between Ceratiosicyos and Acharia and Guthriea
A comparison of ovule and seed characters in
Ceratiosicyos with those of Guthriea and Acharia (Table
1) shows that the three genera are fundamentally very
similar in characters usually regarded as of taxonomic
importance (see No. 1, 2, 8-12, 14, 16 & 17). The struc-
ture of the integuments is also comparable to a large
degree, although Ceratiosicyos lacks the peculiar zigzag
micropyle (see No. 5) that characterizes ovules and seeds
of Guthriea and Acharia. The short outer integument in
the Ceratiosicyos ovule possibly does not denote an
important structural difference with the other two genera,
because this integument overtops the inner after fertiliza-
TABLE 1 . — A comparison of ovule and seed characters in Achariaceae
208
Bothalia 32,2 (2002)
tion. The inner integument of Ceratiosicyos bulges into
the micropyle canal (Figure 2A). The bulging cells might
have been mistaken for a nucellar beak by Dahlgren &
Van Wyk (1988). The present investigation also showed
that the mechanical seed coat layer in Ceratiosicyos is,
like those of the other two genera, of dual origin
(endotestal-exotegmic) and not purely exotegmic as pre-
viously reported (Dahlgren & Van Wyk 1988).
Many of the differences between Ceratiosicyos and
the other two genera can possibly be attributed to specif-
ic adaptations for seed dispersal and germination (see
No. 6, 7, 13, 15, 18 & 19). Ceratiosicyos is not myrme-
chorous, the ovule therefore lacks the pronounced raphe
that, in Acharia and Guthriea, eventually forms a ridge-
like part of the sarcotestal elaiosome (see No. 6 & 19).
The presence of unicellular hairs on the seed surface of
Guthriea and Acharia (see No. 15) possibly also relates
to myrmechory, since openings left by broken-off tri-
chome bases would allow ant-attracting substances to
rapidly reach the seed surface (Steyn et al. in press).
Instead of a pronounced raphe, ovules and seeds of
Ceratiosicyos might have developed long funicles as an
adaptation to seed dispersal (see No. 7). By varying the
lengths of the funicles, the seeds can be manipulated into
a single, vertical row in the narrow, elongated locule.
This arrangement may be necessary for rapid splitting of
the fruit by distributing pressure on the valves evenly
along the length of the locule. In the two diplochorous
(autochorous + myrmecochorous) genera (see No. 19)
the seeds are sessile in the short-cylindrical capsules and
pressure on the valves is applied by the swollen elaio-
somes. Also, the capsules remain within the covering of
persistent corolla tubes to protect developing seeds with
their soft elaiosomes (Steyn et al. in press). Ceratiosi-
cyos is autochorous; its seeds need less protection and
developing capsules rapidly outgrow the protective cov-
ering of the corolla tubes.
Seed size in genera of the Achariaceae does not differ
significantly, but Ceratiosicyos has a much larger
embryo than the other two genera (see No. 11 & 13). We
propose that the small embryo has been the causal factor
for the formation of the unique seed lid in the other two
genera. This device would allow water and air to enter
the seed through the unsclerified cells in the rim of the
lid during the slow maturation (12 weeks) of the embryo
in the hydrated seed (Steyn et al. in press). The much
larger embryo of Ceratiosicyos possibly did not require
such an adaptation.
Martin (1946) reasoned that smallness in embryos, as
compared to size of the endosperm, is representative of a
primitive state in seeds of angiosperms, and, conversely,
that embryos which become well developed before dor-
mancy, reflect a higher evolutionary rank. For dicotyle-
dons, Corner (1976: 48) also regarded small embryos as
primitive and considered simplification through loss of
structures or cell layers as an indication of an advanced
state in seeds. However, considered on its own, it is usu-
ally difficult, if not impossible, to tell which of the two
embryo states— small or large — is the more primitive for
a particular taxon. One reason for this difficulty is that
embryo (and seed) size, like so many other characters.
are subject to homoplasy (parallelism, convergence and
character state reversals). Unfortunately homoplasy is
rampant among seed plants, thus considerably limiting
the reliability of outgroup comparisons to establish
polarity (Cronquist 1988). Compared to the other mem-
bers of the family, Ceratiosicyos shows a notable trend
towards simplification of the seed through the loss of
several cell layers and structures, e.g. trichomes, a bi-lay-
ered testa epidermis, a hypodermis, a crystal-containing
layer inside the fringe layer, perisperm and the reduction
of the chalazal region with seed lid. These reductions,
together with the larger embryo, may indicate that
Ceratiosicyos is more advanced than Guthriea and
Acharia. On the other hand, indications are that Kigge-
lariaceae may well be the sister group of Achariaceae
(see further on). Both Kiggelaria africana and Ceratiosi-
cyos laevis share a medium-sized embryo and rather
unspecialized seed, coat, states that may be the ple-
siomorphic ones in Achariaceae. Relatively small em-
bryos and a rather elaborate seed coat occur in Acharia
tragodes and Guthria capensis, both having specialized
myrmecochorous seeds. Therefore Ceratiosicyos laevis,
with its lianeous habit and mesic forest habitat, could just
as well be the more primitive member of the family.
Acharia tragodes (semi-woody shrublet) is confined to
the xerophytic thicket vegetation of the Eastern Cape,
with Guthria capensis (rosulate herb) confined to tem-
perate grassland and karroid vegetation at high altitude.
Achariaceae versus families in Malpighiales
A detailed comparison of ovule and seed characters in
Achariaceae and the 36 families placed in Malpighiales
(including many families traditionally placed in Violales)
by Savolainen et al. (2000), is hampered by a lack of
comparable data for many of the families (Davis 1966;
Johri et al. 1992; Nandi et al. 1998). Achariaceae seem
generally well placed in Malpighiales and fit in comfort-
ably among those families previously regarded as part of
Violales sensu Dahlgren (1980). Similarities include
bitegmic, anatropous, crassinucellate ovules, parietal
nucellus partly formed by nucellus epidermis deriva-
tives, both integuments participating in formation of
micropyle canal, nuclear endosperm becoming copious
in the seed and a medium-sized to large embryo lying
straight in the seed.
Some of the characters observed in Achariaceae are
rare for Malpighiales, namely a bisporic Allium Type
embryo sac, suspensorless Asterad Type embryos, pro-
tective seed layers containing endotestal sclereids and
exotegmic fibres, and a sarcotesta. A number of charac-
ters shared by Guthriea and Acharia, such as the zigzag
micropyle, distally lobed outer integument, perisperm
and small embryo is also uncommon for the order. A
zigzag micropyle and Asterad embryos conforming to
the Penaea variation only occur in Violaceae (Davis
1966), while lobed integuments and perisperm are found
in Scyphostegiaceae (Van Heel 1976; Johri et al. 1992)
and sarcotestal seeds in Passiflorales (Nandi et al. 1998).
A possible phylogenetic link between Achariaceae
and Flacourtiaceae (tribe Pangieae, particularly
Kiggelaria africana) was first suggested by the breeding
behaviour of a butterfly. Several butterflies in the sub-
Bothalia 32,2(2002)
209
tribe Acraeina (subfamily Acraeinae, tribe Acraeini),
including the common garden acraea {Acraea horta), uti-
lize as larval food plants, members of a closely knit
group of plant families traditionally classified in the
order Violates (notably Achariaceae, Flacourtiaceae,
Passifloraceae and Tumeraceae), all containing a unique
group of toxic compounds known as cyclopentenoid
cyanogenic glucosides (Seigler 1975; Dahlgren 1980;
Cronquist 1981; Takhtajan 1997; Kroon 1999). In their
natural habitat larvae of Acraea horta feed mainly on
Kiggelaria africana, a species containing gynocardin as
major cyanogenic glucoside (Jaroszewski & Olafsdottir
1987; Raubenheimer & Elsworth 1988). The larvae
selectively sequester and store some of the gynocardin,
which are passed on to all other stages in the life cycle,
supplemented by apparent self-synthesis (Raubenheimer
1987, 1989). Accumulation of this toxin is believed to
render the insects unpalatable to predators. Previously
gynocardin has been isolated from the seed of
Gynocardia odorata R.Br., another member of the tribe
Pangieae (Cobum & Long 1966). When live plants of
Ceratiosicyos laevis and Guthriea capensis were intro-
duced into the botanical garden at the University of
Pretoria in the mid-1980s, both were immediately select-
ed by Acraea horta for egg deposition; larvae emerged
and butterflies were raised (Dahlgren & Van Wyk 1988).
This observation led to a chemical study of Ceratiosicyos
laevis, the first of its kind on a member of the
Achariaceae, resulting in the identification of gynocardin
as one of the principal cyanogenic glucosides in this
species (Jensen & Nielsen 1986); its presence in
Guthriea capensis is suspected.
Based on evidence from molecular biology. Chase et
al. (1996) also suggested a linkage between the herba-
ceous Achariaceae and the woody tribe Pangieae. This
tribe included amongst others, Gynocardia R.Br., Hyd-
nocarpus Gaertn., Kiggelaria L., Pangium Reinw. and
Trichadenia Thwaites (Lemke 1988). In the circumscrip-
tion of Soltis et al. (2000), Kiggelariaceae include
Pangium, Hydnocarpus, and Kiggelaria. Our results
show that Achariaceae agree closely with the latter two
genera as far as seed development and seed coat structure
are concerned. Van Heel (1979) found the seeds of
Hydnocarpus and Kiggelaria sarcotestal with undulating
endotestal-exotegmic mechanical layers, a dominant
endotestal layer of sclereids, an outer integument that is
initially short, but overtops the inner during seed forma-
tion and a cuticle that disappears early so that the dual
nature of the protective layer is masked and the en'o-
neous impression given that the seeds are pachychalazal.
These characters are so similar to the characters we
found in Achariaceae that they can be regarded as strong
support for a linkage between Achariaceae and Kigge-
lariaceae.
In Kiggelaria the embryo is of medium size, as found
in Ceratiosicyos. It is perhaps noteworthy that seeds with
a small embryo, a fleshy raphe and a conspicuous
notched cuticle between the tegmen and nucellus as
reported for Guthriea (Steyn et al. 2001) also occur in
Berberidopsis Hook.f. (Van Heel 1979). This taxon, pre-
viously also included in the Flacourtiaceae (Lemke
1988), is currently regarded as a relict with an unclear
taxonomic position (Savolainen et al. 2000).
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Bothalia 32,2: 211-231 (2002)
The grasslands and wetlands of the Sekhukhuneland Centre of
Plant Endemism, South Africa
S.J. SIEBERT*\ A.E. VAN WYK*, G.J. BREDENKAMP* and F. DU PLESSIS*
Keywords: biodiversity, Braun-Blanquet, conservation, endemism, grassland, Mpumalanga, norite, phytosociology, Sekhukhuneland, syntaxonomy
ABSTRACT
A hierarchical classification, description, and ecological and floristic interpretations are presented on the vegetation types
of the grasslands and wetlands of the Sekhukhuneland Centre of Plant Endemism. Releves were compiled in 74 stratified
random plots. ATWINSPAN classification, refined by Braun-Blanquet procedures, revealed eight associations, 1 1 subassocia-
tions and four variants. Many new syntaxa are described and ecologically interpreted. For each syntaxon, the species richness,
endemism and conservation status was determined. The floristic and habitat information, proposed classification, general
description and vegetation key are provided to aid future identification of conservation areas, land use planning and further
research. An ordination (DECORANA). based on floristic data, confirmed the relationships that exist between plant commu-
nities and associated habitats and environmental gradients. Much of the plant community diversity and distribution can be
ascribed to a heterogeneous environment, predominantly determined by soil moisture.
INTRODUCTION
Several phytosociological studies have been conducted
on vegetation types along the northeastern escarpment of
the Northern Province and Mpumalanga (Bloem 1988;
Deall et al. 1989; Matthews et al. 1992a, b; Burgoyne et
al. 2000). However, the vegetation of some areas still
remains to be investigated and described, both on recon-
naissance level and in more detail. The undulating norite
hills in the Roossenekal-Tonteldoos region is an area with
very limited information on the vegetation. It is located in
the southern portion of the Sekhukhuneland Centre of
Plant Endemism (SCPE) (Van Wyk & Van Wyk 1997; Van
Wyk & Smith 2001), and comprises the Roossenekal
Subcentre of Endemism (Siebert 1998). The Roossenekal
Subcentre is known for its many plant endemics, the dis-
tributions of which correlate with the diversity in geologi-
cal substrate in the region (Siebert 1998; Siebert et al.
2001).
Various grassland and wetland vegetation types that
show a floristic affinity and relationships with the Roos-
senekal Subcentre (Siebert 1998), have previously been
described from the adjacent Steenkampsberg (Bloem
1988; Burgoyne 1995), Witbank Nature Reserve (Smit et
al. 1997), and the Great Dyke of Zimbabwe (Werger et
al. 1978). Acocks (1988) mapped the vegetation of the
Roossenekal Subcentre as two veld types, namely North-
eastern Sandy Highveld (57) and Bankenveld (61). A
broad-scale classification of the same region’s vegetation
was given by Low & Rebelo (1996), who recognize three
vegetation types, namely Mixed Bushveld (18), Moist
Sandy Highveld Grassland (38) and North-eastern
Mountain Grassland (43). These veld/vegetation types
are closely associated with seasonal fires (Edwards
1984).
* Department of Botany, University of Pretoria, 0002 Pretoria,
t Present address: SABONET, c/o National Botanical Institute,
Private Bag XIOI, 0001 Pretoria.
MS. received: 2001-07-02.
Only Acocks (1988) accurately mapped the high-alti-
tude outcrops of norite (Leolo Mountains) in the north-
ern region of the SCPE as grassland, namely North-east-
ern Sandy Highveld (57). Geologically and floristically
the Leolo Mountains is similar to the Roossenekal
Subcentre, and is treated as part of the study area covered
in this paper (Siebert 1998). However, the vegetation
classification of the entire Sekhukhuneland Centre needs
further attention, as comparatively little is known about
its plant communities, floristic diversity and the relation-
ship between distribution patterns of plants and the
clayey soils derived from the ultramafic norite.
This paper forms part of a comprehensive vegetation
and floristic survey of the Sekhukhuneland area. It is
envisaged that the identification, classification and
description of the various vegetation units (syntaxa) will
contribute to the knowledge of the plant diversity and
biological intricacies of the region. This paper provides
ecological and floristic data of the region’s grasslands
and wetlands, and the associated habitats, by characteriz-
ing and interpreting the vegetation units. Classification is
basic to the formulation of a management policy and for
proper land use planning. An assessment of the plant
species richness, endemism and Red Data List taxa in the
plant communities of the study area is supplied as base-
line data to apply in future conservation actions in the
region and application in ecosystem management. An
adequate database of natural features and other land uses
is essential for effective land use management and imple-
mentation (Kent & Ballard 1988; Bedward et al. 1992;
Rhoads & Thompson 1992).
In a broad overview of the vegetation types of the
SCPE, Siebert et al. (2002) recognized six major vege-
tation units. The vegetation units described in this paper
cover the Therneda triandra-Senecio microglossus Cool
Moist Grassland and the Fuirena piibescens-Schoeno-
plectus corymbosus Wetland Vegetation, the two major
vegetation types of the relatively moist Roossenekal
Subcentre. A vegetation key is presented to aid with the
identification of the proposed syntaxa in the study area.
212
Bothalia 32.2 (2002)
STUDY AREA
The study area is situated in northern Mpumalanga
between latitude 25° 00' and 25° 20' S and longitude
29° 50' and 30° 05' E (southern part of the SCPE) and
southern Northern Province between latitude 24° 30' and
24° 50' S and longitude 30° 00' and 30° 05' E (Leolo
Mountains) (Eigure 1). The area covers ± 1 500 km^ and
comprises a homogeneous geology (Visser et al. 1989),
with a heterogeneous physiography (Land Type Survey
Staff 1987). The Sekhukhuneland Centre is more or less
restricted to the Rustenburg Layered Suite of the Bushveld
Complex (Siebert et al. 2001). The aiea dealt with in this
paper lies on the Upper and Main Zones of the Rustenburg
Layered Suite and is mainly underlain by concentric belts
of norite, and to a lesser degree outcrops of ferrogabbro
(Visser et al. 1989). This norite stratum extends north-
south and rises to fomi the Leolo Mountains ( 1 700 m asl).
The lower reaches of the mountainous areas are character-
ized by many small outcrops of magnetite.
The Roossenekal Subcentre is defined as the grassland
areas in and adjacent to the sixteenth degree grid 2529 BB
and also includes the outlier Leolo Mountains in the
Northern Province. Thirty-four of the Sekhukhuneland
endemics/near-endemic plant species which are largely
restricted to these areas were recorded during the study
(Siebert 1998). The area forms an ecotone between the
Mixed Bushveld (18) and the North-eastern Sandy
Highveld (57) Veld Types (Acocks 1988), but it is proba-
bly more representative of Bankenveld (61), than any of
the aforementioned veld types. The valleys have a sub-
tropical climate with little or no frost in winter, whereas in
the mountains, conditions become more temperate with
increasing altitude. In this region of undulating rocky hills,
the vegetation is characterized by scattered woodlands in
sheltered habitats of footslopes and in valleys, and grass-
land with bush clumps covering the exposed plateaus, hill
slopes and valleys. The difference in altitude between the
two most extreme locations is ± 500 m ( 1 700 m asl on the
Leolo Mountains to 1 200 m asl where the Klip River cuts
through the hills near Roossenekal).
The study area lies in the summer rainfall region, with
an average annual precipitation of about 700 mm (South
African Weather Bureau 1998). Local rainfall patterns
FIGURE 1 . — Localion of the grass-
lands of the study area in the
Northern Province and Mpu-
malanga. South Africa.
Bothalia 32,2 (2002)
213
FIGURE 2. — Relative positions of all the releves along the second and third axis of the ordination of the grassland vegetation of the
Sekhukhuneland Centre of Plant Endemism. Numbers correspond with those of syntaxa in Table la and in the text.
are strongly influenced by the area’s topography and alti-
tude (Siebert 1998), varying from 721 mm in the east, to
607 mm per annum in the west; 710 mm in the south, to
702 mm in the north (Erasmus 1985). Daily average tem-
perature ranges from a minimum of 2.8°C in winter to a
maximum of 25.9°C in summer, with an average annual
temperature of 16.2°C (South African Weather Bureau
1998). Temperatures vary at different localities within
the area, also correlating strongly with physiographic
regions, being higher in low-lying valleys and lower on
high-lying plateaus (Buckle 1996). However, minimum
temperatures of below freezing point are rare, even in the
high-lying areas.
METHODS
A first approximation of a vegetation classification,
based on the total floristic data set of 415 stratified ran-
dom sample plots was obtained by the application of the
Two-Way Indicator Species Analysis (TWINSPAN)
(Hill 1979a). The first step of an objective multivariate
classification identified six major vegetation types for
the Sekhukhuneland Centre (Siebert et al. 2002). These
results were then used to subdivide the data set into five
phytosociological tables. One of these tables, represent-
ing two major vegetation types, namely grasslands and
wetlands, was again subjected to TWINSPAN. Using
Braun-Blanquet procedures in the MEGATAB computer
program (Hennekens 1996a), we further refined the
resultant classification.
Floristic and habitat data of the grasslands and wetlands
were derived from 74 releves. Stratification was based on
terrain type, aspect and vegetation structure. To standardize
the plot size and to counter the bias of different scale
(Jonsson & Moen 1998), plots sampled in the grassland
were fixed at 100 m-. Within each sample plot, all species
were recorded and a cover-abundance value was assigned
to each species according to the Braun-Blanquet scale
(Mueller-Dombois & Ellenberg 1974). Plant species names
conform to those of Retief & Herman (1997), and the
practice followed in the H.G.W.J. Schweickerdt Herbarium
(PRU), University of Pretoria. Terminology to describe
vegetation structure follows Edwards (1983). Environ-
mental data recorded for each sample plot include teirain
type (Land Type Survey Staff 1987), aspect, slope, geolo-
gy (Visser etc//. 1989), soil type (Mac Vicar et Cl/. 1991) and
rockiness of soil surface. Longitude and latitude readings
were also recorded for each sample plot using a GPS. All
releve data are stored in the TURBOVEG database
(Hennekens 1996b), managed by the Department of
Botany, University of Pretoria (Mucina et al. 2000).
Syntaxa names are given in accordance with the Code of
Phytosociological Nomenclature (Weber et al. 2000).
The ordination algorithm Detrended Correspondence
Analysis (DECORANA) (Hill 1979b) (Eigure 2) was
applied to confirm gradients in vegetation and the rela-
tionship between these plant communities and the phys-
ical environment.
To facilitate the identification of areas of high conser-
vation potential, the alpha diversities of the different
plant communities were calculated. The alpha diversity
(plant species richness) is defined as the number of
species per unit area within a homogeneous community
or the total number of species per community (Whittaker
1977). A 100 m- sample plot was taken as the unit area
within a homogeneous community.
The geographical distribution of all the taxa was ver-
ified at the National Herbarium (PRE), Pretoria, to iden-
tify any taxa endemic/near-endemic to the region
(Siebert 1998). All taxa were also checked against the
Red Data List of southern African plants (Hilton-Taylor
1996) to determine their conservation status.
214
Bothalia 32.2 (2002)
RESULTS
Classification of vegetation
The analysis resulted in the identification of 17 plant
communities, ordered as eight associations, 1 1 subassoci-
ations and four variants (Table la, b). These units were
subsequently hierarchically classified. Since the study
area covered by these communities lies in the climatical-
ly uniform moist and cool southern region of the SCPE
(Siebert 1998), no major macroclimatic variation plays a
role in local differentiation of the plant communities. The
major plant communities relate to soil character, rocki-
ness and teiTain type, with aspect and slope also playing
minor roles. Communities were not always distinctive in
the field. This might be attributed to the homogeneity of
grassland physiognomy and the heterogeneity of the envi-
ronmental factors, which resulted in a complex mosaic
distribution pattern of habitats and associated vegetation.
The hierarchical classification of the vegetation rein-
forces the correlation between habitat and plant commu-
nities (Figures 2, 3). The distribution of the SCPE endemic/
near-endemic and Red Data List plant taxa among vari-
ous plant communities is listed in Table 2. A summary of
selected floristic and habitat attributes for each plant
community is supplied in Table 3.
The Themeda triandra-Senecio microglossns Cool
Moist Grassland (Siebert et al. 2002) is interpreted as
belonging to the proposed Tristachya leucothrix-Trachy-
pogon spicatiis Class (Du Preez & Bredenkamp 1991).
The Fiiirena puhescens-Schoenoplectiis corymbosiis
Wetlands (Siebert et al. 2002) is inteipreted as part of the
Miscantlms jiinceus-Schoenoplectus coiytnhosus Com-
munity (Bloem 1988), proposed here as an alliance. The
grassland and wetland plant communities of the Roos-
senekal Subcentre are classified as follows:
I. Tristachya leucothrix-Trachypogon spicatus Class of
moist mountain slopes and plateaus (Du Preez &
Bredenkamp 1991) [= Themeda triandra-Senecio
microglossns Cool Moist Grassland (Siebert et al.
2002)]
1 . Helichryso splendidi-Tristachyetiim leiicothricis
Association
2. Zantedeschio pentlandi-Aloetnm castaneae Associa-
tion
3. Brachiario serratae-Melhanietnni randii Association
3.1. Brachiario serratae-Melhanietinn randii lielicliiy-
setosiun riigiilosi Subassociation
3.1.1. Digitaria eriantha Variant
3.1.2. Alloteropsis semialata Variant
3.2. Brachiario serratae-Melhanietinn randii argyrolo-
hietosiini transvaalense Subassociation
3.2.1. Koeleria capensis Variant
3.2.2. Berkheya seminivea Variant
3.3. Brachiario serratae-Melhanietinn randii gnidieto-
snin capitatae Subassociation
3.4. Brachiario serratae-Melhanietinn randii setarieto-
sinn nigrirostis Subassociation
4. Elionnro mntici-TrachypogoneInni spicati Association
4. 1 . Elionnro nnitici-Trachypogonetinn spicati hewsieto-
snni hiflorae Subassociation
4.2. Elionnro nuitici-Trachypogonetiim spicati acacieto-
snm tortilis Subassociation
5. Jamesbrittenio macranthae-Londetietum sirnplicis
Association
5.1. Jamesbrittenio macranthae-Londetietum sirnplicis
combretetosnm hereroense Subassociation
5.2. Jamesbrittenio macranthae-Londetietum sirnplicis
eucleetosnm linearis Subassociation
II. Miscantlms jiinceus-Schoenoplectus corymbosiis
Alliance of streams (Bloem 1988) [= Fiiirena
piibescens-Schoenoplectus corymbosus Wetland
Vegetation (Siebert et al. 2002)]
6. Fnireno pnbescentis-Schoenetnm nigricantis Associa-
tion
6.1. Fnireno pnbescentis-Schoenetnm nigricantis trira-
phietosnm andropogonoidis Subassociation
6.2. Fnireno pnbescentis-Schoenetnm nigricantis pyc-
nostachetosnm reticnlatae Subassociation
6.3. Fnireno pnbescentis-Schoenetnm nigricantis bnl-
bostylietosnm hispidnlae Subassociation
7. Andropogono encomnsae-Fimbrisyletnm ferrngineae
Association
III. Drakensberg escarpment wetlands (Hilliard &
Burtt 1987)
8. Limosello maioris-Rannncnletnm meyeri Association
Description of plant communities
The Themeda triandra-Senecio microglossns Cool
Moist Grassland and scattered Fiiirena pnbescens-
Schoenoplectns coiymbosns Wetland Vegetation are pre-
dominantly restricted to the valleys, slopes and plateaus
of undulating norite hills (Siebert et al. 2002). Surface
rocks are common and abundant in many of the commu-
nities, with soil clay percentages varying from 25% to
more than 50%. The vegetation can be classified into
herbland and grassland (Edwards 1983). An important
feature of the region is the fact that the mountain and hill
ranges have a north-south orientation; therefore the
grassland communities are mostly restricted to eastern
and western aspects, crests and valleys. Grasslands have
been a long-standing component of the Afromontane
vegetation mosaic (Meadows & Linder 1989; Matthews
et al. 1993), and are therefore seen as primary vegeta-
tion. Plant communities of the grasslands and wetlands
recognized in the SCPE are classified as follows;
I. Tristachya leucothrix-Trachypogon spicatus Class
(Du Preez & Bredenkamp 1991)
1 . Helichryso splendidi-Tristachyetiim leiicothricis ass.
nova hoc loco
Nomenclatural type: releve 408 (holotypus).
Environmental data: a medium altitude grassland,
forms a transition from low to high altitude ( 1 700 m asl).
It lies in a rather moist region (> 600 mm/annum), most-
ly restricted to the summit of the Leolo Mountains on
norite. The habitat is a gentle undulating plateau (1-5°)
with a general east- west aspect (Table 3). The dominant
TABLE la. — Phytosociological table of grasslands of Sekhukhuneland Centre of Plant Endemism
Bothalia 32,2 (2002)
215
< + ■ + a:
• cs: + + Qi
: a: + o' ■ oc ■
■ • + - ■ + + o'
2 5
TABLE la. — Phytosociological table of grasslands of Sekhnkhuneland Centre of Plant Endemism (cont.)
216
Bothalia 32,2 (2002)
8
y.
It! I fir
"S IS J
, I 'J.a ■§ I ^ ^5
TABLE la. — Phytosociological table of gras.sland.s of Sekhukhuneland Centre of Plant Endemism (cont.)
Bothalia 32,2 (2002)
217
+ ai DC oi ai
DC as + + +
DC - ■ DC
t- DC + -- DC
DC ■ + DC DC + + DC
wiQ-coQ-h-'^kjco-fj: c/5CQ>5ftck'>5cs:(
« i ^ s g. ; g.| I “ £ 2 § -S I 1 -| i S “.2 ”
.2 -a g ■§ -s I -g ■§ I £ B = .2 g -a s 2 -a I S ■§ I g .2 -g g
g I I ^ £.:| I g I ^ I'l g I I t :|, I 'll' I ^I’-S g 3 ^
(/joaa.co^tcoto^'^O'a: xi0-*^Q<5“?^‘Oc^-Sa, c/sQ^co
TABLE la. — Phytosociological table of grasslands of Sekhukhuneland Centre of Plant Endemism (cont.)
218
Bothalia 32,2 (2002)
< + Qi
- DS -I
OL. ■ U. ■ CC <
OJ Oi Di ce: —
Qi ■ CC QC ■■
fti - + + - +
+ Qi ■ + Qi ■ CC
■ - cs: a: - + a
os Ctf ■ O' + cc
I ^
ii
•2 5 := Ci.
^ i 1 5 .5
S ^ I "s
=4.“o ^ b
I a ^ s -a a I
rs g ^
TABLE la. — Phytosociological table of grasslands of Sekhukhuneland Centre of Plant Endemism (cont.)
Bothalia 32,2 (2002)
219
cn an oc
- CD a:
po Cc: — CeC <
CD • — + —
• OC • QC — DC • QC
■a: ■ - cti • + - O' • -
ct:cc: + + - -dcoccic • — cq
• DC —
• + — <
— + CD • + -DC
' DC DC DC i
• DC DC ■
— -DCDC ■< •— ■ + • + +DCDC •
b + ■ • — DC+DCDC ■ ■ ■ -DC + + + -DC
+-DC+'DC'--+DC-
— - ck: • - — •— ■— • — DC+— • -DC + -
— DC+-— + •
- - +DC— +DC - -DC -DC ■ • -DC • -DC
-CD • +OCDC • •+DC + + -DC +
• + • ■ +
- DC Ct: +
C - +aC< • — + +DC+DCDC
CD - +< - + •< -< + +DC +
DC DC - - DC
•DC + •< + • -- + + +DC • + + -1
■DC •+DC — + + + + + +DCDCDC
DC DC DC
S -2 ^ = c-
' I i f
2 I. 5 i
a Cl. C -5 t
220
Bothalia 32,2 (2002)
TABLE lb, — Phyto-sociological table of wetlands of Sekhukhuneland Centre of Plant Endemism
soil type is the Mayo Form, a melanic A-horizon over a
lithocutanic B-horizon. Rock size is 300-500 mm in
diameter and rock cover 15-20%.
Diagnostic and dominant/prominent taxa: in the
SCPE this association is characterized by species group
A (Table la). Euryops hrevipapposus, Helichryswn
splendidum and Vernonia myriantha are the most promi-
nent diagnostic species of this syntaxon. Other diagnos-
tic species include the shrub Buddleja saligna and the
herbaceous Lotononis foliosa and Xerophyta viscosa.
There are no diagnostic grasses, but dominant species
include Eragwstis capensis, E. curvida and Tristachya
leucothrix. A prominent forb is Pentanisia prunelloides,
while the shrub Protea cajfra (endemic form) is conspic-
uously present.
Eloristic diversity, this grassland community is
unique for the SCPE and only a slight floristic affinity
exists with other grassland types of the SCPE in species
groups M and X (Table la). The average number of
species per releve is 30, and the total number of species
recorded for the association is 40 (two releves) (Table 3).
Three plant taxa of con.servation significance occur in
this association (Table 2), namely the endemic form of
Protea caffra, the endemic Zantedeschia jucunda that is
classified as Indeterminate in the Red Data List (also
restricted to the association) and Jamesbrittenia sile-
noides, a taxon assessed as Vulnerable in KwaZulu-
Natal.
2. Zantedeschio pentlandi-Aloetum castaneae ass. nova
hoc loco
Nomenclatural type: releve 100 (holotypus)
Environmental data: a short herbland on rocky flats,
areas where bedrock is exposed at ground level. The
plant community is associated with rock outcrops of
norite, a rock type mined extensively as dimension stone
in the region. It is situated on gentle slopes (0-5°) and
with a very high surface rock cover of 50-70% (Table 3).
Solid exposed rock sheets can cover areas of 25 m^. The
dominant soil type is the Mispah Eorm, indicating very
shallow soils over hard rocks, often restricted to crevices.
The habitat is situated on the midslopes, scarps and
crests of undulating hills.
Diagnostic and dominant/prominent ta.xa: character-
istic species are represented by species group B (Table
la). Diagnostic tree.s/shrubs of the association include
Apodytes dimidiata, Canthium suherosnm, Halleria luci-
Bothalia 32,2 (2002)
221
TABLE 2. — Endemic/near-endemic and Red Data List plant taxa of grasslands and wetlands associated with Sekhukhuneland Centre of Plant
Endemism
Family Syntaxa
1 2
3.1.1 3.1.2 3.2.1 3.2.2 3.3
Jamesbrittenia macrantha
J. silenoides
Jasmimim quinatwn
Melhania randii
Nuxia gracilis
SCHR ^
SCHR |~Nr~|
OLEA .
STER .
LOGA .
I K#1 K#T
Endemism: $, endemic; #, near-endemic.
Red Data List; I. Indeterminate; K. Insufficiently Known; R. Rare; N, Not threatened i
Abundance in communities: 1. abundant; +. frequent; r. rare; . absent.
Collectors: P = P.S. Swartz; S = S.J. Siebert; W = A.E. van Wyk.
Blocks represent community/syntaxon specific taxa.
northern provinces of South Africa, but in other areas of southern Africa.
da and Olinia emarginata. Diagnostic forbs are Thesium
burkei, the succulents Crassula sarcocaulis and Aloe
pretoriensis, and the geophytes Boophone disticha and
Zantedeschia pentlandii. Aristida junciformis and
Cyrnbopogon excavatus are the diagnostic grasses of this
association. Prominent plants are the succulent Aloe cas-
tanea, the shrubby Rhoicissisus tridentata and the grass-
es Eragrostis pseudosclerantha, Themeda triandra and
Tristachya leucothrix.
Floristic diversity, this association exhibits a typical
floristic relationship with other grasslands of the SCPE
(Table la). The average number of species encountered
per sample plot is 36, with 105 species the total number
recorded for the association (five releves) (Table 3).
Fifteen taxa of conservation significance are present in
the association (Table 2), five are SCPE endemics, six
are SCPE near-endemics and seven are Red Data List
taxa. This association has a high number of plant taxa
with conservation status in the southern region of the
SCPE. The association also has a high number of plant
taxa with conservation status restricted to it, such as
Asclepias sp. {Siebert 27) (endemic), Eucomis montana
(Rare), Xerophyta retinervis form (endemic) and
Zantedeschia pentlandii (endemic. Rare).
3. Brachiario serratae-Melhanietum randii ass. nova
hoc loco
Nomenclatural type: releve 321 (holotypus).
Environmental data: short, dense grassland associated
with plateaus or terraces on rocky undulating hills. The
association is found on all aspects, on gentle to moderate
slopes (5-15°) of footslopes, midslopes, scarps and
crests (Table 3). Soils are characterized by a melanic A-
horizon underlain by hard rock (Milkwood Form) or a
soft carbonate horizon (Steendal Form). Rock cover is
15-75% and rock diameter 0.1-1 m (Table 3).
Diagnostic and dominant/prominent taxa: character-
istic species of the association are represented by species
group C (Table la). The dominant diagnostic forbs are
Callilepis leptophylla. Dicoma zeyheri, Gnidia capitata,
Melhania randii and Vemonia oligocephala. Woody
species typical of the association include the geoxylic
222
Bothalia 32,2 (2002)
TABLE 3. — Environmental factors and selected attributes associated with different plant communities
* C, crest; S. scarp: M. midslope; F. footslope; V, valley.
** Ms, Mispah; Ar, Arcadia; Sn, Sleendal; My, Mayo; Mw. Milkwood; Sd. Shortlands.
suffrutices Elephantorrhiza elephantina and Rhus wilm-
sii, and the small trees Acacia cajfra. Protea cajfra and
Vitex obovata subsp. wilmsii. Other prominent forbs
include Clerodendnim triphyllum, Senecio latifolius and
Tephrosia purpurea. Prominent grasses for the associa-
tion are Aiidropogon chinensis, Brachiaria serrata,
Setaria sphacelata, Themeda triandra, Trachypogon spi-
catus and Tristachya leucothrix.
Floristic diversity, the average number of species
encountered per sample plot in this association is 48,
with the total number of plant species being a minimum
of 1 19 taxa (29 releves) (Table 3). There are 25 plant taxa
of conservation value in the association, of which four
are restricted to it. These include taxa such as the near-
endemics, Argyrolobium wilmsii and Pachycarpus trans-
vaalensis (Table 2). Red Data List taxa include Callilepis
leptophylla (status is Rare in KwaZulu-Natal), Melhania
randii (status is Insufficiently Known in the northern
provinces) and Scilla natalensis (status is Vulnerable in
the Free State and KwaZulu-Natal). This association har-
bours the highest number of plant taxa with conservation
status in the southern region of the SCPE and, together
with plant community 2, the highest number of Red Data
List taxa.
3.1. Brachiaria serratae-Melhanietum randii helichry-
setosum rugulosi subass. nova hoc loco
Nomenclatural type: releve 321 (holotypus).
Environmental data: short rocky grassland communi-
ties on clay soils such as Mayo and Milkwood Forms,
occurs on midslopes, scarps and crests of undulating
norite hills on slopes of 3-9° on all aspects. Rock cover
is 20-50% with rock size of 0.15-1 m in diameter (Table
3).
Diagnostic and dominant/prominent taxa: species
group F contains the diagnostic species for this subasso-
ciation, with the taxa Helichrysum rugulosum, Pimpi-
nella cajfra and Tephrosia elongata (Table la). Other
characteristic species include the forbs Acalypha puncta-
ta, Leonotis ocyrnifolia and Senecio lygodes, as well as
the grasses Panicum natalense and Tristachya biseriata.
Trees that are prominent are Euclea crispa and Vitex obo-
vata subsp. wilmsii. Predominant grasses are Themeda
triandra and Tristachya leucothrix.
Eloristic diversity, a strong floristic affinity exists
with all the subassociations of the association (Table la).
The average number of species encountered per sample
plot in this subassociation is 49, with the total number of
plant species being a minimum of 130 taxa (14 releves)
(Table 3). Two plant taxa of conservation value are
restricted to the subassociation (Table 2).
3.1.1. Digitaria eriantha Variant
Environmental data: rocky grassland on shallow clay
soils of moderately sloped hill scarps and crests (Table
3). Rock size is 600 mm in diameter and cover is 25%
(Table 3).
Diagnostic and dominant/prominent taxa: diagnostic
taxa include the forbs Agapanthus inapertus, Berkheya
densifolia, Cyanotis speciosa, Indigofera hedyantha,
Monsonia attenuata and Tephrosia longipes and the
grass Digitaria eriantha (species group D; Table la).
Other frequently occurring grasses are Andropogon
schirensis, Brachiaria serrata and Setaria sphacelata.
Eloristic diversity: a strong floristic similarity exists
with plant community 3.2.1, probably due to the similar-
ity in their rock size and cover (species group H; Table
la & Table 3). Five SCPE endemics, five near-endemics
and seven Red Data List taxa are found in this variant
(Table 2). Of the 112 taxa recorded for the variant (seven
releves), only 15 are of conservation value (one is
restricted to it). The average number of species encoun-
tered per sample plot is 47 (Table 3).
3.1.2. Alloteropsis semialata Variant
Environmental data: rocky grasslands on shallow clay
soils occurring on relatively steep midslopes of hills.
Rock cover is 35%, with rocks 350 mm in diameter
(Table 3).
Diagnostic and dominant/prominent taxa: Berkheya
onopordifolia, Helichrysum albilanatum, H. nudifolium
and the succulent Kalanchoe rotundifolia, are the diag-
nostic forbs, with Alloteropsis semialata and Sporobolus
pectinatus the diagnostic grasses species (species group
E; Table la). Dominant dwarf shrubs are Clutia pulchel-
la and Rhus discolor.
Eloristic diversity: the community shares a floristic
identity within various species groups (Table la). Five
SCPE endemics, and seven near-endemics and seven
Red Data List taxa are found in this variant (Table 2). It
has 17 taxa of conservation value, the highest number for
this paper, with only one taxon restricted to it. The aver-
age number of species recorded per sample plot is 51
(richest plant diversity of all the study area’s grassland
Bothalia 32,2 (2002)
223
communities), with a total number of 130 plant taxa
(seven releves) (Table 3).
3.2. Brachiario serratae-Melhanietum randii argyrolo-
bietosum transvaalense subass. nova hoc loco
Nomenclatural type: releve 8 (holotypus).
Environmental data: a relatively tall, rocky grassland
of midslopes and crests of undulating norite hills, occurs
on soils of the Milkwood and Steendal Forms. It lies on
relatively steep, sloped areas (3-15°). Rock cover is
25-75%, with rock diameter 100-950 mm (Table 3).
Diagnostic and dominant/prominent taxa: diagnostic
species for this vegetation type is Berkheya seminivea
and Drimiopsis atropurpurea. This subassociation is
characterized by group G (Table la). Prominent taxa that
occur in this vegetation unit are the forbs Acalypha punc-
tata, Argyrolobium transvaalense, Barleria ovata,
Pachycarpus transvaalensis, Rhynchosia spectabilis,
Tephrosia purpurea and Vernonia natalensis. Important
grasses for this vegetation type are Bracliiaria serrata,
Diheteropogon amplectens, Setaria sphacelata and
Themeda triandra. Prominent woody species include the
geoxylic suffrutices Elephantorrhiza elephantina and
Rhus discolor.
Eloristic diversity: the community exhibits a strong
floristic affinity with all the grasslands of the study area
(Table la). The average number of species encountered
per sample plot in this subassociation is 49, with the total
number of plant species being a minimum of 119 taxa
(nine releves) (Table 3). No plant taxa of conservation
value are restricted to it (Table 2).
3.2.1. Koeleria capensis Variant
Environmental data: relatively tall rocky mountain
grassland on east-west aspects of steep midslopes and
crests occurs on shallow clay soils, is covered by ± 30%
rock, with a relatively large rock diameter (on average
650 mm) (Table 3).
Diagnostic and dominant/prominent taxa: no diag-
nostic species occur in this variant. Character species
include the prominent forbs Crabbea hirsuta and
Ipomoea obscura, and prominent grasses are Eulalia vil-
losa and Koeleria capensis (species group G; Table la).
Other dominant plants include the geoxylic suffrutex
Rhus discolor and the small tree Euclea crispa. Grass
cover is dense and species rich.
Eloristic diversity: a strong floristic similarity is
shared with plant community 3.1.1, probably due to sim-
ilarity in rock size and cover (species group H; Table la
& Table 3). Of the 14 taxa of conservation value in this
variant, five are SCPE endemics, six near-endemics and
five Red Data List taxa (Table 2). The average number of
species encountered per sample plot is 48, with a total of
119 plant species (6 releves), the second richest plant
diversity in the study area’s grasslands (Table 3).
3.2.2. Berkheya seminivea Variant
Environmental data: rocky grassland communities on
shallow clays of moderate midslopes and crests that are
situated on east-west aspects (Table 3). Average rock size
is 200 mm and cover is 40% (Table 3).
Diagnostic and dominant/prominent taxa: there are
no diagnostic species, but characteristic species for this
variant include the forbs Berkheya seminivea,
Drimiopsis atropurpurea, Helichrysum cephaloideum
and Thesium magalismontanum the most frequent
(species group G; Table la). Other important taxa are
Andropogon schirensis, Argyrolobium transvaalense,
Eragrostis chloromelas and Rhynchosia spectabilis.
Eloristic diversity: this variant is floristically typical
of its association, but is characterized by the absence of
the sister variant’s character species (species group H;
Table la). Of its 13 taxa of conservation value, five are
SCPE endemics, four near-endemics and six Red Data
List taxa (Table 2). The average number of species
encountered per sample plot is 50, with the total being 95
taxa (three releves) (Table 3).
3.3. Brachiario serratae-Melhanietum randii gnidieto-
sum capitatae subass. nova hoc loco
Nomenclatural type: releve 86 (holotypus).
Environmental data: rocky grassland communities on
black clay soils. The habitat is found on footslopes and mid-
slopes of undulating norite hills. The gentle slopes vary from
5-9°, with an east-west aspect the norm. Soils characteristic
of these slopes is the Milkwood Eorm. Rock cover varies
from 35^0% and rock size from 400-750 mm in diameter.
Diagnostic and dominant/prominent taxa: species
group J contains the characteristic species for this subas-
sociation, with prominent forbs such Becium obovatum.
Convolvulus sagittatus, Gerbera ambigua and Vernonia
galpinii (Table la). The forbs Senecio microglossus and
S. latifolius, and the grasses Themeda triandra and
Tristachya leucothrix are the most dominant in this sub-
association. Other important taxa are the woody species
Elephantorrhiza elephantina and Euclea crispa, succu-
lents Aloe castanea and A. greatheadii, and grass species
such as Eragrostis superba, Setaria sphacelata. Sorghum
bicolor and Tristachya rehmannii.
Eloristic diversity: a strong floristic affinity exists
with certain plant communities of the association
(species group J), but excludes plant communities 3.1.1
and 3.4 (Table la). The average number of species
encountered per sample plot in this subassociation is 46,
the total number of plant species being 109 (six releves)
(Table 3). Although 15 taxa with conservation value
occur in this subassociation (Table 2), namely five SCPE
endemics, six near-endemics and six Red Data List taxa,
no plant taxa with conservation value are restricted to it.
3.4. Brachiario serratae-Melhanietum randii setarieto-
sum nigrirostis subass. nova hoc loco
Nomenclatural type: releve 64 (holotypus).
Environmental data: moist rocky mountain grassland
on black turf soils lies on the lower part of gentle, sloped
footslopes, 3-5°. It is found predominantly on soils of the
Arcadia Eorm. Rock cover is ± 15-30%, rocks, with a rel-
atively small average size of 100-250 mm in diameter
(Table 3).
Diagnostic and dominant/prominent taxa: diagnostic
species are represented by species group L (Table la).
224
Bothalia 32,2 (2002)
Diagnostic forbs are Lotononis adpressa and Scleria
dieterlenii, and the grass Setaria nigrirostis. No tree
species are diagnostic, but Protea cajfra is extremely
prominent in this subassociation. Other important forbs
are Bulbostylis contexta, Lotononis macrosepala and
Pegolettia lanceolata. Berkheya insignis, Hermannia
ontonii, Hypoxis rigidula, Melhania randii, Senecio
microglossus and Thesium gracilentum are prominent
forbs. Prominent grasses of the subassociation are
Elionurus muticus and Eragrostis nindensis. Grasses
found frequently are Brachiaria serrata, Dihetewpogon
amplectens, Themeda triandra, Trachypogon spicatus
and Tristachya leucothrix.
Floristic diversity, the subassociation shows its strong
floristic affinity within association 3 in species groups C
and M (Table la). More detailed future studies could
suggest its upgrading to the level of association. The
average number of species encountered per sample plot
in this subassociation is 44, with 77 plant species the
total number (four releves) (Table 3). Twelve taxa with
conservation value occur in this subassociation (Table 2),
namely four SCPE endemics, five SCPE near-endemics
and five Red Data List taxa. No plant taxa with conser-
vation value are restricted to it.
4. Elionuro mutici-Trachypogonetum spicati ass. nova
hoc loco
Nomenclatural type: releve 112 (holotypus).
Environmental data: this rocky mountain grassland
occurs on deep black turf soils (500-750 mm) which lie
on moderately sloped, lower footslopes and valley bot-
toms, 5-15°. It is found predominantly on vertic Arcadia
and Steendal Forms. Rock cover is 10^0% and rocks
have a small average size of 100-150 mm in diameter
(Table 3).
Diagnostic and dominant/prominent taxa: diagnostic
species are presented in species groups N (Table la) and
are characterized by the small trees Rhanmus prinoides
and Rhus rogersii, the grasses Brachiaria eruciformis
and Setaria incrassata, and the forbs Indigofera evan-
siana, Kohautia caespitose and Striga elegans. Forbs
are abundant in the vegetation unit and prominent
species include Albuca setosa, Berkheya onopordifolia,
Felicia rnuricata, Helichrysum albilanatum, H. nudifoli-
um, Justicia anagalloides, Kalanchoe rotundifolia and
Rubia horrida. Prominent grasses of the association are
Alloteropsis semialata and Sporobolus pectinatus, and
dominant grasses such as Brachiaria serrata, Eragrostis
chloromelas, Heteropogon contortus and Panicwn
natalense.
Floristic diversity: the average number of species
encountered per sample plot in this association is 44,
with the total number of plant species being a minimum
of 84 taxa (12 releves) (Table 3). There are 1 1 plant taxa
of con.servation value in the association (Table 2), of
which two are restricted to it, including Rhus rogersii, a
shrub assessed as Insufficiently Known for Swaziland in
the Red Data List.
4. 1 . Elionuro mutici-Trachypogonetum spicati bewsieto-
sum biflorae subass. nova hoc loco
Nomenclatural type: releve 42 (holotypus).
Environmental data: moist, cool mountain grassland
on turf soils, lies on gentle, sloped footslopes, 3-5°,
restricted to deep Steendal soils. Rock cover is ±
20-25% and rocks have an average diameter of 50-150
mm (Table 2).
Diagnostic and dominant/prominent taxa: diagnostic
species are represented by species group O (Table la).
Only one woody species, the shrubby Rhus tumulicola
var. meeuseana, is diagnostic of the subassociation, with
Acacia karroo being a dominant and extremely common
tree. Argyrolobium amplexicaule, Conyza podocephala,
Helichrysum spp., Rhynchosia albissima and Senecio
spp. are the diagnostic forbs. Bewsia biflora is the only
diagnostic grass. Other prominent forbs include
Pearsonia grandifolia, Senecio microglossus and Striga
elegans. Dominant grasses include Brachiaria eruci-
formis, B. serrata, Diheteropogon amplectens, Elionurus
muticus and Themeda triandra.
Floristic diversity: a floristic link with plant commu-
nity 4.2 in species group N, indicates the relationship
between these two subassociations on turf (Table la),
although the environmental data differ to a great extent.
The average number of species encountered per sample
plot is 46. The total number for this subassociation is 84
(four releves) (Table 3). One plant taxon with conserva-
tion value, the near-endemic Rhus tumulicola var.
meeuseana, is restricted to it. Nine taxa with conserva-
tion value occur in this subassociation and comprise four
SCPE endemics, three near-endemics and two Red Data
List taxa (Table 2).
4.2 Elionuro mutici-Trachypogonetum spicati acacieto-
sum tortilis subass. nova hoc loco
Nomenclatural type: releve 112 (holotypus).
Environmental data: wooded grassland communities
on black turf soils. These units occur in valley bottoms
between undulating norite hills. The gentle slope varies
from 1-3° and eastern or western aspects are predomi-
nant. Soil characteristic of these slopes is the Arcadia
Form. Rock cover is 5-10% and rock diameter is 50-100
mm (Table 3).
Diagnostic and dominant/prominent taxa: species
group P contains the diagnostic species for this associa-
tion (Table la). Two trees are diagnostic for the associa-
tion, namely Acacia tortilis and Dichrostachys cinerea.
These taxa are often dominant in the Panico
maximi-Acacietea tortilis class (Winterbach et al. 2000),
a class representing microphyllus savanna in the central
Bushveld of South Africa. Diagnostic grasses for the
subassociation are Digitaria sanguinalis, Panicwn
deustum, P. maximum and Sporobolus fimbriatus. The
association is dominated by forbs such as Chrysanthe-
moides monilifera, Indigastrum burkeanum, Jasminum
quinatum and Kyphocarpa angustifolia. Other prominent
taxa include the woody species Rhanmus prinoides and
Acacia karroo, the herbaceous species Asparagus suave-
olens, Berkheya insignis, Gnidia caffra and Senecio micro-
glossus, and the grasses Cymbopogon validus, Dihetero-
Bothalia 32,2 (2002)
225
pogon amplectens, Elionurus muticus, Hyparrhenia fili-
pendula, Loudetia simplex, Setaria spliacelata and
Themeda triandra.
Floristic diversity: a floristic link exists with the more
herbaceous plant community 4.1, in species group N
(Table la), but a difference in vegetation structure is
apparent. The average number of species encountered
per sample plot is 41 (Table 3). The total number of plant
species for this subassociation is 65 (four releves). No
plant taxa with conservation value are restricted to the
community, although three SCPE endemics, three SCPE
near-endemics and two Red Data List taxa were record-
ed.
5. Jamesbrittenio macranthae-Loudetietum simplicis
ass. nova hoc loco
Nomenclatural type: releve 47 (holotypus).
Environmental data: wooded grassland associated
with anomalous habitats (Siebert 1998) occurs on red
clay and white loam soils of predominantly the Mispah
Form that are characterized by serpentiniferous chemical
compositions. It lies on moderate footslopes of 5-9° on
all aspects of undulating hills. Rock cover is ± 20-70%
and with large rocks 100^00 mm in diameter (Table 3).
Diagnostic and dominant/prominent taxa: characteris-
tic species are represented by species group S (Table la).
Herbs are diagnostic of this community, namely
Helichrysum uninerx’ium, Jamesbrittenia macrantha,
Lotononis wilmsii. Polygala hottentotta and Rhynchosia
kornatiensis. Elephantorrhiza praetermissa is the charac-
teristic woody species and Aristida adscensionis and
Melinis repens the characteristic grasses. Prominent
forbs are Dicoma anomala, Gnidia cajfra, Melhania
prostrata and Thesiiim midtiramulosum. Grasses of
importance include Aristida congesta, Diheteropogon
amplectens, Elionurus muticus, Eragrostis nindensis,
Loudetia simplex, Themeda triandra and Tristacliya leu-
cothrix. Protea cajfra and Vitex obovata subsp. wilmsii
are prominent trees and Rhus wilmsii is a prominent
geoxylic suffrutex of the association.
Floristic diversity: the average number of species
encountered per sample plot is 36, with the total number
of plant species being a minimum of 72 taxa (nine
releves) (Table 3). Fifteen taxa of conservation value
occur and the association (Table 2) has the highest num-
ber of taxa with conservation status restricted to it
(seven), and includes plant species such as the near-
endemics Helichrysum uninervium and Lotononis wilm-
sii, and the endemic and Red Data-listed Jamesbrittenia
macrantha.
5.1. Jamesbrittenio macranthae-Loudetietum simplicis
combretetosum hereroense subass. nova hoc loco
Nomenclatural type: releve 47 (holotypus).
Environmental data: wooded grassland of rocky foot-
slopes with clay soils. It lies on moderate slopes of 7-9°
on north-south aspects of rocky ridges of magnetite.
Soils are predominantly the Mispah Form. Rock cover is
± 60-70% and large rocks, 100-400 mm in diameter
(Table 3).
Diagnostic and dominant/prominent taxa: the diag-
nostic species are represented by species group T (Table
la), and include one tree species, Combretum herero-
ense. The forbs Aneilema longirrhizum, Chamaecrista
comosa, Hemizygia petrensis, Ledebouria marginata,
Lotononis calycina, Phyllanthus incurxms and Triumfetta
sonderi are diagnostic. The most important dominant
grasses include Loudetia simplex, Themeda triandra,
Tristacliya leucothrix and T. rehmannii. Other dominant
taxa include Protea cajfra, Rhus wilmsii and Senecio
microglossus.
Floristic diversity: the subassociation shows a floris-
tic affinity with the turf grasslands of the Roossenekal
Subcentre in species groups V and W (Table la). A spe-
cific link with the grasslands of the study area, which
includes its sister subassociation, is indicated in species
group X. The average number of species encountered per
sample plot is 39, with the total number for this subasso-
ciation being 70 (four releves) (Table 3). Five SCPE
endemics, six near-endemics and three Red Data List
taxa are found in this subassociation (Table 2). Of its 1 1
taxa of conservation value, only the near-endemic
Aneilema longirrhizum is restricted to it.
5.2. Jamesbrittenio macranthae-Loudetietum simplicis
eucleetosum linearis subass. nova hoc loco
Nomenclatural type: releve 106 (holotypus).
Environmental data: wooded, rocky grassland on
white loam soils and is restricted to exposed norite sur-
faces. It lies on moderately sloped footslopes of 5-7° on
all aspects. Soils are predominantly the Mispah Form.
Rock cover is ± 20-70% and large rocks, 100-200 mm in
diameter (Table 3).
Diagnostic and dominant/prominent taxa: species
group U (Table la) represents the diagnostic species of
the subassociation. The diagnostic grass species of the
subassociation is Andropogon chinensis. Six forbs are
diagnostic, namely Indigofera tristoides, Ipomoea bathy-
colpos var. sinuatodentata, Jamesbrittenia burkeana,
Phyllanthus glaucophyllus. Polygala sp. nov. (Siebert
449) and Seddera capensis. Euclea linearis is the diag-
nostic woody species of the subassociation. Dominant
plants in the community are the forbs Dicoma anomala
and Helichrysum uninervium, and grasses Aristida
adscensionis, Diheteropogon amplectens and Elionurus
muticus. Protea cajfra, Rhus wilmsii and Vitex obovata
subsp. wilmsii are dominant woody species.
Floristic diversity: a slight floristic affinity is visible
between this subassociation and plant communities 4.2
and 5.1 in species group V (Table la). The average num-
ber of species encountered per sample plot is 33, with the
total number for this subassociation being 72 (five
releves) (Table 3). It is one of the syntaxa with the high-
est numbers of SCPE endemics and SCPE near-
endemics. It also has three Red Data List taxa (Table 2).
Of its 14 taxa of conservation value, four taxa, namely
the SCPE near-endemics Rhus keetii and Euclea linearis
(form), and the SCPE endemics Ipomoea bathycolpos
var. sinuatodentata and Polygala sp. (Siebert 449), are
restricted to it. This number is the second highest for any
community in the Roossenekal Subcentre.
226
Bothalia 32,2 (2002)
II. Miscanthus junceus-Schoenoplectus corymbosus
Alliance (Bloem 1988)
6. Fuireno pubescentis-Schoenetum nigricantis ass. nova
hoc loco
Nomenclatural type: releve 73 (holotypus).
Environmental data: within the Roossenekal and
Leolo Subcentres, this association represents hygro-
philous vegetation of mountain streams and seepage
areas and occurs on wet sites where rocks of ± 150 mm
diameter cover ± 25% of the soil surface. These areas
have gentle slopes (3°) and the soils are typically clay on
solid rock, with a sandy alluvial layer on the surface.
Diagnostic and dominant/prominent taxa: species
group AA (Table lb) contains the diagnostic species.
Dense stands of the diagnostic sedge Schoenus nigricans
and the diagnostic grass Hyparrhenia tamba dominate
the vegetation. Other diagnostic forbs are Chironia pur-
purascens, Equisetum ramosissimum and Hypoxis
argentea. Sedges such as Fuirena pubescens and
Schoenoplectus corymbosus are also prominent, while
Andropogon eucomis, Cymbopogon validus, Hyparr-
henia filipendula and Imperata cylindrica are prominent
grasses. Acacia karroo and Rhus leptodictya are the
woody species that may occur in the association.
Floristic diversity, a strong floristic affinity with the
wetland community 7 is indicated in species group AI
and some relationships exist with the grasslands (species
group Y) (Table lb). The azonal wetland vegetation is
not as rich in plant diversity as the grasslands. The aver-
age number of species encountered per sample plot is
only 27, with the total number of plant species being a
minimum of 52 taxa (11 releves) (Table 3). There are,
however, four taxa of conservation value that occur in
this association (Table 2), and two taxa with conserva-
tion status restricted to it, namely the Red Data listed
orchid Disa rhodantha (Insufficiently Known), and the
shrub, Nuxia gracilis (Insufficiently Known).
6.1. Fuireno pubescentis-Schoenetum nigricantis trira-
phietosum andropogonoidis subass. nova hoc loco
Nomenclatural type: releve 73 (holotypus).
Environmental data: sparsely wooded, moist herbland
and grassland along non-perennial mountain streams in
rocky areas lying on gentle to moderate midslopes
(5-7°). Rock size 350-750 mm in diameter and rock
cover is 35^5%.
Diagnostic and dominant/prominent taxa: species
group AB (Table lb) contains the diagnostic species for
this subassociation, which include the woody species
Acacia karroo, Nuxia gracilis and Rhus leptodictya.
Diagnostic forbs are the fern Adiantum capillus-veneris,
and the sedges Coleochloa setifera, Dittrichia graveolens
and Juncus punctorius. Triraphis andropogonoides is a
diagnostic grass. Prominent forbs are the sedges Euirena
pubescens, Schoenoplectus corymbosus and Schoenus
nigricans. Andropogon eucomis, Aristida bipartita and
Cymbopogon validus are the most dominant grasses.
Floristic diversity: a notable relationship exists with
plant community 6.2 in species group AC (Table lb).
The average number of species encountered per sample
plot is 28, with the total number for this subassociation
being 51 (four releves) (Table 3). The association has
three taxa of conservation value, including one SCPE
endemic, one SCPE near-endemic and two Red Data List
taxa (Table 2).
6.2. Fuireno pubescentis-Schoenetum nigricantis pyc-
nostachetosum reticulatae subass. nova hoc loco
Nomenclatural type: releve 23 (holotypus).
Environmental data: dense moist herbland (reedbed)
and grassland along permanent mountain streams occurs
on gentle midslopes and footslopes of 1-3°. Rock cover
is ± 5-15% and small rocks, 50-150 mm in diameter
(Table 3).
Diagnostic and dominant/prominent taxa: character-
istic species are represented by species group AC and AG
(Table lb). No woody or grass species are diagnostic of
this community. Characteristic forb species include the
sedges Cyperus sexangularis and Kyllinga erecta, and
the forb Berula erecta. Dominant grass species are
Miscanthus junceus and Phragmites australis, the forbs
Pycnostachys reticulata and Senecio gerrardii, and the
sedges Fuirena pubescens, Schoenoplectus corymbosus
and Schoenus nigricans. Andropogon eucomis, Cymbo-
pogon validus and Hyparrhenia fdipendula are promi-
nent grasses.
Floristic diversity: a floristic affinity exists with the
adjacent grassland plant community 3.4 (species group
L) (Table la), and with the wetland plant community 7
(species group AG) (Table lb). The average number of
species encountered per sample plot is 27, with the total
number for this subassociation being 42 (three releves)
(Table 3). Only two taxa of conservation value, namely
Red Data List taxa, are found in this subassociation
(Table 2).
6.3. Fuireno pubescentis-Schoenetum nigricantis bulbo-
stylietosum hispidulae subass. nova hoc loco
Nomenclatural type: releve 34 (holotypus).
Environmental data: herbland and grassland occurs in
moist valley bottoms or on mountain crests, usually in
non-permanent seepage areas along streams. It is associ-
ated with gentle slopes of 1-3°. Scattered stones cover ±
10-20% of the soil surface with an average diameter of
50-100 mm (Table 3).
Diagnostic and dominant/prominent taxa: diagnostic
species are presented in species group AD (Table lb),
including the forbs, Alepidea amatymbica and Bulbo-
stylis hispidula, and the grasses, Bothriochloa insculpta
and Microchloa cajfra. The woody Rhus leptodictya is a
prominent species. Predominant forbs include Artemisia
afra and Clijfortia nitidula, and the sedges Mariscus
congestus and Schoenus nigricans. Andropogon euco-
mis, Cymbopogon validus, Hyparrhenia filipendula,
Ischaemum fasciculatum and Miscanthus junceus are the
dominant grasses.
Floristic diversity: a strong floristic affinity exists
with the grassland community 4.1 (species group O), but
Bothalia 32,2 (2002)
227
a distinct affinity also exists with the wetland plant com-
munity 7 in species group AH (Table lb). In this subas-
sociation the sedges are less prominent and should be
seen as a moist grassland-wetland ecotone. The average
number of species encountered per sample plot is 25,
with 52 taxa the total number for this subassociation
(four releves) (Table 3). There are four taxa of conserva-
tion value that include one SCPE endemic, one SCPE
near-endemic and three Red Data List taxa (Table 2).
7. Andropogono eucomusae-Fimbristyletumferrugineae
ass. nova hoc loco
Nomenclatural type: releve 192 (holotypus).
Environmental data: wooded herbland and grassland
along larger rivers in valleys, such as the Steelpoort
River. The vegetation covers the zone directly adjacent
to streams, on permanently moist soils. The area has a
gentle slope of 1-3°. Rock cover is ± 20-30% and rock
diameter is 150-200 mm (Table 3).
Diagnostic and dominant/prominent taxa: diagnostic
species are represented by species group AF (Table lb).
The vegetation unit is dominated by diagnostic forbs,
which include the prominent weedy aliens Conyza bonar-
iensis and Flaveria bidentis, the sedges Cyperus margina-
tus, Mariscus rehmannianus, M. sumatrensis, and the forb
Polygonum meisnerianum. The presence of weeds is the
result of disturbance caused by annual floods. Salix
mucronata is the diagnostic woody species and Eragrostis
gummiflua the diagnostic grass. Hygrophilous grasses and
sedges such as Miscanthus junceus and Schoenoplectus
corymbosus are dominant. Acacia karroo and Rhus lepto-
dictya are common small trees of the association.
Andropogon eucomis, Cymbopogon validus, Hemarthria
altissima, Hyparrhenia hirta and Imperata cylindrica are
dominant grasses of the association.
Floristic diversity: a strong floristic affinity exists
with the wetland plant community 6 (species group AI)
and a weaker affinity with the grasslands (species group
Y) (Table lb). The average number of species encoun-
tered per sample plot is 27, with the total number of plant
species being 52 taxa (four releves) (Table 3). This asso-
ciation has the lowest number of taxa with a conservation
status (Table 2), including the Red Data List taxon
Eucomis autumnalis subsp. clavata, which is classified
as Rare in the Free State and Vulnerable in KwaZulu-
Natal, and the SCPE endemic form of Acacia karroo.
III. Drakensberg escarpment wetlands (Hilliard &
Burtt 1987)
8. Limosello maioris-Ranunculetum meyeri ass. nova
hoc loco
Nomenclatural type: releve 410 (holotypus).
Environmental data: this rare association represents
dense herbland of moist seepage areas, recorded only on
the summit plateaus of the Leolo Mountains around
1 800 m and is usually associated with black, marshy
clay soils. A short, dense cover of nongrassy forbs dom-
inates the vegetation. It lies on gentle slopes of 1-3° and
± 10% of the soil surface is covered by small stones with
an average diameter of 50 mm (Table 3).
Diagnostic and dominant/prominent taxa: diagnostic
species are represented by species group AJ (Table lb).
The most predominant diagnostic taxa of the association
are the small forbs Limosella maior. Ranunculus meyeri
and R. multifidus. Sporobolus centrifugus is the diagnos-
tic grass. Schoenoplectus corymbosus is a dominant
sedge in the association.
Floristic diversity: a very slight floristic affinity exists
with the other wetland plant communities of the area
(species group AK) and also with the grasslands of the
study area (species group Y) (Table lb). It is much relat-
ed to similar wetland communities below the Drakens-
berg escarpment above 1 800 m (Hilliard & Burtt 1987).
The average number of species encountered per sample
plot is 20, with the total number of plant species being 24
taxa (two releves) (Table 3). Together with plant com-
munity 7, this association has the lowest number of taxa
with a conservation status, namely two. These include
the Red Data List species Eucomis autumnalis subsp.
clavata, and the undescribed SCPE endemic Tulbaghia
sp. {Siebert 1304), which may be a new genus of the
Alliaceae.
Vegetation key
A dichotomous vegetation key is presented to facili-
tate identification of the various syntaxa found in the
study area (Table 4). The definitions are broad indica-
tions of the syntaxa and should be seen as a guideline,
rather than precise descriptions. A diagnostic characteris-
tic of the vegetation or habitat is given, followed by the
most diagnostic and conspicuous species of a particular
syntaxon. The first species listed is restricted to the spe-
cific syntaxon only, and the second is dominant in the
syntaxon, but may occur in other syntaxa. Where one
species is given, no species were restricted to the partic-
ular syntaxon only.
Ordination
A scatter diagram displaying the distribution of the
releves along the second and third ordination axes is
presented for both grassland and wetlands in Figure 2
(eigen values: axis 2 = 0.471; axis 3 = 0.325) and Figure
3 (eigen values: axis 2 = 0.458; axis 3 = 0.256) respec-
tively. Vegetation units are represented as clusters, their
distribution on the scatter diagram corresponding with
certain physical environmental conditions. The gradient
described by the x-axis in both instances, is related to
drainage and hence, soil moisture. In the grasslands, the
communties with the highest available soil moisture are
situated at the left of the diagram (Figure 2), but com-
munities of the wetlands with poor drainage on water-
logged soils are to the right (Figure 3). In addition, com-
munities of the grasslands on clay soils of steep slopes
are situated to the left of the diagram and communities
on gravel soils of steep slopes to the right (Figure 2).
The clay soils have the highest moisture availability,
although the run-off is high. Grassland communities of
turf soils on moderate slopes are positioned in the cen-
tre of the diagram, because these soils have a high soil
moisture percentage, which is unavailable due to reten-
228
Bothalia 32,2 (2002)
TABLE 4. — Key to syntaxa of grasslands and wetlands of undulating norite hills of Sekhukfiuneland Centre of Plant Endemism
Leads/description
Go to/syntaxon
la Grassland (Tristachya leucothrix & Senecio microglossus)
b Wetland {Schoenoplectus corymbosus & Cymbopogon validus)
2a Mispah soils on foolslopes {Jamesbrittenia macrantha & Dicoma anomala)
b Other soils and terrain types [Tristachya leucothrix)
3a High-altitude seepage [Ranunculus meyeri & Schoenoplectus corymbosus)
b Streams/rivers [Fuirena pubescens & Schoenoplectus corymbosus)
4a Gentle sloped [Euclea linearis & Loudetia simplex)
b Larger rock size and cover [Combretum hereroense & Brachiaria serrala)
5a Higher altitudes [Helichrysum splendidum & Pentanisia prunelloides)
b Lower altitudes [Acacia caffra)
6a Valley rivers [Mariscus rehmannianus & Andropogon eucomis)
b Mountain streams [Schoenus nigricans & Miscanthus junceus)
7a Larger rock size and cover [Zantedeschia pentlandii & Aloe castanea)
b No Mispah soils [Trachypogon spicatus)
8a Rocky streams [Cyperus sexangularis & Hyparrhenia tamba)
b Stream seepage [Bulbostylis hispidula & Chironia purpurascens)
9a Vertic A-horizon [Rhamnus prinoides & Setaria sphacelata)
b Melanie A-horizon [Rhynchosia spectabilis & Protea caffra)
10a Level slope [Cyperus sexangularis & Berula erecta)
b Sleeper slope, more rocky [Triraphis andropogonoides & Kyllinga erecta)
1 la Lower rock cover percentage [Acacia tortilis & Hyparrhenia filipendula)
b Steeper slope [Pearsonia grandifolia & Senecio microglossus)
12a All aspects [Helichrysum rugulosum & Clerodendrum triphyllum)
b East-west aspects [Vemonia oligocephala)
1 3a Steendal soils [Bewsia biflora & Tephrosia purpurea)
b Arcadia soils [Setaria nigrirostris & Callilepis leptophylla)
14a Milkwood soils, midslopes (Alloteropsis semialata & Hyparrhenia hirta)
b Mayo soils, scarps and crests [Digitaria eriantha & Tetraselago wilmsii)
15a Footslopes/midslopes [Vernonia galpinii)
b Midslopes/crests [Berkheya seminivea)
16a Higher rock cover [Argyrolobium transvaalense)
b Larger rock size [Koeleria capensis)
2
3
4
5
8 Limosello maioris-Ranunculetum meyeri
6
5.2 Jamesbriuenio macranthae-Loudetietum simplicis eucleetosum linearis
5.1 Jamesbriuenio macranthae-Loudetietum simplicis combretetosum hereroense
1 Helichryso splendidi-Tristachyetum leucothricis
1
1 Andropogono eucomusaes-Fimbristyletum ferrugineae
8
2 Zantedeschio pentlandi-Aloetum castaneae
9
10
6.3 Fuireno puhescentis-Schoenetum nigricantis bulbostylietum liispidulae
11
12
6.2 Fuireno puhescentis-Schoenetum nigricantis pycnostachetosum reticulatae
6. 1 Fuireno pubescentis-Schoenetum nigricantis triraphietosum andropogonoidis
4.2 Elionuro mutici-Trachypogonetum spicati acacietosum tortilis
13
14
15
4. 1 Elionuro mutici-Trachypogonetum spicati bewsietosum biflorae
3.4 Brachiario serratae-Melhanietum randii setarietosum nigrirostis
3.1.2 Brachiario serratae-Melhanietum randii heUchrysetosum rugulosi. Alloteropsis semialata variant
3.1.1 Brachiario serratae-Melhanietum randii heUchrysetosum rugulosi. Digitaria eriantha variant
3.3 Brachiario serratae-Melhanietum randii gnidietosum capitatae
16
3.2.2 Brachiario serratae-Melhanietum randii argyrolobietosum transvaalense, Berkheya seminivea variant
3.2.1 Brachiario serratae-Melhanietum randii argyrolobietosum transvaalense, Koeleria capensis variant
tion by the soil particles. The gradient shown by the y-
axis for communities of wetlands is that of topographic
position (Figure 3). Here, the water systems of mountain
slopes with a faster run-off are placed at the top of the
scatter diagram. Perennial seepage systems of plateaus
are centred in the middle of the diagram, and communi-
ties of permanent, slow-flowing rivers are located at the
bottom.
The gradient along the first axis of Figure 2 is also an
indication of the species diversity in the grasslands, with
the species diversity at the left of the diagram being high-
er than that of the communities at the right. This phe-
nomenon can be attributed to the heterogeneous environ-
ment experienced by most Brachiario serratae-Melha-
nietum randii rocky grassland communities.
The gradients that have been identified are associat-
ed with each other and have a strong influence on the
vegetation. The three most dominant and conspicuous
taxa of each growth form (trees/shrubs/suffrutices,
forbs/sedges and grasses) are given for each of the eight
major vegetation types depicted in the scatter diagram
(Table 5).
FIGURE 3. — Relative positions of all the releves along the second and third axis of the ordination of the wetland vegetation of the
Sekhukhuneland Centre of Plant Endemism. Numbers correspond with those of syntaxa in Table lb and in the text.
Bothalia 32,2 (2002)
229
TABLE 5. — Three most dominant and conspicuous plant taxa of each growth form recorded for the major vegetation types depicted in
DECORANA scatter diagrams (Figures 2, 3)
DISCUSSION
Plant diversity and geographic variations over south-
ern Africa are best explained by the geology (soils) and
climate, with topographic diversity explaining much of
the remaining variance (Holland 1978; Schulze &
McGee 1978). Much of the distribution of the syntaxa in
the study area can be attributed to fire, and the vegetation
dynamics and historic evolution of the flora of the region
(Siebert 1998), which is influenced by the above-men-
tioned three factors.
The TWINSPAN classification and its subsequent
refinement by Braun-Blanquet procedures resulted in the
delineation of 17 syntaxa. These plant communities can
be related to certain environmental factors, the gradients
of which are illustrated in the DECORANA scatter dia-
grams. The major gradient relates to soil moisture, which
in turn is a direct consequence of run-off and drainage in
the case of grasslands, permanent water bodies/sources
in the case of wetlands, and temperature (drought stress)
in both cases. Soil moisture availability is determined by
the soil type, which in turn was determined by the topo-
graphical position of the substrate during pedogenesis.
The gradients of the diagram are therefore a direct con-
sequence of topography and climate.
In the SCPE, soils of ultramafic origin have caused
the development of syntaxa adapted to the specific soil
conditions, with high concentrations of certain elements,
notably heavy metals. Such a plant community (syntax-
on)-soil association on ultramafic substrates has previ-
ously been identified for southern Africa, on the Great
Dyke of Zimbabwe (Werger et al. 1978) and the
Barberton Greenstone Belt in South Africa (Morrey et al.
1989). Because of the ultramafic nature of the norites of
the Roossenekal and Leolo Subcentres, many taxa of the
study area are uncommon or absent in other grassland
areas of southern Africa.
Among the 17 plant communities, 44 plant species/
infraspecific taxa were identified as of conservation sig-
nificance. fifteen were SCPE endemics and 19 SCPE
near-endemics. Of the 44, 17 were listed in the southern
African Red Data List for plants (Hilton-Taylor 1996).
This number compares well with serpentine sites else-
where in the world, namely 1 8 rare plants and endemics in
central Queensland, Australia (Batianoff et al. 1995), 20
endemics on the Great Dyke, Zimbabwe (Wild 1965) and
22 endemics in southern Mpumalanga, South Africa
(Balkwill et al. 1995).
Species richness of Sekhukhuneland grassland com-
munities is high when compared with the grassland
species richness of other parts of South Africa.
Sekhukhuneland grasslands have a species richness of 20
to 51 species per 100 m^ with a mean of 38 species
(Table 3). This mean is higher than that recorded for the
grasslands of the northeastern Drakensberg (Mpuma-
langa) and southern Drakensberg (Eastern Cape), but is
lower than that of the Highveld (Gauteng/Mpumalanga)
(Hoare & Bredenkamp 2001). Floristically the Grass-
lands of the study area are related to those described by
Bloem (1988), Deall et al. (1989), Matthews et al.
(1992a), Burgoyne (1995) and Smit et al. (1997).
Throughout the SCPE, the remaining natural popula-
tions of plants and animals are under intense pressure
from exploitative land uses. There are certain areas with
specific syntaxa that should be considered as a priority
for conservation purposes. Some plant endemics of the
southern region of the SCPE are restricted to specific
syntaxa and these habitats therefore require urgent atten-
tion for conservation as a result of the rapid expanding
230
Bothalia 32,2 (2002)
mining industry, a common threat to southern Africa’s
rich plant diversity (Dold & Johnson 1997). Biodiversity
increases ecosystem stability by promoting diversity
among species in their responses to environmental fluc-
tuations (Naeem & Li 1997; Grime 1998; Ives et al.
1999) and hence, is imperative for successful conserva-
tion initiatives. The establishment of nature reserves or
protected areas is one solution to this problem of biodi-
versity loss; this is best achieved through land use strati-
fication derived from a holistic overview. Such a strate-
gic environmental assessment, which includes an ade-
quate database of natural features and other land uses
(Bedward et al. 1992), should be a priority for conserva-
tion agencies in the region.
The priority vegetation types for conservation
(hotspots) are associations 2, 3 and 4, of which subasso-
ciation 3.1, the Brachiario serratae-Mellianietum randii
helichrysetosum nigulosi rocky grassland on the scarps
and crests of norite hills, is probably the most important
and threatened. It should also be noted that the wetlands
are sensitive systems that are easily disturbed and should
be protected (Doust & Doust 1995; Van Wyk et ah 2000).
Important wetlands to conserve are the seepage areas,
namely the Limosello maioris-Rammculetum meyeri
Association and the Fuireno pubescentis-Schoenetum
nigricantis bulbostylietosum hispidulae Subassociation.
Alien species primarily invade areas of high indige-
nous species richness (Lonsdale 1999; Stohlgren et al.
1999), such as the grasslands of southern Africa. Acacia
dealbata, an alien tree from Australia, has invaded large
areas of the Roossenekal Subcentre and especially the
adjacent Steenkampsberg. Removing these trees is total-
ly dependent on the owner of the land, with no large-
scale projects being implemented. It is an ironic situa-
tion, with people 50 km to the north extensively chop-
ping down indigenous bushveld trees for firewood.
This description and classification of the syntaxa of
the study area is a contribution towards the understand-
ing of the vegetation and flora in the southern parts of the
SCPE and the region as a whole. The information sup-
plied here should be applied in the management and con-
servation of the vegetation and habitats, especially the
rocky grasslands that are threatened by the mining indus-
try and inappropriate forms of land use, specifically
overgrazing by domestic stock. This paper provides a
basis for proper and sound assessment of the region’s
vegetation, as it includes aspects such as species rich-
ness, rarity and habitat preference. Detailed site-specific
studies, using this paper as a reference, remain a prereq-
uisite before suggestions concerning mining or conserva-
tion can be made. The region boasts a rich natural and
human history (Pollock et al. 1963), and representative
ecologically viable portions of the grasslands of the
region, as well as the wetlands, should be protected and
conserved for future generations.
ACKNOWLEDGEMENTS
Our thanks to Martie Dednam, H.G.W.J. Schweikerdt
Herbarium (PRU), University of Pretoria, for the pro-
cessing of plant specimens, and the Curator and staff of
the National Herbarium (PRE) in Pretoria, for assistance
with plant identification. The National Research
Foundation, University of Pretoria, Edward Mellon
Foundation and Department of Environmental Affairs
and Tourism financially supported this project. The
South African Weather Bureau provided the rainfall and
temperature data.
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Bothalia 32,2; 233-249 (2002)
Miscellaneous notes
POACEAE
CHROMOSOME STUDIES ON AFRICAN PLANTS. 1 7. THE SUBFAMILIES ARUNDINOIDEAE AND DANTHONIOIDEAE
Cytogenetic studies in our laboratory focused on the
former grass subfamily Arundinoideae and we have pub-
lished chromosome numbers of 422 specimens, repre-
senting 12 genera and 54 species (Du Plessis & Spies
1988; Spies & Du Plessis 1988; Spies et al. 1990, 1992;
Du Plessis & Spies 1992; Spies et al. 1994; Visser &
Spies 1994b, d, e; Klopper et al. 1998; Spies & Roodt
2001). This subfamily was recently subdivided into two
separate subfamilies, the Arundinoideae and Dantho-
nioideae by the Grass Phylogeny Working Group
(GPWG 2001): the Arundinoideae, represented by
Styppeiochloa De Winter, has basic chromosome numbers
of 6, 9 and 12 and the Danthonioideae, represented by
Chaetobromus Nees, Karroochloa Conert & Tiirpe, Mefvc-
muellera Conert, Pentameris P.Beauv., Pentaschistis Stapf,
Pseudopentameris Conert, Schismus P.Beauv. and Tribo-
liiim Desv., has x = 6, 7 and 9.
This is a chromosome number report for the two sub-
families.
MATERIALS AND METHODS
Cytogenetic material of identical plants of a popula-
tion was collected and fixed in the field. Voucher speci-
mens, listed in Table 1, are housed in the Geo Potts
Herbarium, Department of Botany and Genetics, Uni-
versity of the Free State, Bloemfontein (BLFU) or in the
National Herbarium, Pretoria (PRB).
Anthers were squashed in aceto-carmine and meiotical-
ly analysed — at least 20 cells per meiotic stage were studied
(Spies et al. 1996). Only gametic chromosome numbers are
presented to conform to previous papers on chromosome
numbers in this journal (Spies & Du Plessis 1986).
RESULTS AND DISCUSSION
One hundred and nineteen plants, representing 38
species and 9 genera, were studied (Table 1).
Arundinoideae
Three specimens of the monospecific genus Styp-
peiochloa, S. gynoglossa, were studied. Two specimens
had n = 2x = 12, and one octaploid specimen (n = 4x =
24) was found (Figure lA). These specimens represent
the first known chromosome number reports for this
genus and six is the original basic chromosome number.
Twelve is probably a secondarily derived basic chromo-
some number.
FIGURE 1. — Meiotic chromosomes in Styppeiochloa and Karroochloa. A, Styppeiochloa gynoglossa, Saayman 79, 2n = 8x = 48, with chromo-
somal laggards; B, Karroochloa curva. Spies 4518, 2n = 2x = 12, diakinesis with 6n C-E, K. purpurea: C, D, Spies 2473, 2n = 4x = 24+2-5B;
C, diakinesis with 3iv6nand 5B (two B chromosomes paired); D, metaphase I with B chromosomes not on metaphase plate; E, Spies 2477,
2n = 2x = 12, metaphase I. F, G, K. schismoides: F, Spies 3371, 2n = 2x = 12, metaphase I; G, Spies 3382, 2n = 2x = 12, cell fusion resulting
in 12 bivalents and two micronuclei during diakinesis. H, Karroochloa species. Spies 5192, 2n = 2x = 12, diakinesis with 6n. Scale bar: B, 5 pm;
B, C, E, H, 8 pm; D, F, G, 15 pm.
234
Bothalia 32,2 (2002)
TABLE 1. — Gametic chromosome numbers (n) of representatives of subfamilies Arundinoideae and Danthonioideae (Poaceae) in southern Africa
with their voucher specimen numbers and specific localities. Species are listed alphabetically and localities presented according to
Edwards & Leistner (1971)
Taxon n Voucher Locality
Arundinoideae
MPUMALANGA. — 2530 (Lydenburg): in the Steenkampsberge, 6 km
from Goede Hoop to Roossenekal, (-AA).
SWAZILAND. — 2631 (Mbabane): Moimba beacon, 16 km from
Mbabane to Oshoek, (-AD).
MPUMALANGA. — 2430 (Pilgrim’s Rest): 3 km from Graskop to
Bosbokrand, (-DD).
NORTHERN CAPE. — 2917 (Springbok): 78 km from Steinkopf to Port
Nolloth, (-BA).
WESTERN CAPE. — 3420 (Bredasdorp): 25 km from Swellendam to
Ashton, (-AB).
NORTHERN CAPE. — 2918 (Gamoep): 26 km southeast from Spring-
bok to Garies, (-CA).
WESTERN CAPE. — 3319 (Worcester): 69 km from Montagu to Touws-
river, (-DB).
WESTERN CAPE. — 3320 (Montagu): 61 km from Montagu to Touws-
river, (-CD).
EASTERN CAPE. — 3126 (Queenstown): Penhoek Pass, (-BC).
NORTHERN CAPE.— 2917 (Springbok): 34 km from Port Nolloth to
Steinkopf. (-CA),
NORTHERN CAPE.— 2917 (Springbok): 25 km east of Port Nolloth.
(-AC).
NORTHERN CAPE.— 2917 (Springbok): 36 km from Port Nolloth to
Kleinsee, (-AC).
NORTHERN CAPE. — 2917 (Springbok): 13 km from Springbok on
road to Hondeklipbaai, (-DB).
NORTHERN CAPE. — 2918 (Gamoep): 26 km southeast from
Springbok to Garies, (-CA).
NORTHERN CAPE.— 3017 (Hondeklipbaai): 42 km west of Garies to
Groenrivier, (-DC).
WESTERN CAPE.— 31 18 (Vanrhymsdorp): 15 km north of
Vanrhynsdorp, (-AD).
NORTHERN CAPE. — 3119 (Calvinia): 35 km from Vanrhynsdorp to
Nieuwoudtville, (-AC).
EASTERN CAPE. — 3324 (Steytlerville): 25 km from Patensie to
Willowmore, (-DA).
EASTERN CAPE. — 3424 (Humansdoip): 16 km from Humansdoip to
Cape St Frances, (-BB).
WESTERN CAPE. — 3219 (Wuppertal): 3 km from Algeria to Citrusdal,
(-AC).
WESTERN CAPE. — 3419 (Caledon): 21 km from Franschoek to
Villiersdorp, (-AA).
WESTERN CAPE. — 3420 (Bredasdorp): 8 km south from Ouplaas to
De Hoop Nature Reserve, (-AD).
EASTERN CAPE. — 3027 (Lady Grey): 43 km from Barkley East to
Lady Grey, (-CA),
EASTERN CAPE. — 3028 (Matatiele): 12 km from Rhodes to Naude’s
Nek, (-CC).
EASTERN CAPE. — 3028 (Matatiele): 22 km from Rhodes on road to
Barkley East, (-CC).
WESTERN CAPE. — 3319 (Worcester): Du Toitskloof next to tunnel, (-AC).
EASTERN CAPE. — 3027 (Lady Grey): 37 km from Rhodes via
Luncheon's Nek, (-DD).
EASTERN CAPE.— 3027 (Lady Grey): 50 km from Rhodes via
Luncheon’s Nek, (-DD),
EASTERN CAPE, — 3027 (Lady Grey): 65 km from Barkley East to
Lady Grey via Joubert’s Pass, (-CA).
EASTERN CAPE — 3028 (Matatiele): 16 km from Rhodes via Naude’s
Nek, (-CC).
WESTERN CAPE— 3219 (Wuppertal): top of Uitkyk Pass, (-AC).
NORTHERN CAPE. — 31 19 (Calvinia): 41 km from Vanrhynsdorp on
Vanrhyns Pass, (-AC).
WESTERN CAPE.— 3218 (Clanwilliam): Uitkyk Pass, (-AC).
WESTERN CAPE.— 3219 (Wuppertal): 22 km from Clanwilliam to
Calvinia, (-AA),
Localities unknown.
NORTHERN CAPE.— 31 18 (Vanrhynsdorp): Gitberg Pass, (-DC).
NORTHERN CAPE. — 3119 (Calvinia): 41 km from Vanrhynsdorp on
Vanrhyns Pass, (-AC).
Bothalia 32.2 (2002)
235
TABLE 1. Gametic chromosome numbers (n) of representatives of subfamilies Arundinoideae and Danthonioideae (Poaceae) in southern Africa
with their voucher specimen numbers and specific localities. Species are listed alphabetically and localities presented according to
Edwards & Leistner (1971) (cont.)
Taxon n Voucher Locality
WESTERN CAPE. — 3218 (Clanwilliam): Uitkyk Pass, (-AC).
WESTERN CAPE. — 3219 (Wuppertal): 2 km from Houdenbek to
Katbakkies Pass, (-DC).
WESTERN CAPE. — 3319 (Worcester); 4 km from Franschoek turn-off
on Villiersdorp-Grabouw road, (-CC).
WESTERN CAPE. — 3419 (Caledon); Swartberg Pass, (-AA).
WESTERN CAPE. — 3419 (Caledon): 18 km from Botriver to Villiers-
dorp via Vanderstel Pass, (-AA).
Localities unknown.
WESTERN CAPE. — 3220 (Sutherland): 15 km from Sutherland to
Matjiesfontein, (-BC).
WESTERN CAPE. — 3322 (Oudtshoorn): Montagu Pass, (-CD).
WESTERN CAPE. — 3218 (Clanwilliam): 40 km from Clanwilliam to
Lamberts Bay, (-AB).
WESTERN CAPE. — 3218 (Clanwilliam): 32 km from Clanwilliam to
Nieuwoudtville at Klawer turn-off, (-BB).
WESTERN CAPE. — 3318 (Cape Town): Table Mountain, (-CD).
NORTHERN CAPE. — 3018 (Kamieskroon): 5 km from Kamieskroon
to Leliehoek. (-AC).
FREE STATE. — 2828 (Bethlehem): Bethlehem. Golden Gate National
Park. (-DA)
WESTERN CAPE. — 3218 (Clanwilliam): 32 km from Clanwilliam to
Nieuwoudtville. at Klawer turn-off, (-BB).
WESTERN CAPE. — 3218 (Clanwilliam): 14 km from Clanwilliam to
Nieuwoudtville. (-BB).
WESTERN CAPE. — 3118 (Vanrhynsdorp): On top of Gifberg Pass, (-DC).
WESTERN CAPE.— 3218 (Clanwilliam): Uitkyk Pass, (-AC).
WESTERN CAPE. — 3319 (Worcester): 21 km from Franschoek to
Villiersdorp, (-CC).
WESTERN CAPE. — 3322 (Oudtshoorn): Swartherg Pass, (-AC).
WESTERN CAPE. — 3419 (Caledon): Galgeberg, (-BA).
WESTERN CAPE. — 3219 (Wuppertal): on top of Nieuwoudfs Pass, (-AC).
EASTERN CAPE.— 3028 (Matatiele): Barkley East. Naude's Nek, (-CC).
WESTERN CAPE. — 3418 (Simonstown): 7 km from Betty's Bay to
Onrus. (-BD).
WESTERN CAPE. — 3218 (Clanwilliam): 13 km from bridge over
Olifants River, (-DB).
NORTHERN CAPE. — 2917 (Springbok): 24 km from Springbok to
Hondeklipbaai. (-DB).
WESTERN CAPE.— 31 19 (Calvinia): on Vanrhyns Pass, (-AC).
WESTERN CAPE. — 3420 (Bredasdorp): 1 km north of De Hoop
Nature Reserve, (-AD).
WESTERN CAPE. — 3418 (Simonstown): 7 km from Betty’s Bay to
Onrus, (-BD).
WESTERN CAPE. — 3319 (Worcester); FM tower on Matroosberg. (-BC).
WESTERN CAPE. — 3219 (Wuppertal): 5 km from Algeria to Citrus-
dal. Nieuwoudt's Pass. (-AC).
WESTERN CAPE. — 3219 (Wuppertal): 5 km from Algeria to Citrus-
dal, Nieuwoudt's Pass, (-AC).
WESTERN CAPE.— 3219 (Wuppertal): Uitkyk Pass, (-AC).
WESTERN CAPE.— 3219 (Wuppertal): Uitkyk Pass, (-AC).
WESTERN CAPE. — 3219 (Wuppertal): on top of Nieuwoudt’s Pass, (-AC).
WESTERN CAPE.— 3219 (Wuppertal): Uitkyk Pass, (-AC).
NORTHERN CAPE. — 3017 (Hondeklipbaai): 7 km from Kamieskroon
to Leliefontein. (-BB).
NORTHERN CAPE. — 3017 (Hondeklipbaai): 7 km from Kamieskroon
to Leliefontein, (-BB).
NORTHERN CAPE. — 3119 (Calvinia): 7 km from Nieuwoudtville to
Clanwilliam, (-AC).
NORTHERN CAPE. — 3119 (Calvinia): on top of Vanrhyns Pass, (-AC).
WESTERN CAPE. — 3320 (Montagu): 24 km from Montagu to Touws
River. Burger’s Pass, (-AC).
WESTERN CAPE. — 3420 (Bredasdorp): 5 km from Ouplaas to Malgas,
(-AD).
WESTERN CAPE. — 3219 (Wuppertal): on top of Nieuwoudt’s Pass, (-AC).
WESTERN CAPE. — 3322 (Oudtshoorn): Robinson’s Pass, (-CC).
WESTERN CAPE.— 3218 (Clanwilliam): 21 km from Clanwilliam to
Nieuwoudtville, (-BB).
WESTERN CAPE.— 3318 (Cape Town); Table Mountain, (-AB).
236
Bothalia 32,2 (2002)
TABLE 1. — Gametic chromosome numbers (n) of representatives of subfamilies Arundinoideae and Danthonioideae (Poaceae) in southern Africa
with their voucher specimen numbers and specific localities. Species are listed alphabetically and localities presented according to
Edwards & Leistner (1971) (cont.)
Taxon n Voucher Locality
GAUTENG. — 2528 (Pretoria): Pretoria, (-CA).
NORTHERN CAPE. — 2917 (Springbok): 2 km from Springbok to
Pofadder, (-DB).
NORTHERN CAPE. — 2917 (Springbok): 13 km from Springbok to
Hondeklipbaai, (-DB).
FREE STATE. — 2925 (Jagersfontein): Petrusburg, (-AB).
NORTHERN CAPE. — 3017 (Hondeklipbaai): 24 km from Soebatsfon-
tein to Kamieskroon, (-BA).
NORTHERN CAPE. — 3018 (Kamiesberg); 28 km from Leliefontein to
Garies, (-AB).
WESTERN CAPE — 3119 (Calvinia): 55 km from Nieuwoudtville to
Clanwilliam, (-CC).
WESTERN CAPE. — 3219 (Wuppertal): 44 km from Clanwilliam to
Calvinia, (-AA).
WESTERN CAPE. — 3318 (Vanrhynsdorp): Vanrhynsdorp, (-DA).
WESTERN CAPE. — 3320 (Montagu): 24 km from Montagu to Touws
River, (-CC).
WESTERN CAPE — 3320 (Montagu): 38 km from Montagu to Touws
River, (-CC).
NORTHERN CAPE. — 3017 (Hondeklipbaai): 18 km from Kamieskroon
to Leliefontein, (-BA).
NORTHERN CAPE. — 3018 (Kamiesberg): 3 km from Leliefontein to
Garies in Groenkloof, (-AB).
WESTERN CAPE. — 3318 (Cape Town): Langebaan, (-DC).
WESTERN CAPE. — 3421 (Riversdale): 2 km south of Vermaaklikheid
on road to Puntjie, (-AC).
WESTERN CAPE. — 3420 (Bredasdorp): 1 km from Waenhuiskrans to
Bredasdorp, (-CA).
Locality unknown.
NORTHERN CAPE. — 3121 (Fraserburg): 55 km from Loxton to Fraser-
burg, (-DB).
NORTHERN CAPE. — 3220 (Sutherland): 2 km from Sutherland to
Calvinia, (-BC),
WESTERN CAPE. — 3319 (Worcester): 5 km from Gouda to Porterville,
(-AC).
WESTERN CAPE. — 3420 (Bredasdorp): 1 km north of De Hoop Nature
Reserve, (-CA).
EASTERN CAPE. — 3424 (Humansdorp): 30 km from Humansdorp to
Knysna, (-AA).
Locality unknown.
WESTERN CAPE. — 3218 (Clanwilliam): 7 km from Clanwilliam in
Pakhuis Pass, (-BB).
Locality unknown.
The two tetraploid specimens, Spies 1485, 2642, with
x-values of 0.829 and 1 respectively, agree with the 2:2
model of Kimber & Alonso (1981) (Table 2). The high
x-value of Spies 1485 indicates segmental alloploidy,
with a tendency towards alloploidy. The other specimen.
Spies 2642, has a value of 1, suggesting an alloploid ori-
gin. In both specimens mostly bivalents were observed.
In Spies 1485, multivalent formation was rarely
observed, but in Spies 2642 only rod and ring bivalents
TABLE 2. — Genomic relationships in tetraploid Slyppeiochloa speci-
mens analysed according to Kimber & Alonso (1981). Values
indicated represent the sums of squares calculated for the four
possible tetraploid models, x-values are indicated in parenthe-
ses. Model best suited for each specimen is indicated in bold
were observed, evidence of alloploidy. The fonnation of
mainly bivalents indicates an alloploid or segmental
alloploid origin, thus suggesting a hybrid origin for this
monotypic genus.
Danthonoideae
A single specimen of Chaetohromiis involucratus
subsp. dregeamis was investigated with n = x = 6. This
confinns a basic chromosome number of six for this
genus, endemic to the semi-arid and arid western Cape
region of South Africa and the extreme southwestern cor-
ner of Namibia (Du Plessis & Spies 1988; Spies & Du
Plessis 1988; Spies et al. 1990; Verboom & Linder
1997). Studies of the genus Chaetohromus by Spies et al.
(1990) suggested a polyploid complex, ranging from
diploid to duodecaploid. Polyploidy occurs as segmental
alloploidy or occasionally as alloploidy (Spies et al.
1990). Spies et al. (1990) reported on extensive morpho-
logical, cytogenetical and anatomical variation in
Chaetohromus, indicating hybridization and polyploidy.
Bothalia 32,2 (2002)
237
FIGURE 2. — Meiotic chromosomes in Merxmiiellera. A, M. cincta. Spies 3504, 2n = 6x = 36, diakinesis with 1 8n, three bivalents are not in focus
on this photograph. B, C, M. decora'. B, Spies 4407. 2n = 8x = 48, metaphase I; C, Spies 4458, 2n = 8x = 48, metaphase I. D, M. drak-
ensbergensis. Spies 4687, 2n = 6x = 36, early anaphase I 18-18 segregation; E, M. htpuUna. Spies 4601, 2n = 8x = 48, diakinesis with 24n;
F, M. macowanii. Spies 4727, 2n = 8x = 48, metaphase I; G, M. ritfa. Spies 4402, 2n = 8x = 48, diakineses. H-K, M. stricta. Spies 3140,
2n = 9x = 54: H, J, metaphase I with various univalents; 1, K, anaphase 1 with numerous laggards. Scale bar: A, E, F, G, J, 5 pm; B, C, D,
H, I. K, 9 pm.
Diploid chromosome numbers (n = x = 6) were
observed for all of the Karroochloa specimens studied,
(Figure IB-H) namely Karroochloa cw\’a, K. schis-
moides and K. tenella. One K. cun’a specimen was stud-
ied cytogenetically and a low percentage of univalents
{Spies 4518, Figure IB) was observed. Two to five B
chromosomes were observed in a single K. purpurea
specimen (Spies 2475, Figure 1C, D) and this was also
the only tetraploid specimen found. Further K. purpurea
specimens exhibited normal meiosis (Figure IE, F). In
K. chismoides meiosis was normal (Figure IF, H), but
cell fusion was observed in some specimens {Spies 3382,
Figure IG) which could lead to polyploidy. The basic
chromosome number of six was confirmed for this genus
(Du Plessis & Spies 1988; Spies & Du Plessis 1988).
Chromosome numbers are reported for eight of the 1 8
southern African species of the genus Merxmiiellera. The
genus has a basic chromosome number of six (Du Plessis
& Spies 1988; Spies & Du Plessis 1988). A single M.
cincta specimen was investigated and found to be hexa-
ploid (Figure 2A). All three M. decora specimens were
octaploid (n = 4x = 24) (Figure 2B, C). Meiosis was
abnormal with a high percentage of laggards during
anaphase I (Figure 2C) and micro-nuclei during
telophase I. In M disticha meiosis was very abnormal
with a high percentage of univalents during metaphase I,
anaphase I laggards and micro-nuclei. This is the first
report for M. drakensbergensis, both specimens being
hexaploid (n = 3x = 18) (Figure 2D). Only one M. lupuli-
na specimen was investigated for the first time with n =
4x = 24 (Figure 2E). All four M. macowanii specimens
were octaploid (Figure 2F). An octaploid chromosome
count for M. riifa is a first for this species (Figure 2G).
Several multivalents are evident in various cells of this
specimen. A large number of the specimens were M.
stricta. Mostly tetraploid and hexaploid numbers, or
deviations thereof, {Spies 3140, 2n = 6x = 51) (Figure
2H-K) were evident. Meiotic irregularities such as uni-
valents (Figure 2H, J), laggards (Figure 21, K) and
micro-nuclei as well as cytomixis were observed. This
abnormal meiotic behaviour in the genus and especially
M. stricta would suggest these specimens to be of hybrid
origin. Many M. stricta specimens were found to contain
a high number of chromosomal laggards, and it is a vari-
able perennial species.
238
Bothalia 32,2 (2002)
FIGURE 3. — Meiotic chromosomes in Pentaschistis and Pseudopentameris. A, Pentaschistis pallida. Spies 6208, 2n = 4x = 28, anaphase I with
14-14 segregation; B, P. rigidissima. Spies 6243, 2n = 2x = 14, diakinesis with 7n; C, P. tomentella. Spies 6344, 2n = 2x = 14+0-2B, diaki-
nesis with 1\\, D, E, P. tortuosa. Spies 6214, 2n = 8x = 56, early anaphase I with chromosomes starting to segregate. F, G, Pseudopentameris
macrantha. Spies 3431, 2n = 2x = 12, diakinesis with 6n in each cell. Scale bar: B, F, G, 6 pm; A, C-E, 10 pm.
Chippindall (1955) states that ‘There is considerable
variation in the plants referred to as Danthonia stricta
(M. stricta), and it is possible that they comprise more
than one variety’. Ellis (1980) divided M. stricta into
four ‘forms’: the ‘typical form’ (M. stricta), the
‘Drakensberg form’ (M. stricta), the ‘Cathedral Peak
form’ (M guillarmodiae Conert) and the ‘alpine form’
(M guillarmodiae). Each of these forms exhibits distinct
epidermal structure and leaf anatomy. In the Drakens-
berg region, two Pentaschistis species displayed remark-
able anatomical similarities with M. stricta. These are
Pentaschistis tysonii Stapf and an unnamed Pentaschistis
species. Anatomically they seem to show greater affinity
with the M. stricta group than with Pentaschistis (Ellis
1980). This raises the issue as to whether hybridization
occurred, or is still occurring between the different M.
stricta forms, or between the Pentaschistis species and
M. stricta, and whether this could clarity the possible
hybrid nature of M. stricta.
One diploid Pentameris thuarii specimen was studied
(n = X = 7). This confirms a basic chromosome number
of seven for the genus (Spies & Roodt 2001).
In the genus Pentaschistis, 14 of the 57 indigenous south
African species (Gibbs Russell et al. 1990) were studied.
This is the first report for numbers in P. acinosa (n = x = 7),
P. galpinii (n = x = 7) and P. veneta (n = 2x = 14). Diploidy
was observed in 21 of the 39 specimens (Figure 3B, C).
Polyploidy is frequently encountered in Pentaschistis. In the
study by Du Plessis & Spies ( 1 992), 59% of the species
investigated were polyploid, or had different polyploid lev-
els. In this study tetraploidy was observed in P. airoides, P.
pallida (Spies 5737, 5917, 6208) (Figure 3A), P. tomentella
(Spies 530], 5738), P. veneta and P. viscudula. Hexaploid
(P. rupestris), octaploid (P. densifolia, P. lima, P. tortuosa)
(Figure 3D, E), decaploid (P. rupestris, P. viscidula) and up
to 14-ploid levels (P. rupestris) were also observed. Higher
ploidy levels in some cells of specimens can be caused by
cell fusion (Spies & Van Wyk 1995). It is not an uncommon
phenomenon in this genus.
Polyploidy is frequent in Pentaschistis. Klopper et al.
(1998) reported on the existence of young polyploid
complexes in 17 species. Twelve species were found to
be old polyploid complexes, but were not adequately
studied and the age of the complexes should be verified.
Furthermore, Klopper et al. (1998) suggest that the genus
Pentaschistis is a young polyploid hybrid complex.
B chromosomes were mostly encountered at the
diploid level (33% of the diploid specimens having 1 to
4 B chromosomes present in some cells). In Penta-
schistis, the number of B chromosomes per cell varied
within the same species and even within the same spec-
imen. This is evident in P. curvifolia (Spies 4456, 6169,
6236, 6315) where the number of B chromosomes var-
ied from 0^ in the different specimens. B chromosomes
were also encountered in the tetraploid P. viscidula
specimen (0-2B) but were absent from higher polyploid
levels.
Only four genera, namely Merxmuellera, Pentameris,
Pentaschistis and Prionanthium Desv., share the basic
chromosome number of seven. Davidse et al. (1986) sug-
gested that X = 7 is a primitive number, as in the genus
Pentaschistis, and that x = 13, which also occurs in the
genus, was secondarily derived through an aneuploid
reduction from x = 14. In this study no specimens with a
basic chromosome number of 1 3 were observed.
Bothalia 32,2 (2002)
239
FIGURE 4. — Meiotic chromosomes in Schismiis and Thbolium. A, 5. barhatus. Davidse 34033, 2n = 2x = 12, early metaphase I. B-D, S. scaber-
rimus: B, C, Spies 4660, 2n = 2x = 12, metaphase I; D, Spies 4661, 2n = 2x = 12, diakinesis with 6n E, T. obtusifoliwn, Davidse 34049,
2n = 6x = 36, diakinesis with 18n; F, T. piisillum, Davidse 34022, 2n = 2x = 12, diakinesis with 6n. Scale bar: A, F, 5 pm; B-E, 10 pm.
One single specimen in the genus Pseudopentameris
is reported here for the first time, P. macrantha with 2n
= 2x = 12 (Figure 3F, G), and a basic chromosome num-
ber of six for the genus.
Three of the four endemic Schismus species were
investigated cytogenetically. With the exception of a sin-
gle S. inermis specimen (Davidse 33804), all the speci-
mens were diploid, 2n = 2x = 12, (Figure 4A-D) and con-
firms the basic chromosome number of six for the genus
(Du Plessis & Spies 1988; Spies & Du Plessis 1988).
Du Plessis & Spies (1988) reported a polyploid com-
plex in Schismus barhatus, based on the meiotic behav-
iour of one diploid, three tetraploid and one hexaploid
specimen. However, all 14 5. barhatus specimens of this
study were diploid.
The genus Tribolium was thoroughly investigated
cytogenetically over the last couple of years (Spies et al.
1992; Visser & Spies 1994a-e). It was concluded that six
is the basic chromosome number for the genus. Six addi-
tional specimens representing five species were investi-
gated. Diploidy 2n = 2x = 12 was encountered in the
species T. hispidum and T. pusillum (Figure 4F).
Tribolium acutiflorum, T. hispidum and T. uniolae were
found to be tetraploid and a single hexaploid specimen,
T. obtusifolium was found (Figure 4E). We and Visser &
Spies (1994b) found diploid and tetraploid specimens in
T. hispidum. In Tribolium polyploidy is common. This
led Visser & Spies (1994b, d, e) to conclude that the
genus Tribolium is a polyploid complex. Visser & Spies
(1994d) reported on a T uniolae hybrid swarm in which
polyploidy is frequent and various meiotic abnormalities
occur. 0-2 B chromosomes were observed, but only in
the single T. uniolae specimen.
The basic chromosome numbers of six and seven for
the Danthonioideae and six for Styppeiochloa (Arundi-
noideae) is confirmed.
ACKNOWLEDGEMENTS
The University of the Free State and the Foundation
for Research and Development are thanked for financial
assistance. Gerrit Davidse (Missouri Botanical Garden,
USA) and Peter Linder (Institut fur Systematische
Botanik, Zurich) are thanked for providing some of the
meiotic material used.
REFERENCES
CHIPPINDALL, L.K.A. 1955. A guide to the identification of grasses
in South Africa. In C. Meredith, The grasses and pastures of
South Africa: 1-527. Central News Agency, Cape Town.
DAVIDSE, G.; HOSHINO, T. & SIMON, B.K. 1986. Chromosome
counts of Zimbabwean grasses and an analysis of polyploidy in
the grass flora of Zimbabwe. South African Journal of Botany
52:521-528.
DU PLESSIS, H. & SPIES, J.J. 1988. Chromosome studies on African
plants. 8. Bothalia 18: 119-122.
DU PLESSIS, H. & SPIES, J.J. 1992. Chromosome numbers in the genus
Pentaschistis (Poaceae, Danthonieae). Ta.xon 41: 709-720.
EDWARDS, D. & LEISTNER, O.A. 1971. A degree reference system
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der Botanischen Staatssammlung, Miinchen 10: 501-509.
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ELLIS, R.P. 1980. Leaf anatomy of the South African Danthonieae
(Poaceae). III. Merxmuellera stricta. Bothalia 13: 191-198.
GIBBS RUSSELL, G.E., WATSON, M„ KOEKEMOER, M., SMOOK,
L., BARKER, N.P., ANDERSON, H.M, & DALLWITZ, M.J.
1 990. Grasses of southern Africa. Memoirs of the Botanical
Survey of South Africa No. 58.
GRASS PHYEOGENY WORKING GROUP (GPWG). 2001. Phylo-
geny and subfamilial classification of the grasses (Poaceae).
Annals of the Missouri Botanical Garden 88: 373—457.
KIMBER. G. & ALONSO, L.C. 1981. The analysis of meiosis in hybrids.
111. Tetraploid hybrids. Canadian Journal of Genetics and
Cyro/og>' 23: 235-254.
KLOPPER, K.C., SPIES, J.J. & VISSER, B. 1998. Cytogenetic studies
in the genus Pentaschistis (Poaceae: Arundinoideae). Bothalia
28: 231-238.
SPIES, J.J., DAVIDSE, G. & DU PLESSIS, H. 1992. Cytogenetic stud-
ies in the genus Tribolium (Poaceae: Arundineae). American
Journal of Botany 79: 689-700.
SPIES, J.J. & DU PLESSIS, H. 1986. Chromosome studies on African
plants. I. Bothalia 16: 87, 88.
SPIES, J.J. & DU PLESSIS, H. 1988. Chromosome studies on African
plants. 6. Bothalia 18: 111-123.
SPIES, J.J., DU PLESSIS, H., BARKER, N.P. & VAN WYK, S.M.C.
1990. Cytogenetic studies in the genus Chaetobromus
(Poaceae: Arundineae). Genome 33: 646-658.
SPIES, J.J., LINDER, H.P., LABUSCHAGNE, I.F. & DU PLESSIS, H.
1994. Cytogenetic evidence for the species delimitation of
Pentaschistis airoides and P. patula (Poaceae: Arundineae).
Proceedings from the XI lit h Plenary Meeting of AETFAT.
Zomha, Malawi 1: 373-383.
SPIES, J.J. & ROODT, R. 2001. The basic chromosome number of the
genus Pentameris (Poaceae: Amndinoideae). Bothalia 31:145, 146.
SPIES, J.J., SPIES, S.K., VAN WYK, S.M.C., MALAN, A.E. & LIEBEN-
BERG, E. J.L. 1 996. Cytogenetic studies of the subfamily Pooideae
( Poaceae) in South Africa. 1 . The tribe Aveneae, subtribe Aveninae.
Bothalia 26: 53-61.
SPIES, J.J. & VAN WYK, S.M.C. 1995. Cell fusion: a possible mech-
anism for the origin of polyploidy. South African Journal of
Botany 61 : 60-65.
VERBOOM, G.A. & LINDER. H.P. 1998. A re-evaluation of species limits
in Chaetobromus (Danthonieae: Poaceae). Nordic Journal of Botany
18:57-77.
VISSER, N.C, & SPIES, J.J. 1994a. Cytogenetic studies in the genus
Tribolium (Poaceae: Danthonieae). II. A report on embryo sac
development, with special reference to the occurrence of apomixis
in diploid specimens. South African Journal of Botany 60: 22-26.
VISSER, N.C. & SPIES, J.J. 1994b. Cytogenetic studies in the genus
Tribolium (Poaceae: Danthonieae). III. Section Tribolium.
South African Journal of Botany 60: 31-39.
VISSER, N.C. & SPIES, J.J. 1994c. Cytogenetic studies in the genus
Tribolium (Poaceae: Danthonieae). I. A taxonomical overview.
South African Journal of Botany 60: 127-13 1 .
VISSER, N.C. & SPIES, J.J. I994d. Cytogenetic studies in the genus
Tribolium (Poaceae: Danthonieae). IV. Section Uniolae. South
African Journal of Botany 60: 279-284.
VISSER, N.C, & SPIES, J.J. 1994e. Cytogenetic studies in the genus
Tribolium (Poaceae: Danthonieae). V, Section Acutiflorae, related
genera, and conclusions. South African Journal of Botany 60:
285-292.
R. ROODT*, J.J. SPIES*, A.F. MALAN*, F. HOLDER*
and S.M.C. VAN WYK*
* Department of Plant Sciences: Genetics (62), University of the Free
State, P.O. Box 339, 9300 Bloemfontein.
MS. received: 2002-02-27.
CHROMOSOME STUDIES ON AFRICAN PLANTS. 18. THE SUBFAMILY CHLORIDOIDEAE
The subfamily Chloridoideae comprises ± 150 genera
and 1 360 species and occurs mainly in arid regions (Hilu
& Alice 2001). The plants probably originated in Africa
(Hartley 1964), hence the great representation of the sub-
family in Africa and especially southern Africa, with +
5 1 genera and 235 species (Gibbs Russell et al. 1990). It
is currently divided into five tribes by the Grass
Phylogeny Working Group (GPWG 2001): Cynodonteae
Dumort., Eragrostideae Stapf, Leptureae Dumort.,
Orcuttieae Reeder and Pappophoreae Kunth. The genus
Centropodia Reich, and the species Merxmuellera
range! (Pilg.) Conert, previously included in the
Arundinoideae, are now included in the Chloridoideae.
They have not previously been included in any of the
recognized tribes (GPWG 2001).
The aim of this study is to investigate chromosome
numbers, meiotic chromosome behaviour and polyploid
levels of some southern African representatives of this
subfamily.
MATERIALS AND METHODS
Cytogenetic material of identical plants of a popula-
tion was collected and fixed in the field. Voucher speci-
mens, listed in Table 3, are housed in the Geo Potts
Herbarium, Department of Plant Sciences, University of
the Free State, Bloemfontein (BLEU), or in the National
Herbarium, Pretoria (PRE).
Anthers were squashed in aceto-carmine and meiotical-
ly analy.sed (Spies et al. 1 996) — at least 20 cells per meiot-
ic stage were studied. Only gametic chromosome numbers
are presented to conform to previous papers on chromo-
some numbers in this journal (Spies & Du Plessis 1986).
RESULTS AND DISCUSSION
Seventy-nine plants, representing 42 species and 19
genera, were studied (Table 3). They represent three of the
recognized tribes, namely Cynodonteae, Eragrostideae and
Pappophoreae, as well as the unplaced genus Centropodia.
Tribe Cynodonteae
In the genus Chloris Sw., a single C. virgata specimen
was investigated and found to be diploid (n = x = 10)
FIGURE 5. — Meiotic chromosome.s. A, Chloris virgata. Spies 6616,
2n = 2x = 20. diakine.ses with 20n; B, Cynodon dactylon. Spies
2549, 2n = 4x = 36, metaphase I. Scale bar: 5 pm.
Bothalia 32,2 (2002)
241
TABLE 1 . — Gametic chromosome numbers (n) of representatives of subfamily Chloridoideae (Poaceae) in southern Africa with their voucher
specimen numbers and specific localities. Species listed alphabetically and localities presented according to Edwards & Leistner (1971)
242
Bothalia 32,2 (2002)
TABLE 1. — Gametic chromo.some numbers (n) of representatives of subfamily Chloridoideae (Poaceae) in southern Africa with their voucher
specimen numbers and specific localities. Species listed alphabetically and localities presented according to Edwards & Leistner (1971)
(cont.)
(Figure 5A). This confirms the basic chromosome num-
ber of 10 for this genus (Darlington & Wylie 1955;
Pienaar 1955; Ornduff 1967, 1968, 1969; Federov 1969;
Moore 1970, 1971, 1972, 1973, 1974, 1977; Goldblatt
1981, 1983, 1985, 1988; Goldblatt & Johnson 1990,
1991, 1998, 2000). Polyploid levels do occur in the
genus and triploid and tetraploid numbers have been
observed in South African specimens by previous
authors (Moffet & Hurcombe 1949; De Wet 1954; Spies
& Du Plessis 1987). The genus is known to contain many
aneuploid deviations from the basic chromosome num-
ber of ten (Fish 2000), but none have been observed in
South African specimens before (Hunter 1934; Moffet &
Hurcombe 1949; De Wet 1954; Spies & Du Plessis 1987;
Spies & Jonker 1987; Strydom & Spies 1994).
Cynodon dactylon is an introduced species in tropical
and wann temperate areas throughout the world, but is
indigenous to southern Africa (Gibbs Russell & Spies
1988). Two specimens were investigated, both with
2n = 4x = 36 (Figure 5B). Tetraploidy is by far the most
numerous polyploid level present in the genus (Darlington
& Wylie 1955; Pienaar 1955; Ornduff 1968, 1969; Federov
1969; Moore 1970, 1972, 1973, 1974, 1977; Goldblatt
Bothalia 32,2 (2002)
243
FIGURE 6. — Meiotic chromosomes in Harpochloa falx. A, Spies 3986, 2n = 4x = 36, metaphase I. B, C, Spies 4695, 2n = 4x = 36: B, metaphase
I; C, desynapsis of bivalents during metaphase I. D. Spies 4729, 2n = 4x = 36, metaphase I; E, Spies 5113, 2n = 4x = 36, metaphase I; F,
Spies 6955, 2n = 4x = 36, diakineses with 18u. Scale bar: C, E, 8 pm; A, B, D. F, 10 pm.
1981, 1983, 1985; Goldblatt & Johnson 1990, 1994, 1996,
1998). De Silva & Snaydon (1995) concluded that the dif-
ferences in the polyploid level in C dactylon could be
related both to different climatic regions and different eco-
logical habitats. The populations growing in arid, dry and
intermediate regions were tetraploid and those from wetter
regions consisted entirely of diploid plants. De Silva &
Snaydon (1995) also related these polyploid levels to soil
acidity and alkalinity (tetraploid = pH > 6.5; diploid = pH
< 5.0). These findings might explain the predominance of
tetraploidy in South Africa with its dryer, more arid
regions. A basic chromosome number of x = 9 is confirmed
for this species and genus, although 10 has also been
reported in a few instances (Omduff 1968, 1969; Moore
1970, 1972, 1974, 1977; Goldblatt 1981, 1983, 1985;
Goldblatt & Johnson 1990, 1994, 1996, 1998).
Chromosome numbers for the genus Enteropogon
Nees have only been reported once and a basic chromo-
some number of 10 published for the genus (Darlington
& Wylie 1955). One E. macrostachyiis specimen investi-
gated in this study was found to be tetraploid (n = 2x =
20). This is the first report on a South African specimen
in the genus, the previous one being from India.
Twelve specimens representing the species Harpo-
chloa falx were studied. Nine of these were tetraploid
and three were hexaploid. All specimens had multiples of
nine (n = 2x = 18; n = 3x = 54) (Figure 6A-F). This rep-
resents a new basic chromosome number for this genus
and could imply that Harpochloa also has two basic
chromosome numbers (De Wet 1958; Spies & Du Plessis
1986; Spies et al, 1991; Strydom & Spies 1994).
Furthermore, no known reports of any diploid specimens
in the genus exist, which could indicate the existence of
an older polyploid complex.
The genus Tragus Haller is widespread throughout the
tropics, but mainly in Africa. It is especially common in
disturbed areas (Clayton & Renvoize 1986; Fish 2000).
One Tragus specimen was investigated and found to be
diploid with n = x = 10. Only diploid, as in this study, or
tetraploid chromosome numbers are known for this
genus, based on a basic chromosome number of 10
(Darlington & Wylie 1955; Omduff 1967, 1968, 1969;
Federov 1969; Moore 1970, 1972, 1973, 1974, 1977;
Goldblatt 1981, 1983, 1988; Goldblatt & Johnson 1990,
1994, 1998).
Tribe Eragrostideae
The genus Bewsia Gooss. is monotypic and a single
Bewsia biflora specimen was investigated. This speci-
men was tetraploid (n = 2x = 20) (Figure 7 A), which
confirms a basic chromosome number of ten, based on
previous reports by De Wet & Anderson ( 1956) of 2n =
3x = 30 and Davidse et al. (1986) of 2n = 45 from
Zimbabwe. Davidse et al. ( 1986) reported on very irreg-
ular meiosis in the particular specimen. Results present-
ed in this study are the third known report for this genus.
Cladoraphis Franch. comprises two species C. cyper-
oides and C. spinosa. De Winter (1955) included this
genus in Eragrostis Wolf, but later authors (Phillips
1982; Clayton & Renvoize 1986; Gibbs Russell et al.
1990; Watson & Dallwitz 1992) retained its separate
generic status. It has a very specific habitat and occurs in
sandy desert (C. spinosa) and coastal dunes (C. cyper-
oides) (Clayton & Renvoize 1986), mainly in the western
regions of Namibia and Northern and Western Cape
(Fish 2000). Five specimens were investigated represent-
ing both species. All four C. cyperoides specimens were
diploid, with C. spinosa being tetraploid (n = 2x = 20)
244
Bothalia 32,2 (2002)
FIGURE 7. — Meiotic chromosomes. A, Bewsia hiflora. Spies 1531, 2n = 4x = 40, diakineses with 20n, three chromosomes are not in focus on
this photograph. B, C, Cladoraphis cyperoides. Spies 4894, 2n = 2x = 20: B, metaphase I; C, anaphase I, 10-10 segregation. D, C. cyper-
oides. Spies 5356, 2n = 2x = 20, diakinesis; E, C. spinosa. Spies 4885, 2n = 4x = 40, diakineses with 20n; F, Dactyloctenium aegyptium.
Spies 2403, 2n = 4x = 40, metaphase 1. Scale bar: 5 pm.
(Figure 7B-E). This is, to the best of our knowledge, the
first reports for chromosome numbers in this genus.
One specimen of the widespread tropical weed
(Gould & Soderstrom 1974) Dactyloctenium aegyptium
was investigated. It was found to be tetraploid
(n = 2x = 20) (Figure 7F). Three basic chromosome
numbers are recorded for this genus, x = 9, 10, 12
(Darlington & Wylie 1955 (x = 10, 12); Pienaar 1955
(x = 9, 12); Ornduff 1967 (x = 10), 1968 (x = 12); Moore
1971 (x-9), 1972 (x = 9), 1977 (x= 12); Goldblatt 1981
(X = 9, 10, 12), 1983 (X = 9, 10, 12), 1985 (x - 12), 1988
(x = 9, 10); Goldblatt & Johnson 1990 (x = 9, 10, 12),
1991 (X = 10), 1994 (X = 12), 1998 (x = 10, 12). This is
one of the genera in the Chloridoideae (as is Sporoholus),
with the most variation in basic chromosome number.
The generic status of Diplachne P.Beauv. has long
been in doubt (McVaugh 1983; Peterson et al. 1997),
with some authors preferring to unite this genus
with the closely related genus Leptochloa P.Beauv.
(McNeill 1979; Phillips 1982). These two genera have
traditionally been kept distinct by Old World taxono-
mists where these genera are quite distinct, whereas
the position of the genera from the Americas are very
confused with intergrading taking place (Phillips
1982). For this study, the accepted name Leptochloa
will be used.
Eight Leptochloa fusca specimens were investigated
and all were diploid (n = x = 10) (Figure 8A-E), which
confirms the basic chromosome number of ten for this
genus (Darlington & Wylie 1955; Ornduff 1968; Federov
1969; Moore 1977; Goldblatt & Johnson 1990, 1991,
1994, 1998). Previous studies have mostly reported
tetraploids and this is the first study with such a large
number of diploids. The specimens investigated were
largely from Northern and Western Cape, and due to the
widespread distribution of this species, the total variation
present might not be represented.
Eleiisine Gaertn. is predominantly an African genus,
with six of the nine species confined to tropical and sub-
tropical Africa (Phillips 1972). Eleusine coracana (L.)
Gaertn. is widely grown in Africa, India and China, and
used as a cereal. It is derived from E. indica (L.) Gaertn.,
a diploid cosmopolitan weed (subsp. indica, 2n = 18),
which has a tetraploid race in Africa (subsp. africana).
The moiphological characters of the two races overlap
greatly, and this leads to their inclusion in a single species
(Clayton & Renvoize 1986). Eleusine coracana subsp.
africana {- E. indica subsp. africana) is native to Africa,
where it is widespread along the eastern highlands and the
highlands of the southern African plateau (Phillips 1972).
Three E. coracana subsp. africana specimens were
investigated. The two specimens from Cape Vidal were
Bothalia 32,2 (2002)
245
diploid (Figure 9 A, B) and the specimen from Koeke-
naap in Western Cape was tetraploid (n = 2x=18)
(Figure 9C). This indicates that diploid and tetraploid
forms of this species are present in South Africa and con-
firm a basic chromosome number of nine for this genus
(Darlington & Wylie 1955; Pienaar 1955; Ornduff 1967,
1968, 1969; Federov 1969; Moore 1970, 1971, 1972,
1974, 1977;Goldblatt 1981, 1983, 1985, 1988; Goldblatt
& Johnson 1990, 1991, 1994, 1998, 2000).
Eragrostis Wolf is the largest genus in the subfamily
Chloridoideae. It has a worldwide occurrence in the
warmer regions where it is found in most habitats, show-
ing a preference for open sites, poor dry soil and weedy
places (Clayton & Renvoize 1986). The genus exhibits
the full range of morphological and anatomical variation
found in the subfamily (Van den Borre & Watson 1994).
Eragrostis is the largest grass genus in southern
Africa with ± 90 species. In this study 27 specimens were
investigated, representing 17 species. Twenty of the
specimens were tetraploid (Figure lOE, F, H). Only five
FIGURE 8. — Meiotic chromosomes
in Leptochloafiisca. A, B, Spies
3794, 2n = 2x = 20: A. ana-
phase I with 10-10 segrega-
tion; B, metaphase I. C, Spies
3932, 2n = 2x = 20, metaphase
I; D, Spies 43 16, 2n = 2x = 20,
diakineses with 10n; E, Spies
5200, 2n = 2x = 20, metaphase
I. Scale bar: 5 pm.
specimens investigated were diploid and two were hexa-
ploid (Figure lOD, G). Polyploidy is frequent in this
genus as can be seen from the results presented.
Tetraploidy, as in this study, is the most frequent poly-
ploid level observed, followed by diploidy (Darlington &
Wylie 1955; Pienaar 1955; Ornduff 1967, 1968, 1969;
Moore 1970, 1971, 1972, 1973, 1974, 1977; Goldblatt
1981, 1983, 1985, 1988; Goldblatt & Johnson 1990,
1991, 1994, 1998, 2000).
Seven specimens were investigated for E, capensis. All
but one was tetraploid (Figure lOA-C). Spies 4696 was
diploid and is the second report for this species (De Wet
1958) where tetraploidy (Avdulov 1931; Pienaar 1953;
Davidse et al, 1986; Spies & Du Plessis 1986; Spies et al,
1991) and hexaploidy (Moffet & Hurcombe 1949; Spies
& Voges 1988) have previously been observed.
De Winter (1955) regards E, curxnda as the most vari-
able species in the genus in southern Africa, with a great
many morphological forms. This was corroborated by
large-scale cytogenetic studies by Vorster & Liebenberg
EIGURE 9. — Meiotic chromosomes
in Eleusiite coracana subsp.
africana. A, B. Spies 2365,
2n = 2x=18, metaphase 1;
C, Spies 2783, 2n = 4x = 36,
diakineses with 18u. Scale bar:
5 pm.
246
Bothalia 32,2 (2002)
FIGURE 10. — Meiotic chromosomes. A-C, Eragrostis capensis: A, Spies 3483, 2n = 4x = 40, diakineses with 20u; B, Spies 3498, 2n = 4x = 40,
metaphase I; C, Spies 5069, 2n = 4x = 40, metaphase I. D, E. cunmla. Spies 1137, 2n = 6x = 60; E, E. echinocloidea. Spies 2799, 2n = 4x
= 40, diakineses with 20n; F, E. inainoena. Spies 2392, 2n = 4x = 40, diakineses with 20a; G, E. planiculmis, Du Plessis 116, 2n = 6x =
60+0-4B, diakineses with 30a, two B chromosomes are indicated; H, E. superba, Du Plessis 136, 2n = 4x = 40, diakineses with 20a. Scale
bar; B, G, 10 pm; A, C-F, H, 12 pm.
(1977). Several species are known to intergrade with E.
curvula through hybridization: E. barbinodes, E. caesia
Stapf, E. chloromelas, E. lehmaniana Nees, E. planicul-
mis and E. rigidor Pilg. (Smook 1990). Proof of
hybridization in this species indicates a collapse of iso-
lating mechanisms between different species in the Era-
grostis curvula complex, resulting in a large-scale hybrid
swarm, with continuous variation of characters between
parental extremes. The variation in morphological char-
acters is an indication of the extent of hybridization
(Spies 1984). Four specimens in this complex were cyto-
genetically investigated and tetraploid (E. barbinodes
and E. chloromelas) and hexaploid {E. cunmla and E.
planiculmis) levels were observed (Figure lOD, G). In all
but E. barbinodes, various univalents were observed
which resulted in laggards and later formed micro-
nuclei. According to Church (1929), the presence of
unpaired or univalent chromosomes is one of the most
prominent suggestions that a plant is of hybrid origin
(Church 1929) and therefore, in this complex with its
large-scale hybridization, these phenomena will be very
prevalent.
This is the third report for the species E. heteromera
with 2n = 4x = 40 (De Wet 1958; De Wet 1960). As far
as is known only two reports for E. tef exist ( Avdulov
FIGURE 1 1 . — Meiotic chromo.somes. A, B, Eingerhuthia africana. Spies 2947, 2n = 4x = 40: A, late anaphase I; B, metaphase I. C, Sporobolus
albicans. Spies 3141, 2n = 6x = 54, metaphase I; D, S. africanus. Spies 2369, 2n = 4x = 36, metaphase I; E, F, S. virginicus, Du Plessis
122, 2n = 2x = 18, diakineses with lOn (3, H, Stiburus conrathii, Du Plessis 19, 2n = 2x = 20: G, metaphase I; H, diakineses with lOu.
,Scale bar: 8 pm.
Bothalia 32,2 (2002)
247
1931; Moffet & Hurcombe 1949) and corroborates
tetraploidy present in this species. Previously only
tetraploid levels were observed in the species E. tenuifo-
lia (Omduff 1967; Moore 1973, 1977; Goldblatt 1983).
Here we report on a diploid specimen from Siteki in
Swaziland. Univalents were observed in some cells in
this specimen. The tetraploid chromosome count for E.
trichophora reported here is the second for the species
(Davidse et al. 1986) and a hexaploid specimen was
reported by Spies & Jonker (1987). One previous report
of 2n = 4x = 40 for E. racemosa (Ornduff 1967) is con-
firmed in this study where three tetraploid specimens
were found.
Two Eingerhuthia africana specimens were investi-
gated and were tetraploid (n == 2x = 20) (Figure 1 1 A, B).
Previous reports by Spies & Du Plessis (1987) and De
Wet ( 1958, 1960) also found tetraploidy in the genus, but
diploidy has been reported (De Wet 1958, 1960) as well.
Odyssea Stapf is a xerophytic grass genus with two
species, one indigenous to southern Africa (Clayton &
Renvoize 1986). It has a very distinct, much-branched,
spiny habit, which is an adaptation to its specialized
sandy and saline habitats (Phillips 1982). This is a first
report for the genus. The specimen, O. paucinenns, was
tetraploid (n = 2x = 18) and, therefore the basic chromo-
some number is 9.
The genus Sporobolus R.Br. is cytogenetically com-
plex and different basic chromosome numbers, x = 6, 9
and 10 may be present (Davidse et al. 1986). Five
Sporobolus species were investigated. Three diploid (S.
africanus. Spies 4508, S. ioclados, S. virginicus), one
tetraploid (S. africanus. Spies 2369) and one hexaploid
(5. albicans) specimen were found (Figure 1 IC-F). They
all displayed multiples of nine and this confirms x = 9 as
FIGURE 12. — Meiotic chromosomes
in Enneapogon. A, B, E.
cenchroides. Spies 2709,
2n = 6x = 60: A, diakinesis
with 30ii; B, metapha.se 1. C,
D, E. pretoriensis. Spies
3716, 2n = 2x = 20, diakine-
sis with lOii. Scale bar; A, C,
D, 5 pm; B, 8 pm.
the basic chromosome number for the genus (Darlington
& Wylie 1955; Pienaar 1955; Omduff 1967, 1968, 1969;
Federov 1969; Moore 1970, 1971, 1972, 1973, 1974,
1977; Goldblatt 1981, 1983, 1985, 1988; Goldblatt &
Johnson 1990, 1991, 1994, 1996, 1998). A chromosome
number for S. albicans has not previously been published
and this is the first report for the species (Figure IIF).
Large-scale studies in this genus are still necessary to
investigate the different basic chromosome numbers pre-
sent and their possible phylogenetic relationships.
One specimen each of the two species Stiburus
alopecuroides and 5. conrathii was examined. Both were
diploid (n = X = 10) (Figure IIG, H) and this confirms
chromosome counts based on multiples of 10 for the
genus (Spies & Du Plessis 1986).
A single Trichoneura grandiglumis specimen was
found with a chromosome count of n = x = 10. Moffet &
Hurcombe (1949) and De Wet & Anderson (1956) also
reported on diploid specimens.
Tribe Pappophoreae
The genus Enneapogon Desv. ex P.Beauv. is a very
uniform genus in which most species (28 in total) close-
ly resemble one another. Four specimens representing
two species and one unidentified specimen were examin-
ed in this study (Figure 12A-D). One specimen, Ennea-
pogon cenchroides was tetraploid with n = 2x = 20, but
another specimen was hexaploid (Figure 12A, B). Spies
5532, the unidentified specimen, was also found to be
tetraploid. A single E. pretoriensis specimen had
n = X = 10 (Figure 12C, D). Two basic chromosome
numbers are reported for this genus, x = 9, 10
(Darlington & Wylie (x = 9, 10); Omduff 1968 (x = 10),
1969 (x = 10); Moore 1970 (x = 10), 1977 (x = 10);
248
Bothalia 32,2 (2002)
FIGURE 13. — Meiotic chromosomes in Centropo-
dia glauca: A, B, Spies 5706, 2n = 8x = 48,
metaphase I. Scale bar: 8 pm.
Goldblatt 1981 (x = 10), 1985 (x = 10); Goldblatt &
Johnson 1991 (x = 10), 1998 (x = 10). The majority of
the studies support a basic chromosome number of ten.
Only three studies, De Wet (1954), Thomas unpublished
(listed Darlington & Wylie 1955) and De Wet & Ander-
son (1956) ever reported x = 9. They found one diploid
(De Wet & Anderson 1956) and five tetraploid (De Wet
1954; Thomas (Darlington & Wylie 1955); De Wet &
Anderson 1956) specimens based on x = 9. Davidse etal.
(1986) also reported on aneuploidy in the genus.
The genus Schmidtia Steud. ex J.A. Schmidt compris-
es only two species, both widespread in southern Africa.
Two specimens were examined representing S. pap-
pophoroides and an unidentified Schmidtia species. Both
were tetraploid but Spies 5536 had 0^ B chromosomes
present in some cells. This confirms the basic chromo-
some number of 9 for the genus (De Wet & Anderson
1956; De Wet 1958), although Reeder & Singh (1968)
reported on a basic chromosome number of ten.
Unplaced
Centropodia Rchb. was formerly recognized as an
arundinoid genus but has recently (GPWG 2001) been
included in the subfamily Chloridoideae. A single speci-
men of this genus was investigated and found to be octa-
ploid (n = 4x = 24) (Figure 13A, B). This confirms the
basic chromosome number of the genus as six (Du
Plessis & Spies 1988; Hoshino & Davidse 1988).
CONCLUSIONS
Chromosome numbers are reported for three of the
five tribes of the subfamily Chloridoideae. Basic chro-
mosome numbers of x = 9 and 10 occur in all the tribes.
A basic chromosome number of six is also corroborated
for the genus Centropodia. The high incidence of poly-
ploidy (65% in this study) in Poaceae and especially the
southern African grasses are once again confirmed by
this study.
ACKNOWLEDGEMENTS
The University of the Free State and the Foundation
for Research and Development are thanked for finan-
cial assistance during this study. Henriette Du Plessis
(formerly from the National Botanical Institute,
Pretoria) and Johan Venter (UFS, Bloemfontein) are
thanked for providing some meiotic material used dur-
ing this study.
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MS. received: 2002-02-27.
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Bothalia 32.2: 251.252(2002)
Book Reviews
REGIONS OF FLORISTIC ENDEMISM IN SOUTHERN AFRICA.
A REVIEW WITH EMPHASIS ON SUCCULENTS, by ABRAHAM
E. VAN WYK & GIDEON F. SMITH. 2001. Umdaus Press. P.O. Box
11059. 0028 Hatfield. Pretoria. South Africa. Pp. viii + 199. fron-
tispiece + 393 colour figures. 25 distribution maps. 36 tables. Hard
cover: ISBN 1-919766-18-9. price R290.00.
This is one of the exceptional botanical books published in 2001 . A
work of this nature has long been overdue as there is an extraordinary
floristic diversity in South Africa, extending far beyond the Cape
Floristic Region. Various books, symposia and workshop proceedings
on the diversity of the flora and vegetation of southern Africa were pub-
lished in the 1980s (Huntley 1989) and in the 1990s (Cowling &
Hilton-Taylor 1994. 1997). Cowling et al. (1997) pinpointed hotspots
of southern Africa, but this new scholarly, yet popular account, high-
lights the importance of floristic endemism. In his foreword to this
book. Prof Edward O. Wilson of Harvard University emphasized that
’one of the hottest spots is surely southern Africa'. The concept of
hotspots as centres of plant endemism was developed by Myers ( 1988.
1990). and extended to southern Africa in a paper by Cowling &
Hilton-Taylor ( 1994). This book has taken these ideas a step further and
systematically treated the entire region and examined the centres of
plant endemism in detail. The authors come up with 13 centres, some
of which are grouped into different floristic regions such as the
Succulent Karoo Region. Overall there are 19 different centres dis-
cussed in the book. To these, some 1 1 000 species and infra-specific
taxa are assigned. Since sponsorship was provided by the Lesley Hill
Succulent Karoo Trust, a greater emphasis is placed on succulent
species — this amounting to some 2 300 succulents in these endemic
regions.
The significance of the book lies in the descriptions of the different
regions and the centres that they contain. However, before we discuss
centres in detail, a review of the introduction should be given. It
includes a description of the science of plant geography, definitions of
vegetation, biomes and plant communities, and how the classification is
used in terms of the geographical range of taxa. This leads to the defin-
itions of centres of endemism as it has been used in the literature (White
1983) or centres of plant diversity (WWF & lUCN 1994). The authors
discuss classifications of the region based on growth form and patterns
of endemism and geographical range of taxa. Temis such as near-endemic,
disjunct distributions, habitat continuity, floristic elements are all very
clearly explained in the text. The recognition and demarcation of centres
of endemism in this book is not a subjective process to which the
authors applied their own ideas. Rather, they have used the wealth of
data contained within the National Herbarium. Pretoria. (PRE). and the
computerized data bank (PRECIS) which is available in that herbarium.
They have chosen and selected centres of endemism based on percep-
tion or intuitive discernment, but have taken this further by analysing
the distribution of species within these ranges. Perhaps future studies
will apply a more critical objective approach, but at this level of study,
the regions and centres of plant endemism appear to be sound and con-
form to the concepts of other authors. The map of the distribution of the
principal regions of plant endemism in South Africa, is likely to become
a classic in teaching and research literature.
Discussions of each floristic region and centre is covered in the
same way with information on its name and history, the boundaries,
topography and climate, geology and soils, vegetation and flora, (par-
ticularly succulents), including information on conservation and future
research. The Cape Floristic Region, however, is not covered in great
detail, but there is already a wealth of literature available on the Cape
flora, so this does not distract from the value of the book. Since succu-
lents form the main focus of the book, the Succulent Karoo Region is
an important component thereof with five different centres discussed
within that region. Although the Succulent Karoo is well known, these
centres are discussed in some detail in this book, and the representative
endemic succulents will become better known and the centres better
defined as a result of this study.
The remaining centres: Maputuland, Pondoland. Albany. Drakens-
berg Alpine, Barberton. Wolkberg, Sekhukhuneland. Soutpansberg.
Chimanimani-Nyanga, Great Dyke. Kaokoveld and Griqualand West
are less known and often these are areas of great interest previously not
well described. For example, the Great Dyke Centre of Zimbabwe is
shown as a dark red line on the map dividing Zimbabwe in two.
Although one of the smallest and least known centres, interesting infor-
mation on the distribution of plants along this geological feature, will
surely emphasize the importance of geology and distribution of plants.
Some of the centres have been poorly studied, yet are under poten-
tial threat of development. For example, the Pondoland Centre is
threatened with the proposed development of a toll road through the
Transkei, yet, on the other hand, it may still be subject to conservation
in the event that the Pondoland Coastal National Park becomes a real-
ity. Likewise, the Sekhukhuneland Centre has been under threat of min-
ing for platinum and iron for many years. The flora of the.se mountains
may be lost if we do not emphasize the importance of conservation in
these areas.
As an ageing ecologist 1 find it unfortunate that this book was not
published earlier in my career. With its magnificent photographs,
detailed maps of the centres and lists of species which are endemic to
these regions, it would have stimulated a wider interest in the botany of
southern Africa. We need to emphasize to would-be developers and the
lay public in southern Africa that plants should not come second to ani-
mals in conservation, since they provide the habitat and food for the
animals, as well as being conservationally important in their own right.
There is still much to be learnt about tbe detailed evolution of these flo-
ras. the fragile links between centres, and the additional aesthetic
appeal of the plants, which is of paramount interest to botanists and lay-
men alike.
REFERENCES
COWLING. R.M. & HILTON-TAYLOR. C. 1994. Patterns of plant
diversity and endemism to soutbern Africa: an overview. In B.J.
Huntley. Botanical diversity in southern Africa. Strelitzia 1:
31-52.
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: 43-61. Cambridge
University Press. Cambridge.
COWLING. R.M., RICHARDSON, D.M. & PIERCE, S.M. (eds). 1997.
Vegetation of southern Africa. Cambridge University Press,
Cambridge.
HUNTLEY. B.J. (ed.). 1989. Biotic diversity in southern Africa: con-
cepts and conservation. Oxford University Press, Cape Town.
MYERS. N. 1988. Threatened biotas: 'hotspots' in tropical forests.
Environmentalist 8: 187-208.
MYERS. N. 1990. The biodiversity challenge: expanded hotspot analy-
ses. Environmentalist 10: 243-256.
WHITE. F. 1983. The vegetation of Africa: a descriptive memoir to
accompany the Unesco/AETEAT/UNSO vegetation map of
Africa. Natural Resources Research XX: 1-356. Unesco, Paris.
WWF & lUCN 1994. Centres of plant diversity. A guide and strategy
for their consenxition. 3 volumes. lUCN Publications Unit,
Cambridge. UK.
R.A. LUBKE*
* Department of Botany. Rhodes University, P.O. Box 94, 6140 Grahams-
town. South Africa.
BULBS. Revised edition by JOHN E. BRYAN. 2002. Timber Press,
133 S.W. Second Ave. Suite 450. Portland, Oregon 97204-3527, USA.
Pp. 524. Hard cover: ISBN 0-88192-529-2, price US$ 89.95,
This edition provides an account of 3 1 5 bulbous plant genera, many
in cultivation, illustrated by 1 128 colour photographs, some black-and-
white drawings and 43 colour reproductions from Curtis’s Botanical
Magazine.
This monumental work is an extensively revised and enlarged edi-
tion of John Bryan's American Horticultural Society, award-winning
first edition of Bulbs which was published in 1989. The North
American edition now comprises one volume instead of two. The text
252
Bothalia 32.2 (2002)
is in slightly larger, easier-to-read font and with some changes in for-
matting and layout. Text pages number 524 (previously 450), largely
due to an increase in the number of genera discussed from 251 to 3 1 5.
The number of pages of colour plates has increased to 374 (previously
300) with I 171 colour photographs and reproductions (previously
832), individually numbered and referenced in Chapter 9. The text and
illustrations in the first edition are retained, but wherever possible the
scientific names and other information have been updated. Headings of
the introductory and nine numbered chapters are unchanged in the new
edition, but there has been some revision of the appendices, with the
old Appendix D to J now combined into one. Appendix C. Two new
appendices, B and D, are respectively bulbs around the World (previ-
ously on endpapers) and a useful conversion table for readers in metri-
cated countries. The old Appendix B is now replaced by an index of
common names, which follows the glossary and bibliography, while
the old list of slide contributors, about the botanical drawings and com-
bined index have been dispensed with altogether. It is nevertheless easy
to find a plant by its scientific or common name, and that is surely
essential! A feature of both editions is that authorities for scientific
names are regrettably dispensed with, which decreases the book’s use-
fulness to botanists.
Enough of comparisons. I will now review the new edition in
greater detail. The first eight, brief chapters include an overview, histo-
ry, botany and classification of bulbs, propagation, cultivation, bulbs in
the landscape, growing bulbs out of season (i.e. ‘forcing’), and pests
and diseases. The final chapter. Chapter 9, comprising the bulk of the
book (pp, 64-480), is an alphabetical listing of the genera. It is fol-
lowed by four appendices, which are mentioned in the preceding para-
graph. Finally a glossary, bibliography, and index of common names
complete the book.
The book is essentially a horticultural reference work, but due to its
encyclopaedic coverage of the subject, will be of great value to home
gardeners, professional horticulturists and botanists alike. The author’s
concise but humorous writing style makes this an enjoyable read. Mr
Bryan’s fascination with bulbs is evident throughout; this is clearly a
work of love. Additionally, his more than 50 years’ experience as hor-
ticulturist, director of botanical gardens and garden tour guide both
locally and internationally, have resulted in an authoritative text. As a
botanist 1 particularly appreciated his painstaking efforts to use the cor-
rect scientific names for plants, and to explain the mysteries of plant
classification and nomenclature to lay readers. The brief discussion of
conservation issues (Chapter 5), in these days when virtually every
plant is threatened directly by over-exploitation or indirectly by habitat
destruction, is apposite.
Turning to Chapter 9. the attributes by which any particular genus
qualifies for inclusion are unclear. A ‘bulb’ is defined (Chapter I ) as an
underground storage organ (true bulb, corm, tuber, rhizome, swollen
root or roots) which enables the plant to survive unfavourable climatic
conditions; generally the aerial parts are deciduous. This definition
includes monocotyledonous and dicotyledonous families without
drawing this fundamental distinction {Ceropegia: Apocynaceae, is next
to Chamaeliriiim: Melanthiaceae), which is surely of importance to
gardeners and horticulturists. Although many are ‘bulbous’, orchids are
specifically excluded by the author as being specialist subjects,
Zingibcraceae, too, are excluded as being mostly tropical and therefore
outside the bounds of this book, yet the small temperate genus Roscoea
is to be found here, as well as the evergreen, rhizomatous Clivici
(Amaryllidaceae), Several of the genera added in this edition are South
African, and 1 cannot help wondering why such a genus as Neodregea
would be included, especially since, quoting Bryan ‘it is unlikely that
this plant is in cultivation’. The author answers the question in his
introduction where he says ‘The selection ... is my choice alone ...’
resulting in the astounding total of 315 genera! I did not even attempt
to count the number of species, subspecies, cultivars and hybrids. Also
included in the alphabetical listing are intergeneric hybrids, and so-
called ‘blank’ entries which provide a cross reference (in the absence of
a general index) to the genus where the relevant plant names are now
to be found as a result of a change in classification. This practice could
be misleading as in the case of Cryploslephanii.s (Amaryllidaceae)
which is only to be found as the old genus for Cyrtanthus herrei,
whereas it is a good and horticulturally interesting tropical African
genus of three species, related to Cliviu. It appears, too. that while older
name changes are mostly accepted and reasonably well explained,
recent ones such as the sinking of Homeria (Iridaceae) into Moraea
have been dismissed somewhat cavalierly! While on the subject of
Homeria (now Moraea), it should be mentioned that several species
have become serious weeds (due to their toxicity) in countries other
than their native lands, primarily through garden introductions, and
here would have been an opportunity for the author to caution garden-
ers about this problem. Another mystery is Spiloxene (Hypoxidaceae).
the species of which are, in this new edition, returned to Hypoxis with-
out explanation, a move for which 1 could find no grounds in recent
taxonomic literature.
For each genus the family name is given, then general common
names are listed, followed by several general paragraphs discussing the
derivation of the name, origin of the genus, medicinal usage, horticul-
tural attributes and so on. Genera (e.g. Liliiim, Tiilipa) which have con-
tributed a vast number of important garden plants are naturally dis-
cussed in some detail, while less important genera are allocated less
space. Then follows paragraphs on culture, pests and diseases, propa-
gation. species and finally, synonyms. Incidentally the ‘species’ list
also includes categories that are not strictly species, such as hybrids and
cultivars. Following each of these names is the specific common name
which is linked to the index of common names at the back of the book.
Each ‘species’ has, where known, a date of introduction to cultivation
(chiefly in Europe), a brief description, specific cultural requirements
and a reference to a colour photograph or reproduction, if provided.
The book is profusely illustrated, the 1 171 colour photographs and
reproductions having extensive captioning. The general standard of the
colour photographs is reasonable, but some are poor, e.g. Plate 927,
Ornithogahan seineri is out of focus, whereas the identification of
some is questionable e.g. Plate 236, identified as Blandfordia nohilis:
is this not the Madagascan weed Bryophyllum delagoense (Crassu-
laceae)? and Plate 1012 is certainly not Scilla natalensis: it is rather
Ledebouria revoluta. I enjoyed seeing the reproductions of the exquisite
plates from Curtis's Botanical Magazine, but it needs to be pointed out
that the colours of the pigments have over time undergone some altera-
tion which detracts from their usefulness for identification puiposes.
The book is written primarily with Northern Hemisphere users in
mind. Because genera of diverse origin are included, there is need for
a mention of the differences in seasons between the hemispheres; there
could be included also a brief discussion of the process by which a
Southern Hemisphere bulb is adapted to the growing season in the
Northern Hemisphere. Likewise the season in which the rain is
received in the country of origin is of great importance and more
emphasis could be placed on how winter rainfall bulbs as a group (e.g.
from Western Cape, South Africa) need to be treated differently to
those from summer rainfall regions (e.g. from Eastern Cape, South
Africa). Because this important difference has been somewhat glossed
over, some confusion may result. An example is in the treatment of
Nerine where in Chapter 1 it states that ‘Nerines, coming from a part of
the world that enjoys winter rainfall . . . ’ . However, N. howdenii, one of
the species discussed in Chapter 9, is quite decidedly a summer rainfall
species. There is not always agreement between the plate caption and
the body text, such as Eriospernnnn ahyssinicinn (now E. flagelli-
fonne): on plate 480 it is said to be 'extremely rare’ whereas in the text
on p. 223 it is described as ‘common’ and indeed it is very common in
summer rainfall South Africa and northwards in tropical Africa right up
to the Central African Republic and Kenya.
Considering the scope of the book, however, these imperfections
are minor ones, and this is certainly a volume I would like to have on
my bookshelf
C. ARCHER*
* National Botanical Institute, Private Bag XlOl, OOOl Pretoria.
Bothalia 32,2: 253-267 (2002)
National Botanical Institute South Africa:
administration and research staff 31 March 2002,
publications 1 April 2001-31 March 2002
Compiler: B.A. Momberg
CHIEF DIRECTORATE (ADIR)
CAPE TOWN— PEARSON HOUSE
Huntley, Prof. B.J. M.Sc. Chief Executive Officer
Adams, Ms M.A. Principal Typist I. Receptionist
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MARKETING AND COMMUNICATION SUBDIRECTORATE (AMAR)
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Clerk III. Admin, support
PRETORIA
Mthembi, Ms C. Dip. (Market. & Pub. Relations), Dip. (Market. Manag.).
Senior Communications Officer. Marketing & pub. relations
GRAPHIC SERVICES— CAPE TOWN (AMAR)
Loedolff, Mrs J. B.Sc.dnd.Tech.). Chief Industrial Technician. Photographer/Illustrator (part time)
VISITORS’ CENTRE— CAPE TOWN (AMAR)
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Officer. Centre Manager
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Clerk II. Facilities Co-ordinator
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Malan, Ms C. B. Sc. (Hons). Principal Communication Officer. Tour Co-ordinator
HUMAN RESOURCES DIRECTORATE (APER)
CAPE TOWN
Tlali, P.L.N. B.Iuris Lib. Director. Human Resources
Liebenberg, Mrs E.J.L. Acting Manager: Admin, and Human Resources — Pretoria (RPTA)
Kriel, Mrs G.A. Dip. Sec. Senior Secretary IV
Finca, Ms N.F. Cleaner II
Haupt, Mrs C.S. Cleaner II. Guest house
Woodward, Ms Y.J. Senior Provisioning Admin. Clerk III. Secretary to NBI Board
254
Bothalia 32,2 (2002)
PERSONNEL— CAPE TOWN
Van Zyl, J.M. M.Econ.(lnd. Psych.). Assistant Director: Organizational Development & Training
Engelbrecht, B. N.Dip.(Hort.), N.Dip.(PRM), Dip.(Eorestry). Assistant Director: Human resources
management and health & safety co-ordination
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Claassen, Ms G. Senior Telecom Operator III. Admin,
support
Crowie, A.C. Senior Registry Clerk I
Dollie, Mrs N.J. Cleaner II
Du Toit, Ms R. H.E.D. Senior Training Officer. Training
Co-ordinator
Fredericks, C.H. Driver II. Courier services
Mtyalela, D.M.S. Senior Training Officer
Nicholas, Mrs W.L. Senior Photocopy Machine Operator
Petersen, R.E. Principal Personnel Officer. Employment
equity
Sass, Ms D. Senior Provisioning Admin. Clerk I. Leave
& lOD admin.
Sotshononda, Ms P. Telecom Operator II. Reception HO
Williams, Mrs L.R. Dip. Sec. Senior Personnel Officer II.
Human resources management
ADMINISTRATION AND HUMAN RESOURCES— PRETORIA (RPTA)
Liebenberg, Mrs E.J.L. Acting Manager
Gbtzel, Mrs A. Senior Telecom. Operator III
Khumalo, N.P. Principal General Foreman. Supervisor:
Office services
Ledwaba, Mrs D.M. Senior Registry Clerk I
Makgobola, Mrs M.R. Cleaner II
Malefo, Mrs R.P. Cleaner II
Maphuta, Mrs M.S. Cleaner II
Mamie, P.M. Senior Handyman
Nkosi, Mrs M.R Cleaner II
Phaahla. M.C. Cleaner II
Pretorius, Ms M.A. Senior Provisioning Admin. Clerk II
Randima, G. Cleaner I
Sithole, A.M. Cleaner I
Tloubatla, J.M. Driver II. Courier services
FINANCE DIRECTORATE (AFIN)
CAPE TOWN
Hughes, W.S.G. B.Com., C.A.(SA). Director. Chief Financial Officer
Rawoot, N.A. B.Acc.Sc. Deputy Director. Financial management
Maholwana, S. B.Com. Assistant Director. Financial management
Neuwirth, Ms E.V. Assistant Director. Staff benefits
Bagus, Ms E.J. Senior State Accountant. Financial admin.:
General ledger
Cassiem, Ms S. Senior Accounting Clerk III. Staff bene-
fits & salaries
Cooper, S.K. Accounting Clerk II. Creditors
Geduldt, D.C. Senior Accounting Clerk II. Debtors
Goodman, Mrs I.W. Chief Accounting Clerk. Supervisor:
Creditors
Jacobs, F.H. Senior Accounting Clerk II. Staff benefits &
salaries
Koyana, Ms Z. Senior Accounting Clerk I. Salaries
Mcontsi, Ms N. N. Dip. (Accounting). Senior Accounting
Clerk I. Creditors
Mirkin, Ms Y.A. Senior Secretary II
Ogunniyi, Mrs E. Internal Auditor (contract worker)
Paulse, Mrs D.W.S. Senior Accounting Clerk III. Credi-
tors
September. Ms M. Senior Accounting Clerk I. Debtors
BIODIVERSITY POLICY AND PLANNING DIRECTORATE
Njobe, Ms K. M.Sc. Director. Biodiversity and Action Planning
Golding, Ms J. B. Sc. (Hons). Manager: Threatened Plants Programme
GARDENS DIRECTORATE— ADMIN STAFF (GDIR)
PRETORIA
Willis, C.K. M.Sc. (Cons. Biol.) — Director: Gardens and Horticultural Services
Winter, J.H.S. N.Dip.(Hort.). Deputy Director: Horticultural Services (Cape Town)
Davis, G.W. Ph.D. Assistant Director: Project management and fund raising (Cape Town)
Heilgendorff, J.P. N.H.Dip.(Hort.). Gardens IT Manager
Mkefe, T..I. S.P.E.D. Principal Communications Officer. Public affairs & liaison
Bothalia 32.2 (2002)
255
INTERPRETATION
Joffe. Mrs H. B.Sc. Chiet Agricultural Development Technician. Interpretation (part time)
Roff, J. Communication Officer. Interpretation (Pietermaritzburg)
CURATORS
Behr, Ms C.M. Curator: Harold Porter NBG (Betty’s Bay)
Blitz. R.M. Curator: Lowveld NBG (Nelspruit)
Le Roux. PH. Deputy Director. Curator: Kirstenbosch NBG (Cape Town)
Mogale, A.O. Curator: Tree State NBG (Bloemfontein)
Oliver. I.B. Curator: Karoo Desert NBG (Worcester)
Tarr, B.B. Curator: Natal NBG (Pietermaritzburg)
Turner. Ms S.L. Curator: Witwatersrand NBG (Roodepoort)
PLANNING. MAINTENANCE & DEVELOPMENT— CAPE TOWN (BPMD)
Linde. D.C. N.T.C.IIKCivil & Structural: Building) N.T.C.III (Inspector of Works: Building).
M.S.A.I.D. Cert. Estate Agency. Control Inspector of Works
Arendse. D.S. Eactotum. Building maintenance
Manasse. S.P. Dip. (Masonry). Artisan Eoreman. Building maintenance
Peck, W.I. Artisan’s Assistant II. Building maintenance
HAROLD PORTER NBG— BETTY’S BAY (GHPG)
Behr, Ms C.M. B.Sc. (Hons). Control Agricultural Technician. Curator
Abrahamse, S.J. Principal Eoreman. Garden
Ai'endse, Ms M. Auxiliary Services Officer II. Access control
Bezuidenhout. Mrs H.M. Provisioning Admin. Officer
Carolus, Ms B.J. N.H.Dip.(Hoit.) Agricultural Develop-
ment Technician. Horticulture
Eorrester, Ms J.A. N.TC.IIKHort.). Chief Agricultural
Development Technician. Horticulture
October. Ms R.P. Dip. (Education). Senior Auxiliary
Services Officer. Plant records and asset register
Oliver, R.C. Senior Specialist Groundsman
Samuels, Ms D.C. Cleaner II
Simpson. H.A.S. Auxiliary Services Officer II. Access
control
Van Wyk. A.B. Artisan. Building maintenance
KAROO DESERT NBG— WORCESTER (GKAR)
Oliver, I.B. N.Dip.(Hort.), N. Dip. (Public Relations & Admin). Control Agricultural Technician. Curator
Ashworth, Mrs E.H. Senior Provisioning Admin. Clerk
III. Admin, support
Mtetwa. A. Artisan. General maintenance
Makubalo, E.N. Senior Foreman. Nursery
Mpeke, Ms E.N. Cleaner II
Sibozo, N.E. Driver (HMV) II. Plant sales
Simani, D.K. Principal Foreman. Garden
Van Tonder, M. (volunteer)
Viljoen. D.M. N.Dip.(Hort.). Chief Agricultural Develop-
ment Technician. Collections
Voigt, W.E. N.Dip.(Hort.). Agricultural Development Tech-
nician. Garden development
Williams, C. (student)
KIRSTENBOSCH NBG— CAPE TOWN (GKBC)
Le Roux, PH. Dip. (Forestry), N.Dip.(Hort.), N. Dip. (Parks & Recr.), Cert. (Turf Management).
Deputy Director: Garden Management. Curator
Goldschmidt, S.M. B.A.(Soc.)(Hons.). Assistant Director. Personnel Manager
Morkel, A.T. N. Dip. (Nat. Cons.). Control Agricultural Technician. Estate Manager
Hitchcock. A.N. N.H.Dip.tHort.). Control Agricultural Technician. Nursery Manager
Notten, Ms A.L. B.Sc., N.Dip.(Hort.). Chief Agricultural Development Technician. Supervisor: Seed room
Trautman, C.E. Artisan. Supervisor: Workshop
Adams, T.D. B.Tech.(Hort.). Agricultural Development Technician. Supervisor: Greenhouse
Adams, G. Specialist Groundsman. Nursery maintenance^ Arends, Ms S.J. Principal Auxiliary Services Officer. Plant
Adonis, A. Principal Foreman. Dell & ericas records
Adonis, S.J. Senior Foreman. Aliens vegetation control August, T. Senior Foreman. Visitors’ Centre. Infonnation
256
Bothalia 32,2 (2002)
services
Brown, B.M. N.Dip.(Hort.). Agricultural Development
Technician. Nursery
Crowie, R.W. Senior Foreman. General garden
Davids, I.M. Specialist Groundsman. Bulbs
Difoloko, J.A. (student)
Dlela, Ms D.I. (student)
Duncan, G.D. N.Dip.(Hort.). Chief Agricultural Develop-
ment Technician. Bulbs
Engelbrecht, F. Provisioning Admin. Clerk II. Stores
Engelbrecht, Mrs L.D. Control Auxiliary Services Offi-
cer. Plant records
Eredericks, N.C.E. Senior Auxiliary Services Officer.
Visitors’ Centre. Information services
Grace, T. Senior Provisioning Admin. Clerk III. Stores &
admin, support
Hope, C. Senior Handyman. Maintenance services
Jacobs, A.P. Chief Auxiliary Services Officer. Visitors’
Centre. Information services
Jacobs, H.C. Senior Eoreman. Nursery maintenance
Jamieson, Mrs H.G. N. Dip. (Parks & Rec.). Chief Agri-
cultural Development Technician. Restio/ Aspara-
gus (part time)
Jodamus, Ms N.L. N.Dip.(Hort.). Senior Agricultural
Development Technician. Annuals
Kamalie, Ms S. Typist II. Receptionist
Kayster, G.J. Senior Foreman. Construction
Kuscus, G.W. Senior Foreman. General maintenance
Eawrence, E. N.Dip.(Hoi1.). Senior Agricultural Develop-
ment Technician. Dell & ericas
Eewin, T.B. Specialist Groundsman. Nursery
Manuel, I.P. Senior Foreman. Seed room
Marthinus, E. Senior Specialist Groundsman. Succulents
Mathys, Mrs S.S.B. Senior Accounting Clerk III. Reve-
nue and visitor statistics
Matthews, I.N. Senior Eoreman. Estate & trails
Mbambezeli, N.G. N.Dip.(Hort). Agricultural Develop-
ment Technician. Trees & shrubs
Morris, J.N.M. Senior Eoreman. Proteas
Newman, W. Senior Handyman. Mechanical workshop
Petersen, A.D. Specialist Groundsman. Cycads
Picane, Ms S. Auxiliary Services Officer II. Tissue culture
Prins, E.B. Security Officer III
Rudolph, A. Security Officer III
Shaide, Ms A.C. N.Dip.(Hort.). Principal Communica-
tions Officer. Outreach Programme
Smith, Mrs A. Senior Provisioning Admin. Clerk I. Admin,
support
Solomons, T.C. Senior Security Officer II
Tamboer, J.S. Senior Foreman. Nursery services
Van Gusling, E.J. Senior Foreman. Mowing unit
Van der Walt, Mrs L.E. N.Dip.(Hort.). Chief Agricultural
Development Technician. Herbaceous collections
(part time)
Van Jaarsveld, E.J. M.Sc., N.Dip.(Hort.). Control Agri-
cultural Technician. Succulents
Van Rooy, K. Senior Eoreman. Annuals
Van Wyk, E. Senior Auxiliary Services Officer II. Eable
Maker
Viljoen, Ms C.C. N.Dip.(Hort.). Senior Agricultural
Development Technician. Plant production
Williams, G.C. Senior Accounting Clerk I. Access control
LOWVELD NBG— NELSPRUIT (GLOW)
Britz, R.M. N.Dip.(Eorestry). Control Agricultural Technican. Curator
Froneman, W.C.F. N.T.C.IIKHort.), N. Dip. (Nature Cons.
& Man.), N. Dip. (Parks & Rec. Admin.) Control
Agricultural Technician. Nursery management &
garden development
Hurter, P.J.H. B. Sc. (Hons). Control Agricultural Techni-
cian. Cycad conservation
Maqungo, Ms V.L.B. Senior Accounting Clerk I. Kiosk
Mathe, B.X. (student)
Mathebula, Ms N.R. Accounting Clerk II. Admin, support
Musweli, K.J. Senior Foreman. New development
Ndlovu, L.D. Senior Handyman. Maintenance
Ngwenya, PS. Auxiliary Services Officer II. Kiosk
Sibanyoni, Ms S.M. Cleaner I
Van der Walt, Mrs G.A.M. Senior Provisioning Admin.
Clerk III
Xozumti, M.M. Senior Foreman. Supervisor
Zuma, R.S. (student)
NATAL NBG— PIETERMARITZBURG (GNAT)
Tarr, B.B. N. Dip. (Parks & Rec. Admin.), Advanced Dip. (Adult Educ.). Control Agricultural Technician. Curator
Dlungwane, T.R. Senior Eoreman. Garden maintenance
Johnson, Ms I. M.Sc., H.E.D. Senior Agricultural Develop-
ment Technician
Nkabini, A.B. Senior Specialist Groundsman. Garden
Nonjinge, S.H.B. N.T.C.IIKHort.). Chief Agricultural
Development Technician
Sibiya, Ms C.P.T. Cleaner I
Van der Merwe, Mrs M.E.H. Senior Provisioning Admin.
Clerk III
Xaba, PA. (student)
Zimu, M.J. Senior Specialist Groundsman. Garden
FREE STATE NBG— BLOEMEONTEIN (GESG)
Mogale, A.O. N.Dip.dlort.). Control Agricultural Technician. Curator
Eysele, Mrs J.P. Senior Provisioning Admin. Clerk HI. ment Technician. Nursery
Admin, support Harris, Ms S. N.Dip.(Hort,). Agricultural Development
Gavhi, M.P. N.Dip.tHort.). Senior Agricultural Develop- Technician
Bothalia 32,2 (2002)
257
Lekhetho, T.S. Specialist Groundsman. Nursery Sebolai, R.P.A.N. Senior Foreman. General maintenance
Nakanyane, R.B. Principal Foreman. Garden Thaele, Mrs M.E. Cleaner II
Raditlhare, Mrs E.M. Cleaner II Tsiloane, FM. Driver (HMV) 1
PRETORIA NBG (GPTA)
Klapwijk, N.A. N.Dip.(Hort.), N. Dip. (Plant Prod.), N. Dip. (Diesel Fitting). Control Agricultural Technician.
Acting Curator
Baloyi, K.J. Senior Auxiliary Services Officer II. Garden
records
Baloyi, M.S. Dip. (IBM), Dip.(PTM), Dip. (Payroll Admin.).
Senior Provisioning Admin. Clerk I. Personnel
records and H.R. support
Creighton, Ms D.D. Senior Provisioning Admin. Clerk
III. Admin, support
Eyssel, Ms A. Senior Agricultural Development Tech-
nician. Garden
Keyter, B.A. Senior Security Officer I
Kutama. B.T Foreman. Garden development
Makena, N.S. Specialist Groundsman. General maintenance
Makgobola, Ms M.R. Auxiliary Services Officer II.
Reception & admin, support
Makoeng, TP Foreman Production nursery
Mariri, Ms M.A. Cleaner II
Matshika, S.P. Groundsman II. Cook
Modisha, M.D. Cleaner II
Noku, Y.A. Senior Foreman. Transport
Sete, B.L. Senior Foreman. Garden maintenance
Shilubane, E. Senior Eoreman. Scientific collections
Shirindi, J.R. Specialist Groundsman. Cycads
Solomons, Ms C.V. Senior Auxiliary Services Officer.
Records clerk
Swartz, Ms PP M.Sc. Chief Agricultural Development
Technician. Scientific and horticultural curation of
living collections; inteipretation, garden planning
and development; tour groups and visitors
Venter, W.A. N.T.C.II. Artisan. Workshop and general
maintenance
WITWATERSRAND NBG— ROODEPOORT (GWIT)
Turner, Ms S.L. B. Sc. (Hons), N.Dip.(Hoi1.). Control Agricultural Technician. Curator
Aubrey. Mrs A.E. N.Dip.(Hort.). Chief Agricultural De-
velopment Technician. Plant records, inteipreta-
tion, information
Baloyi, S.J. Handyman
Hankey, A.J. B.Tech.(Hort.). Control Agricultural Tech-
nician. Garden, estate, collections, nursery
Head, Mrs S.E. Dip. (Shorthand & Typing). Provisioning
Admin. Officer
Mamosebo, M.A. Factotum
Manjati, Mrs N.L. Senior Provisioning Admin. Clerk I
Manyikana, TM. Factotum
Mmola, Mrs B.E. Cleaner II
Mutshinyalo, TT. N.Dip.(Hort.) Agricultural Develop-
ment Technician. Nursery, garden
Ndou, Ms N.L. Auxiliary Services Officer II
Ndzondo, Ms N.L. Senior Provisioning Admin. Clerk I
Ndzondo, Mrs P.G. Cleaner II
Nedambale, M.P. Senior Eoreman
Nemalili, M.E. Senior Eoreman. Garden
Nenugwi, M.S. Senior Eoreman. Nursery
Tebeile. Ms Z.M. Senior Provisioning Admin. Clerk I.
Receptionist
RESEARCH DIRECTORATE (RDIR)
PRETORIA
Smith. Prof. G.F Ph.D., F.L.S. Director: Research
Rutherford, M.C. Ph.D., Dip.(Datamet.). Deputy Director; Ecology and conservation (Cape Town)
Wolfson, Mrs M.M. Ph.D. Deputy Director; Environmental education and research support
Klopper, Ms R. M.Sc. Technician (contract worker) Marais, Mrs A.C. Senior Secretary IV
Meyer, Mrs N.L. B. Sc. (Hons) Technician (contract Steyn, Mrs E.M. A. D.Sc. Specialist Scientist. Embry-
worker) ology, anatomy, taxonomy
PLANT SYSTEMATICS SUBDIRECTORATE
PRETORIA
Smith, Prof. G.F Ph.D., F.L.S. Systematics of succulents and rosulate, petaloid monocots
Arnold, T.H. Head: Data Management (Pretoria)
Crouch, N.R. Head; Ethnobotany Unit (Durban)
Koekemoer, Ms M. Curator: National Herbarium (Pretoria)
Leistner, O.A. D.Sc. E.L.S. Scientist (contract worker)
Rourke, J.P Curator: Compton Herbarium (Cape Town)
Siebert. S.J. Regional Project Co-ordinator: SABONET (Pretoria)
Williams, Ms R. Curator; Natal Herbarium (Durban)
258
Bothalia 32,2 (2002)
COMPTON HERBARIUM— CAPE TOWN (RHEC)
Rourke, J.P Ph.D., F.M.L.S., F.R.S.S.Af. Assistant Director.
Systematics of southern African Proteaceae, Stilbaceae
Baatjes, Ms A. Data Capturer (SABONET contract worker)
Beyers, Mrs J.B.P. Ph.D. Principal Agricultural Scientist.
Assistant Curator; Collections. Taxonomy of the
Gnidieae (Thymelaeaceae)
Chesselet, Ms P.C.M. M.Sc. Senior Agricultural Scientist
Conrad, Ms C. M.Sc. Agricultural Scientist. Molecular
systematics laboratory
Cupido, C.N. M.Sc. Senior Agricultural Scientist
Cupido, Mrs C.S. Senior Auxiliary Services Officer II.
Technical Assistant
Engelbrecht, Ms M. (SABONET contract worker)
Foster, Mrs S.E. Senior Secretary IV
Kurzweil, H. Ph.D. Specialist Scientist. Systematics of
southern African terrestrial orchids
Leith, Mrs J. Senior Provisioning Admin. Clerk III
Manning, J.C. Ph.D. Specialist Scientist. Systematics of
Iridaceae and Hyacinthaceae; anatomy
Marinus, Ms E.D.A. Chief Auxiliary Services Officer.
Herbarium Assistant
Oliver, E.G.H. Ph.D. Specialist Scientist. Taxonomy of
the Ericoideae (Ericaceae)
Oliver, Mrs I.M. (contract worker)
Parker, Ms F. B. Sc. (Hons). (SABONET contract worker)
Paterson-Jones, Mrs D.A. (nee Snijman) Ph.D. Principal
Agricultural Scientist. Systematics of Amarylli-
daceae; cladistics
Reeves, Ms G. Ph.D. Molecular systematics (contract
worker)
Roux, J.P. N.T.C.IIKHort.), F.L.S., Ph.D. Principal Agri-
cultural Scientist. Systematics of Pteridophyta
Williams, Mrs V.J. Data Capturer (SABONET contract
worker)
NATAL HERBARIUM— DURBAN (RHED)
Williams, Ms R. B. Sc. (Hons), H.E.D. Control Agricultural Technician. Curator
Dimon, Ms Z.Y. B.Sc. Data Capturer (SABONET con-
tract worker)
Govender, Mrs N. B.Sc. (Hons) Data Capturer. Chironia
systematics (SABONET contract worker)
Hlongwane, Mrs N.C. Cleaner II & messenger
Mbonambi, B.M. Groundsman II.
Nathoo, Ms M. B.Sc. Data Capturer (SABONET con-
tract worker)
Ngwenya, A.M. Agricultural Development Technician.
Herbarium Officer. Plant identification and infor-
mation, Zulu Botanical Knowledge Project
Mazibuko, J.V.G. Auxiliary Services Officer II. Herba-
rium Assistant
Noble, Mrs H-E. Senior Provisioning Admin. Clerk III
Singh, Ms Y. M.Sc., H.E.D. Senior Agricultural Scien-
tist. Taxonomy of Araceae, Hypoxidaceae
ETHNOBOTANY UNIT— DURBAN (RETH)
Crouch, N.R. Ph.D. Assistant Director. Ethnobotany of southern African flora
NATIONAL HERBARIUM— PRETORIA (RHEN)
Koekemoer, Ms M. Ph.D. Deputy Director. Herbarium management.
Taxonomy of Asteraceae: Gnaphalieae
Bredenkamp, Mrs C.L. M.Sc. Principal Agricultural Scientist. Assistant Curator: Public relations.
Taxonomy of Vitex, Passenmi, Rhamnaceae, Sterculiaceae and other related families
Herman, PPJ. M.Sc. Principal Agricultural Scientist. Assistant Curator; Personnel. Taxonomy of
Asteraceae, Flora of Transvaal
Anderson, Ms H.M. Ph.D. Principal Agricultural Scien-
tist. Palaeobotany, palaeogeography
Anderson, J.M. Ph.D. Specialist Scientist. Palaeobotany,
palaeogeography
Archer, R.H. Ph.D. Senior Agricultural Scientist. Taxon-
omy of mainly Celastraceae, Euphorbiaceae
Archer Mrs C. M.Sc. Senior Agricultural Scientist. Taxon-
omy of Cyperaceae, Restionaceae, Orchidaceae
Burgoyne, Ms PM. M.Sc. Principal Agricultural Scien-
tist. Mesembryanthemaceae
Fish, Mrs L. B.Sc. Principal Agricullural Scientist. Taxon-
omy of Poaceae. Plant collecting programme;
supervising mounters
Glen, H.F Ph.D. Specialist Scientist. Taxonomy of trees,
herbarium for cultivated plants, and botanical col-
lectors
Glen, Mrs R.P M.Sc. Chief Agricultural Development
Technician. Taxonomy of ferns, water plants
Jordaan, Mrs M. M.Sc. Principal Agricultural Scientist.
Taxonomy of Casuarinaceae-Connaiaceae, Maytenus
Kgaditsi, TW. Senior Auxiliary Services Officer. Speci-
men mounter, general assistant in cultivated plants
section
Klein, R.G. Plant Collector, Bioprospecting Project (con-
tract worker)
Makgakga, M.C. Senior Auxiliary Services Officer. Her-
barium Assistant, Wing B
Makgakga, K.S. Senior Auxiliary Services Officer. Her-
barium Assistant. Encoding plant specimens
Manamela, Ms M.T B.Sc. Agricultural Development
Technician. Information Officer
Maserumule, M.K. Auxiliary Services Officer, Wing B
Bothalia 32.2(2002)
259
Masombuka, Ms A.S. Senior Auxiliary Services Officer.
Herbarium Assistant. Wing A
Meyer, J.J. H.E.D. Senior Agricultural Development Tech-
nician. Bioprospecting Project
Mmakola, E.K. Data Capturer (SABONET contract worker)
Moeng, E.T. B. Sc. (Hons). Agricultural Development Tech-
nician. Wing A
Mothapo, M.A. Data Capturer (SABONET contract worker)
Mothogoane, M.S. Senior Auxiliary Services Officer.
Herbarium Assistant
Naicker, K. Dip.( Bookkeeping). Sales & Marketing Manage-
ment Certificate, H.Cert.(Prac. Accounting). Senior
Provisioning Admin. Clerk II. Administrative sup-
port
Netnou, Ms N.C. M.Sc. Senior Agricultural Develop-
ment Technician. Wing D
Nkoana, L.S. B.Sc. Senior Agricultural Development
Technician. SABONET project
Nkoane, Ms G.K. Auxiliary Services Officer II. Herbarium
Assistant. Parcelling, pressing, general assistance
Nkonki, Mrs T. B.Sc. Senior Agricultural Development
Technician. Wing B
Perold, Mrs S.M. Ph.D. Taxonomy of Hepaticae (con-
tract worker)
Phahla. T.J. Senior Auxiliary Services Officer. Specimen
mounter of bryophytes and vascular plants
Ready, Mrs J.A. N.Dip.(Hort.). Principal Auxiliary
Services Officer. Herbarium Assistant. Wing D
Rampho, Ms E.T. Data Quality Controller (SABONET
contract worker) SECOSUD
Retief, Miss E. M.Sc. Principal Agricultural Scientist.
Pollen studies of Boraginaceae. Taxonomy of Bora-
ginaceae, Verbenaceae, Lamiaceae, Asteraceae,
Rubiaceae
Sebothoma, P.N. Senior Auxiliary Services Officer II.
Plant identifications co-ordinator
Smithies, Mrs S.J. M.Sc., Dip. Ed. (Moray House). Chief
Agricultural Development Technician. Taxonomy
of Scrophulariaceae, Selaginaceae, Lobeliaceae
Steyn, Ms C.C. Principal Auxiliary Services Officer.
Label typist
Van Rooy, J. Ph.D. Principal Agricultural Scientist. Taxon-
omy and biogeography of mosses
Van Wyk, E. M.Sc. Agricultural Development Techni-
cian. Seedbank Manager, Kew Millenium Seed-
bank Project (contract worker)
Victor, Ms J.E. M.Sc., H.Dip.(Journ.). Principal Agri-
cultural Scientist. Taxonomy of Rutaceae, Ascle-
piadaceae
Welman, Ms W.G. M.Sc. Principal Agricultural Scientist.
Taxonomy of Convolvulaceae, Solanaceae, Cucur-
bitaceae, Campanulaceae, Asteraceae, Acanthaceae
DATA MANAGEMENT— PRETORIA (RPDC)
Arnold, T.H. M.Sc. Assistant Director. Computer database application especially in taxonomy
Botha, Mrs A.G. Chief Auxiliary Services Officer. Admin.
Assistant (part time)
De Wet, Mrs B.C. B.Sc. (Computer Science), B.A.,
H.D.L.S. Principal Agricultural Datametrician
Harris, Mrs B.J. Chief Auxiliary Services Officer. En-
coding, quality control
Neveling, Mrs V.H. Principal typist 1
Smit, G.C. NT Workstation 4, NT Server 4. Principal
Network Controller
Snyman, Mrs E.E. B.Sc. N. Dip. (Comp. Data Proc.)
Senior Agricultural Development Technician
Steyn, Ms H.M. Botanical Information Officer (contract
worker)
Swelankomo, N. Agricultural Development Technician.
Quality control
SABONET
PRETORIA
Siebeit, S.J. Ph.D. Regional Co-ordinator (contract worker)
Haasbroek, Ms C.M. B.Com. Einancial Officer (contract Management (contract worker)
worker) Noko, Ms N.R. Admin. Officer (contract worker)
Mossmer, Ms M. B.Sc. (Hons). Publications and Website
EDUCATION AND RESEARCH SUPPORT— PRETORIA (EDIR)
Wolfson, Mrs M.M. Ph.D. Deputy Director. Physiology/Ecophysiology of Poaceae, carbon uptake
metabolism, allocation in response to environmental and management stress
Liebenberg, Mrs E.J.L. Manager: Research Support Services, Publications
Potgieter, Mrs E. Principal Librarian
260
EDUCATION (EDIR)
GOLD FIELDS CENTRE— CAPE TOWN (EECT)
Bothalia 32,2 (2002)
Gaffoor, Ms M. H.E.D. Principal Communication Officer Co-ordinator Environmental education
Cupido, Ms M. Senior Provisioning Admin. Clerk II. Admin, support
Hitchcock, Mrs W.A. H.E.D. , B. Sc. (Hons). Principal Communications Officer. Environ-
mental education
Mgodeli, W.M. Driver (HMV) I
Tyhokolo, Ms S.E. S.P.E.D. Senior Communications Officer. Environmental education
NORTHERN GARDENS (GENT/HO) AND PRETORIA (GENT/GP)
Symonds, Ms A.M. N. Dip. (Nature Cons.), H.E.D. Assistant Director. Environmental education
Adams, Ms E.M. Senior Provisioning Admin. Officer. Environmental Education Manager
De Bruyn, Ms A.J. B.Sc. (Zoo., Bot. & Mammalogy).
Principal Communication Officer. Environmental
education
Mathaba, T.C. Environmental Education Officer
Nkomo, Mrs A. Assistant Admin. Officer (contract worker)
Novellie, Mrs E. H.E.D. B.Sc. (Hons) (Zool. & Mam-
mology) Senior Environmental Education Officer.
Environmental education
WITWATERSRAND
Molefe, Ms K.E. Senior Environmental Education Officer
Moore, Mrs J.M. Senior Provisioning Admin. Clerk II. Admin, support (part time)
Van der Westhuizen, Mrs S. M. Sc. (Bot.). Principal Communications Officer
RESEARCH SUPPORT SERVICES AND PUBLICATIONS— PRETORIA (RPUB)
Liebenberg, Mrs E.J.L. M.Sc. Chief Agricultural Development Technician. Cytotaxonomy. Manager
Brink, Mrs S.S. Dip. (Typing). Chief Typesetter. Type-
setting, layout, word processing
Condy, Ms G.S. M.A. Chief Industrial Technician. Bota-
nical artist
Du Plessis, Mrs E. B.Sc. (Hons), S.E.D. Principal Com-
munications Officer. Technical editor. Editing, trans-
lating, layout
Germishuizen, G. M.Sc. Assistant Director. Editor
Mapheza, T.P. Senior Provisioning Admin. Clerk III.
Bookshop Manager
Momberg, Mrs B.A. B.Sc. (Entomology & Zoology).
Principal Language Practitioner. Technical editor.
Editing, layout (part time)
Maree, Ms D.J. H.E.D. Senior Computer Operator.
Nkosi, PB. Provisioning Admin. Clerk I. Bookstore
Romanowski, Mrs A.J. Dip. (Photography). Chief Indus-
trial Technician (Photography). Scientific photog-
rapher
Turck, Mrs S. B.A. (Information Design). Senior Indus-
trial Technician. Graphic design
MARY GUNN LIBRARY— PRETORIA (RLBP)
Potgieter, Mrs E. B.Libr. Principal Librarian
Eourie, Mrs A. B.A., H.Dip.Libr.Sci. Principal Librarian (part time)
Shipalana, Ms K.M. N.Dip. Library and Information Studies (student contract worker)
ECOLOGY AND CONSERVATION SUBDIRECTORATE (RREL)
CAPE TOWN
Rulherford, M.C. Ph.D., Dip.(Datamet.). Deputy Director: Research
Morkel, Ms L. N.Dip. (Office Admin.). Senior Provisioning Admin. Clerk 111.
Personal Assistant to Deputy Director: Research
Parenzee, Ms H.A. Dip. (Ed.) Senior Provisioning Admin. Clerk III
Powrie, L.W. M.Sc. Senior Data Technologist. Spatial modelling, databases
Bothalia 32,2 (2002)
261
CLIMATE CHANGE
Rutherford, M.C. Ph.D., Dip.(Datamet.). Chief Specialist Scientist. Modelling, global change
Arnolds, Ms J.L. Principal Auxiliary Services Officer. Laboratory
Kgope, B.S. M.Sc. Agricultural Scientist. Plant ecophysiology
Midgley, G.F. Ph.D. Principal Specialist Scientist. Plant ecophysiology, modelling
Millar, Ms D.L. M.Sc. Research Coordinator (contract worker)
Motete, Ms N. B.Sc.(Bot. & Ed.), M.Sc.(Envir. Biol.), Dip. (Science Ed.). Agricultural Scientist
Musil, C.E. Ph.D. Senior Specialist Scientist. Ecophysiology, modelling
Snyders, S.G. Senior Auxiliary Services Officer II. Greenhouse, maintenance
CONSERVATION BIOLOGY
Donaldson, J.S. Ph.D. (Zoology) Principal Specialist Scientist. Supervisor:
Conservation farming. Cycad biology
Bosenberg, J. de Wet. B. Sc. (Hons). Chief Agricultural Development Technician. Cycad biology.
Conservation farming
Ebrahim, I. N.Dip.(Hort.). Assistant. Protea Atlas Project (contract worker)
Marinus, E.M. N. Cert. (Building & Structures). Principal Auxilliary Services Officer. Conservation farming
Rebelo, A.G. Ph.D. (Zoology) Principal Agricultural Scientist. Protea Atlas Project
Smit, W.J. M.A. Geographic Information Assistant. Protea Atlas Project (contract worker)
CONSERVATION FARMING PROJECT
Nanni, Ms I. B.Sc., H.E.D. Chief Agricultural Development Technician. Project Co-ordinator
Breebaart, Ms L. M.Sc. (Range and Eorage Resources). Researcher. Production benefits of different
grazing systems (Nama Karoo) (contract worker)
Leonhart, Ms A. N. Dip. (Nature Cons.). Research Assistant. Assessments of biodiversity and eco-
system services (contract worker)
Segers, Ms A. Senior Provisioning Admin. Clerk III (contract worker)
Skowno, A. M.Sc. Research Officer. Biodiversity risk analysis in farm landscapes (contract worker)
Theron, L.J. M.Sc. (Zoology). Research Assistant (contract worker)
DESERTIFICATION
Petersen, Ms A. B.Sc. (Hons). Senior Agricultural Development Technician. Land use and vegetation mapping
HORTICULTURAL RESEARCH
Brown, N.A.C. Ph.D. Specialist Scientist. Seed research
Botha, PA. N.H.Dip.(Hort.). Chief Agricultural Development Technician. Seed research
INFORMATION TECHNOLOGY
Evans, N. Control Network Controller. Technical IT Manager
Pekeur, Ms B.E. IT Admin. Clerk (contract worker)
SUPPORT SERVICES
Bardien-Overmeyer, Ms S. B.A.tPharm.). Provisioning Admin. Officer. Admin. Manager
Bowler, Mrs M. Cleaner II. Assistant: teas and functions
De Witt, D.M. Senior Artisan. Maintenance
Boonzaaier, I. Groundsman II
HARRY MOLTENO LIBRARY (RRLC)
Jagger, B.W. B.A. PG Dip. Lis. Librarian
Ovens, Dr C.S.H. Ph.D.dnf.Sc.) Dip.Datametrics. (contract librarian)
NBI WEBSITES (AMWS)
Reynolds, Ms P.Y. M.A.(Inf.Sc.), B.Proc., Dip.Datametrics, NBI Web Site Manager
262
PUBLICATIONS BY THE STAFF
1 April 2001-31 March 2002
Bothalia 32,2 (2002)
ADAMS, T. & NOTTEN, A. 2001-08. Veltheimia hracteata Harv. ex Bak.
(Hyacinthaceae). Internet 3 pp.
http://www.plantzafrica.com/planttuv/veltheimbract.htm.
ARCHER, C. 2001 , JodreUia fistulosa, a new generic record for the Flora
of southern Africa region (Asphodelaceae). Bothalia 31: 39, 40.
ARCHER. C. 2002. News from South Africa. Report back from Cape
Town internship. SABONET News 7: 68.
ARCHER, C. & KATIVU, S. 2001. The correct name in Chlorophytum
for Anthericum longistylum. Bothalia 31: 40, 41.
ARCHER. R.H. CARTER. S. ’& CONDY, G, 2001, Euphorbia venteri.
Flowering Plants of Africa 57: t. 2176, 86-90.
ARCHER, R.H., SNIJMAN, D.A. & BRUMMITT, R.K. 2001. (1478)
Proposal to conserve the name Boophone Herbert with that
spelling (Amaryllidaceae) Taxon 50: 569-571.
ARNOLD, T.H., PRENTICE, C.A., HAWKER, L.C., SNYMAN, E.E.,
TOMALIN, M„ CROUCH, N.R. & POTTAS-BIRCHER, C.
2002. Medicinal and magical plants of southern Africa: an
annotated checklist. Strelitzia 13. National Botanical Institute,
Pretoria,
AUBREY, A. 2001-04. Clivia gardenii Hook. (Amaryllidaceae). Inter-
net 2 pp. http://www.plantzafrica.com/plantcd/cliviagarden.htm,
AUBREY. A. 2001-05. Dombeya autumnalis l.Verd. (Sterculiaceae). Inter-
net 2 pp. http://www.plantzafrica.com/plantcd/dombeyaut.htm.
AUBREY, A. 2001-06. Tetradenia riparia (Hochst.) Codd (La-
miaceae), Internet 2 pp,
http://www.plantzafrica.com/planttuv/tetradenripar,htm.
AUBREY, A. 2001-07. Aloe ferox Mill. (Aloaceae). Internet 2 pp.
http://www.plantzafrica.com/plantab/aloeferox.htm.
AUBREY, A. 2001-08. Clivia miniata (Lindl.) Regel (Amaryllida-
ceae). Internet 3 pp.
http://www.plantzafrica.com/plantcd/cliviaminiata.htm.
AUBREY, A. 2001-09. Acacia caffra (Thunb.) Willd. (Fabaceae:
Mimosoideae). Internet 2 pp.
http://www.plantzafrica.com/plantab/acaciacaffra.htm,
AUBREY, A. 2001-09. Dombeya rotundifolia Hochst. (Sterculiaceae).
Internet 2 pp.
http://www.plantzafrica.com/plantcd/dombeyrotund.htm.
AUBREY, A. 2001-11. Zantedeschia aethiopica (L.) Spreng. (Ara-
ceae). Internet 3 pp.
http://www.plantzafrica.com/plantwxyz/zantedeschaeth.htm.
AUBREY, A. 2001-12. Plumbago auriculata Lam. (Plumbaginaceae)
Internet 4 pp.
http://www.plantzafrica,com/plantnop/pluinbago.htm,
AUBREY, A. 2002-01. Acacia karroo Hayne. (Fabaceae: Mimosoi-
deae). Internet 4 pp.
http://www.plantzafrica.com/plantab/acaciakar.htm.
BEYERS, J.B.P. 2001a. A new species of Oedera from Western Cape,
South Africa (Asteraceae). Bothalia 31: 41, 42.
BEYERS, J.B.P. 2001b. The Cape genus Lachnaea (Thymelaeaceae): a
monograph. Strelitzia 1 1 . National Botanical Institute, Pretoria.
BREDENKAMP, C.L. & VAN WYK, A.E. 2001a. Leaf anatomy of the
genus Passerina (Thymelaeaceae): taxonomic and ecological
significance, Bothalia 3 1 : 53-70.
BREDENKAMP, C.L. & VAN WYK, A.E. 2001b. Taxonomic signifi-
cance of inflorescences, floral morphology and anatomy in
Passerina (Thymelaeaceae). Bothalia 31: 213-236.
BROWN, B, & NOTTEN, A. 2001-08. Hermannia saccifera (Turcz.)
K.Schum. (Sterculiaceae). Internet 2 pp.
http://www.plantzafrica.com/planthij/hermansaccifer.htm.
BROWN, L. & VAN DER WALT, L. 2001. In the garden. White
Christmas. HL December 2001: 89-93.
BROWN, N.A.C., KOTZE, G.J, & BOTHA, PA. 2001 . Grow proteas,
edn 2. Kirstenbosch Gardening Series, National Botanical
Institute, Cape Town.
BRUMMEIT, R.K.. CASTROVIEJO, S., CHIKUNl, A.C., ORCHARD,
A.E., SMITH, G.F. & WAGNER, W.L, 2001. The Species
Plantarurn Project, an international collaborative initiative for
higher plant taxonomy. Taxon 50: 1217-1230.
BURCOYNE, P.M. 2001 . Fascinating Frilhia — notes on a summer rain-
fall window-plant. Cactus and Succulent Journal (US) 73: 12-19.
CARf)I,US, B. 2001-05. Olinia cinarginataThunb. (Oliniaceae). Inter-
net 2 pp. http://www.plantzafrica.com/plantnop/oliniaemarg.htm.
CHESSELET, P. 2001, A new combination in Aniiminia (Mesem-
bryanthcrnaceae). /1/w 38: 17.
CHESSELET, P. & SMITH. G.F. 2001 . Aniiminia aristidata comb, nov.
(Mesembryanthemaceae Fenzl); a heterophyllous succulent
from Cape Town, South Africa. Bradleya 19: 121-123.
CHESSELET, P, SMITH, G.F, & VAN WYK. A.E. 2001. A new trib-
al classification for the Mesembryanthemaceae Fenzl based on
characters of the floral nectary. Aloe 38: 25-28.
CHIMPHANGO. S.B.M., MUSIL, C.F. & DAKORA, ED. 2001.
Growth and symbiotic response of cowpea (Vigna unguiculata)
to elevated levels ofUV-B radiation. Proceedings of 13th Inter-
national Congress on Nitrogen Fixation, Hamilton, Ontario,
Canada. 2-7 July, 2001,
CONDY, G. & ROURKE, J.P. 2001. Concise dictionary of South
African botanical artists. In M. Arnold, South African botanical
art — peeling back the petals'. 185-207. Fernwood Press, Vlae-
berg. Cape Town.
CRAIB. C. & CONDY, G. 2001a. Pelargonium nephrophvllum (Gera-
niaceae). Curtis’s Botanical Magazine 18: t. 422, 149-152.
CRAIB, C. & CONDY, G. 2001b. Geophytic pelargoniums. Field and
cultivation studies o/Pelargonium .section Hoarea. Umdaus Press.
Pretoria.
CROUCH, N.R. 2001. The role of horticulture in threatened species
conservation. FSA ethnomedicinal taxa in focus. SABONET
News 6: 185-188.
CROUCH, N.R., LOTTER. M„ KRYNAUW, S. & POTTAS-BIRCHER,
C. 2001 (2000). Siphonochilus aethiopicus (Zingiberaceae), the
prized indungulu of the Zulu — an overview. Herbertia 55:
115-129.
CROUCH, N.R. & SMITH, G.F. 2001. Ga.steria batesiana faces dam
construction threat, again. British Cactus and Succulent
Journal 19: 21-24.
CROUCH, TE. & CROUCH, N.R. 2001. Insect infestations on leaves
of the ethnomedicinal pepper-bark tree, Warburgia salutaris
(Canellaceae). Durban Museum Novitates 26: 1-5.
CUPIDO, C.N, & CONRAD, F. 2001 . Lobelias in South Africa. Veld &
Flora 87: 118. 119.
DOUWES, E., CROUCH. N.R. & SYMMONDS, R. 2001a. Blue
squill in the red. Scilla natalensis as a conservation charge.
PlantLife 24: 14-19.
DOUWES, E., CROUCH, N.R. & SYMMONDS, R. 2001b. The Moth-
fruit, Acridocarpus natalltius, in the service of gardeners, tradi-
tional healers and the two-pip policeman. PlantLife 25: 30-33.
DREWES, S.E., CROUCH, N.R., MASHIMBYE, M.J., DE LEEUW,
B.M. & HORN, M.M. 2001. A phytochemical basis for the
potential use of Warburgia salutaris (pepper-bark tree) leaves
in place of bark. South African Journal of Science 97: 383-386.
DUNCAN, G.D. 2001a. Chasmanthe. Notes on their cultivation and
propagation, with particular reference to Chasmanthe bicolor.
Veld & Flora 87: 108-111.
DUNCAN, G.D. 2001b. Review: Geophytic pelargoniums, by Charles
Craib. Artist: Gillian Condy, 2001. Veld Flora 87: 191.
DUNCAN, G.D. 2001c. Spectacular, rewarding Scado.xus. Veld & Flora
87: 60-63.
DUNCAN, G.D. 2001d. Magnificent flowering bulbs at Kirstenbosch.
Cape Tourist Guide’s Association'. 4, 5. (August newsletter).
Email publication.
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,
-' « M4 ' -'’ f
' '« h ^ '•» *• *
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'♦ 4^1
i* •>,■». # *^- '"•.jsjti*. ■ I
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.• • >’ -J ,:K:. - ^ - , Ji
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. t .'.V ;,, i]fe.. »t. V, '• ' «.T,
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■ >1. *('<-»•♦ ♦•■ ,«• «,. .1. • . lliiM|l ’
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Bothalia 32.2: 269-278 (2002)
Guide for authors to Bothalia
This guide is updated when necessary and includes an
index. Important points and latest additions appear in
bold type.
Bothalia is named in honour of General Louis Botha,
first Premier and Minister of Agriculture of the Union of
South Africa. This house journal of the National
Botanical Institute, Pretoria, is devoted to the furtherance
of botanical science. The main fields covered are taxon-
omy, ecology, anatomy and cytology. Two parts of the
journal and an index to contents, authors and subjects are
published annually.
1 Editorial policy
1.1 Bothalia welcomes original papers dealing with
flora and vegetation of southern Africa and related sub-
jects. Full-length papers and short notes, as well as book
reviews and obituaries of botanists, are accepted. The edi-
tor should be notified that an article is part of a series of
MSS; please submit a list of the parts of a series; all parts
should preferably be published in one journal.
1.2 Submission of a manuscript to Bothalia implies
that it has not been published previously and is not being
considered for publication elsewhere.
1.3 Authors whose first language is not English are
requested to have their MS edited by an English speaker
before submission.
1.4 Articles are assessed by referees, both local and
overseas. Authors are welcome to suggest possible refer-
ees to judge their work. Authors are responsible for the
factual coiTectness of their contributions. Bothalia main-
tains an editorial board (see title page) to ensure that
international standards are upheld.
1.5 Page charges: as stated in our notification includ-
ed in volume 23,1 (May 1993), MSS submitted for pub-
lication in Bothalia are subject to payment of page
charges of R1 25,00 per printed page, VAT included. The
following are exempt from these charges: 1, NBI mem-
bers; 2, persons/institutions who have been granted
exemption by the Executive Committee of the NBI; 3,
authors of contributions requested by the Editor; 4. con-
tributors to the column "FSA contributions’. The Editor's
decision on the number of pages is final. An invoice will
be sent to the author, who must arrange for payment as
soon as possible to NBI, Publications Section, Private
Bag XlOl, Pretoria 0001.
1.6 Deadline dates for submission of MS: for possi-
ble inclusion of the MS for the May issue — August of
the previous year, and for the October issue — March
of the same year.
2 Requirements for a manuscript
2.1 The original manuscript should be typed on one
side of A4-size paper, double line spacing throughout
(including abstract, tables, captions to figures, literature
references, etc.) and have a margin of at least 30 mm all
round. Three photocopies (all pages photocopied on both
sides of the paper, including figures, to reduce weight for
postage) of all items, including text, line drawings, tables
and lists should be submitted, and the author should
retain a complete set of copies. Three photographs (or
high quality photocopies) of each photograph/photo-
graph mosaic should be submitted for review puiposes.
The electronic version should be submitted with the final
(accepted) manuscript (see 3).
2.2 Papers should conform to the general style and lay-
out of recent issues of Bothalia (from volume 26 onwards).
2.3 Material should be presented in the following
sequence; Title page with title, name(s) of author(s), key-
words, abstract (and information that should be placed in
a footnote on the title page, such as address(es) of
author! s) and mention of granting agencies).
2.4 The sequence continues with Introduction and
aims. Contents (see 8), Material and methods. Results,
Interpretation (Discussion), Specimens examined (in revi-
sions and monographs). Acknowledgements, References,
Index of names (recommended for revisions dealing with
more than about 15 species). Tables, Captions for figures
and figures. In the case of short notes, obituaries and book
reviews, keywords and abstract are superfluous.
2.5 All pages must be numbered consecutively begin-
ning with the title page to those with references, tables,
captions for figures and figures.
2.6 Special characters; use your own word or code that
is unique and self-explanatory, enclosed between
ANGEE BRACKETS, e.g. <mu>m for pm. Please sup-
ply us with a list of the codes.
2.7 Use a non-breaking space (in MS Word — Ctrl,
shift, space) to keep two elements together on the same
line, e.g. 3 500.
2.8 DO NOT JUSTIEY LINES.
2.9 Do not break words, except hyphenated words.
2.10 A hyphen is designated as one dash, with no space
between the letter and the dash. e.g. ovate-lanceolate.
See also 17.6.
2.11 An N-dash is typed in MS Word code (alt + 0150)
or as three hyphens with no space between the letter and
the hyphen, e.g. 2 5 mm (typeset, it looks like this,
2-5 mm).
2.12 An M-dash is typed in MS Word code (alt + 0151 )
or as two hyphens with no space between the letter and
the hyphen, e.g. computers- -what a blessing! (typeset, it
looks like this: computers — what).
2.13 Do not use a double space anywhere between
words, after commas, full stops, colons, semicolons or
exclamation marks.
270
Bothalia 32,2 (2002)
2.14 Use lower case x as times sign, with one space on
either side of the x, e.g. 2x3 mm.
2.15 Use single (not double) opening and closing
quotes, e.g. the so-called ‘stiffy’ refers to a rigid diskette.
2. 1 6 Keys — put only three leader dots before number of
taxon (with one space before and after each dot), regard-
less of how far or near the word is from the right margin,
e.g. ... 1. /?. ovata (see 13.18).
3 Requirements for diskettes/stiffies
(text to be submitted only with final/accepted version)
3. 1 USE NORMAL STYLE ONLY.
3.2 Electronic files can be provided on 1.4 MB stiffie
disks, Iomega zip diskettes, 640 MB optical disks for
Apple Mac or on CD.
3.3 Data must be IBM compatible and written in
ASCII, or in Word 97 for Windows 95/98. An rtf file is
preferable because it retains the formatting.
3.4 All lines, headings, keys, etc., should start flush at
the margin, therefore NO INDENTATIONS, EOOT-
NOTES, TABS OR STYLES of any kind.
3.5 In MS Word, italics and bold should be used where
necessary.
3.6 Paragraphs and headings are delineated by a car-
riage return (ENTER) but no indentation.
3.7 Graphics i.e. drawings, graphs or photographs:
submit in a separate file, must not be included in the text.
3.8 Image files with a bigger file size than 1MB can-
not be e-mailed as the NBI has a 1MB limitation on the
network’s firewall at Head Office.
3.9 If any image file was originated in CorelDraw ver-
sions 3-9, please provide the image file as a CDR file (please
include fonts). The conversion to TIF or other file extensions
will be accommodated by the NBI (see 12.2, 12.3).
3.10 If extensive changes to image files are proposed by
the editor, the author will be contacted and the specific
image file will have to be re-submitted after the indicat-
ed corrections have been implemented.
3.11 Tracked changes must not be included when
submitting a MS on diskette or electronically.
4 Author(s)
When there are several authors, the covering letter
should indicate clearly which of them is responsible for
correspondence and, if possible, telephonically available
while the article is being processed. The contact address
and telephone number should be mentioned if they differ
from those given on the letterhead.
5 Title
The title should be as concise and as informative as
possible. In articles dealing with taxonomy or closely
related subjects, the family of the taxon under discussion
(see also 13.2) should be mentioned in brackets but
author citations should be omitted from plant names (see
also 13.6).
6 Keywords
Up to 10 keywords (or index terms) should be pro-
vided in English in alphabetical sequence. The following
points should be borne in mind when selecting key-
words:
6. 1 Keywords should be unambiguous, internationally
acceptable words and not recently coined little-known
words.
6.2 They should be in a noun form and verbs should be
avoided.
6.3 They should not consist of an adjective alone;
adjectives should be combined with nouns.
6.4 They should not contain prepositions.
6.5 The singular form should be used for processes and
properties, e.g. evaporation.
6.6 The plural form should be used for physical ob-
jects, e.g. augers.
6.7 Location (province and/or country); taxa (species,
genus, family) and vegetation type (community, veld
type, biome) should be used as keywords.
6.8 Keywords should be selected hierarchically where
possible, e.g. both family and species should be included.
6.9 They should include terms used in the title.
6.10 They should answer the following questions:
6.10.1 What is the active concept in the document
(activity, operation or process).
6.10.2 What is the passive concept or object of the
active process (item on which the activity, operation or
process takes place).
6. 10.3 What is the means of accomplishment or how is
the active concept achieved (technique, method, appara-
tus, operation or process).
6.10.4 What is the environment in which the active
concept takes place (medium, location).
6.10.5 What are the independent (controlled) and de-
pendent variables?
6.11 Questions 6.I0.I to 6.10.3 should preferably also
be answered in the title.
7 Abstract
7.1 Abstracts of no more than 200 words should be
provided. Abstracts are of great importance and should
convey the essence of the article.
Bothalia 32.2 (2002)
271
7.2 They should refer to the geographical area con-
cerned and, in taxonomic articles, mention the number of
taxa treated. They should not contain information not
appearing in the article.
7.3 In articles dealing with taxonomy or closely related
subjects all taxa from the rank of genus downwaids should
be accompanied by their author citations (see also 13.6).
7.4 Names of new taxa and new combinations should
not be italicized but put in bold. If the article deals with
too many taxa, only the important ones should be men-
tioned.
8 Table of contents
A table of contents should be given for all articles
longer than about 60 typed pages, unless they follow the
strict format of a taxonomic revision.
9 Acknowledgements
Acknowledgements should be kept to the minimum
compatible with the requirements of courtesy. Please
give all the initials of the person! s) you are thanking.
10 Literature references
In text
10.1 Literature references in the text should be cited as
follows: ‘Jones & Smith (1986) stated...’, or ‘...(Jones &
Smith 1986)’ or (Ellis 1988: 67) when giving a reference
simply as authority for a statement. For treatment of lit-
erature references in taxonomic papers see 14.
10.2 When more than two authors are involved in the
paper, use the name of the first author followed by et al.
10.3 When referring to more than one literature refer-
ence, they should be arranged chronologically and sepa-
rated by a semicolon, e.g. (Nixon 1940; Davis 1976;
Anon. 1981, 1984).
10.4 Titles of books and names of journals should
preferably not be mentioned in the text. If there is good
reason for doing so, they should be treated as described
in 10.12 & 10.13.
10.5 Personal communications are given only in the
text, not in the list of references. Please add the person’s
full initials to identify the person more positively, e.g. C.
Boucher pers. comm.
In References at end of article
10.6 References of the same author are arranged in
chronological sequence.
10.7 Where two or more references by the same author
are listed in succession, the author’s name is repeated
with every reference, except in an obituary, where the
name of the deceased in the list of publications (not in
the references) is replaced by an N-dash.
10.8 All publications referred to in the text, including
those mentioned in full in the treatment of correct names
in taxonomic papers, but no others, and no personal com-
munications, are listed at the end of the manuscript under
the heading References.
10.9 The references are airanged alphabetically accord-
ing to authors and chronologically under each author, with
a, b, c, etc. added to the year, if the author has published
more than one work in a year. This sequence is retained
when used in the text, in'espective of the chronology.
10.10 If an author has published both on his own and as
a senior author with others, the solo publications are list-
ed first and after that, in strict alphabetical sequence,
those published with one or more other authors.
10.1 1 Author names are typed in capitals.
10.12 Titles of journals and of books are written out in
full and are italicized as follows: Transactions of the
Linnean Society of London 5: 171-217, or Biology and
ecology of weeds: 24.
10.13 Titles of books should be given as in Taxonomic
literature, edn 2 by Stafleu & Cowan and names of jour-
nals as in the latest edition of World list of scientific peri-
odicals.
10.14 Examples of references:
Collective book or Flora
BROWN. N.E. 1909. Asclepiadaceae. In W.T. Thiselton-Dyer. Flora
copensis 6.2: 518-1036. Reeve. London.
CUNNINGHAM. A.B. 1994. Combining skills; participatory
approaches in biodiversity conservation. In B.J. Huntley. Botanical
diversity in southern Africa. Strelitzia 1: 149-167. National Botanical
Institute. Pretoria.
Book
DU TOIT, A.L. 1966. Geology of South Africa, edn 3. S.M. Haughton
(ed.). Oliver & Boyd. London.
HUrrCHINSON. J. \946. A botanist in southeni Africa. Gawthom. London.
Journal
DAVIS. G. 1988. Description of a proteoid-restioid stand in Mesic
Mountain Fynbos of the southwestern Cape and some aspects of its
ecology. Bothalia 18: 279-287.
SMOOK. L. & GIBBS RUSSELL. G.E. 1985. Poaceae. Memoirs of the
Botanical Sitn'ey of South Africa No. 51: 45-70.
STEBBINS. G.L. Jr 1952. Aridity as a stimulus to plant evolution.
American Naturalist 86: 35^4.
In press, in preparation
TAYLOR. H.C. in press. A reconnaissance of the vegetation of
Rooiberg State Forest. Technical Bulletin, Department of Forestry.
VOGEL, J.C. 1982. The age of the the Kuiseb river silt terrace at
Homeb. Palaeoecology of Africa 15. In press.
WEISSER. P.J.. GARLAND. J.F. & DREWS. B.K. in prep. Dune
advancement 1937-1977 and preliminary vegetation succession
chronology at Mlalazi Nature Reserve. Natal. South Africa. Bothalia.
Thesis
KRUGER. F.J. 1974. The physiography and plant communities of the
Jakkalsrivier Catchment. M.Sc. (Forestry) thesis. University of Stellenbosch.
272
Bothalia 32.2 (2002)
MUNDAY, J. 1980. The genus Monechma Hochst. (Acanthaceae tribe
Justiciae) in southern Africa. M.Sc. thesis. University of the
Witwatersrand. Johannesburg.
Miscellaneous paper, report, unpublished article, techni-
cal note, congress proceedings
ANON, no date. Eetbare plante van die Wolkberg. Botanical Research
Unit. Grahamstown. Unpublished.
BAWDEN. M.G. & CARROL. D.M. 1968. The land resources of
Lesotho. Land Resources Study No, 3, Land Resources Division,
Directorate of Overseas Surveys, Tolwoith.
BOUCHER. C. 1981. Contributions of the Botanical Research
Institute. In A.E.F. Heydorn, Proceedings of workshop research in
Cape estuaries: 105-107. National Research Institute for Oceanology,
CSIR. Stellenbosch.
NATIONAL BUILDING RESEARCH INSTITUTE 1959. Report of
the committee on the protection of building timbers in South Africa
against termites, woodboring beetles and fungi, edn 2. CSIR Research
Report No. 169.
1 1 Tables (also digital submissions)
11.1 Each table should be presented on a separate sheet
and be assigned an Arabic numeral, i.e. the first table
mentioned in the text is marked ‘Table 1’.
1 1 .2 In the captions of tables the word ‘TABLE" is writ-
ten in capital letters. See recent numbers of Bothalia for
the format required.
11.3 Avoid vertical lines, if at all possible. Tables can
often be reduced in width by interchanging primary hor-
izontal and vertical heads.
12 Figures (also digital submissions)
12.1 Eigures should be planned to fit, after reduction,
into a width of either 80, 118 or 165 mm, with a maxi-
mum vertical length of 230 mm. Allow space for the cap-
tion in the case of figures that will occupy a whole page.
12.2 Line drawings, including graphs and diagrams,
should be the same size as it will appear in the journal,
if appropriate, and should be in Jet-black Indian ink,
preferably on fine Schoellers Hammer Parole or similar
paper, 200 gsm, or tracing film. Lines should be bold
enough and letters/ symbols large enough to stand
reduction. If submitted electronically, provide each
drawing as a separate TIF, BMP or JPG file at 600 dots
per inch (dpi) and a hard copy of the fignre.
12.3 Photographs should be of excellent quality on
glos.sy paper with clear detail, moderate contrast and
clear lettering, and they should be the same size as
required in the Journal. If submitted electronically, pro-
vide as a TIF, BMP or JPG file at 300 dpi and not as a
doc file. Include a hard copy of good quality.
12.4 Photograph mosaics should be submitted com-
plete, the component photographs mounted neatly on a
white Hexible card base (can be curved around drum of
scanner) leaving a narrow gap of uniform width (2 mm)
between each print. Note that grouping photographs of
markedly divergent contrast results in poor reproduc-
tions.
12.5 Lettering on photograph mosaics, in capital letters,
should be put on a small white disk ± 7 mm in diameter,
if the background is dark, and placed in the lower left
hand corner of the relevant photo.
12.6 If several illustrations are treated as components of
a single composite figure they should be designated by
capital letters.
12.7 Note that the word ‘Figure’ should be written out
in full, both in the text and the captions and should begin
with a capital ‘F’ (but see 14.7 for taxonomic papers).
12.8 In the text the figure reference is then written as in
the following example: ‘The stamens (Figure 4A, B) ai'e..."
12.9 In captions, ‘FIGURE’ is written in capital letters.
Magnification of figures should be given for the size as
submitted.
12.10 Scale bars or scale lines should be used on fig-
ures.
12.11 In figures accompanying taxonomic papers, vouch-
er specimens should be given in the relevant caption.
12.12 Figures are numbered consecutively with Arabic
numerals in the order they are referred to in the text.
These numbers, as well as the author’s name and an indi-
cation of the top of the figure, must be written in soft
pencil on the back of all figures.
12.13 Captions of figures must not be pasted under the
photograph or drawing and must also not be included in
any electronic version.
12.14 Authors should indicate in pencil in the text where
they would like the figures to appear.
12.15 Authors wishing to have the originals of figures
returned must inform the editor in the original covering let-
ter and must mark each original ‘To be returned to author’.
12.16 Authors wishing to use illustrations already pub-
lished elsewhere must obtain written permission before
submitting the manuscript and inform the editor of this fact.
12.17 Captions for figures should be collected together
and typed at the end of the MS and headed Captions for
figures.
12.18 It is strongly recommended that taxonomic arti-
cles include dot maps as figures to show the distribution
of taxa. The dots used must be large enough to stand
reduction to 80 mm (recommended size; 5 mm diameter).
1 2. 1 9 Blank distribution maps of southern Africa, Afri-
ca and the world are available from the Bookshop, NBI
Pretoria.
12.20 A dot map PC programme for distribution of
taxa in South Africa, called MAPPIT is available for pur-
chase from the Data Section, National Botanical
Institute, Pretoria.
12.21 AreView GIS maps are acceptable. The lay-
out representing all the appropriate themes (includ-
ing gridlines) should be exported as a PostScript New
(EPS) file at 600 dpi.
Bothalia 32.2 (2002)
273
12.22 Colour figures are permitted only if a) it will
clarify the article and b) the cost of reproduction and
printing is borne by the author.
1 3 Text
13.1 As a rule, authors should use the plant names (but
not of all authors of plant names — see 13.6) as listed in
PRECIS (National Herbarium PREtoria Computerised
Information System).
13.2 Names of genera and infrageneric taxa are usually
italicized, with the author citation (where relevant; see
13.6) not italicized. Exceptions include names of new
taxa in the abstract, correct names given in the synopsis
or in paragraphs on species excluded from a given
supraspecific group in taxonomic articles; in checklists
and in indices, where the position is reversed, coiTect
names are not italicized and synonyms are italicized.
13.3 Names above generic level are not italicized.
13.4 In articles dealing with taxonomy, the complete
scientific name of a plant (with author citation) should be
given at the first mention in the text. The generic name
should be abbreviated to the initial thereafter, except
where intervening references to other genera with the
same initial could cause confusion (see 16.6).
13.5 In normal text, Latin words are italicized, but in the
synopsis of a species, Latin words such as nom. nud. and
et al. are not italicized (see 14.3, 16.4, 17.9).
13.6 In accordance with Garnock-Jones & Webb ( 1996)
in Taxon 45: 285, 286. authors of plant names are not to
be added to plant names except in taxonomic papers.
Names of authors of plant names should agree with the
list published by the Royal Botanic Gardens, Kew, enti-
tled, Authors of plant names, edited by R.K. Brummitt &
C.E. Powell (1992).
13.7 Modern authors not included in the list should use
their full name and initials when publishing new plant
names. Other author names not in the list should be in
agreement with the recommendations of the Code.
13.8 Names of authors of publications are written out in
full, without initials, except in the synonymy in taxo-
nomic articles where they are treated like names of
authors of plant names.
13.9 Names of plant collectors are italicized whenever
they are linked to the number of a specimen. The collec-
tion number is also italicized, e.g. Acocks 14407.
13.10 Surnames beginning with ‘De', 'Du' or ‘Van’
begin with a capital letter unless preceded by an initial.
13.11 For measurements use only units of the Inter-
national System of Units (SI). In taxonomic papers only
mm and m, should be used; in ecological papers cm or m
should be used.
13.12 The use of ‘±’ is preferred to c. or ca (see 17.7).
13.13 Numbers ‘one’ to ‘nine’ are spelt out in nomial text,
and from 10 onwards they are written in Arabic numerals.
13.14 In descriptions of plants, numerals are used
throughout. Write 2.0^. 5 (not 2^.5). When counting
members write 2 or 3 (not 2-3), but 2^.
13.15 Abbreviations should be used sparingly but con-
sistently. No full stops are placed after abbreviations end-
ing with the last letter of the full word (e.g. edition = edn;
editor = ed.); after units of measure; after compass direc-
tions; after herbarium designations; after countries, e.g.
USA and after well-known institutions, e.g. CSIR.
13.16 Apart from multi-access keys, indented keys
should be used with couplets numbered la- lb, 2a-2b,
etc. (without full stops thereafter).
13.17 Keys consisting of a single couplet have no num-
bering.
13.18 Manuscripts of keys should be presented as in the
following example:
la Leaves closely an'anged on an elongated stem; a sub-
merged aquatic with only the capitula exserted ... lb. L.
setaceum van piiniilian
lb Leaves in basal rosettes; stems suppressed; small
marsh plants, ruderals or rarely aquatics:
2a Annuals, small, fast-growing pioneers, dying when
the habitat dries up; capitula without coarse white setae;
receptacles cylindrical;
3a Anthers white .. .2. E. cinerewn
3b Anthers black .. .2. E. nigrum
2b Perennials, more robust plants; capitula .sparsely to
densely covered with short setae:
13.19 Herbarium voucher specimens should be referred
to wherever possible, not only in taxonomic articles.
14 Species treatment in taxonomic papers
14.1 The procedure to be followed is illustrated in the
example ( 17.9), which should be referred to, because not
all steps are described in full detail.
14.2 The correct name (bold, not italicized) is to be fol-
lowed by its author citation (italicized) and the full liter-
ature reference, with the name of the publication written
out in full (not italicized).
14.3 Thereafter all literature references, including those
of the synonyms, should only rellect author, page and
year of publication, e.g. C.E.Hubb. in Kew Bulletin 15:
307 (I960); Boris et ah; 14 (1966); Boris; 89 (1967);
Sims; t. 38 (1977); Sims: 67 (1980).
14.4 The description and the discussion should consist
of paragraphs commencing, where possible, with itali-
cized leader words such as flowering time, etymology,
diagnostic characters, distribution and habitat.
14.5 When more than one species of a given genus is dealt
with in a paper, the correct name of each species should be
274
Bothalia 32.2 (2002)
prefixed by a sequential number followed by a full stop.
Infraspecific taxa are marked with small letters, e.g. lb.,
12c., etc.
14.6 Names of authors are written as in 13.6, irrespec-
tive of whether the person in question is cited as the
author of a plant name or of a publication.
14.7 The word ‘figure’ is written as ‘fig.’, and ‘t.’ is used
for both ‘plate’ and ‘tablet’ (but see 12.7 for normal text).
14.8 Literature references providing good illustrations
of the species in question may be cited in a paragraph
commencing with the word Illustrations followed by a
colon. This paragraph is given after the last paragraph of
the synonymy, see 17.9.
14.9 When new combinations are made, the full litera-
ture reference must be given for the basionym, e.g.;
Antimima saturata (L.Boliis) H.E.K. Hartmann,
comb. nov.
Riischia saturata L. Bolus in Notes on Mesembrianthemum and
allied genera, part 2: 122 (1929). Mesemhryanthemum atrocinctum
N.E.Br.: 32 (1930). Type: Pillans BOL18952 (BOL, holo.!).
15 Citation of specimens
15.1 Type specimen in synopsis: the following should be
given (if available): country (if not in RSA), province, grid
reference (at least for new taxa), locality as given by origi-
nal collector, modern equivalent of collecting locality in
square brackets (if relevant, e.g. Port Natal [now Durban]),
quarter-degree square, date of collection (optional), collec-
tor’s name and collecting number (both italicized).
15.2 The abbreviation s.n. {sine luimero) is given after
the name of a collector who usually assigned numbers to
his collections but did not do so in the specimen in ques-
tion (see 15.11), or the herbarium number can then be
cited with no space between the herbarium and its num-
ber e.g. Marloth SAM691 (see 17.9). The herbaria in
which the relevant type(s) are housed are indicated by
means of the abbreviations given in the latest edition of
Index Herhariorum.
15.3 The holotype (holo.) and its location are mentioned
first, followed by a semicolon, the other herbaria are
arranged alphabetically, separated by commas.
15.4 Authors should indicate by means of an exclamation
mark (!) which of the types have been personally examined.
15.5 If only a photograph or microfiche was seen, write
as follows: Anon. 422 (X, holo.-BOL, photo.!).
15.6 Lectotypes or neotypes should be chosen for cor-
rect names without a holotype. It is not necessary to lec-
totypify synonyms.
15.7 When a lectotype or a neotype are newly chosen,
this should be indicated by using the phrase ‘here desig-
nated’ (see 17.9). If reference is made to a previously
.selected lectotype or neotype, the name of the designating
author and the literature reference should be given. In
cases where no type was cited, and none has subsequent-
ly been nominated, this may be stated as ‘not designated’.
15.8 In brief papers mentioning only a few species and a
few cited specimens the specimens .should be airanged
according to the grid reference system: Provinces/countries
(typed in capitals) should be cited in the following order:
Namibia, Botswana, Limpopo (previously Northern
Transvaal, Northern Province), North-West (previously
northeastern Cape and southwestern Transvaal), Gauteng
(previously PWV), Mpumalanga (previously Eastern
Transvaal), Free State (previously Orange Free State),
Swaziland, KwaZulu-Natal (previously Natal), Fesotho, and
Northern Cape, Western Cape and Eastern Cape (Figure 1 ).
1 5.9 Grid references should be cited in numerical sequence.
15.10 Focality records for specimens should preferably
be given to within a quarter-degree square. Records from
the same one-degree square are given in alphabetical
order, i.e (-AC) precedes (-AD), etc. Records from the
same quarter-degree square are ari'anged alphabetically
according to the collectors’ names; the quarter-degree
references must be repeated for each specimen cited.
15.11 The relevant international code of the herbaiia in
which a collection was seen should be given in brackets
after the collection number; the codes are separated by com-
mas. The following example will explain the procedure:
KWAZULU-NATAL. — 2731 (Louwsburg): 16 km E of Nongoma.
(-DD). Pelser 354 (BM. K. PRE); near Dwansrand. Van der Merwe
4789 (BOL. M). 2829 (Hamsmith): near Groothoek, (-AB). Smith 234:
Koffiefontein. (-AB). Taylor 720 (PRE); Cathedral Peak Forest
Station. (-CC), Marriot s.n. (KMG); Wilgeifontein. Rota 426. Grid ref.
unknown: Sterkstroom. Stiydom 12 (NBG).
15.12 For records from outside southern Africa authors
should use degree squares without names, e.g.:
KENYA. — 0136: Nairobi plains beyond race course. Napier 485.
15.13 Monographs and revisions: in the case of all
major works of this nature it is assumed that the author
has investigated the relevant material in all major
herbaria and that he has provided the specimens seen
with determinavit labels. It is assumed further that the
author has submitted distribution maps for all relevant
taxa and that the distribution has been described briefly
in words in the text. Under the heading ‘Vouchers’ no
more than five specimens should be cited, indicating
merely the collector and the collector's number (both
italicized). Specimens are alphabetically arranged ac-
cording to collector’s name. If more than one specimen
by the same collector is cited, they are arranged numeri-
cally and separated by a comma. The purpose of the cited
specimens is not to indicate distribution but to convey
the author’s concept of the taxon in question.
15.14 The herbaria in which the specimens are housed
are indicated by means of the abbreviation given in the lat-
est edition of Index Herhariorum. They are given between
brackets, arranged alphabetically and separated by com-
mas behind every specimen as in the following example:
Vouchers: Arnold 64 (PRE); Fisher 840 (NH. NU, PRE); Flanagan 831
(GRA, PRE), 840 (NH, PRE); Marloth 4926 (PRE, STE); Schelpe
6161. 6163. 6405 (BOL); Svhlechter 4451 (BM, BOL, GRA, K. PRE).
15.15 If long lists of specimens are given, they must be
listed together before Acknowledgements under the head-
ing Specimens examined. They are airanged alphabetically
by the collector’s name and then numerically for each col-
lector. The species is indicated in brackets by the number
that was assigned to it in the text and any infraspecific taxa
Bothalia 32.2 (2002)
275
by a small letter. If more than one genus is dealt with in a
given mlicle, the first species of the first genus mentioned
is indicated as 1.1. This is followed by the international
herbaiium designation. Note that the name of the collector
and the collection number are italicized:
Acocks 12497 (2.1b) BM. K. PRE; 14724 (1.13a) BOL. K. P. Archer
1507 a A) BM. G. Burchell 2847 (2.8c) MB. K. Bunnan 2401 (3.3)
MO. S. B.L. Burn 789 (2.6) B. KMG. STE.
1 6 Synonyms
16. 1 In a monograph or a revision covering all of south-
ern Africa, all synonyms based on types of southern
African origin, or used in southern African literature,
should be included.
16.2 Illegitimate names are designated by nom. illeg.
after the reference, followed by non with the author and
date, if there is an earlier homonym.
16.3 Nomina nuda {noin. mid.) and invalidly published
names are excluded unless there is a special reason to
cite them, for example if they have been used in promi-
nent publications.
16.4 In normal text, Latin words are italicized, but in the
synopsis of a species Latin words such as nom. mid., et
al. are not italicized (see 13.5, 14.3, 17.9).
16.5 Synonyms should be arranged chronologically into
groups of nomenclatural synonyms, i.e. synonyms based
on the same type, and the groups should be an'anged
chronologically by basionyms, except for the basionym
of the coiTect name which is dealt with in the paragraph
directly after that of the connect name.
16.6 When a generic name is repeated in a given syn-
onymy it should be abbreviated to the initial, except
where intervening references to other genera with the
same initial could cause confusion (see 13.4).
1 7 Description and example of species treatment
17.1 Descriptions of all taxa of higher plants should,
where possible, follow the sequence: Habit; sexuality:
underground parts (if relevant). Indumentum (if it can be
easily described for the whole plant). Stems/branches. Bark.
Leaves: arrangement, petiole absent/present, pubescence;
blade: shape, size, apex, base, mai'gin; midrib: above/below,
texture, colour; petiole; stipules. Inflorescence: type, shape,
position; bracts/bracteoles, involucral bracts: inner, outer.
Flowers: shape, sex. Receptacle. Calyx. Corolla. Disc.
Androeciiim. Gynoeciiim. Fruit. Seeds. Flowering time. Chro-
mosome number (reference). Conseiration .stanis. Figure
(word written out in full) number.
17.2 Asa mle, shape should be given before measurements.
17.3 In general, if an organ has more than one of the parts
being described, use the plural, otherwise use the singular,
for example, petals of a flower but blade of a leaf.
17.4 Language must be as concise as possible, using
participles instead of verbs.
17.5 Dimension ranges should be cited as in 17.9.
17.6 Care must be exercised in the use of dashes and
hyphens. A /;yp/;cn is a short stroke joining two syllables of
a word. e.g. ovate-lanceolate or sea-green, with no space
between the letter and the stroke. An N-dash (en) is a
longer stroke commonly used instead of the word ‘to’
between numerals, ‘2-5 mm long’ (do not use it between
w'ords but rather use the word ‘to’, e.g. ‘ovate to lanceo-
late’; it is produced by typing three hyphens next to each
other, or in MS Word the code is alt -i- 0150. An M-dash
iem) is a stroke longer than an N-dash and is used various-
ly, e.g. in front of a subspecific epithet instead of the full
species name; it is produced by typing two hyphens next to
one another, or in MS Word the code is alt -i- 015 1 .
1 7.7 The use of ‘±‘ is prefemed to c. or ca when describ-
ing shape, measurements, dimensions, etc. (see 13.12).
17.8 The decimal point replaces the comma in all units
of measurement, e.g. leaves 1.0-1. 5 mm long.
17.9 Example:
1. Engierophytum magalismontanum (Sond.)
T.D.Penn. The genera of Sapotaceae: 252 (1991). Type:
Gauteng, Magaliesberg, Zeyher 1849 (S. holo.-BOL,
photo.!).
Beqiiaertiodendron magalismontanum (Sond.) Heine & Hemsl.: 307
(1960); Codd: 72 (1964); Elsdon: 75 (1980).
Chrysophyllitm magalismontanum Sond.: 721 (1850); Harv.: 812
(1867); Engl.: 434 (1904); Bottmar: 34 (1919). Zeyherella magalis-
montana (Sond.) Aubrev. & Pellegr.: 105 ( 1958): Justin: 97 ( 1973).
Cluysophyllum argyrophyllum Hiem: 721 (1850); Engl.: 43 (1904).
Boivinella argyropliylla (Hiern) Aubrev. & Pellegr.: 37 ( 1958); Justin et
al.: 98 (1973). Types: Angola. Wehvitsch 4828 (BM!. lecto.. here des-
ignated; PRE!); Angola, Welwitsch s.ii. (BM!).
Chrysophytluin wilmsii Engl.: 4. t. 16 (1904); Masonet: 77 (1923);
Woodson: 244 ( 1937). Boivinella wilinsii (Engl.) Aubrev. & Pellegr.: 39
(1958); Justin: 99 (1973). Type: without locality and collector [B.
holo.t: K!. P!. lecto.. designated by Aubrev. & Pellegr.: 38 (1958).
PRE!. S!, W!, Z!].
Beqiiaertiodendron fruticosa De Wild.: 37 (1923). non Bonpl.: 590
(1823); D.Bakker: 167 (1929): H.Fr.: 302 (1938); Davy: 640 (1954);
Breytenbach: 117 (1959): Clausen: 720 (1968): Palmer: 34 (1969).
Type: Mpumalanga, Tzaneen Dist.. Granville in Herb. Pillans K48625
(K. holo.!; G!, P!. PRE!. S!).
B. fragruns auct. non Oldemann: Glover: 149. t. 19 (1915); Henkel:
226 ( 1934); Stapelton: 6 ( 1954).
Illustrations: Harv.: 812 (1867); Henkel: t. 84 (1934?); Codd: 73
(1964); Palmer: 35 (1969).
Woody perennial; main branches up to 0.4 m long,
erect or decumbent, grey woolly-felted, leafy. Leaves lin-
ear to oblanceolate. 3-10(-23) x 1.0-1. 5(^.0) mm,
obtuse, base broad, half-clasping. Heads heterogamous,
campanulate, 7-8 x 5 mm, solitary, sessile at tip of axil-
lary shoots; involucral bracts in 5 or 6 series, inner
exceeding flowers, tips subopaque, white, very acute.
Receptacle nearly smooth. Flowers ± 23-30, 7-1 1 male,
16-21 bisexual, yellow, tipped pink. Achenes ± 0.75 mm
long, elliptic. Pappus bristles very many, equalling corol-
la, scabridulous. Flowering time: September. Chromo-
some number: 2n = 22. Figure 23B.
276
Bothalia 32,2 (2002)
18 New taxa
18.1 The name of a new taxon must be accompanied by
at least a Latin diagnosis. Authors should not provide
full-length Latin descriptions unless they have the
required expertise in Latin at their disposal.
1 8.2 It is recommended that descriptions of new taxa be
accompanied by a good illustration (line drawing or pho-
tograph) and a distribution map.
18.3 Example:
109. Helichrysum jubilatum Hilliard, sp. nov., H.
alsinoidei DC. affinis, sed folds ellipticis (nec spatu-
latis), inflorescentiis compositis a folds non circumcinc-
tis, floribus femineis numero quasi dimidium hermaph-
roditorum aequantibus (nec capitulis homogamis vel
floribus femineis 1-3 tantum) distinguitur.
Herba annua e basi ramosa; caules erecti vel decum-
bentes, 100-250 mm longi, tenuiter albo-lanati, remote
foliati. Folia plerumque 8-30 x 5-15 mm, sub capitulis
minora, elliptica vel oblanceolata, obtusa vel acuta,
mucronata, basi semi-amplexicauli, utrinque cano-lana-
to- arachnoidea. Capitula heterogama, campanulata,
3. 5-4.0 X 2.5 mm, pro parte maxima in paniculas
cymosas terminates aggregata; capitula subterminalia
interdum solitaria vel 2 vel 3 ad apices ramulorum nudo-
rum ad 30 mm longorum. Bracteae involitcrales 5-seri-
atae, gradatae, exteriores pellucidae, pallide stramineae,
dorso lanatae, seriebus duabus interioribus subaequal-
ibus et flores quasi aequantibus, apicibus obtusis opacis
niveis vix radiantibus. Receptacidiim fere laeve. Flores ±
35-41. Aclienia 0.75 mm longa, pilis myxogenis praedi-
ta. Pappi setae multae, corollam aequantes, apicibus
scabridis, basibus non cohaerentibus.
TYPE. — Northern Cape, 2817 (Vioolsdrif): Richters-
veld, (-CC), ± 5 miles E of Eekkersing on road to Stink-
fontein, kloof in hill south of road, annual, disc whitish, 7-
11-1962, Nordenstam 1823 (S, holo.; E, NH, PRE).
19 New provinces of South Africa (Oct. 1996)
FIGURE 1. — 1, Western Cape; 2, Eastern Cape; 3, Northern Cape; 4,
Free .State (previously Orange Free State); 5. KwaZulu-Natal
(previously Natal); 6, North-West (previously tiortheastern Cape
and southwestern Transvaal); 7, Gauteng (previously PWV); 8,
Mpumalanga (previously Eastern Transvaal); 9. Limpopo (pre-
viously Northern Transvaal, Northern Provinee).
20 Proofs
Only page proofs are normally sent to authors. They
should be coirected in red ink and be returned to the edi-
tor as soon as possible. Do not add any new information.
21 Reprints
Authors receive 100 reprints free. If there is more than
one author, this number will have to be shared between
them.
22 Documents consulted
Guides to authors of the following publications were
made use of in the compilation of the present guide:
Annals of the Missouri Botanic Garden, Botanical
Journal of the Linnean Society, Flora of Australia.
Smithsonian Contributions to Botany, South African
Journal of Botany (including instructions to authors of
taxonomic papers). South African Journal of Science.
23 Address of editor
Manuscripts should be submitted to: The Editor,
Bothalia, National Botanical Institute, Private Bag XI 01,
Pretoria 000 1 .
24 FSA contributions
24.1 Eigures and text must conform to Bothalia format.
24.2 These articles will be considered as a full contri-
bution to the Flora of southern Africa and will be listed
as published in the 'Plan of Flora of southern Africa',
which appears in all issues of the ESA series.
INDEX
abbreviation, 13.4, 13.15, 15.2, 15.14, 16.6
abstract, 2.1, 2.3, 7, 13.2
acknowledgements, 9
address of
authors, 2.3, 4
editor, 23
alphabetical. 6, 10.9, 10.10, 15.3, 15.10, 15.13, 15.14, 15.15
Apple Mac, 3.2
Arabic numerals, 11.1, 12.12, 13.13
Arc View GIS maps, 12,21
ASCII. 3.3
author(s), 1.3, 2.1,4, 10.14, 12.14-12.16
address, 2.3, 4
citation, 5, 7.3, 13.2, 13.4, 14.2
first, 10.2
names, 2.3, 10.2, 10.7, 10.9, 10.11, 12.12, 13.7, 13.8, 14.3, 14.6,
15.7
names of plant names, 5, 7.3, 13.1. 13.2, 13.6, 13.7, 13.8, 14.6
senior. 10.10
BMP file, 12.2, 12.3
book reviews, 1.1, 2.4
books, 10.4, 10.12, 10.13, 10.14
Bothalia. 1 , 2.2, 1 1 .2, 24. 1
brief taxonomic articles, 15.8
BRUMMITT, R.K. & POWELL, C.E. (eds) 1992. Authors of plant
names. Royal Botanic Gardens, Kew, 13.6
c„ 13,12, 17.7
ca, 13.12, 17.7
Cape. 15.8, 18.3, 19
capitals, 11.2, 12.5, 12.6, 12.9, 15.8
Bothalia 32,2 (2002)
111
captions, 2,1, 2.4, 2.5, 11.2, 12.7. 12.9, 12.11,
CD. 3.2
CDR file, 3.9
checklist. 13.2
chromosome number, 17.1, 17.9
chronological sequence. 10.3. 10.6, 10.9, 16.5
citation
author. 5, 7.3. 13.2, 13.4. 14.2
of specimens. 15
cm. 13.11
collection
date. 15.1
number, 13.9, 15.1. 15.2. 15.11, 15.13. 15.
collective book, 10.14
collector. 13.9, 15.1. 15.2, 15.10. 15.13, 15.15
colon. 2.13. 14.8
colour figures. 12.22
comma, 2.13, 15.3, 15.11, 15.13. 15.14, 17.8
compass directions. 13.15
composite figure. 12.6
congress proceedings. 10.14
contents, 8
CorelDraw 3-9, 3.9
correspondence. 4
countries. 6.7, 15.8
deadline dates for submission of MS, 1.6
decimal point. 17.8
description and example of species treatment,
determinavit labels. 15.13
diagrams. 12.2
digital submissions of graphics, tables, 11. 12
discussion. 2.4, 14.4
diskette, 3. 3.2
distribution maps. 12.18, 12.19, 12.20. 15.13.
documents consulted. 22
dot maps. 12.18. 12.19. 12.20. 15.13. 18.2
double
line spacing, 2.1
space, 2.13, 2.16
dpi (dots per inch). 12.3. 12.21
drawing paper. 12.2
drawings. 3.7, 12.2
Eastern Transvaal, see Mpumalanga
edition, 13.15
editor. 13.15. 23
editorial
board. 1.4
policy. 1
electronic files. 3.2. 3.8-3.10, 12.2, 12.3
e-mail. 3.8
EPS file. 12.21
eta\.. 10.2. 13.5, 14.3, 17.9
example of
new taxa. 18.3
species treatment, 17.9
exclamation mark. 2.13. 15.4
family name. 5. 6.7
fig.. 14.7
figure(s), 12. 14.7, 17.1
colour. 12.22
digital submissions, 12
reduction of. 12.1. 12.2. 12.18
returned. 12.15
file
extensions, 3.9. 12.2. 12.3
BMP. 12.2. 12.3
CDR. 3.9
EPS. 12.21
JPG, 12.2, 12.3
RTE, 3.3
TIE, 3.9. 12.2. 12.3
firewall, 3.3
first author, 10.2
first language. 1 .3
flora, 10.14
Flora of southern Africa. 24
footnote, 2.3, 3.4
Free State (previously Orange Free State), 15.
FSA contributions, 24
12.13, 12.17 full stop, 2.13, 13.15, 13.16, 14.5
GARNOCK-JONES, P.J. & WEBB, C.J 1996. The requirement to
cite authors of plant names in botanical journals. Taxon 45: 285, 286.
13.6
Gauteng (previously PWV), 15.8, 17.9, 19
genera, 13.2
generic name. 13.3, 13.4, 16.6
geographical area. 7,2
granting agencies, 2.3
graphics, 3.7
digital submissions. 12
graphs. 3.7. 12.2
15 grid reference system. 15.1, 15.8. 15.9, 15.11
headings, 3.4, 3.6
sequence of. 2.3. 2.4
herbaria, 15.2, 15.3. 15.11, 15.13. 15.14
herbarium
code, 15.11
designations. 13.15, 15.15
numbers, 15.2
voucher specimens. 12.11. 13.19
here designated, 15.7, 17.9
holo.. 15.3, 15.5. 17.9, 18.3
holotype, 15.3, 15.6
homonym. 16.2
hyphenated words, 2.9
hyphen, 2.10-2.12. 17.6
17 IBM compatible, 3.4
illegitimate names (nom. illeg.), 16.2
illustrations. 12.2, 12.6. 12.16, 14.8, 17.9
previously published, 12.16
image files, 3.8-3.10
indentations. 3.4, 3.6
18.2 Index Herbarionini. 15.2, 15.14
index of names, 2.4
indices, 13.2
infrageneric taxa, 13.2
initials, 9, 10.5, 13.7
in prep.. 10.14
in preparation, 10.14
in press. 10.14
International
Code of Botanical Nomenclature, 13.7
System of Units (SI), 13.11
invalidly published names. 16.3
Iomega zip diskette. 3.2
italics, 7.4, 10.12. 13.2. 13,3, 13.5, 13.9. 14.2, 15.1. 15.13. 15.15
journals. 10.4, 10.12. 10.14
names of, 10.4. 10.13
JPG file. 12.2, 12.3
justify. 2.8
keys, 2.16, 3.4, 13.16, 13.17, 13.18
keywords, 2.3, 2.4, 6
KwaZulu-Natal (previously Natal). 15.8, 19
language, 1.3
Latin. 13.5. 16.4
descriptions, 18.1
layout. 2.2
lecto.. 15.6. 15.7. 17.9
lectotype, 15.6, 15.7, 17.9
lettering. 12.5
Limpopo (previously Northern Transvaal. Northern Province)
line
drawings, 2.1, 12.2, 18.2
spacing, 2,1
literature
references. 2.1. 10. 10.7, 14.2, 14.3, 14.8. 14.9
within synonymy. 14,8
localities outside southern Africa, 15,12
locality. 15.1, 15.10
location, 6.7
m, 13.11
magnification of figures, 12.3, 12.9
manuscript
language. 1.3. 17.4
requirements, 1.1. 1.2, 1.3, 2
sequence, 2.3, 2.4
8, 19 map
Arc View GIS. 12.21
278
Bothalia 32,2 (2002)
map (cont.)
distribution. 12.19, 15.13, 18.2. 19
dot, 12.18, 12.20
MAPPIT. 12.20
M-dash. 2.12, 17.6
mm, 13.11
margin. 2.1, 2.16, 3.4. 17.1
material, 2.3, 2.4
measurements, 13.11. 17.2, 17.7, 17.8
methods. 2.4. 6.10.3
microfiche. 15.5
miscellaneous paper. 10.14
monograph, 2.4, 15.13. 16.1
Mpumalanga (previously Eastern Transvaal), 15.8. 19
MSWord. 2.7. 2.11, 2.12, 3.5
name(s)
collector’s, 15.10
illegitimate. 16.2
in validly published, 16.3
of author(s), 2.3, 10.2, 10.7. 10.9, 10.11. 12.12. 13.7, 13.8, 14.3,
14.6. 15.7
of authors of plant names, 5.7.3, 13.1. 13.2, 13.6. 13.7, 13.8. 14.6
of publications, 13.8
Natal, see KwaZulu-Natal. 15.8, 19
N-dash. 2.11. 17.6
neotype, 15.6. 15.7
new
combinations, 7.4. 14.9
provinces of South Africa (Oct. 1996, April 2002), 15.8, 19
taxa, 7.4, 13.2, 13.7, 15.7, 18
nom. illeg., 16.2
nom. nud.. 13.5. 16,3, 16.4
non-breaking space, 2.7
normal style. 3.1
Northern
Province, see Limpopo. Northern Transvaal, 15.8, 19
Transvaal, see Northern Province. Limpopo. 15.8, 19
North-West, 15.8, 19
notes, 1, 2.4
technical. 10.14
number
chromosome, 17.1, 17.9
herbarium. 15,2
numbering, 13.13
figures. 12.12. 17.1
keys. 13.16. 13.17
pages. 2.5
taxa. 14.5. 15.15
numerals. Arabic. 11.1. 12.12, 13.3
obituaries, 1.1. 2.4. 10.7
optical disk, 3.2
Orange Free State, see Free State. 15.8, 19
page charges, 1 ,5
paragraphs, 3.6
PC diskettes, 3
pers, comm., 10,5, 10.8
personal communications (pers. comm,), 10.5, 10.8
photocopies, 2.1
photograph, 3.7, 12.3. 12.4. 12.13, 15.5, 18,2
mosaic, 2.1, 12.4, 12.5
plant
collectors. 13.9
name, 5, 13.4, 13.6, 13.7, 13.8, 14.6
plate (t.), 14.7
PRECIS (National Herbarium PREtoria Computerised Information
System). 13,1
prepositions. 6.4
proceedings. 10.14
proofs, 20
provinces, 6.7, 15.1, 15.8
of South Africa. 15.8. 19
publications, 10.8, 13.8. 14.3
name of, 14.2
solo, 10,10
year of. 10,9, 14.3
PWV, see Gauteng, 15.8, 19
quarter-degree squares. 15.1, 15.10
quotes, 2.15
reduction of figures, 12.1, 12.2, 12.18
referees. 1.4
reference, 2.4, 10.6, 10.7-10.9, 10.14
figure, 12.8
grid, 15.1, 15.8, 15.9. 15.11
list, 10.5. 10.8. 10.9
literature. 2.1. 10
report. 10.14
reprints, 21
requirements for
diskette, 3
manuscript, 2
results, 2.4
revision. 2.4, 8, 15.13, 16.1
RTF file, 3.3
scale bar, 12.10
semicolon, 2.13. 10.3, 15.3, 15.13
senior author, 10.10
sequence of headings, 2.3, 2.4
short notes, 1.1, 2.4
space
double, 2. 13
non-breaking, 2.7
one, 2, 16
special characters, 2.6
species treatment in taxonomic papers, 1 4
specimens examined, 2.4, 15.15
square brackets, 15.1. 17,9
STAFLEU, F.A. & COWAN. R.S. 1976-1988. Taxonomic liTeratiire.
Vols 1-7, 10.13
stiffy/stiffies, 3.2
style(s), 3.1, 3.4
submission of MS, 1.2, 1.6
surnames. 13.10
synopsis. 13.2, 13.5, 15.1, 16.4
synonymy, 13.8, 14.8, 16.6
t.. 14.3, 14.7, 17.9
table(s), 2.1, 2.4, 2.5, 11
digital submissions. 1 1
of contents, 8
tablet (t.), 14,7
tabs, 3.4
taxa
name of, 5, 7.4, 10.8, 13.2, 13.3
new. 7.4, 13.2, 13.7, 15.7, 18
numbering of, 14.5, 15,15
taxonomic
articles/papers, 7.2, 10,8, 12.11, 12.18, 13.2, 13,6, 13.8, 14
revision, 8
taxonomy, 5, 7.3, 13.4
technical note, 10,14
text, 2.1, 3.7, 10,1, 10.4, 10.5, 10.8, 10.9, 11.1, 12.7, 12.8, 12.12, 12.14,
13, 15.13, 15.15, 16.4
thesis, 10.14
TlFfile, 3.9, 12.2, 12.3
times sign. 2.14
title, 2.3. 5. 6.9, 6.11
of books, 10.4, 10.12, 10.13, 10.14
of journals, 10.4, 10.12. 10.13, 10.14
page. 2.3. 2.5
tracked changes, 3.1 1
Transvaal, 15.8. 17.9, 19
type, 15.2, 15.4, 15.7, 16.1, 16,5, 17.9
here designated. 15.7, 17.9
not designated, 15.7
specimen, 15.1
units of measure, 13.11, 13.15, 17.8
unpublished article, 10.14
voucher(s) specimens, 12.11. 13.19. 15,13. 15.14
Word for Windows, 3.3
World li.sl of scientific periodicals. 10. 1 3
year of publication, 10.9, 14.3
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BOTHALIA
Volume 32,2
CONTENTS
Oct. 2002
1. Systematics of the genus Daubenya (Hyacinthaceae; Massonieae). J.C. MANNING and A.M. VAN
DERMERWE 133
2. The genus Trichodesma (Boraginaceae: Boraginoideae) in southern Africa. E. RETIEF and A.E. VAN
WYK 151
3. Six new species and one new subspecies of Erica (Ericaceae) from Western Cape, South Africa E.G.H.
OLIVER and I.M. OLIVER 167
4. Studies in the liverwort family Aneuraceae (Metzgeriales) from southern Africa. 4. Riccardia obtusa.
S.M. PEROLD 181
5. Notes on African plants;
Amaryllidaceae. The typification of Cyrtanthus smithiae Watt ex Harv. J.P. ROURKE 197
Asteraceae. A distinctive new species of Felicia (Astereae) from Western Cape, South Africa.
J.C. MANNING and P. GOLDBLATT 193
Asteraceae. A new species of Arctotheca from Northern Cape, South Africa. J.B.P. BEYERS. . 185
Asteraceae-Anthemideae. Reduction of Foveolina albida to Foveolina dichotoma. J.B.P.
BEYERS 185
Hyacinthaceae-Massonieae. A new species of Lachenalia from Namaqualand, South Africa.
G. D. DUNCAN & T.J. EDWARDS 190
Mesembryanthemaceae. Mesembs with nut-like schizocarpic fruit and Ruschianthemum Fried-
rich sunk under Stoeberia Dinter & Schwantes. P. CHESSELET and A.E. VAN WYK . . 187
Portulacaceae. The synonymy of Ceraria namaquensis (Sond.) H. Pearson & E.L. Stephens.
H. F. GLEN 196
Pteridophyta. A new fern record for the Flora of southern Africa region. J.E. BURROWS and
S.M. BURROWS 195
6. A study of ovule-to-seed development in Ceratiosicyos (Achariaceae) and the systematic position of
the genus. EM. A . STEYN, A.E. VAN WYK and G.F. SMITH 201
7. The grasslands and wetlands of the Sekhukhuneland Centre of Plant Endemism, South Africa. S.J.
SIEBERT, A.E. VAN WYK, G.J. BREDENKAMP and F. DU PLESSIS 211
8. Miscellaneous notes:
Poaceae. Chromosome studies on African plants. 17. The subfamilies Arundinoideae and
Danthonioideae. R. ROODT, J.J. SPIES, A.E. MALAN, F. HOLDER and S.M.C. VAN
WYK 233
Poaceae. Chromosome studies on African plants. 18. The subfamily Chloridoideae. R. ROODT
and J.J. SPIES 240
9. Book reviews 251
10. National Botanical Institute South Africa: administration and research staff, 31 March 2002, publica-
tions 1 April 2001-31 March 2002. Compiler: B.A. Momberg 253
11. Guide for authors to 269
Abstracted, indexed or listed in • AETFAT Index • AGRICOLA • AGRIS • BIOSIS: Biological Ab.stracts/RRM • CABS • CABACCESS • CAB
ABSTRACTS • ISI: Current Content.'!, Scisearch, Re.tearch Alert • Kew Record of Taxonomic Literature • Taxon: reviews and notices.
ISSN 006 8241
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