ISSN 0006 8241 = BothaUa
Bothalia
A JOURNAL OF BOTANICAL RESEARCH
Vol. 37,1
May 2007
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BOTHALIA
Bothaha 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 South African National Biodiversity 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 sub-
jects are published annually.
Three booklets of the contents (a) to Vols 1-20, (b) to Vols 21-25 and (c) to Vols 26-30, are available.
STRELITZIA
A series of occasional publications on southern African flora and vegetation, replacing Memoirs of
the Botanical Survey of South Africa and Annals of Kirstenbosch Botanic Gardens.
MEMOIRS OF THE BOTANICAL SURVEY OF SOUTH AFRICA
The memoirs are individual treatises usually of an ecological nature, but sometimes dealing with
taxonomy or economic botany. Published: Nos 1-63 (many out of print). Discontinued after No.
63.
ANNALS OF KIRSTENBOSCH BOTANIC GARDENS
A series devoted to the publication of monographs and major works on southern African flora.
Published: Vols 14-19 (earlier volumes published as supplementary volumes to tho, 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 South African National Biodiverity 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 Lefty (over 700 plates),
Claire Linder-Smith and Ellaphie Ward-Hilhorst. The Editor is pleased to receive living plants of
general interest or of economic value for illustration.
From Vol. 55, twenty plates are published at irregular intervals.
An index to Vols 1^9 is available.
FLORA OF SOUTHERN AFRICA (FSA)
A taxonomic treatise on the flora of the Republic of South Africa, Lesotho, Swaziland, Namibia
and Botswana, the FSA contains descriptions of families, genera, species, infraspecific taxa, keys
to genera and species, synonymy, literature and limited specimen citations, as well as taxonomic
and ecological notes.
Contributions to the FSA also appear in Bothalia.
PALAEOFLORA OF SOUTHERN AFRICA
A palaeoflora on a pattern comparable to that of the Flora of southern Africa. Much of the informa-
tion is presented in the form of tables and photographic plates depicting fossil populations. Now
available:
Molteno Formation (Triassic) Vol. 1. Introduction. Dicroidium, 1983, by J.M. & H.M.
Anderson.
Molteno Fonnation (Triassic) Vol. 2. Gymnosperms (excluding Dicroidium), 1983, by J.M. &
H.M. Anderson.
Prodromus of South African Megafloras. Devonian to Lower Cretaceous, 1985, by J.M. & H.M.
Anderson. Obtainable from:A.A. Balkema Marketing, Box 317, Claremont 7735, RSA.
Towards Gondwana Alive. Promoting biodiversity and stemming the Sixth Extinction, 1999, by
J.M. Anderson (ed.).
BOTHALIA
A JOURNAL OF BOTANICAL RESEARCH
Volume 37,1
Scientific Editor: G. Germishuizen
Technical Editor: B.A. Momberg
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ISSN 0006 8241
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Editorial Board
D.F. Cutler
B.J. Huntley
P.H. Raven
M.J.A. Werger
Royal Botanic Gardens, Kew, UK
South African National Biodiversity Institute, Cape Town, RSA
Missouri Botanical Garden, St Louis, USA
University of Utrecht, Utrecht, The Netherlands
Acknowledgements to referees
Archer, Mrs C. South African National Biodiversity Institute, Pretoria, RSA.
Archer, Dr R.H. South African National Biodiversity Institute, Pretoria, RSA.
Boucher, Dr C. Botany Dapartment, University of Stellenbosch, RSA.
Brandham, Dr P.E. Jodrell Laboratory, Royal Botanic Gardens, Kew, UK.
Brown, Prof. L.R. Department of Nature Conservation, UNISA, Florida, RSA.
Bruyns, Dr P. Department Mathematics, University of Cape Town, RSA.
Burrows, J. P.O. Box 710, 1120 Lydenburg, RSA.
Cincotta, Dr R.P. Population Action International, Washington DC, USA.
Daemane, M.E. Arid Ecosystems Research Unit, Golden Gate National Park, RSA.
Foden, Ms W. South African National Biodiversity Institute, Pretoria, RSA.
Goldblatt, Dr P. Missouri Botanical Garden, St Louis, USA.
Goyder, Dr D.J. The Herbarium, Royal Botanic Gardens, Kew, UK.
Greuter, Prof Dr W. Botanical Garden and Botanical Museum, Berlin, Germany.
Hutchings, Ms A. P.O. Box 1362, 3880 Empangeni, RSA.
Kativu, Dr S. Dept. Biological Sciences, University of Zimbabwe, Harare, Zimbabwe.
Klopper, Ms R.R. South African National Biodiversity Institute, Pretoria, RSA.
Krige, Ms A. South African National Biodiversity Institute, Pretoria, RSA.
LiedeSchumann, Prof Dr S. Department of Systematic Botany, University of Bayreuth, Germany.
Linder, Prof H.P. University of Zurich, Zurich, Switzerland.
Manning, Dr J.C. South African National Biodiversity Institute, Cape Town, RSA.
McNeill, Dr J. Royal Botanic Garden, Edinburgh, Scotland, UK.
Meerow, Dr A.W. National Gennplasm Repository, Miami, Florida, USA.
Meve, Dr U. Department of Plant Systematics, University of Bayreuth, Germany.
Nordenstam, Prof R.B. Naturhistoriska Riksmuseet, Stockholm, Sweden.
Oliver, Dr E.G.H. Department of Botany and Zoology, University of Stellenbosch, RSA.
Paton, Dr A. The Herbarium, Royal Botanic Gardens, Kew, UK.
Retief, Dr E. South African National Biodiversity Institute, Pretoria, RSA.
Ross, Dr J.H. Royal Botanic Gardens, Melbourne, Australia.
Schrire, Dr B. Royal Botanic Gardens, Kew, UK.
Thulin, Dr M.L. Department of Systematic Botany, University of Uppsala, Uppsala, Swedeh.
Timberlake, Dr J.R. Royal Botanic Gardens, Kew, Richmond, England, UK.
Van Wyk, Prof A.E. University of Pretoria, RSA.
Venter, Prof H.J.T. Department of Plant Science, University of Free State, Bloemfontein, RSA.
Wallndfer, Dr B. Naturhistorisches Museum, Vienna, Austria.
Werger, Prof M.J.A. Department Plant Ecology and Evolutionary Biology, Utrecht, The Netherlands.
Winter, P.J.D. South African National Biodiversity Institute, Pretoria, RSA.
CONTENTS
Bothalia 37, 1
1 . Notes on new and misunderstood taxa of Cyrtanthus (Amaryllidaceae: Cyrtantheae) from the Western
Cape, Eastern Cape and KwaZulu-Natal, South Africa. D.A. SNIJMAN 1
2. New species and notes on the genus Cliffortia (Rosaceae). C.M. WHITEHOUSE and A.C. FELLING-
HAM 9
3. Notes on African plants:
Apocynaceae. Transfer of Schizoglossum iimbelhiliferiim to Stenostelma, and its neotypification
(Asclepiadoideae). S.R BESTER and A. NICHOLAS 48
Apocynaceae. A new species of Huernia (Asclepiadoideae-Ceropegieae) from southern Angola.
P.V. BRUYNS 23
Asphodelaceae. Trachyandra arenicola and T. montana, two new species from South Africa. J.C.
MANNING and P. GOLDBLATT 26
Asteraceae. A key to Dicoma taxa (Dicomeae) in southern Africa. N.C. NETNOU and B-E. VAN
WYK 55
Boraginaceae. Lectotypification of the basionym, Echiiim glaucophyllum. M.H. BUYS and B.
NORDENSTAM 25
Burseraceae. Commiphora kimeneana, a new species from the Kaokoveld, Namibia. W. SWANE-
POEL 40
Fabaceae. Aspalathus theresae, a new species from Western Cape, South Africa. C.N. CUPIDO . . 34
Hyacinthaceae. A new pyrophytic Lachenalia species (Massonieae) from Western Cape, South
Africa. G.D. DUNCAN and T.J. EDWARDS 31
Lamiaceae. Rabdosiella leemannii, a new species from the Limpopo Province of South Africa. N.
HAHN and G.J. BREDENKAMP 37
Leguminosae. Vigna verdcourtii (Papilionoideae), a new species from eastern Africa. R.S. PAS-
QUET 51
4. A reconnaissance survey of the woody flora and vegetation of the Catapu logging concession, Cheringoma
District, Mozambique. M. COATES PALGRAVE, A.E. VAN WYK, M. JORDAAN, J.A. WHITE
and P. SWEET 57
5. The concept of 'Musa-pelo and the medicinal use of shrubby legumes (Fabaceae) in Lesotho. A.
MOTEETEE and B-E. VAN WYK 75
6. Can anthropogenic variables be used as threat proxies for South African plant richness? M. KEITH and
M. WARREN 79
7. Threatened Limestone Fynbos plant communities of Andrew’s Field and Tsaba-Tsaba Nature Reserve,
Western Cape. M.M. ZIETSMAN and G.J. BREDENKAMP 89
8. Miscellaneous notes:
Amaryllidaceae. Chromosome studies on African plants. 20. Karyotypes of some Cyrtanthus
species. A. STRYDOM, R. KLEYNHANS and J.J. SPIES 103
9. The handling of the proposal to conserve the name Acacia at the 1 7th International Botanical Congress —
an attempt at minority rule. G. MOORE 109
10. Obituary: Elsie Elizabeth Esterhuysen (1912-2006). E.G.H. OLIVER (Compiler), J.P. ROURKE and
H.P. LINDER 119
11 Book review 129
12 Notice on Change of policy for reprints of Bothalia articles 131
New combinations, species, statuses and varieties in Bothalia 37,1 (2007)
Aspalathus theresae C.N.Cupido, sp. nov., 35
Cliffortia anthospermoides Fellingham, sp. nov., 9
Cliffortia bolusii Diels ex C. Whitehouse, sp. nov., 20
Cliffortia cruciata C. Whitehouse, sp. nov., 11
Cliffortia cuneata Dryaud. var. cylindrica C. Whitehouse, var. nov., 18
Cliffortia dentata Willd. var. gracilis (Hat'v.) C. Whitehouse, comb, et stat. nov., 14
Cliffortia ferricola C. Whitehouse, sp. nov., 12
Cliffortia lilifolia L.f. var. arenaria C. Whitehouse, var. nov., 20
Cliffortia gracillima C. Whitehouse, sp. nov., 14
Cliffortia perpendicularis C. Whitehouse, sp. nov., 15
Cliffortia sparsa C. Whitehouse, sp. nov., 16
Cliffortia weimarckii C. Whitehouse, sp. nov., 17
Commiphora kuneneana Swanepoel, sp. nov., 42
Cyrtanthus aureolinus Snijman, sp. nov., 1
Cyrtanthus mackenii Hook.f. subsp. cooperi (Baker) Snijman, stat. et comb, nov., 6
Huernia lopanthera Bruyns, sp. nov., 23
Lachenalia lutzeyeri G.D. Duncan, sp. nov., 31
Rabdosiella leemannii N.Hahn, sp. nov., 37
Stenostelma umbelluliferum (Schltr.) S.P.Bester & Nicholas, comb, nov., 50
Trachyandra arenicola J.C. Manning & Goldblatt, sp. nov., 27
Trachyandra moniana J.C. Manning & Goldblatt, sp. nov., 27
Vigna verdcourtii Pasquet, sp. nov., 53
IV
Bothalia37,l: 1-8 (2007)
Notes on new and misunderstood taxa of Cyrtanthus (Amaryllidaceae:
Cyrtantheae) from the Western Cape, Eastern Cape and KwaZulu-
Natal, South Africa
D.A. SNIJMAN*
Keywords: Amaryllidaceae, Cyrtantheae, Cyrtanthus, new species, subspecies. South Africa, taxonomy
ABSTRACT
Cyrtanthus aureolinus Snijman is a new, rare species of fire lily, which is localized in a vlei on the northern slopes of the
Groot Swartberg, Western Cape. The upright or slightly spreading, yellow to cream-coloured flowers and the perigone tube
which gradually widens to the throat suggest that it is closely related to the Western Cape endemic, C. ochroleucus (Herb.)
Burch, ex Steud., and C. mackenii Hook.f., a variable species from southern KwaZulu-Natal and Eastern Cape. The species
differs mainly by the shape, size and position of the tepals and the length of the filaments. Cyrtanthus mackenii var. cooperi
(Baker) R. A. Dyer is raised to subspecies rank as C. mackenii subsp. cooperi (Baker) Snijman. Its hysteranthous leaf habit
and grassland habitat differ from the riverine habitat of the evergreen C. mackenii subsp. mackenii. Described in detail are C.
aureolinus, C. ochroleucus, and C. mackenii.
INTRODUCTION
Found in southern and East Africa, Cyrtanthus Aiton,
comprising about 55 species, is one of the largest gen-
era in the family Amaryllidaceae. Throughout its distri-
bution, Cyrtanthus has species adapted to cope with fire
and because of this specialization they are commonly
known as fire lilies.
In the Cape Floristic Region, some of the first plants
to flower in response to summer fires and sufficient
moisture are species of Cyrtanthus which characteris-
tically produce their inflorescences in advance of the
leaves. In the heat of summer, however, their flowering is
often fleeting, hence many populations of Cyrtanthus in
the Cape Fold Mountains are infrequently seen or prob-
ably yet to be recorded. This became evident after a wild
fire burnt a large tract of arid fynbos on the dry north-
ern slopes of the Groot Swartberg, east of Oudtshoom,
Western Cape, in December 2004. Two to three weeks
later, a large population of Cyrtanthus appeared in a
blackened seepage zone on the lower slopes and the
stand of yellow flowers promptly attracted attention
from local nature conservationists and botanists. Field
and morphological studies of the newly discovered popu-
lation in relation to other species of Cyrtanthus with
tubular, yellow flowers have confirmed that the plants
are distinct from all the currently known species of the
genus and they are described here as a new species, C.
aureolinus.
Due to the discovery of C. aureolinus, its putative
close allies, C. ochroleucus (Herb.) Burch, ex Steud. and
C. mackenii Hook.f, were examined in detail. The mor-
phological variability within and among populations of
the two varieties of C. mackenii was re-assessed in rela-
tion to phenology and habitat. Consequently, the rank of
C. mackenii var. cooperi (Baker) R.A.Dyer is raised to
that of subspecies as C. mackenii subsp. cooperi (Baker)
* Compton Herbarium, South African National Biodiversity Institute,
Private Bag X7, 7735 Claremont, Cape Town.
MS. received: 2006-08-17.
Snijman. To accommodate the taxonomic changes and
new data, C. mackenii and C. ochroleucus are described
and mapped in full.
Cyrtanthus aureolinus iSnzytwor?, sp. nov., floribus
tubularibus et aureis ad C. ochroleucum (Herb.) Burch,
ex Steud. et C. mackenum Hook.f accedens, differt fila-
mentis exterioris 5 mm longis et filamentis interioris 7
mm longis, staminibus perigonio exserto, et florifero
tante post ignem.
TYPE. — Western Cape, 3322 (Oudtshoom); Swartberg
Mountains, N side of mountain on property directly W of
Meiringspoort and abutting Swartberg Nature Reserve,
(-BC), 14-1-2005, J.H.J.Vlok & A.L.Schutte 505 (NBG,
holo.; PRE).
Semi-evergreen, bulbous geophyte, 220-300 mm tall
when flowering. Bulb solitary, hypogeal, ovoid, up to
30^5 X 25-30 mm, narrowed to a slender neck 40-60
X 8-15 mm; outer tunics brown and papery; inner tunics
cream-coloured and fleshy. Leaves developing sequen-
tially, either absent or 2 or 3 newly emerging at flower-
ing, finally up to 4, strap-shaped, up to 300 x 5-9 mm,
suberect to recurved, ± channelled, glabrous; abaxial
surface with 3 or 4 prominent median veins, not keeled;
margin smooth. Inflorescence 4-10-flowered, solitary or
occasionally 2 per bulb; scape erect, up to 250 mm long,
5-8 mm diam. at base, tapering distally up to 4-6 mm
at apex, round to slightly compressed in t/s, green with
a reddish blush towards base, hollow; spathe valves 2,
outer sheathing inner at base, lanceolate, exceeding the
pedicels, up to 45-65 x 7-9 mm (at base), membranous,
suberect and flushed ox-blood red when fresh, later turn-
ing brown and reflexing; bracteoles several, linear, up to
15 mm long; pedicels suberect to slightly spreading, up
to 40 X 1.5 mm at anthesis, lengthening up to 50 mm and
becoming more rigid when fruiting, green to pale brown.
Flowers erect to spreading slightly above the horizontal,
opening sequentially, sulphur-yellow, sometimes with 6
diffuse green median bands from perigone base to tepal
tips, firm, unscented, apparently without nectar; peri-
gone tube dilated gradually to throat, straight or slightly
2
Bothalia37,l (2007)
curved distally, 29^0 mm long, ± 3 mm diam. at base,
widening gradually to 6-8 mm diam. at throat; tepals
narrowly ovate, slightly spreading, neither reflexed nor
rolled back, 12-15 x 5-7 mm, subacute at apex; outer
tepals sometimes slightly narrower than inner, shortly
mucronate. Stamens biseriate, regular, attached slightly
below and in perigone throat, both whorls shortly
exserted; outer filaments ± 5 mm long, attached ± 1 mm
lower than inner; inner filaments ± 7 mm long; anthers
dorsifixed, ± 4.5 mm long before dehiscing, yellow; pol-
len yellow. Ovary ellipsoidal, somewhat 3-angled, ± 5 x
2 mm, green; ovules axile, ± 25 per locule; style usually
pressed against upper tepals, as long as or slightly longer
than perigone, reaching up to 40 mm; stigmatic branches
3, spreading, slender, ± 2.0 x 0.5 mm, truncate, shortly
papillate towards apex. Capsule ovoidal, ±15x8 mm.
Seeds unknown. Plate 1 .
Phenology’ and pollination', the first flowers appear
two to three weeks after summer fires. Individual flowers
fade after a few days, but a many-flowered inflorescence
may remain attractive for approximately one week. The
response of individual plants is also staggered, so the
entire flowering period lasts approximately three weeks.
Although most individuals produce at least one leaf
while flowering, the full complement of leaves is pres-
ent only later in the season. Thereafter the plants remain
evergreen until burnt by another fire.
Cyrtanthus aureolinus shows the typical fire lily
response of not flowering in the seasons between fires.
Observations in the early morning indicate that the flow-
ers are visited by honeybees which alight on the tepals
but do not enter the perigone tube. In the absence of nec-
tar, pollen appears to be their only floral reward.
Diagnostic features: in its floral characters, particu-
larly the form, position and colour of the flowers,
Cyrtanthus aureolinus resembles C. ochroleucus from
the Langeberg, Western Cape, and to a lesser degree the
Eastern Cape and KwaZulu-Natal species, C. mackenii.
All three species have somewhat tubular flowers with
± equally long perigone tubes that widen gradually
from the base to the throat and have small tepal lobes.
Although the flowers of C. mackenii and C. ochroleucus
vary in colour from yellow to cream-coloured or rarely
white, they resemble the clear yellow flowers of C. aure-
olinus in being upright to somewhat spreading, and are
seldom horizontally spreading.
Despite their many similarities, these species can
be distinguished florally mainly by the shape, size and
position of the tepals, and by the length of the fila-
ments (Table 1). The narrowly ovate to oblong tepals in
Cyrtanthus aureolinus and C. ochroleucus are more than
twice as long as broad and slightly spreading, unlike
those of C. mackenii which are ovate, less than twice as
TABLE 1 . — Morphological characters that distinguish Cyrtanthus mackenii, C. aureolinus, and C, ochroleucus
Bothalia 37,1 (2007)
3
long as broad and patent or rolled back at anthesis. The
filaments, however, are most diagnostic. In C. aureoli-
nus both the outer and inner whorls of filaments (5 and
7 mm long respectively) extend beyond the perigone
throat. Those of C. ochroleucus are slightly shorter (± 2
and ± 4 mm respectively) and only the inner filaments
extend beyond the throat. In contrast, both the outer and
inner filaments in C. mackenii are very short (± 1 and ±
1 mm respectively) and the stamens remain included in
the perigone throat or within the upper part of the tube.
If the features of the tepals and stamens that charac-
terize the species are also significant for determining
affinities, then Cyrtanthus aureolimts is closest to C.
ochroleucus. The perigone throat of C. aureolimis, how-
ever, is wider than that of C. ochroleucus and its leaves
are broad and smooth, unlike the narrow leaves of C.
ochroleucus which are minutely papillate on the margin
and midvein. Both species are found in the fynbos of the
Western Cape where they occupy different habitats and
display dissimilar flowering habits.
Distribution and habitat: Cyrtanthus aureolimis is
confined to damp habitats on the northern foothills of the
Groot Swartberg, west of Meiringspoort (Figure 1). The
only known population, covering an area of ± 1 5 x 40 m
in size, is found in a permanent seepage zone in arid fyn-
bos, where sandstone-derived soils meet a band of loam-
rich soils at ± 800 m. In winter the area is waterlogged
but it remains sufficiently wet in summer to support a
population of the clicking stream frog, Strongylopus
grayii (Smith). In periods between fires (of ± 18^0
years) the seepage zone is dominated by large clumps
of restioids which form dense, fibrous tussocks at their
base. Most individuals of C. aureolinus grow between
these clumps, but in the wetter parts of the vlei, they are
found on the raised tussocks. They are absent from the
wettest parts of the site.
Other specimen examined
WESTERN CAPE. — 3322 (Oudtshoom): Farm Wilgermond, north-
ern slopes of Groot Swartberg, W of Meiringspoort, ± 800 m, (-BC),
19-1-2005, Snijman 1980 (K, NBG, PRE).
FIGURE 1. — Distribution of Cyrtanthus ochroleucus, •; and C. aure-
olinus, ■, in Western Cape Province; and C. mackenii subsp.
mackenii, ▼ ; and C. mackenii subsp. cooperi, A , in KwaZulu-
Natal and Eastern Cape Province.
Re-assessment o/ Cyrtanthus mackenii and
C. ochroleucus
The discovery of Cyrtanthus aureolinus, which
is putatively most closely related to C. ochroleucus,
makes it necessary to focus attention on the question-
able identity of a group of hysteranthous-leaved plants
from the grasslands of the Eastern Cape, northeast of
King William’s Town — a problem raised by Reid &
Dyer (1984) and to some extent by Turrill (1960). Reid
& Dyer (1984) pointed out that these populations are
found in the same area as C. mackenii var. cooperi, but
remarked that the plants might possibly represent out-
liers of C. ochroleucus, a species originally collected
by William Burchell whose travels in the Cape never
extended northeast of the Great Fish River (McKay
1943). The inconsistent naming of these particular
plants is reflected in Batten & Bokelmann (1966: 31,
t. 25, fig. 4) who referred to the plants from near King
William’s Town as Cyrtanthus sp., and those from near
East Eondon as C. ochroleucus (t. 24, fig. 1 ). In contrast,
Manning (2001: 74, fig. 1 & 81, fig. 4) referred to both
as C. mackenii.
In an attempt to resolve the relationships of the
Eastern Cape plants to Cyrtanthus mackenii and C.
ochroleucus, all the South African herbarium mate-
rial (BOL, GRA, NBG, NH, PRE, SAM) belonging to
this complex was studied together with the types of C.
ochroleucus, C. mackenii var. mackenii and C. mackenii
var. cooperi.
The types of Cyrtanthus mackenii var. cooperi —
Cooper 1806 (K, NH, PRE) collected in British Kaffraria,
the eastern part of today’s Eastern Cape Province — all
lack leaves which strongly suggests that the leaves are
hysteranthous. Furthermore, the pressed inflorescences
(on the sheets at K and PRE) closely resemble those of
the hysteranthous-leaved plants from the Eastern Cape
referred to by Reid & Dyer (1984) with regard to flower
number, perigone tube length, size of the perigone throat
(4 mm wide), and shape and size of the tepals (ovate and
4 X 3.5 mm). This close similarity is in contrast to the
more distant likeness found between C. ochroleucus and
the hysteranthous-leaved Eastern Cape plants which are
in question. Despite their shared hysteranthous-leaved
state, they differ in leaf width (± 3 vs 7-1 1 mm); tepal
shape (narrowly ovate to oblong vs ovate); tepal size (7-
1 1 X 3-4 vs 3. 5-6.0 x 2. 5-4.0 mm); and flowering time
(Oct.-Dec. vs Aug.-Oct.). Moreover, no intermediates
between the two elements are known and their distribu-
tion ranges are disjunct, separated by a gap of ± 750 km.
Based on the above comparison, the resemblance of
the hysteranthous-leaved plants from the Eastern Cape
to the type of Cyrtanthus mackenii var. cooperi is con-
sidered to be sufficiently close to warrant their inclusion
in C. mackenii var. cooperi. This is in agreement with
Baker’s (1896) circumscription of the taxon which he
originally named C. lutescens var. cooperi Baker.
Following the inclusion here of the Eastern Cape’s
populations of hysteranthous-leaved plants into Cyrtan-
thus mackenii var. cooperi, is the need to re-assess the
classification of C. mackenii into two varieties which
4
Bothalia 37,1 (2007)
Dyer (1939) and Reid & Dyer (1984) distinguished pri-
marily by the possession of yellow to cream-coloured
rather than white flowers. To do so, this comparison has
incorporated field data on flowering and leafing phenolo-
gies and habitat which are often missing from herbarium
records.
The descriptions of Cyrtanthus mackenii (Dyer 1939;
Turrill 1960; Reid & Dyer 1984) mention that the spe-
cies reproduces vegetatively from bulblets. Field data,
however, indicate that this habit is confined to the typi-
cal variety and leads to the fomiation of large clumps of
bulbs with several shiny, evergreen leaves. Furthermore,
the flowers of C. mackenii var. mackenii are sweetly
scented, the perigone has a fairly broad throat (6-8 mm
wide), and the tepals (5-10 x 3. 5-5.0 mm) are ovate and
vary from patent to rolled back at anthesis. Distributed
from near Port Shepstone in southern KwaZulu-Natal
southwards along the Eastern Cape coast, the plants
grow most often on the shaded banks of permanent
streams, in patches of remnant forest which seldom or
never bum. They flower mainly in winter, from June to
September, or rarely as late as November.
In contrast, Cyrtanthus mackenii var. cooperi is
deciduous and has solitary bulbs that produce two to five
glaucous leaves each season. The flowers have a strong
spicy scent with a 4. 0-5. 5 mm wide perigone throat,
and short, ovate tepals (3. 5-6.0 x 2. 5^.0 mm) that are
slightly recurved to rolled back at anthesis. Populations
belonging to this taxon are plentiful in seasonally
damp places in open, rolling grassland but occasional
plants can be found in drier situations. They have been
recorded from the Midlands of southern KwaZulu-Natal,
near Kokstad, through to King William’s Town, Eastern
Cape, and close to East London where they approach
the coast. Although the plants do not depend on fire to
flower, they are known to flower most prolifically in the
spring (August to October) following winter fires. With
few exceptions, the bulbs produce new leaves slightly
later in the season.
Despite the subtle nature of the floral features that
separate these taxa, they may play a role in attracting
a slightly different range of pollinators. This, together
with the slightly asynchronous flowering times and their
discrete habitats, may impose some level of reproductive
isolation upon the taxa. It is apparent, nevertheless, that
these morphological, phonological and habitat differ-
ences reflect a level of discontinuity that warrants more
than varietal delimitation, especially as the populations
of the two taxa occupy different, albeit partially sym-
patric, geographical ranges, namely the coast versus the
Midlands of southern KwaZulu-Natal and the Eastern
Cape. The only places where populations of the two taxa
co-occur are where patches of forest extend inland into
the temperate grasslands along major river courses. This
interpretation, consequently, excludes the coastal, ever-
green plants with yellow flowers that Dyer (1939) and
Reid & Dyer (1984) previously considered to belong to
C. mackenii var. cooperi.
Based on the above conclusions, C. mackenii var.
cooperi is elevated to subspecies rank, C. mackenii
subsp. cooperi. The full descriptions of C. mackenii and
C. ochroleucus are given below.
Cyrtanthus mackenii Hook.f. in The Gardener’s
Chronicle and Agricultural Gazette: 641 (1869); Baker:
57 (1888); Baker: 225 (1896). Type: Natal [KwaZulu-
Natal], P.C. Sutherland s.n. in Herb. Hookerianum, July
1864 (K000366180, lecto.; designated here, image only
seen).
Bulbous geophyte, 180-530 mm tall when flowering.
Bulb hypogeal, up to 20-50 x 15-35 mm, narrowed to
a neck 10-80 x 7.5-15.0 mm; outer tunics brown and
papery; inner tunics cream-coloured and fleshy. Leaves
narrowly lanceolate, up to 400 x 7-18 mm, suberect to
recurved, slightly channelled, glabrous; abaxial surface
with 3 prominent median veins. Inflorescence 3-8(9)-
flowered; scape erect to curved, up to 450 x 5-8 mm
diam. at base, tapering distally up to 3-5 mm at apex,
round to slightly compressed in t/s, hollow; spathe valves
2, outer sheathing inner at base, lanceolate, shortly
exceeding pedicels, up to 25-60 x 4-8 mm (at base),
membranous, suberect and flushed pinkish when fresh,
later reflexing and turning brown; bracteoles several,
linear, up to 20 mm long; pedicels suberect to slightly
spreading, up to 27 x ± 1.5 mm at anthesis, lengthening
up to 30 mm and becoming more rigid when fruiting.
Flowers opening ± simultaneously, white to yellow often
suffused with apricot-pink; nectar unknown; perigone
tube dilated gradually to throat, slightly to distinctly
curved; tepals ovate, outer shortly mucronate. Stamens
biseriate, regular; filaments ± 1 mm long; anthers dorsi-
fixed, 2. 0-2. 5 mm long before dehiscing, yellow; pollen
yellow. Ovary ellipsoidal, somewhat 3-angled, 4-5 x 3
mm; ovules axile, ± 25 per locule; style usually pressed
against upper tepals, as long as or rarely slightly longer
than perigone tube, reaching up to ± 45 mm; stigmatic
branches 3, spreading, slender, ± 1 mm long, truncate,
shortly papillate towards apex. Capsule ovoidal, 10-20
X 7-10 mm. Seeds 5x3 mm, winged, dark shiny brown.
subsp. mackenii
Dyer: 79 (1939); Reid & Dyer: 16 (1984).
C. mackenii var. cooperi (Baker) R. A. Dyer, pro parte, excluding
the type of this taxon: 79 (1939); Reid &. R. A. Dyer, pro parte, exclud-
ing the type of this taxon: 17 (1 984).
Illustrations: Phillips: t. 33 (1921); Turrill: t. 368 (1960); Batten
& Bokehnann: t. 24, fig. 6 [as C. mackenii var. cooperi (Baker) R.A.
Dyer] (1966); Manning: 75, fig. 1 top right (2001).
Evergreen, often semi-aquatic bulbous geophyte.
Bulbs clumped, fonning numerous offsets. Leaves pres-
ent at flowering, 4 or 5, shiny green; margin smooth.
Inflorescence solitary per bulb; scape green often with a
reddish blush towards base; pedicels green. Flowers 3-
6(-8), suberect to slightly nodding, clear yellow, cream-
coloured or white, sweetly scented; perigone tube 36-45
mm long, ± 2 mm diam. at base, widening gradually to
6-8 mm diam. at throat; tepals ovate, patent to rolled
back, 5-10 X 3. 5-5.0 mm, obtuse at apex; outer tepals as
wide as inner. Stamens included in perigone throat and
upper part of perigone tube; outer filaments attached 3^
mm below throat; inner filaments attached in perigone
throat, rarely up to 2 mm below throat. Ovary green.
Phenology, flowering starts in June and usually lasts
until September, but can extend into early November.
The leaves are evergreen.
Bothalia 37,1 (2007)
5
Diagnostic features: the plants, commonly known
as Ifafa lilies, are clump-forming and produce sev-
eral leaves which persist throughout the year. The usu-
ally sweet-smelling flowers have ovate tepals that are
patent to rolled back at anthesis and the stamens are
characteristically included in the fairly broad perigone
throat (Table 1). Populations of white-flowered plants
are fairly localized and have been recorded ifom the Port
Shepstone District in KwaZulu-Natal near the mouths
of the Umzimkulu and Umtamvuna Rivers, and in the
Eastern Cape near the Great Kei River mouth. Elsewhere
in the distribution range, populations occasionally have
cream-coloured flowers, but most often the flowers are
yellow.
Distribution and habitat: Cyrtanthus mackenii subsp.
mackenii is distributed along the Indian Ocean coastal
belt from Port Shepstone southwards to Port Edward,
KwaZulu-Natal, and from Port St Johns to just south of
the Great Kei River, Eastern Cape. Scattered populations
have also been recorded a short distance inland along
the banks of large rivers (Figure 2). The plants favour
partially shaded habitats, amongst rocks on the edge of
watercourses in remnant patches of Scarp Forest and
Southern Mistbelt Forest (sensu Mucina et al. 2005).
Specimens examined
KWAZULU-NATAL. — 2930 (Pietermaritzburg): Malvern, (-DD),
22-7-1951, Kent s.n. (NH40424). 3029 (Kokstad): Umzimkulu, (-BD),
Aug. 1967, Gardner s.n. (NH57299). 3030 (Port Shepstone): Isipingo,
(-BB), Jun. 1882, Wood 1336 (K, NH); Ifafa, (-BC), Feb. 1885, Knox
2103 (PRE), Aug. 1916, Lansdell s.n. {NH16127); Natal coast, Ifafa
Dist., (-BC), May 1883, Tyson 2103 (BOL, K, SAM); The Valleys,
(-CB), 8-7-1952, Martin 936 (NBG), 3-10-1937, Mogg 13945 (PRE);
Oribi Gorge, (-CB), 16-8-1980, Schrire 1850 (NH); Umzimkulwana
River, (-CB), July 1916, Thode 2515 (NH); Umtamvuna Nature
Reserve, (-CC), 31-7-1982, Abbott 170 (PRE), 11-7-1977, Nicholson
1784 (PRE), 15-6-1969, Strey 8745 (PRE), 15-7-1975, Van Wyk 1669
(PRE), 14-7-1976, Venter 1004 (PRE), 18-7-1995, Victor 1381 (PRE).
Inexact locality; Natal, Gerrard s.n (K000366179).
EASTERN CAPE. — 3129 (Port St Johns): Mbotyi, on river banks,
50 ft [15 m], (-BC), 9-9-1961, Batten s.n. {NBG60489)-, Mbotyi, on
river banks, 100 ft [30 m], (-BC), 10-9-1962, Batten s.n. {NBG66279);
Mbotyi, on river banks, 200 ft [60 m], (-BC), 12-8-1963, Batten s.n.
(NBG69947); Magwa Estates, (-BC), 7-9-1979, Germishuizen 1193
(PRE); Ntsubane Forest Station, Fraser’s Falls, (-BC), 24-8-1976,
Venter & Vorster 101 (PRE); 2 km from forestry station on road to [van
bosbou stasie op pad na] Mboyti, (-BC), 12-7-1976, Venter 886 (PRE);
Coffee Bay, (-CC), Oct. 1919, Tyson s.n. (PRE TM20573); ± 2.5 km
from Umzimvubu bridge at Port St Johns, (-DA), 20-8-1998, Abbott
7296 (NBG). 3228 (Butterworth): Tsolorha. Bashee River, (-BA), Van
Jaarsveld 16903 (NBG); Cwebe Forest, (-BB), 14-9-1998, Winter s.n.
(NBG179726); between Qora and Mazeppa Bay, (-BC), Winter 513
(NBG); Dwesa Nature Reserve, (-BD), Winter 496 (NBG); Kei River
mouth, margins of streams, (-CB), Aug. 1889, Flanagan 300 (BOL,
PRE, SAM); banks of Kei River, 200 ft [60 m], (-CB), July 1894,
Flanagan 2335 (BOL, NBG); Kentani Dist., (-CB), July 19fl, Pegler
28 (PRE); Nyutura, (-CB), 12-7-1966, Strey 6641 (NH, PRE), 12-7-
1966, Ward 5724 (PRE); half mile [0.8 km] E of Nenga Post Office
(-CC), 17-7-1955, Codd 9275 (PRE); Farm Hill and Dale, near Haga
Haga, in stream bed and banks, 60 m, (-CC), 8-8-2001, McMaster s.n.
(NBG177935): near Haga Haga, (-CC), McMaster s.n. (NBG182889).
subsp, cooperi (Baker) Snijman, stat. et comb. nov.
C. mackenii var. cooperi (Baker) R. A. Dyer, syn. nov.: 79 (1939).
C. lutescens var. cooperi Baker, Handbook of the Amaryllideae: 58
(1888); Baker: 225 (1896). Type: British Kaffraria [Eastern Cape],
1860, TCooper 1806 (K000400308, lecto.; [designated by Reid &
Dyer: 17 (1984)], image only seen; NH!, PRE!).
C. lutescens sensu Hook.f: 89: t. 5374 (1863), non Herb.: 129. t.
33, fig. 14 (1837) [= C. ochroleucus (Herb.) Burch, ex Steud.]
Illustrations: Batten & Bokelmann: t. 24, fig. 1, [as C. ochroleucus
(Herb.) Burch, ex Steud.]; t. 25, fig. 4 [as C. sp.] (1966); Manning: 75,
fig. 1 top left & 81, fig. 4 (2001).
Deciduous, bulbous geophyte, 180-530 mm tall when
flowering. Bulb solitary. Leaves developing sequentially,
absent at flowering, rarely 1 or 2 emerging at end of flow-
ering period, finally up to 5, green or glaucous; margin
smooth or minutely rough. Inflorescence solitary or occa-
sionally 2 per bulb; scape glaucous with a pinkish blush.
Flowers 3-8(9), suberect to spreading horizontally, cream-
coloured to dull yellow, often suffused with apricot-pink on
tube; scent spicy, usually bittersweet; perigone tube 25^7
X ± 2 mm diam. at base, widening gradually up to 4.0-5. 5
mm diam. at throat; tepals ovate, slightly recurved to rolled
back, 3. 5-6.0 x 2. 5^.0 mm; outer tepals slightly narrower
than inner, with somewhat subacute and shortly mucro-
nate tips; inner tepals with round tips. Stamens included
in upper part of perigone tube and throat, outer attached ±
5 mm below throat, inner attached ± 2 mm below throat.
Ch’aty green or pinkish grey.
Phenology: the flowering period which often follows
winter fires starts at the end of July at low altitudes (30
m) and peaks in September, but a few individuals con-
tinue flowering until November at high altitudes ( 1 800
m). The plants flower in the seasons between fires so
are not true fire lilies. The leaves are absent during the
main flowering period in September and begin to emerge
from scattered individuals in October. Leafing contin-
ues throughout summer until the onset of the dry season
when the bulbs become dormant over winter.
Diagnostic features: the plants are solitary and decidu-
ous, and the leaves mostly reach maturity after flowering.
The perigone tube has a relatively narrow throat with
small, ovate tepals (Table 1). The flowers are most often
dull yellow but in populations from around Maclear,
Umtata, Stutterheim, and King William’s Town, the peri-
gone tube is pinkish cream to apricot. Often the flowers
have a spicy scent.
Distribution and habitat: populations of Cyrtanthus
mackenii subsp. cooperi are usually concentrated in sea-
sonally damp places in open grassland where they are
most prolific, but occasional individuals are also found
in dry grassland. The distribution extends through the
Sub-Escarpment Grassland Bioregion in East Griqualand
and the Transkei (sensu Mucina et al. 2005), below the
Amathole Mountains around King William’s Town and
Stutterheim and to grassy flats in the vicinity of East
London. In the northern parts of the range, populations
are found at altitudes of up to 1 400 m but in the south
they reach the coast (Figure 2).
Specimens examined
KWAZULU-NATAL. — 3028 (Matatiele): between Swartberg and
Matatiele, (-BD), 29-9-1962, Strey 4315 (PRE). 3029 (Kokstad): Farm
Thomham, Kokstad, (-CB), 10-10-1972, Coleman 633 (NH); Kokstad
golf course, (-CB), Oct. 1966, Germishuizen 100 (PRE); Kokstad, (-
CB), Sep. 1940, Sister Mildred 267 (NBG); circa Kokstad, (-CB), Oct.
1883, Tyson 1550 (BOL); Harding, 3000 ft [914 m], (-DB), Sep. 1928,
Oliver 19 (NH).
Bothalia37,l (2007)
EASTERN CAPE.— 3127 (Lady Frere): Elliot, (-BD), 30-9-1935,
Reynolds s.n. {PRE37783), 5-10-1935, Reynolds s.n. (BOL22397); 12
miles [19 km] N of Gala, (-BD), 16-10-1953, Theron s.n. (PRE37793);
3 km on road to McLeantown, (-DA), 30-9-1976, Stirton 6289 (PRE);
10 miles [16 km] S of Cala Pass on road to Engcobo, (-DB), Sep.
1940, Reynolds 3605 (PRE). 3128 (Umtata): near Ugie, (-AA), 6-10-
2001, Adendorff s.n. (NBG); Maclear commonage, 1 470 m, (-AB),
30-8-1993, Bester 715 (NH); Maclear commonage, 1 380 m, (-AB),
13-10-1993, Bester 1201 (NH); 1 mile [1.6 km] W of Maclear railway
station, (-AB), Reynolds 3033 (PRE); Farm Borva, ± 12 km SSE of
Xuxa drift siding, 1 410 m, (-AC), 1-9-1993, Bester 754 (PRE), 755
(NH); Tsolo Agricultural College, 900 m, (-BD), 9-10-1990, Cloete
508 (NH); Bazeia, in low moist places, 2000 ft [609 m], (-CB), Bauer
242 (GRA); Umtata, 2500 ft [625 m], (-DB), 1-9-1962, Bokelmann
s.n {NBG62902); Umtata, (-DB), 13-9-1997, Singh & Baijnath 243
(NH, PRE). 3226 (Fort Beaufort): Katberg, 6000 ft [1 829 m], (-BC),
Nov. 1926, Dyer 760 (GRA, PRE); Katberg, no date, H.Hutton s.n.
(K000400306). 3227 (Stutterheim): Stutterheim Div., Dohne Research
Station, (-CB), 21-8-1942, Acocks 9022 (PRE); Stutterheim Dist.,
towards Kologha, 880 m, (-CB), 28-9-1997, McMaster 30 (NBG);
Stutterheim Dist., adjacent to Van Rensburgdorp, 860 m, (-CB), 28-
9- 1997, McMaster 31 (NBG); grassy valley above Komgha, 4000 ft
[1218 m], (-DB), Sep. 1889, Flanagan 294 (BOL, GRA, PRE);
Berlin, King William’s Town Dist., (-DC), 14-9-1954, Marais
445 (GRA); Gonubie, grassy flats, (-DD), 20-5-1961, Batten s.n.
(NBG60457). 3327 (Peddie): Bufiffalo Pass, East London, (-BB), 7-1-
1945, Barker 3520 (NBG); East London, 20-100 ft [61-30 m], (-BB),
10- 9-1962, Batten s.n. (NBG62903)', 1 mile [1.6 km] from Bonza
Bay, (-DD), 20-7-1955, Comins 1255 (PRE). Inexact locality: British
Kaffraria, Cooper 3225 (K000400310), C.Hutton s.n. (K000400310).
Cyrtanthus ochroleucus (Herb.) Burch, ex Steud.,
Nomenclator botanicus 1,4 (edn 2): 475 (1840). Monella
ochroleitca Herb.; 29 (1821). Type: [Western Cape]
Riversdale Div., on or near the Langebergen, WJ.Biirchell
7144 (K, lecto.; here designated, image only seen).
C. lutescens Herb. var. hitescens: 129, t. 33, fig. 14 (1837), nom.
superfl.
Deciduous, bulbous geophyte, 130-360 mm tall when
flowering. Bulb hypogeal, ovoidal, 30^0 x 30-35 mm,
narrowed to a short neck up to 10 x 7-10 mm; outer
tunics brown and papery; inner tunics cream-coloured
and fleshy. Leaves developing sequentially, either absent
or 1 newly emerging at flowering, 2 or more, linear, at
least 100 X up to 3 mm, suberect; abaxial surface with
a prominent median vein, keeled; margin and midvein
minutely papillate. Inflorescence 2-6-flowered, soli-
tary per bulb; scape erect, up to 320 x 5 mm (at base),
tapering distally up to ± 3 mm at apex, round to slightly
compressed in t/s, dull green, hollow; spathe valves 2,
outer sheathing inner at base, lanceolate, exceeding
pedicels, up to 25 x ± 3.5 mm (at base), membranous,
suberect and pinkish when fresh, later reflexing and
turning brown; bracteoles several, linear, up to 15 mm
long; pedicels suberect to slightly spreading, dull green,
up to 12 X ± 1.5 mm at anthesis, lengthening up to 15
mm and becoming more rigid when fruiting. Flowers
erect to slightly spreading, opening ± simultaneously,
cream-coloured to dull yellow, suffused with pink or
green on perigone tube sometimes extending to tepals,
with a bittersweet scent; nectar unknown; perigone tube
dilated gradually to throat, slightly curved, 33-50 mm
long, ± 2 mm diam. at base, widening gradually up to
5-6 mm diam. at throat; tepals narrowly ovate to oblong,
slightly spreading, neither reflexed nor rolled back,
7-1 1 X 3^ mm, outer tepals ± as wide as inner tepals,
shortly mucronate at apex; inner tepals obtuse. Stamens
biseriate, regular; outer stamens included; inner stamens
shortly exserted; outer filaments ± 2 mm long, attached
± 3 mm below perigone throat; inner filaments ± 4 mm
long, attached in perigone throat; anthers dorsifixed, 2
mm long before dehiscing, dark yellow; pollen yellow.
Ovary ellipsoidal, ±5x3 mm, dull green; ovules axile,
± 25 per locule; style usually pressed against upper tepal,
slightly shorter than perigone, reaching up to 47 mm;
stigmatic branches 3, short, spreading, slender, ± 1 mm
long, truncate, shortly papillate towards apex. Capsule
unknown.
Phenology, the plants flower from October to
December, mostly in advance of the leaves, but in a few
individuals one newly emerging leaf may be present.
After flowering, the leafing habit remains poorly known.
Diagnostic features: Cyrtanthus ochroleucus is dis-
tinguished by its hysteranthous leaves which are nar-
row (± 3 mm wide) and minutely papillate on the mar-
gin and midvein, by the suberect tubular flowers which
are cream-coloured to dull yellow, and by the short (± 2
mm) outer filaments and longer (± 4 mm) inner filaments
so that only the inner stamens are exserted from the peri-
gone throat. The similarities and differences between C.
ochroleucus and its close allies, C. aureolinus and C.
mackenii, are summarized in Table 1 .
Distribution and habitat: this species is only known
from fynbos on the lower northern slopes of the
Langeberg, and on the Albertinia coastal plain. Western
Cape (Figure 2). Populations have been found in stony
or sandy habitats and, although the bulbs flower after
fire, they also flower in the intervals between fires. The
plants grow singly and are scattered in open patches.
Nomenclatural note: Herbert (1821) based his origi-
nal description of Monella ochroleuca Burch, (the basi-
onym of C. ochroleucus) on the herbarium collections of
Burchell, but he failed to- cite any particular specimen.
The original diagnosis of M. ochroleuca states that it
is allied to M. odora (Ker Gawl.) Burch, (based on C.
odorus Ker Gawl.) but that it is distinguished by having
yellow-white flowers. Cyrtanthus odorus itself is a dis-
tinctive species which is characterized by hysteranthous
leaves, suberect, tubular flowers with small tepals and, in
contrast to C. ochroleucus, has dark red flowers.
Herbert ( 1 837) later adopted the name C. lutescens for
M. ochroleuca, which he explicitly referred to as a syno-
nym, and he cited Burchell 7144 as the only preserved
specimen. There can be little doubt that the diagnosis of
M. ochroleuca clearly distinguishes it from other taxa
and that the name is validly published. Thus C. lutes-
cens is considered to be nomenclaturally superfluous and
is illegitimate. The lectotype of C. ochroleucus that has
been designated here is Burchell 7144 (K) which con-
sists of leafless, flowering bulbs with narrowly tubular
flowers that appear to be held upright.
Specimens examined
WESTERN CAPE. — 3321 (Ladismith): Garcia’s Pass, (-CC),
Oct. 1904, H. Bolus s.n. (BOL111388, PRE); Langeberg above Corente
River, (-CC), Nov. 1908, Muir 191 (PRE); Corente River Farm,
Riversdale Dist., (-CC), Nov. 1908, Muir 5379 (PRE); Oudenbosch
at foot of Langeberg, stony slope recently burned, (-CC), 16-12-
1979, Oliver 7550 (NBG, PRE); Langeberg, NE of Garcia’s Pass, W
iiMi
PLATE 1. — Cyrtanthus aureolinus, Snijman 1980. A, bulb; B, leaves and inflorescence; C, young infructescence; D, half flower laid open. Scale
bar: A-C, 10 mm; D, 15 mm. Artist: Vicki Thomas.
Bothalia37,l (2007)
of Welgemoed, in old fire break, 2150 ft [655 m], (-CD), 13-12-1984,
Oliver 8639 (NBG, PRE); Langeberg at Bergfontein, 350 m, (-DC),
14-12-1979, Oliver 7529 (NBG). 3421 (Riversdale): Farm Vogelstruis,
in coastal fynbos, (-BC), Horstmann s.n. (NBG167767).
ACKNOWLEDGEMENTS
I would like to thank William Pulles and Colin
Paterson-Jones for assistance in the field; Leon and Tilla
Nell and Jan and Anne Lise Vlok for their observations
on C. aureolinus; Cameron and Rfioda McMaster and
Jaco Adendorff for their field data on C. mackenir, and
Vicki Thomas for the illustration of C. aureolinus. The
Western Cape Nature Conservation Board granted a per-
mit to collect specimens, and K, PRE and NH kindly
made their collections available for this study.
REFERENCES
BAKER, J.G. 1888. Handbook of the Amaryllideae. Bell, London.
BAKER, J.G. 1896. Cyrtanthus. Flora capensis 6: 218-228. Reeve,
Ashford, Kent.
BATTEN, A. & BOKELMANN, FI. 1966. Wild flowers of the Eastern
Cape Province. Books of Africa, Cape Town.
DYER, R.A. 1939. Description, classification and phylogeny. A review
of the genus Cyrtanthus. Herbertia 6: 65-103.
HERBERT, W. 1821. Appendix. Ridgway, London.
HERBERT, W. 1837. Amaryllidaceae. Ridgway, London.
HOOKER, J.D. 1863. Cyrtanthus lutescens. Curtis's Botanical
Magazine 89: t. 5374.
HOOKER, J.D. 1 869. Cyrtanthus mackenii. The Gardener 's Chronicle
and Agricultural Gazette: 641.
MANNfNG, J. 2001. Eastern Cape. South African Wild Flower Guide
1 1 . Botanical Society of South Africa & National Botanical
Institute, Cape Town.
McKAY, H.M. 1943. Sketch map of BurchelFs trek. Journal of South
African Botany 9: 27-78.
MUCINA, L., RUTHERFORD, M.C. & POWRIE, L.W. (eds). 2005.
Vegetation map of South Africa, Lesotho and Swaziland, 1:
1 000 000 scale sheet maps. South African National Biodiversity
Institute, Pretoria.
PHILLIPS, E.P. 1921. Cyrtanthus mckenii. The Flowering Plants of
South Africa 1: t. 33.
REID, C. & DYER, R.A. 1 984. A review of the southern African species
of Cyrtanthus. American Plant Life Society, La Jolla, California.
STEUDEL, E.T. 1840. Nomenclator Botanicus 1,4, edn 2. Cottae,
Stuttgart.
TURRILL, W.B. 1960. Cyrtanthus mackenii. Curtis’s Botanical
Magazine 173: t. 368.
Bothalia37,l: 9-22 (2007)
New species and notes on the genus Cliff ortia (Rosaceae)
C.M. WHITEHOUSE* and A.C. FELLINGHAM**
Keywords: Cape Floristic Region, CUffortia L., endemic, new combinations, new species, Rosaceae
ABSTRACT
Seven new species of CUffortia L, endemic to the Cape Floristic Region (CFR) are described, C. anthospermoides, C.
cruciata, C. ferricola, C. gracillima, C. perpendicularis, C. sparsa and C. weimarckii. A further species from the Graaff-
Reinet area, described by Weimarck but not formally named, is here given the name C. bolusii. New varieties C. cuneata var.
cylindrica and C. filifolia var. arenaria are also described, and C. gracilis Harv. is recombined as C. dentata var. gracilis.
CUffortia discolor and C. hermaphroditica Weim. are reduced to synonyms of C. odorata L.f. and C. juniperina L.f
respectively. There are now 132 species recognized in CUffortia, 124 of which are found in the CFR and 109 endemic to the
region.
INTRODUCTION
CUffortia L. is one of the ten largest genera in the
Cape Floristic Region (CFR) (Goldblatt & Manning
2000). In Fellingham’s (2000) enumeration of the spe-
cies in the CFR, 1 1 1 described species were recognized.
Five other species had been described that occurred
exclusively outside the CFR (Oliver & Fellingham 1994;
Whitehouse 2004a), including one, C. nitidula (Engl.)
R.E.Fr. & T.C.E.Fr. (incorporating C. aequatorialis
R.E.Fr. & T.C.E.Fr.), found as far as Kenya (Graham
1960). Since then, eleven more species have been
described (Fellingham 2003; Whitehouse 2004a, b).
In this paper, eight more species are described, as well
as new varieties of CUffortia cuneata Dryand. and C.fiU-
folia L.f. One species, C. gracilis Harv., is considered to
be a variety of C. dentata Willd., while a further two spe-
cies, C. discolor Weim. and C. hermaphroditica Weim.,
are reduced to synonymy of more widespread species.
Seven of the new species are endemic to the CFR, but
the eighth species is only known from the Nardouwsberg
on the Great Escarpment east of Graaff-Reinet. This
brings the total number of species in the genus CUffortia
to 132.
The following species were listed as endemic to the
CFR by Fellingham (2000), but have collections from
outside the region: CUffortia amplexistipula Schltr.
(Namaqualand, Fellingham 1993), C. erectisepala
Weim. (Grahamstown, Whitehouse 2004a), C. erio-
cephalina Cham. (Graaff-Reinet, see C. bolusii Diels
ex C. Whitehouse below, and Amatole Mtns, Phillipson
1987), C. graminea L.f. (Grahamstown, Weimarck 1934),
C. ilicifolia L. (Grahamstown, Weimarck 1934), C. mon-
tana Weim. (see C. bolusii below and Graaff-Reinet,
Weimarck 1934) and C. ruscifolia L. (Namaqualand,
Weimarck 1934). The presence of C. juniperina L.f
north of Nieuwoudtville is doubtful, whereas C. repens
Schltr. is regarded now as being found only outside the
CFR (Whitehouse 2004a). Therefore, the total number
* Bolus Herbarium, University of Cape Town, 7700 Rondebosch.
Present address: The Royal Horticultural Society Garden, Wisley,
Woking, Surrey GU23 6QB, England, UK.
** Bolus Herbarium, University of Cape Town, 7700 Rondebosch.
Present address: 25 Bon Esperance, La Vie est Belle, 7550 Sonstraal
Heights, South Africa.
MS. received: 2006-01-18.
of species found within the CFR is 124, of which 109
are endemic to the area.
1 . CUffortia anthospermoides Fellingham, sp. nov.,
C. ramosissimae Schltr. affinis, sed intemodiis brevibus,
brachyblastis imbricatis, staminibus 9-12 notabilis.
TYPE. — Western Cape, 3419 (Caledon): Caledon Dis-
trict, Walker Bay area, Grootbos Private Nature Reserve,
slight northern slope, 150 m, (-CB), 31 Aug. 1996,
Fellingham 1691 (BOL, holo.!; K!, NBG!, PRE!).
C. sp. 2 sensu Fellingham: 614 (2000).
Medium, erect shrub, up to 1 m high, monoecious
but with alternating male and female phases; young
branches reddish with indumentum of long white hair,
turning greyish brown, becoming glabrous, hair bases
remaining as pustules, bark splitting and peeling, ageing
to a smooth glabrous reddish brown surface with persis-
tent patent, woody sheaths of fallen leaves with central
nerves prominent, giving appearance of thorny petioles;
intemodes ± 10 mm long, shorter than leaves, giving
plant a leafy appearance. Leaves trifoliate; leaflets 5.5-
7.0 X 0. 9-1.0 mm, straight to falcate to twisted giving an
untidy appearance; margins scabrid to minutely denticu-
late; lamina bright green, glabrous adaxially, with occa-
sional long hairs on margin and abaxially; sheath 1-2
mm long below central leaflet, diminishing distally, gla-
brous except for a few long cilia on margins, 3-nerved,
medially herbaceous, laterally pinky white membranous;
stipules represented by the membranous edges of sheaths
forming rectangular ‘wings’. Flowers solitary, ebracte-
ate, in axils of vegetative leaves separated by long inter-
nodes. Male flowers', bracteoles ovate-lanceolate, 2.2 x
1.5 mm, glabrous, scarious, whitish; pedicel and recepta-
cle 1 mm long, glabrous; sepals 3(or 4), linear-lanceolate,
3. 7^. 3 X 2. 6-3.0 mm, green with occasional maroon
longitudinal lines, prominently mucronate even in bud,
adaxially with obvious locking mechanism consisting
of subapical tuft of crisped papillae; stamens (9)10-12;
filaments 5 mm long, maroon, glabrous; anthers 0.7-0. 8
mm long, maroon-red with white connective. Female
flowers', bracteoles ovate-lanceolate, 3. 4-3. 6 x 2. 0-2. 3
mm, membranous, translucent, sheathing fruit, inner
bracteole with apical one fifth part reflexed, outer with a
tuft of retrorse hair at base; pedicel absent up to 0.6 mm
long in fruit, glabrous; sepals 3, broadly elliptic, 1. 9-2.0
10
Bothalia37,l (2007)
FIGURE 1. — Cliffortia anthospermoides. A, branch: Aa, female flower in situ; Ab, fruit in situ; Ac, male flower in situ. B, leaf; C, female flower;
D, fruit; E, fruit with sepals and bracts in situ; F, male flower; A, D, E, Fellingham I545\ B, C, Fellingham 1529\ F, Fellingham 1634. Scale
bars: A-E, I mm; F, 3 mm. Artist: A.C. Fellingham.
Bothalia37,l (2007)
11
X 1.5-1. 6 mm, spreading, mucronate, glabrous but for
papillate locking mechanism ventrally near apex, fuga-
cious; immature receptacle ellipsoid, 1.5 mm x 0.8 mm,
pale green with occasional longitudinal reddish lines;
style solitary, 0.2-0. 3 mm, pale green, glabrous; stigma
1.8-2. 2 mm long, deep maroon-red, fimbriate. Achene
ellipsoid, 1.9-2. 1 x 1.5-1. 6 mm, apex with small inden-
tation containing staminodes and ridged by remains of
fallen sepals, apical two thirds with ± 12 low longitudi-
nal ridges, green, tinged with maroon where exposed to
sun, lower third smooth, base concave with central pedi-
cel scar surrounded by a ring of stiff hairs. Figure 1.
Diagnostic characters and affinities', at first sight,
this species can be mistaken for a member of the genus
Anthospermnm (Rubiaceae). However, on closer exami-
nation the alternate arrangement of shoots becomes evi-
dent. Morphologically it appears closest to Cliffortia
ramosissima Schltr. but the habit of that species is lower
and spreading, not upright as in C. anthospermoides.
Furthermore, the leaves of C. anthospermoides are larger
and more tightly packed on the stems and the species
generally has more stamens in the mate flowers: 9-12, as
opposed to C. ramosissima, which has fewer than nine.
The distributions of the two species do not overlap.
Superficially the flowers appear to be borne singly in
the axils of normal vegetative leaves, particularly those
at the top of shoots. On closer inspection of the implan-
tation sites of these flowers, however, the presence
of a growth point next to each flower has been noted.
Furthermore, the axils of leaves lower down on the veg-
etative shoots contained well-developed short shoots,
bearing the bracts of a lower (fallen) flower as well as
a second developing flower above them. From the size
of the empty bracts it can be deduced that these sub-
tended a female flower, whereas the second flower was
male. Axils in the intermediate zone contained fruits in
the lower bracts with very young male flowers above,
whereas the axils in the tops of the shoots contained
young male flowers. From the emerging pattern of male
and female flower sites, it can be deduced that the first
flowers of the season to appear on any particular plant,
would be female. As a result of their diminutive size and
obscure arrangement, these would go unnoticed except
for the showy red styles. The shedding of the styles
completely obscures the female phase, whereas the male
phase which follows is patently obvious by its showy
stamens, thus leading a casual observer to the erroneous
conclusion that the plant is a male specimen of a dioe-
cious species.
Habitat, fynbos on brown, sandy soil on slight slopes;
altitude 50-250 m.
Distribution', only known from the Gansbaai area,
Caledon District (Figure 2).
Conservation status: four populations are known.
The population at Woest Arabie has been ± destroyed by
road widening, the one at Danger Point is under threat of
development and the one at Wortelgat has been severely
invaded by Acacia cyclops', only the population in the
Grootbos Nature Reserve is protected.
Etymology: an early Elsie Esterhuysen collection was
annotated by her as having ‘the aspect of Anthospermum
aethiopicum' ', an apt description as the tightly packed
leaves obscure the alternate arrangement of the leaves
which would distinguish it from an Anthospermum.
2. Cliffortia cruciata C.Whitehouse, sp. nov., C.
ramosissimae Schltr. primo aspectu similis, sed nullis
petiolis, sepalis quatuor, costis acheniorum manifesto
quatuor differt.
TYPE. — Western Cape, 3319 (Worcester): Worcester
District, Jonaskop, Wildepaardeberg, just below sandy
plateau, 900 m, (-DC), 15 Sept. 2000, Whitehouse 137
(BOL, holo.!; K!, MO!, NBGI, Zl).
Low, erect shrub, up to 0.3 m high, killed by fire;
densely divaricately branched, forming brachyblasts;
young stems 0.6-0. 9 mm wide, tinged reddish, hairy;
stem hairs adpressed upwards, 0. 2-0.4 mm long.
Leaves trifoliate; leaflets linear, 2. 7-3. 7 x 0.4-0. 6 mm,
base noticeably swollen and bulbous, apex 0. 1-0.2 mm
long, margins rounded but slightly grooved, entire and
smooth; lamina chartaceous, 0.3-0. 5 mm thick, slightly
curved upwards and towards stem, green with two paler
stripes on each side, glabrous; sheath 0. 7-1.0 mm long,
abaxially glabrous, adaxially hairy; stipules 0.3-0. 8 mm
long, free, margins smooth; petiole absent. Flowers soli-
tary at base of undifferentiated leaves; bracteoles hairy
on keel, margins ciliate; sepals 4, glabrous. Male flow-
ers: bracteoles 1.2-1. 5 mm long; pedicel and receptacle
0.5-0. 8 mm long; sepals broadly ovate, 3.4-4.4 x 1.3-
2.2 mm, acute to acuminate at apex, tips not completely
separating at anthesis; stamens 6 or 7; filaments 1. 2-2.0
mm long, red; anthers brownish red. Female flowers:
bracteoles 1.3- 1.7 mm long, shorter than immature
receptacle; sepals 0.8-1. 2 x 0.2-0. 4 mm, erect; carpel 1;
stigma 1.4— 2.6 mm long, red to pinkish, feathery, hidden
within leaves; immature receptacle 1.3-2. 7 x 0. 6-1.1
mm, glabrous, clearly ribbed. Achene broadly ellipsoid,
3. 8^. 7 X 1. 7-2.0 mm, medium brown, glabrous; ribs 4,
rounded, ± 0.6 mm wide, slightly tuberculate. Flowering
time: around September. Figure 3.
Diagnostic characters and affinities: closely allied to
Cliffortia subsetacea (Eckl. & Zeyh.) Diels ex Bolus &
Wolley Dod, but this is only clearly discernible by exam-
ination of the achenes, which are very similar except
that the ribs on C. cruciata are not curved. In general
appearance the species looks more similar to C. ramo-
sissima with its short, but comparatively broad and flat,
very slightly curved leaves. Apart from the achene, the
two species can be easily separated because C. cruciata
lacks any evidence of a petiole and the flowers have four
sepals and four prominently ribbed achenes.
FIGURE 2. — Known distribution of Cliffortia anthospermoides, 'tV; C.
ferricola. A; and C. perpendicularis, O, in South Africa.
12
Bothalia37,l (2007)
FIGURE 3. — Cliffortia cmciata. A, branch with achenes, x 2; B, branch
with female flowers, x 4; C, achene, x 10.
Habitat: fynbos in full sun, on well-drained soils on
deep sandy plateau; acid sands from Table Mountain
Series; altitude 900-1 000 m.
Distribution: a very narrow endemic restricted to the
sandy plateau of the Wildepaardeberg on the northern
slopes of Jonaskop in the Riviersonderend Mtns (Figure 4).
FIGURE 4. — Known distribution of Cliffortia cruciata, A; C. sparsa,
o; and C. weimarckii, A, in South Africa.
Conservation status: very narrow endemic, only
known from a very particular habitat on a single moun-
tain, although relatively inaccessible, it could be threat-
ened by over-frequent fires.
Etymology: cruciata, Latin for crosswise, referring
to the shape that the four prominent ribs of the achene
make when viewed in cross section.
3. Cliffortia ferricola C.Whitehouse, sp. nov., ex
affinitate C. ruscifoliae L. et specierum affinium, sed foliis
saepissime profunde bilobis vel trilobis distinguenda.
TYPE. — Western Cape, 3419 (Caledon); Caledon
District Kogelberg, S slopes of Mt Horeb, Kleinmond-
Highlands road, 200 m, (-AC), 27 Sept. 2000, White-
house 145 (BOL, holo.!; NBG!).
C. versiformis C.Whitehouse (2003), manuscript name.
Medium, erect shrub, up to 1.5 m high, resprouting
after fire and spreading clonally; densely divaricately
branched, forming brachyblasts; young stems 1.2-1 .7
mm wide, tinged reddish, hairy; stem hairs adpressed
upwards, 0.2-0. 7 mm long. Leaves usually unifoliate
and deeply toothed, but occasionally bifoliate or trifo-
liate; leaflets broadly ovate to lanceolate, 5.3-10.0 x
1.7-3. 5 mm, apex sharply acuminate to pungent, 0.7- 1.5
mm long, margins turned upwards to rounded, smooth
and entire or more frequently with 2 teeth becoming
deeply 3-lobed, lobes 0.6-^. 9 mm long, straight; lam-
ina 4-7-veined from base, cartilaginous, 0. 3-0.4 mm
thick, midrib prominent abaxially against lamina, curved
downwards and away from stem, glabrous adaxially,
hairy abaxially on very young grov^fih but soon becom-
ing glabrous, hairs 0.3-0. 7 mm long; sheath 1.0-1. 8 mm
long, abaxially glabrous, adaxially scattered adpressed
hairs; stipules 0.7-1. 3 mm long, free, margins smooth
to ciliate; petiole absent. Flowers solitary in axil of
undifferentiated leaves; bracteoles hairy on keel with
margins ciliate or not; sepals 3, hairy abaxially. Male
Bothalia37,l (2007)
13
flowers: bracteoles 4. 0^.6 mm long; pedicel and recep-
tacle 1.1-1. 4 mm long; sepals broadly ovate, 6.4— 6.8 x
2.2- 3. 0 mm, acute to acuminate at apex; stamens 9 or
10; filaments 8.8-10.1 mm long, red; anthers brownish
red. Female flowers: bracteoles 3. 1^.4 mm long, longer
than immature receptacle; sepals ovate, 3.2^. 1 x 0.8-
1.3 mm, recurved; carpel 1; stigma 6. 7-8. 7 mm long,
red, feathery, prominent above leaves; immature recep-
tacle 2.4— 2.9 X 0.9-1. 2 mm, glabrous, clearly ribbed.
Achene narrowly ellipsoid and slightly curved, 3. 5-4. 8 x
1.2- 1. 5 mm, medium to dark brown, glabrous; ribs 9-13,
rounded to acute, 0. 1-0.2 mm wide; elaiosome swollen
around base. Flowering time: September-February, pos-
sibly sporadically all year. Figure 5.
Diagnostic characters and afltnities: Clijfortia ferri-
cola is clearly closely related to the very common and
widespread C. ruscifolia, but has smaller leaflets, which
are often deeply divided, sometimes almost into two or
three leaflets. The habit is also characteristic, often form-
ing erect, strongly ascending stems so that the plants are
narrowly columnar. Plants are more frequently monoe-
cious than C. ruscifolia.
Habitat: well-drained, clayish soil of ferricretes from
shale bands of Table Mountain Series or from Bokkeveld
Series, in full sun; altitude 150-250 m.
FIGURE 5. — Cliffortia ferricola. A, branch with female flowers, x 2; B,
leaf, X 4; C, female flower, x 4; D, achene, x 6.
Distribution: restricted to ferricretes in the Bot River
Valley between Houwhoek, Kleinmond and Hermanns
(Figure 2).
Conservation status: known from only a few localities
in a small area, one of which is near a highway junction
and under threat of development.
Etymology: ferricola, Latin for dweller on iron, refer-
ring to the habitat of this species on ferricrete-derived
soils. The name was suggested by Nick Helme, who has
found this species in several localities around the Bot
River growing on ferricrete.
4. Cliffortia dentata Willd., Species plantarum 2,
edn 4: 842 (1806); DC.: 596 (1825); Harv.: 299 (1862);
Weim.: 20, fig. lA-F (1934); Levyns: 445 (1950);
Fellingham: 612 (2000). Type: Cape of Good Hope,
without further information (B-W, holo.).
Low, sprawling or trailing, semi-herbaceous shrub
up to 0.7 m high, killed by fire; densely divaricately
branched, only forming long branches; young stems 0.3-
0.7 mm wide, hairy; stem hairs adpressed downwards,
0.2-1. 1 mm long. Leaves trifoliate; leaflets obovate,
4.2- 10.3 X 2. 3-6.4 mm, apex acute to rounded, up to
0.4 mm long, margins flat except for apices which are
sometimes recurved, with up to 5 lobes or teeth; outer
leaflets often narrower than middle one and sometimes
untoothed; lamina herbaceous, midrib slightly prominent
abaxially against lamina, curved upwards and towards
stem, glabrous; sheath 0.4— 0.9 mm long, abaxially hairy,
adaxially with a fringe of long straight hairs; stipules
1.9^.0 mm long, partially joined on reverse side of
stem, margins ciliate; petiole absent. Flowers solitary in
axil of undifferentiated leaves; bracteoles glabrous on
keel, margins ciliate; sepals 4, glabrous. Male flowers:
bracteoles 1.6-2. 3 mm long; pedicel and receptacle gla-
brous; sepals broadly ovate, 3. 5-3. 8 x 1.5-2. 0 mm, with
long attentuate apices; stamens 8; filaments 4. 0^.3 mm
long, red; anthers brownish red. Female flowers: bracte-
oles 1.5-3. 4 mm long, longer than immature receptacle;
sepals ovate, 1.2-2. 2 x 0. 5-0.9 mm, strongly recurved;
carpels 2; stigma 2. 1-2.6 mm long, greenish white,
feathery; immature receptacle 0. 9-1.1 x 0. 7-1.0 mm,
glabrous, smooth. Achene slightly flattened, 2. 2-2. 8 x
1.0-1. 2 mm, pale brown, glabrous, unribbed but with
a central groove dividing two carpels. Flowering time:
April-October, possibly as late as December.
4a. var. dentata
Middle leaflet 4. 2-8. 5 x 2.9-6. 1 mm; outer leaflets,
4.3- 10.3 X 2.3-5. 1 mm; all leaflets 3-5-lobed or toothed,
middle lobe 0.6-1. 5 x 0.9-2 mm.
Habitat: shady slopes in gulleys and on damp south-
facing cliffs on well-drained soils from Table Mountain
Series; altitude 300-1 450 m.
Distribution: found between Wemmershoek Mtns
in the north to the Helderberg and Hottentots Holland
Mtns in the south, with an outlying population on the
Cape Peninsula, where it is only known from the eastern
slopes of the saddle between Table Mtn and Devil’s Peak
(Figure 6).
14
Bothalia 37,1 (2007)
Conservation status: an uncommon species but its
localities are well conserved and inaccessible; however,
being a seeding species, too frequent fires on Devil’s
Peak could threaten the Cape Peninsula population.
Etymology: dentata, Latin for toothed, referring to the
apices of the leaves that can appear like the teeth of a
saw.
4b. var. gracilis (Hai'v.) C.Whitehouse, comb,
et stat. nov. Type; Western Cape, 3319 (Worcester):
Worcester District, Du Toit’s Kloof, 610-915 m, (-CA),
1839, Drege 145b (K, holo.!; Bf, L, S, W).
C. gracilis Harv. in Flora capensis 2: 299 (1862); Weim.: 22 (1934)
pro parte; Fellingham: 612 (2000) pro parte.
Middle leaflet 4. 5-8. 4 x 4. 1-6.1 mm; outer leaflets
4. 7-9. 5 X 3. 0-6. 4 mm, untoothed; middle leaflet 3(-5)-
lobed or toothed, middle lobe 0.6-1 x 0.6-1. 2 mm.
Diagnostic characters and affinities: Cliffortia graci-
lis was considered a distinct species by Weimarck.
However, the populations in the Langeberg have con-
sistently smaller leaves and are considerably disjunct
from the Du Toit’s Kloof population (the type local-
ity). In terms of size of the leaves and flowers, the Du
Toit’s Kloof population falls within the range of varia-
tion shown by C. dentata, and only differ on the degree
of dentation at the apex. Furthermore, the discovery of a
population of the latter species from the Wemmershoek
Mtns (C. Whitehoiise 14), a mere 10 km or so from
Du Toit’s Kloof, means that the geographical distance
between the populations is much smaller than previously
thought. Therefore, C. gracilis is here reduced to vari-
etal status and a new name, C. gracillima C.Whitehouse,
given to the Langeberg populations.
Cliffortia dentata and C. gracillima form a pair of
closely related species which create decumbent trail-
ing mats on shady rocky slopes. The trifoliate, glabrous
leaves, along with the general habit, distinguish this
group from any other Cliffortia. Var. gracilis can be dis-
tinguished from the typical variety by the middle leaflet,
which is often only 3-lobed, with the middle lobe being
much narrower than the outer two, and untoothed outer
leaflets.
Habitat: shady slopes in gulleys and on damp south-
facing cliffs on well-drained soils from Table Mountain
Series; altitude 600-1 700 m.
FIGURE 6. — Known distribution of Cliffortia cienlaia var. dentata.
A; C. dentata var. gracilis, A; and C. gracillima, o, in South
Africa.
Distribution: only known from Molenaarsberg and
Upper Wellington Sneeukop to the north of Du Toit’s
Kloof (Figure 6).
Conservation status: only known from a small inac-
cessible area to the north of Du Toit’s Kloof, but it could
be threatened by over-frequent fires as it probably only
survives as seed.
Etymology: gracilis, Latin for thin, slender, referring
to the stems that are too weak to hold the plant upright
and trail over the rocks.
5. Cliffortia gracillima C.Whitehouse, sp. nov., a
C. dentata Willd. var. gracili (Harv.) C.Whitehouse foliis
minoribus differt.
TYPE. — Western Cape, 3321 (Ladismith): Riversdale
District, Garcia’s Pass, 395 m, (-CC), Oct. 1904, Bolus
11271 (BOL, holo.!; PRE!).
C. dentata sensu Muir: 56 (1929), non Willd.
C. gracilis sensu Weim.: 22 (1934) pro parte; Fellingham: 612
(2000) pro parte.
Illustration: Weim.: fig. 1G-L(1934).
Low, sprawling or trailing, semi-herbaceous shrub
up to 0.3 m high, probably killed by fire; densely divari-
cately branched, only forming long branches; young
stems 0.2-0. 6 mm wide, hairy; stem hairs adpressed
downwards, 0.2-0. 8 mm long. Leaves trifoliate; middle
leaflet obovate, 3. 1-5.4 x 2. 2^. 6 mm, apex acute to
mucronate, up to 0.4 mm long, margins flat except for
apices which are sometimes recurved, entire or up to
3-lobed, middle lobe much smaller and narrower than
outer two, 0.4-0. 9 x 0.3-1. 4 mm, outer leaflets elliptic
to obovate, 3.7-6. 1 x 1. 6-3.1 mm, almost always entire;
lamina herbaceous, up to 0.1 mm thick, midrib slightly
prominent abaxially against lamina, curved upwards and
towards stem, glabrous; sheath 0.2-0. 6 imn long, abaxi-
ally hairy, adaxially with a fringe of long straight hairs;
stipules 1. 6-3.1 mm long, partially joined on reverse
side of stem, margins ciliate; petiole absent. Flowers
solitary at base of undifferentiated leaves; bracteoles
glabrous on keel, margins ciliate; sepals 4, glabrous.
Male flowers: pedicel and receptacle glabrous; sepals
broadly ovate with long attentuate apices; filaments red;
anthers brownish red. Female flowers: bracteoles 1.7-
3.2 mm long, longer than immature receptacle; sepals
ovate, strongly recurved; carpels 2; stigma 2. 6-3. 2 mm
long. Achene slightly flattened, 2. 2-3. 8 x 0.9-1. 5 mm,
pale brown, glabrous, unribbed but with a central groove
dividing two carpels. Flowering time: July-October.
Figure 7.
Diagnostic characters and affinities: previously
included as part of C. gracilis, which is now regarded as
a variety of C. dentata. It differs from that species by its
much smaller leaves, as well as having a clear disjunc-
tion in distribution and therefore occurs in areas where
summer rainfall is more common.
Habitat: shady areas in gulleys on well-drained soils
from Table Mountain Series; altitude 250-1 400 m.
Distribution: known from a few localities in the
Langeberg from Leeurivierberg to Garcia’s Pass, with an
Bothalia 37,1 (2007)
15
H Bolus al-stko-Afr^came
Etoft* ^CCUUn^
outlying population on the Waboomsberg near Montagu
(Figure 6).
Consei’vation status: only known from four localities
but areas in between are poorly explored and it is possi-
ble that more populations occur, it grows in inaccessible
areas but might be threatened by too frequent fires.
Etymology: gracillima, Latin for very thin and slen-
der, referring to the stems and allusion to its previ-
ous inclusion within C. gracilis, from which it is much
smaller in all its parts.
6. Cliffortia perpendicularis C. Whitehouse, sp.
nov., ex affinitate C. falcatae L.f. et C. ramosissimae
Schltr., sed ab utraque forma ramifactionis distincta.
TYPE. — Western Cape, 3419 (Caledon): Bredasdorp
District, Elim, Geelkop Nature Reserve, W slope of hill,
120 m, (-DB), 3 Sept. 2001, C. & A. Whitehouse 259
(BOL, holo.!; K!,NBG!).
Sprawling shrub up to 0.5 m high but often semi-
prostrate, killed by fire; divaricately branched and form-
ing brachyblasts that spread at right angles to main stem,
but few long branches developing; young stems 0.9-1. 6
mm wide, tinged reddish, hairy; stem hairs spreading
or adpressed upwards, 0.2-0. 5 mm long. Leaves trifoli-
ate; leaflets oblong to linear, outer ones sickle-shaped,
3. 5-7. 5 X 0.7-1. 1 mm, apex sharply acuminate, 0.3-0. 6
mm long, margins flat, entire and smooth; lamina charta-
ceous, 0.2-0. 5 mm thick, curved upwards and towards
stem, green but with two paler stripes on either side of
midrib abaxially, glabrous; sheath 1. 1-2.0 mm long,
abaxially glabrous, adaxially glabrous except for fringe
of hairs at apex; stipules 0.8- 1.5 mm long, free, margins
ciliate; petiole present, 0.6-1. 4 mm long. Flowers soli-
tary at base of undifferentiated leaves; bracteoles short-
hairy on keel, margins ciliate; sepals 3, glabrous. Male
FIGURE 7. — Type specimen of
Clijfortia gracillima.
flowers unknown. Female flowers: bracteoles 2. 7-3. 8
mm long, longer than immature receptacle; sepals oblong
to linear, 3. 5^. 5 x 0. 5-1.0 mm, spreading; carpel 1;
stigma 2. 5^. 9 mm long, red, feathery, hidden amongst
incurved leaves; immature receptacle 1.1-1. 8 x 0.7-1. 2
mm, sparsely and very short-hairy. Achene ellipsoid to
cylindrical, 3. 5^. 6 x 1.8-2. 6 mm, medium brown but
sometimes appearing greyish on account of being cov-
ered by membranous layer, glabrous or sparsely and very
FIGURE 8. — Cliffortia perpendicularis. A, branch, x 1; B, leaf, x 4.
16
Bothalia37,l (2007)
short-hairy, irregularly acutely ribbed beneath membra-
nous layer. Flowering time: September. Figure 8.
Diagnostic characters and affinities: the leaves of
this species appear most similar to Cliffortia arcuata
Weim., but that species is only found on the mountains
surrounding the Little Karoo. Related species that occur
in the vicinity of C. perpendiciilaris include C. ramosis-
sima and C. falcata L.f The leaves are generally longer
and more curved than C. ramosissima and more sharply
pointed than C. falcata. However, the most distinc-
tive feature about C. perpendicularis is the very sparse
branching (excluding the brachyblasts). It has a spindly,
lax habit, so much so that often only a couple of long
branches exist and the main stem is unable to bear the
weight of the plant, resulting in its sprawling amongst
the surrounding vegetation. This is in contrast to C. arcu-
ata and C. falcata, which have erect ascending branches,
and C. ramosissima which has an intricate divaricate
branching pattern. The distinctive branching pattern is
accentuated by the brachyblasts, which appear to project
from the stem at right angles.
The species is possibly of hybrid origin. In both the
localities where it has been found, C. falcata and C.
ramosissima are found growing within a few kilometres.
However, seed is clearly viable, as the species occurs fre-
quently in the habitats even though the plants are killed
by fire; specific status is considered the most appropriate
treatment for this taxon. Male flowers have not yet been
found, so it is presumed that most of the reproduction by
seed is apomictic and therefore asexual.
The three populations of this species appear ± iden-
tical despite the contrasting soil types (gritty acid sands
from Table Mountain sandstone on the Potberg and clay-
ish silcretes and ferricretes over Bokkeveld shales near
Elim) and they both share the unusual sprawling habit
that is diagnostic for this species. Population level stud-
ies of the species would be needed to determine if they
share a common ancestry or represent a case of two
cryptic apomictic species derived from hybridization.
Habitat: fynbos in full sun on acid sands from Table
Mountain Series or clayish soil of Bokkeveld Series;
altitude 100-300 m.
Distribution: known only from three localities: two
populations close to Elim and one on Potberg (Figure 2).
Conservation status: a poorly known species and only
known from three very small areas, one that is threat-
ened by agriculture and quarrying, while the others are
in danger of invasion by alien vegetation.
Etymology: perpendicularis, Latin for at right angles,
referring to the way the branches and brachyblasts
appear to join the stem at right angles to it.
7. Cliffortia sparsa C. Whitehouse, sp. nov., planta
inter C. atratam Weim. et C. sericeam Eckl. & Zeyh. ali-
quam intermedia sed foliis sparse pubescentibus digno-
scenda.
TYPE. — Western Cape, 3419 (Caledon): Caledon
District, Hottentots Holland Nature Reserve, around
Nuweberg Forest Station, 560 m, (-AA), 25 Oct. 2001,
Whitehouse 2H3 (BOL!; NBG!).
C. pterocarpa sensu Weim.: 62, fig. 16K (1934) pro parte, non
(Harv.) Weim. sensu stricto.
Medium, erect shrub up to 1.2 m high, killed by fire;
densely divaricately branched, forming brachyblasts;
young stems 0. 7-1.0 mm wide, sometimes tinged red-
dish, hairy; stem hairs adpressed upwards, 0.3-0. 6 mm
long. Leaves trifoliate; leaflets linear to needle-shaped,
6.0-10.1 X 0.6-1. 1 mm, apex acute to sharply acumi-
nate, 0. 2-0.4 mm long, margins flat, entire and smooth;
lamina chartaceous, 0.2-0. 5 mm thick, midrib promi-
nent abaxially, ± straight to curved downwards and away
from stem, sparsely hairy adaxially and abaxially, hairs
0. 1-0.6 mm long; sheath 1. 2-2.1 mm long, abaxially
glabrous, adaxially glabrous except for fringe of hairs
at apex; stipules 1. 1-2.5 mm long, free, margins smooth
to serrulate; petiole absent. Flowers solitary in axil of
undifferentiated leaves; bracteoles hairy on keel, mar-
gins serrate to very shortly ciliate; sepals 3, hairy abaxi-
ally. Male flowers: bracteoles 2-A mm long; pedicel and
receptacle 0.6-0. 8 mm long; sepals broadly ovate, 4.8-
6.0 X 1.9-2. 5 mm, acute to acuminate at apex; stamens
10 or 11; filaments 6.4-7. 8 mm long, red or greenish
white; anthers yellow. Female flowers: bracteoles 2.6-
4.2 mm long, longer than immature receptacle; sepals
ovate, 2.2^. 1 x 0.9-1. 5 mm, recurved; carpel 1; stigma
2. 9-5.0 mm long, red to pinkish white, feathery, promi-
FIGURE 9. — Cliffonia sparsa. A, branch with achenes, x 2; B, leaf, x
4; C, achene, x 12.
Bothalia37,l (2007)
17
nent above leaves; immature receptacle 1. 7-2.0 x 0.8-
1.0 mm, glabrous, clearly ribbed. Achene ellipsoid, 3.5-
4.1 X 1.1-1. 5 mm, sometimes slightly curved, medium
to dark brown, glabrous; ribs 9-12, acute, 0. 1-0.3 mm
wide; elaiosome swollen around base. Flowering time:
mainly August-November, once in May. Figure 9.
Diagnostic characters and affinities: specimens of
this species have in the past generally been attributed
to the poorly delimited species, Cliffortia pterocarpa.
Molecular evidence suggests, however, that this species
may be of hybrid origin between C. atrata Weim. and
C. cristata Weim. or C. sericea Eckl. & Zeyh. of sec-
tion Inflexae (Whitehouse 2003). In many respects it is
intermediate between the two putative parents, having
achenes and needle-shaped leaves most similar to C.
atrata, but having the hairs of section Inflexae. However,
it is common where it occurs and being a reseeder is
clearly able to reproduce, even if only apomictically, by
seed. Male and female flowers have been found and fur-
ther introgression with parent species is likely and would
explain the many intermediate forms that are found.
Habitat: well-drained clayish soil from shale bands of
Table Mountain Series in full sun; altitude 250-800 m.
Distribution: common along the lower slopes of the
mountains from Sir Lowry’s Pass to Franschhoek Pass
(Figure 4).
Conservation status: widespread and common through
its range, often occurring in disturbed areas.
Etymology: sparsa, Latin for sparse, referring to the
hairs on the leaves, which although appearing glabrous
are in fact covered with short, sparse hairs.
8. Cliffortia weimarckii C. Whitehouse, sp. nov.,
C. eriocephalinae Cham, similis sed foliis longioribus
angustioribus rigidioribus, apicibus acutis differt.
TYPL. — Western Cape, 3319 (Worcester): Worcester
District, Matroosberg, Spekrivierskloof, stony L slopes
FIGURE 10. — Cliffortia weimarckii. A, branch, x 1; B, leaf, x 4; C,
achene, x 4.
above Ski Hut, 1 950 m, (-BC), 27 Jan. 2001, Whitehouse
757 (BOL, holo.!;NBG!).
Erect shrub up to 0.9 m high, resprouting after fire and
spreading clonally; densely divaricately branched, form-
ing brachyblasts; young stems 1.1-1. 5 mm wide, usually
tinged reddish, densely hairy; stem hairs spreading, 0.9-
1.5 mm long. Leaves trifoliate; leaflets narrowly oblong
to needle-shaped, 6.2-13.4 x 0.8-1. 2 mm, base con-
tracted abruptly to form a pseudopetiolule, apex acute to
acuminate, 0. 1-0.2 mm long, margins inrolled abaxially,
entire; lamina chartaceous to rigid, 0. 1-0.3 mm thick,
midrib prominent abaxially against lamina, ± straight or
curved upwards and towards stem, densely greyish hairy
adaxially, hairs 0.4- 1.7 mm long, densely whitish hairy
abaxially, hairs 0. 2-0.4 mm long, sheath 2.7-3. 1 mm
long, abaxially hairy, adaxially densely hairy; stipules
0.2-0. 4 mm long, free, margins ciliate; petiole absent.
Flowers solitary at base of undifferentiated leaves; brac-
teoles hairy; sepals 3, densely hairy abaxially. Male
flowers: bracteoles 3. 6^.0 mm long; pedicel and recep-
tacle 0.9-1. 3 mm long; sepals broadly ovate, 5. 0-5. 8 x
1.8-2. 3 mm, acute to acuminate at apex; stamens 9-11;
filaments 4. 6-5. 7 mm long, red; anthers brownish red.
Female flowers: bracteoles 2. 8^. 8 mm long, longer
than immature receptacle; sepals linear, 1.5-2. 9 x 0.2-
0.5 mm, erect; carpel 1; stigma 4. 1-6.5 mm long, red,
feathery, prominent above leaves; immature receptacle
1. 6-2.2 X 0.7-0. 9 mm, glabrous, clearly ribbed. Achene
ellipsoid, 3. 2-3.7 x 1.1-1. 4 mm, pale greyish brown,
glabrous; ribs 1 1-15, rounded to acute, less than 0. 1 mm
wide. Flowering time: January, although probably in the
neighbouring months as well. Figure 10.
Diagnostic characters and affinities: this species is
very similar in many respects to Cliffortia eriocepha-
lina but differs on account of its longer, narrower, stiffer
and pointed leaves. The leaves are similar in form to the
unrelated lowland species, C. stricta, but are much more
hairy and grey in appearance. C. eriocephalina and C.
weimarckii grow sympatrically in some places such as
Matroosberg, where they can be easily distinguished.
However, some specimens from the Baviaansberg
(Stokoe 4537, Esterhuysen 29806) and Cederberg
Langberg (Bond 1383) have leaves of intermediate
length, and without field observations on the popula-
tions, they are only tentatively attributed to this species.
Weimarck annotated some collections of this species
indicating that he thought they were a new species. It
therefore seems most appropriate to name this species in
his honour, especially considering his immense contribu-
tion to our current understanding of the genus.
Habitat: high altitude mountain fynbos on well-
drained acid sands from Table Mountain Series in full
sun; altitude 1 200-2 250 m.
Distribution: Hex River Mtns and the highest peaks of
the Koue Bokkeveld and Groot Winterhoek Mtns, possi-
bly also Baviaansberg and Cederberg Langberg (see note
above. Figure 4).
Conservation status: populations are scattered as it is
restricted to such high altitudes, however it is therefore
also not threatened, especially as it is a resprouter.
18
Bothalia37,l (2007)
Etymology: weimarckii is named after Henning
Weimarck who wrote the first full monograph of Cliffortia.
9. Cliffortia cuneata Dryand. in Aiton, Hortus kew-
ensis, edn 1, 3: 413 (1789); DC.: 595 (1825); Weim.: 129,
fig. 37A-F (1934); Fellingham; 609 (2000). Lectotype,
designated by Weimarck (1934): Western Cape, 3418
(Simonstown): False Bay, Robertson s.n. (BM, holo.!).
Medium to tall, erect shrub, up to 3 m high, killed
by fire; densely divaricately branched, forming long
branches and brachyblast when flowering; young stems
1.0-2. 5 mm wide, glabrous or slightly hairy but quickly
becoming glabrous; stem hairs adpressed upwards, 0. 1-
0.4 mm long. Leaves unifoliate; wedge-shaped to oblong,
13-41 X 4.9-13.9 mm, margins flat, only toothed at apex
with 2-6 broad straight teeth, 0.6-2. 9 mm long; lamina
3-7 -veined from base, chartaceous, 0. 1-0.6 mm thick,
midrib very prominent abaxially, curved upwards and
towards stem, slightly glaucous, glabrous; sheath 0.8-2. 3
mm long, abaxially glabrous, adaxially hairy or glabrous
except for fringe at apex; stipules 0.2-1. 6 mm long, free,
margins smooth; petiole absent. Flowers solitary at base
of undifferentiated leaves but often clustered together;
bracteoles hairy or glabrous on keel, margins smooth;
sepals 3, glabrous. Male flowers: bracteoles 6.2-8. 5 mm
long; pedicel and receptacle 1. 2-2.1 mm long; sepals
broadly ovate, 6.6-10.3 x 4. 0-5. 9 mm, acute to acumi-
nate at apex; stamens 27-36; filaments 11-14 mm long,
red; anthers yellow. Female flowers: bracteoles 3.8-10.5
mm long, longer than immature receptacle; sepals ovate
to obovate, 3. 7-5. 9 x 1.4-2. 9 mm, erect; carpel 1; stigma
6.5-11.0 mm long, greenish white, feathery, prominent
at base of the leaves; immature receptacle 1.9-5. 2 x 1.2-
2.9 mm, glabrous, smooth. Achene cylindrical to almost
globose, 4. 5-6. 8 x 1.5-3. 8 mm, medium to dark brown,
glabrous; ribs 6 or 12-14, rounded, 0. 1-0.8 mm wide.
Diagnostic characters and aflinities: the reverse,
long, almost triangular glaucous leaves, toothed only at
the truncate apex, are very distinctive on this medium-
sized species. This character is not only diagnostic but
emphasizes the isolated position that this species holds
within the genus as it has no close relatives.
However, while the typical form of the species has
large, almost globose, achenes with numerous ribs, the
achenes from the populations around Arieskraal are
smaller, more cylindrical in shape, with only six low
ribs. Furthermore, the leaves appear to be consistently
FIGURE 11. — Cliffortia cuneata var. cylindrica: A, branch with
achenes, x 2; B, achene, x 4. C. cuneata var. cuneata: C, achene,
x4.
FIGURE 12. — Known distribution of Cliffortia cuneata var. cuneata. A;
and C. cuneata var. cylindrica, A; C. cuneata var. cuneata and
var. cylindrica, ★, in South Africa.
smaller, although still within the variation shown by the
large-achened form of the species. This form is described
here as a new variety.
The fruit is so distinct that specific rank was consid-
ered. However, as the number of specimens examined
is still low, male flowers are unknown and a population
near Riviersonderend was of questionable status, varietal
rank was considered most appropriate for the present.
Etymology: cuneata, Latin for wedge-shaped or inver-
sely triangular, referring to the distinctive shape of the
leaves.
9a. var. cuneata
Leaves \6-^\ x 5.4—13.9 mm, margins toothed at
apex with 2-6 broad straight teeth, 0.9-2. 9 mm long;
lamina 3-7-veined from base. Male flowers as above.
Female flowers: bracteoles 8.2-10.5 mm long; sepals
ovate to obovate, 4. 2-5. 9 x 1.7-2. 9 mm; immature
receptacle 3. 0-5. 2 x 2. 0-2. 9 mm. Achene broadly ovoid
to almost globose, 6. 2-6. 8 x 2. 9-3. 8 mm; ribs 12-14,
rounded, 0.3-0. 8 mm wide. Flowering time: June-
December. Figure IIC.
Habitat: low slopes in fynbos on well-drained clayish
shale soils in full sun; altitude 0-950 m.
Distribution: found on lower slopes between the Paarl
side of Du Toit’s Kloof, Helderberg and Houw Hoek,
with an outlying population in Boesmanskloof in the
Riviersonderend Mtns (Figure 12).
Conservation status: widespread but scattered locali-
ties of quite extensive populations, several within nature
reserves; threats come from too frequent fires and inva-
sive alien vegetation.
9b. var. cylindrica C. Whitehouse, var. nov., a var.
cuneata acheniis subcylindricis sexcostatis differt.
TYPE. — Western Cape, 3418 (Simonstown): Caledon
District, Arieskraal, south-facing slopes on Bokkeveld
shale above Arieskraal Dam, 215 m, (-BB), 29 Nov.
2000, Whitehouse 166 (BOL!; NBG!).
Leaves 13-23 x 4. 9-7. 5 mm, margins toothed at apex
with 4 broad, straight teeth, 0.6-1. 6 mm long; lamina
3-5-veined from base. Male flowers unknown. Female
flowers: bracteoles 3. 8-6.0 mm long; sepals narrowly
ovate to linear, 3.7-4. 1 x 1. 4-2.0 mm; immature recep-
tacle 1.9-3 .4 X 1.2-1. 4 mm. Achene cylindrical, 4.5-
Bothalia37,l (2007)
19
5.0 X 1.5-1. 7 mm; ribs 6, rounded, 0. 1-0.3 mm wide.
Flowering time'. October-November. Figure llA, B.
Habitat', fynbos on south-facing shale slopes of the
Bokkeveld Series; altitude 250-^00 m.
Distribution', only known from Arieskraal in the
Kogelberg area, but possibly present near Riviersonderend
(see note above, Figure 12).
Conservation status', known from only one or two
very small populations in a very confined area much of
which has already been converted to orchards because
it grows on relatively fertile shale; the only population
that the first author has seen is on a slope too steep to be
used.
Etymology', cylindrica, Latin for cylindrical, referring
to the shape of the ± straight-sided achenes in contrast
with the typical variety, which are broadly ovate.
10. Cliffortia filifolia L.f., Supplementum plan-
tarum: 430 (1782); DC.: 596 (1825); Weim.; 68, fig.
17J-N (1934); Levyns: 447 (1950); Fellingham: 615
(2000). Type: Western Cape, 3318 (Cape Town): Cape
Peninsula, Table Mtn, E slopes, (-CD), Thimberg s.n.
(LINNf, holo.; CP, LD, UPS-THUNB).
Low, often sprawling, shrub, up to 1 m high, killed
by fire; densely divaricately branched, forming closely
overlapping brachyblasts; young stems 0. 5-1.0 mm
wide, sometimes tinged reddish, glabrous. Leaves trifoli-
ate; leaflets needle-shaped, 5.8-17.0 x 0.3-0. 8 mm, apex
sharply acuminate, 0.2-0. 9 mm long, margins rounded,
minutely serrulate to scabrid; lamina chartaceous, 0.2-
0.4 mm thick, with two pale stripes on either side of
midrib, glabrous; sheath 1. 3-3.0 mm long, abaxially gla-
brous, adaxially glabrous except occasionally for a few
hairs at apex; stipules 0.7-3. 1 mm long, free, margins
smooth to serrulate; petiole 0. 2-2.0 mm long. Flowers
solitary in axil of undifferentiated leaves; bracteoles gla-
brous on keel, margins smooth to serrate; sepals 3, gla-
brous. Male flowers', bracteoles 2.4-4. 3 mm long; pedi-
cel and receptacle 0.4-0. 7 mm long, glabrous; sepals
broadly ovate, 3. 2-5. 2 x 1.2-1. 8 mm, acute to acuminate
at apex; stamens 5 or 6; filaments 3. 4-5. 2 mm long, red;
anthers yellow. Female flowers', bracteoles 2. 5^.4 mm
long, longer than immature receptacle; sepals narrowly
ovate to linear, 1.7-2. 7 x 0.4-1. 0 mm, erect to spread-
ing; carpel 1; stigma 1.6-2. 5 mm long, greenish white,
feathery, hidden at base of leaves; immature receptacle
0.8-1. 2 X 0.4-0. 7 mm, glabrous, smooth. Achene ellip-
soid to cylindrical, 2. 8-5. 2 x 1.2-1. 7 mm, greyish to
medium brown, covered by membranous layer, glabrous,
obscurely ribbed; ribs 6-9, rounded, up to 0.1 mm wide.
Habitat', in the west it is found on shale bands of
Table Mountain Series, but east of Cape Agulhas it is
restricted to dune slacks; altitude 0-1 100 m.
Distribution', widespread though scattered from
Piketberg to Knysna.
Consen’ation status', widespread though scattered, but
no serious threats except possibly var. arenaria, which
grows in areas subject to development.
Etymology', filifolia, Latin for thread-leaved, referring
to the fine, needle-like leaves.
10a. var. filifolia
C. leptophylla Eckl. & Zeyh.: 270 (1836). Type: Western
Cape, 3319 (Worcester): Caledon District, Houwhoek Mtn, near
Knofflookskraal, (-AA), July, Ecklon & Zeyher 1752 (Bt, CP, G-DEL,
K!, L, PRE, UPS, S, W, Z).
Young stems 0. 6-1.0 mm wide. Leaflets 6.1-17.0 x
0.4-0. 8 mm, apex 0.2-0. 9 mm long; lamina 0.2-0. 4 mm
thick, curved upwards and towards stem; sheath 1. 3-3.0
mm long; stipules 0.7-3. 1 mm long; petiole 0. 2-2.0
mm long. Female flowers', bracteoles 2. 5^.4 mm long;
sepals 1.7-2. 7 x 0. 4-1.0 mm, erect to spreading; carpel
1; stigma 1.6-2. 5 mm long, greenish white, feathery.
FIGURE 13. — Known distribution of Cliffortia filifolia var. filifolia. A; and C. filifolia var. arenaria, o; C. filifolia var. arenaria and var. filifolia, •;
and C. bolus a, A, in South Africa.
20
Bothalia37,l (2007)
FIGURE 14. — Cliffortia filifolia var. arenaria. A, branch with
achenes, x 2; B, leaf, x 3; C, achene, x 10.
hidden at base of leaves; immature receptacle 0.8-1. 2 x
0.4— 0.7 mm. Achene 2. 8^.4 x 1.2-1. 6 mm. Flowering
time: May-September.
Habitat: well-drained soils on shale bands of Table
Mountain Series rocks in full sun; altitude 0-1 100 m.
Distribution: common but scattered on the lower
slopes of mountains from Piketberg in the north to
Caledon Swartberg in the south, including the Cape
Peninsula, with an outlying population on the Potberg in
De Hoop Nature Reserve (Figure 13).
10b. var. arenaria C.Whitehouse, var. nov., a vari-
etate typica folds plerumque brevioribus saepe deorsum
curvatis differt.
TYPE. — Western Cape, 3422 (Mossel Bay): George
District, sand dunes E of Wilderness, (-BA), 3 Dec.
\95\, Esterhuysen 19336 (BOL!; NBG!, PRE!).
Young stems 0.5-0. 6 mm wide. Leaflets 5. 8-6. 6 x
0.3-0. 6 mm, apex 0. 1-0.3 mm long; lamina 0.2-0. 3 mm
thick, often curved downwards and away from stem;
sheath 0.6-1 .1 mm long; stipules 0. 7-1.0 mm long; peti-
ole 0.3-0. 5 mm long. Female flowers: bracteoles 2. 5-2. 8
mm long; sepals 2. 3-2. 5 x 0.6-0. 8 mm. Achene 4. 7-5. 2
X 1 .6-1 .7 mm. Flowering time: March to August. Figure
14.
Diagnostic characters and affinities: Cliffortia fili-
folia is relatively distinctive amongst the numerous
needle-leaved speeies of Cliffortia. The presence of
a short petiole along with the fine leaflets is diagnos-
tic. However, within C. filifolia, there appear to be two
distinct ecotypes. The western variety, -var. filifolia, has
longer leaflets, which curve upwards towards the stem.
This variety generally occurs on clayish soils derived
from sandstones of the Table Mountain Series. The east-
ern variety described here, var. arenaria, is distinguished
by its shorter leaflets, which often curve in directions
away from the stem. It is almost entirely restricted to
sandy coastal areas, especially dune slacks. In the south-
ern Overberg [3419, Caledon District, Waterford Farm,
(-DA), Whitehouse 222] and Cape Flats [3318, Cape
Town, Penhill, (-DC), Helme 1594], the distinction
between the two varieties and their habitats is less clear,
otherwise species status might have been considered
more appropriate.
Habitat: wind-blown sand of dune slacks in full sun;
altitude 0-250 m.
Distribution: coastal areas from De Hoop Nature
Reserve as far as Knysna, with a possible record from
the sandplains of the Cape Flats (Figure 13).
Etymology: arenaria, Latin meaning growing on sand,
referring to the fact that it is found growing in dune
slacks.
1 1 . Cliffortia bolusii Diels ex C. Whitehouse, sp.
nov., a congeneribus stipulis nullis folds oblongis 3-7
mm longis planis apicibus rotundatis distinguitur.
TYPE. — Eastern Cape, 3224 (Graaff-Reinet); Graaff-
Reinet District, Nardouwsberg, (-BB), Apr. 1873, Bolus
2630 (BOL, holo.; Bf).
C. sp. 1 sensu Weim.; 158 (1934).
Illustration: Weim.: fig. 47A (1934).
Shrub, densely divaricately branched, forming brach-
yblasts; young stems hairy; stem hairs spreading to
curled. Leaves trifoliate; leaflets elliptic to oblong, 3-7
X 0.6-0. 8 mm, apex mucronate to rounded, margins
flat, entire and smooth; lamina chartaceous, straight to
curved downwards and away from stem, hairy adaxi-
ally and abaxially; sheath 1. 5-3.0 mm long, abaxially
hairy, adaxially glabrous except for fringe of hairs at
apex; stipules absent; petiole absent. Flowers unknown
but solitary bracteoles found at base of undifferentiated
leaves; bracteoles hairy. Figure 15.
Diagnostic characters and affinities: the species has
no clear affinities. The leaves bear a resemblance to a
trifoliate version of Cliffortia dichotoma Fellingham, but
are more rounded at the apex. Apart from its isolated
geographic locality there is no further reason to suggest
that it belongs to section Arborea. A solitary hairy brac-
teole was found in a leaf axil, but this does not reveal
any significant systematic information about the species.
Weimarck refrained from naming this species until
more infonnation was available, although he realised
that it was distinctive both taxonomically and biogeo-
Bothalia37,l (2007)
21
graphically. Seventy years on from his monograph, no
further collections have been made, but the trifoliate
leaves, lacking stipules, with flat oblong leaflets, are
diagnostic. Furthermore, it is important to recognize
the taxon formally to encourage further searches for the
species and to ensure that it is included in conservation
assessments for the area.
Habitat, unknown, although the mountain is gener-
ally montane grassland with rocky outcrops and dolerite
cliffs; altitude 1 700-1 800 m (but see note below).
Distribution: known only from a single collection
made on the Nardouwsberg near Graaff-Reinet (Figure
13).
Conservation status: not been collected since it was
first collected in 1873, though the Nardouwsberg is a
large mountain and it may still exist on some rocky out-
crops. The Nardouwsberg is subject to grazing and burn-
ing, making the chance of this species surviving away
from the protection of rocks unlikely.
Etymology: bolusii is named after Harry Bolus,
founder of the Bolus Herbarium at the University of
Cape Town and the only person to have ever collected
this species.
A very poorly known species with only a single sur-
viving fragment remaining of a collection made over
100 years ago. Unfortunately Bolus’s notes describing
his collecting trip to the Nardouwsberg were lost in a
shipwreck, as probably were further specimens account-
FIGURE 15. — Type specimen of Cliffortia bolusii.
ing for the paucity of the remaining fragment at BOL.
(The other specimen seen by Weimarck was at B and is
presumably now destroyed.) More accurate information
on where he collected is therefore unavailable. A cursory
search of the mountain did not reveal this species, but
three other species were found: Clijfortia eriocephalina,
C. montana and C. ramosissima.
The name bolusii is written on the herbarium sheet at
BOL and attributed to Diels, though he never published
it. Although the altitude is given as 5800 ft (± 1 770
m), this is also the altitude given for Bolus’s specimen
of C. eriocephalina (Bolus 851) from the same local-
ity. However, on the recent trip to the Nardouwsberg, C.
eriocephalina was only seen just below the summit, an
altitude of ± 2 300 m.
12. Cliffortia odorata L.f, Supplementum plan-
tarum: 431 (1782); Weim.: 152, fig. 45A-E (1934);
Levyns: 450 (1950); Fellingham: 610 (2000).
TYPE. — South Africa, without precise locality, ‘Cape
of Good Hope’, Thunberg s.n. (LfNNf, holo.; CP, UPS-
THUNB).
C. discolor Weim.: 202, fig. 16d-f (1948); Fellingham: 609 (2000),
syn. nov. Type: Western Cape, 3318 (Cape Town): Cape Peninsula,
Table Mm, 1 000 m, (-CD), Nov. 1884, Marloth 360 (PRE, holo.),
Clijfortia discolor was described by Weimarck
based on a specimen from Table Mtn with very dense
white hairs on the underside of the leaves. However,
field observations of this well-explored locality reveal
a continuous range of variation typical of C. odorata.
Specimens of C. odorata at higher altitudes are often
more densely hairy, especially beneath, and have smaller,
rounder leaves. This is probably a result of increased
exposure and as no further specimens have ever been
attributed to C. discolor, the name is here reduced to
synonymy of C. odorata.
13. Cliffortia juniperina L.f, Supplementum
plantarum: 430 (1782); Weim.: 57, fig. 14A-E (1934);
Fellingham: 615 (2000). Type: South Africa, without
precise locality, Sparrman s.n. (LINNf, holo.)
C. hermaphroditica Weim,: 172, fig. 2a-c, t. 4 (1948); Fellingham:
612 (2000), syn. nov. Type: Western Cape, 3318 (Cape Town):
Stellenbosch District, Jonkershoek. (-DD), 4 Nov. 1943, Compton
15332 (NBG, holo.).
The identification of Cliffortia hermaphroditica is
difficult as the specimen of Compton 15332 supposedly
deposited at NBG has not been found. Two other speci-
mens of the same collection have been seen at BOL and
PRE and both belong to a form of C. juniperina with
close acute ribs. Weimarck placed the BOL specimen
in his concept of C. juniperina var. pilosula on account
of its hairy stems. The photograph in Weimarck l.c., t.
4 (1948) does nothing to suggest that the type was any-
thing other than a slightly abnormal specimen of C. juni-
perina. Certainly the type locality has been very well
collected and nothing else similar has been found, indi-
cating further that it was an atypical form, possibly of
hybrid origin, that arose once and has since disappeared.
22
Bothalia37,l (2007)
ACKNOWLEDGEMENTS
The first author wishes to thank the Leverhulme
Trust, who funded this research with a Study Abroad
Studentship, along with assistance from the University
of Cape Town through a J.W. Jagger International
Scholarship and the Smuts Memorial Fellowship. The
project was supported by Prof H.P. Linder, formerly
of the Bolus Herbarium, now in Zurich, Switzerland,
with much valuable encouragement by Prof. T.A.J.
Hedderson. The curators of BM, BOL, GRA, K, NBG,
NU and PRE are thanked for pennission to examine their
collections or for sending loans.
SPECIMENS EXAMINED
Acocks 16102 (9a) BOL, K; 21529, 22531 (10b) K, PRE; 22426 (10a)
K, PRE. Acocks & Hafstrom 568 (10a) PRE. Andreae s.n. (4a) BOL;
1154 (8) NBG, PRE.
Barker 4583 (10a) NBG; 5950 (9a) NBG. Bean, Vlok & Viviers 2096
(5) NBG, PRE. Berg STE19824 (4a) NBG. Beyers 214 (9a) NBG, PRE.
A. Bolus 1545 (4a) BOL. F. & L. Bolus 13310 (10a) BOL. H. Bolus
2630 (11) BOL; 4077 ( 10a) BOL, K; 6994 (9a) BOL, NBG, PRE; 7987
(4a) BOL, K; 11271 (5) BOL, PRE. Bond 727 ( 9a) NBG. Bosenberg
& Rutherford 337 (1) NBG. Boucher 1749 ( 9b) NBG. Boucher &
Shepherd 4359 ( 1 Oa) NBG. Boucher & Stindt 5282 ( 1 Oa) NBG. Bousse
s.n. (9a) BM. Burchell 5583 (10b) K; 7816 (9a) K. Burgers 1947 (10a)
NBG, PRE. Burke s.n. (10a) K. Buys 64 (9a) NBG.
Compton 6408, 8151, 9219, 16534 (7) NBG; 8359, 15301, 18324 (9a)
NBG; 15828 (10b) BOL, NBG; 16471 (9b) NBG; 17090 (10a) NBG;
17451, 18546 (4a) NBG; 20120 (4b) NBG. Cowling 178 (lOb) GRA.
De Villiers s.n. (9a) NBG. Drege s.n. (9a) K; s.n. (4b) K; a (9a) BM.
Dtyjhout 1557 (9a) NBG. Duthie 957 (9a) BOL.
Ecklon 1751, 1752 (10a) K. Esterhuysen s.n., 1993, 11968, 33622,
33625 (4a) BOL; s.n., 1992 (9a) BOL; 121 (4a) PRE; 2647 (4a) BOL,
K, NBG, PRE; 8312 (8) BOL, K, PRE; 9296, 9368, 9920, 13782,
14183, 14280, 14722, 14832, 14891, 14946, 27575, 27688, 27772,
28090 (8) BOL; 10100, 15242 (4a) BOL, PRE; 11636, 11857, 11947,
14501, 16185, 31353 (10a) BOL; 11658 (XQn) BOL, PRE; 12813 (4b)
BOL; 14062 (4b) BOL, K; 14499, 15325 (10a) BOL, NBG; 14647 (4a)
BOL, NBG; 15745 (10a) BOL, NBG, PRE; 79336 (10b) BOL, NBG,
PRE; 27765 (5) BOL; 29401 (10b) BOL; 30602 (1) BOL, K, NBG,
PRE.
Fellingham 655 (10a) NBG, PRE; 1056 (10a) NBG; 1529 ( 1 ) BOL, K,
NBG, PRE; 1530, 1545, 1546, 1582, 1613, 1626, 1634, 7633 (1) BOL,
NBG, PRE; 1551, 1597, 1601 (1) NBG; 7623 (1) NBG, PRE; 7693 (1)
BOL, NBG. Froembling s.n. (4a) NBG; s.n. (10a) NBG.
Galpin 4002 (5) BOL, PRE. Gamble 22232 (10a) K. Garside 1025
(9a) K. Gillett 712, 1802 (9a) NBG. Goulimis s.n. (10a) BOL. Guthrie
33 (10a) BOL; 2348 (9a) NBG.
Haynes 692 (4a) K, NBG, PRE; 1148 (9a) NBG, PRE. Helme 1594
(10b) NBG; 2939 (3) NBG; 3970 (I) NBG; 39 73 ( 6) NBG. Herb.
Goodenough s.n. (9a) K. Hort. Kew s.n. (9a) BM. Hubbard 176, 396
(9a) NBG. Humbert 9436, 9663 (10a) PRE.
Keet STE13379 (\Qh) NBG; 725 (10b) GRA, PRE, Kerfoot 5412 (9a)
NBG; 5705 (4a) PRE; 6021 (4a) K, NBG, PRE.
Leighton 460 (10a) BOL; 792 (9b) BOL. Levyns 2707(5) BOL; 3364,
3979, 7284, 8500, 11406 (9a) BOL; 3322 (10a) BOL; 3329 (I Oa) BOL,
PRE; 7890, 9155, 10262, 10313, 70343 (10b) BOL,
Maguire 1067 (7) NBG. Marloth 2250 (8) PRE; 4247 (8) NBG; 4347
(4a) PRE. McDonald 651 (4a) K, NBG, PRE, Morley 8 (10b) NBG,
PRE,
O 'Callaghan, Fellingham & Van Wyk 105 (10b) NBG, PRE.
Page s.n. (10a) BOL, GRA. Palmer 78 (lOa) NBG, Pappe II (9a)
BM, Parker 3489 (4a) BOL, K, NBG; 3676, 4281 (9a) BOL, K, NBG.
Penfold 32 {]i)a) NBG. PHlans s.n., 8422 (9a) BOL; 7370 (I Oa) BOL.
Prior s.n. (4a) K, PRE; s.n. (9a) K; s.n. ( lOa) K, PRE.
Robertson s.n. (9a) BM. Rodin 3266 (9a) BOL, K, PRE. Rogers 29245
(9a) BOL, K, PRE. Runnalls 587 (9a) NBG, PRE; 587a (9a) NBG.
Rycroft 3209, 3284, 3318 (9a) NBG.
Schlechter 798 (9a) BOL; 2203 (5) K, NBG, PRE; 7558 (9a) BM, K,
NBG, PRE. Schoeman s.n. (9a) NBG. Sieber 508 (10a) K. Stauffer
5022 (10a) K, NBG, PRE. Stokoe SAM59213 (4a) NBG, PRE;
SAM61481 (9a) PRE; 8619 (9a) BOL, PRE; 8835 (8) BOL.
Taylor 6363 (10a) NBG, PRE; 6370 (10a) K, NBG, PRE; 6529 (10a)
BOL, K, NBG, PRE; 6851 (10a) BOL, NBG, PRE; 6905 (9a) NBG,
PRE. Thode 6174 (4a) NBG; 9181 (4a) NBG. Thompson 176 (3) NBG,
PRE; 994 (4a) K, NBG, PRE.
Van der Merwe 844 (9a) NBG, PRE. Van Jaarsveld & Sardien 7686
(9b) NBG. Van Rensburg 469 (9a) NBG; SKF2033 (9a) K, PRE. Van
7977 (10b) NBG, PRE.
WalHch s.n. (10a) K. Werdermann & Oberdieck 367, 743 (9a) K,
PRE. C Whitehouse 14 (4a) BOL, K, NBG; 27 (9a) BOL, NBG; 34
(4a) BOL, NBG; 34, 111 (10a) BOL, NBG; 98 (5) BOL, NBG; 779 (6)
BOL; 737 (2) BOL, K, MO, NBG, Z; 145 (3) BOL, NBG; 737 (9a)
BOL; 766, 282 (9b) BOL, NBG; 187 (8) BOL, NBG; 279 (1) BOL,
K, NBG, Z; 222 (10a) BOL; 283 (7) BOL, NBG. C. tfe A. Whitehouse
149 (9a) BOL, K, NBG; 273 (10a) BOL; 243 (9a) BOL, NBG; 237 (6)
BOL, NBG; 239 (6) BOL, K, NBG; 304 (10a) BOL, NBG; 318 (10b)
BOL, NBG; 329 (3) BOL, NBG; 337 (7) BOL. Wilms 3182 (4a) K.
Wolley Dod 1072 (\ Oa) K; 263 7 (4a) BOL, K. Wright ^.«. ( 1 Oa) K.
REFERENCES
AITON, W. 1789. Hortus kewensis, edn 1, 3. Nicol, London.
DE CANDOLLE, A.P. 1 825. Prodromus systematis naturalis regni veg-
etabilis 2. Treuttel et Wurtz, Paris.
ECKLON, C.F, & ZEYHER, C. 1836. Enumeratio plantanim Africae
australis extratropicae 2. Hamburg.
FELLINGHAM, A.C. 1993. Cliffortia fasciculata, a superfluous name
for C. amplexistipula. Bothalia 23: 67, 68.
FELLINGHAM, A.C. 2000. Cliffortia. In P. Goldblatt & J.C. Manning,
Cape plants. A conspectus of the Cape flora of South Africa.
Strelitzia 9: 608-616. National Botanical Institute, Cape Town
and Missouri Botanical Garden, St Louis.
FELLINGHAM, A.C. 2003. A new serotinous species of Cliffortia
L. (Rosaceae) from Northern Cape, South Africa and section
Arboreae emended. Bothalia 33: 41-47.
GOLDBLATT, P. & MANNING, J.C. 2000. Cape plants. A conspectus
of the Cape flora of South Africa. Strelitzia 9. National Botanical
Institute, Cape Town and Missouri Botanical Garden, St Louis.
GRAHAM, R.A. 1960. Rosaceae. Flora of tropical East Afi-ica. Crown
Agents for Oversea Governments and Administrations, London.
HARVEY, W.H. 1 862. Rosaceae. In W.H. Harvey & O.W. Sonder, Flora
capensis 2 : 285-305. Hodges & Smith, Dublin.
LEVYNS, M.R. 1950. Rosaceae. In R.S. Adamson & T.M. Salter,
Flora of the Cape Peninsula'. 443^52. Juta, Cape Town and
Johannesburg.
LINNAEUS, C. (fil). 1782 (1781). Supplementum plantanim. Impensis
orphanotrophei, Braunschweig.
MUIR, J. 1929. The vegetation of the Riversdale area. Memoirs of the
Botanical Survey of South Africa No. 13.
OLIVER, E.G.H. & FELLINGHAM, A.C. 'l 994. A new serotinous spe-
cies of Cliffortia (Rosaceae) from the southwestern Cape with
notes on Cliffortia arborea. Bothalia 24: 1 53-162.
PHILLIPSON, P.B. 1987. A checklist of vascular plants of the Amatole
Mountains, eastern Cape Province/Ciskei. Bothalia 17: 237-
256.
WEIMARCK, H. 1934. Monograph of the genus Cliffortia. Hakan
Ohisson, Lund.
WEIMARCK, H. 1948, The genus Cliffortia, a taxonomical survey.
Botaniska Notiser 90: 1 67-203.
WHITEHOUSE, C.M. 2003. Systematics of the genus Cliffortia L.
(Rosaceae). Ph.D. thesis. University of Cape Town.
WHITEHOUSE, C.M. 2004a. The genus Cliffortia (Rosaceae) in
KwaZulu-Natal. Bothalia 34: 1-10.
WHITEHOUSE, C.M. 2004b. New species in the section Multineiwiae
of Cliffortia (Rosaceae). Bothalia 34: 77-85.
WILLDENOW, C.L. 1806. Species plantanim 2, edn 4. Nauk, Berlin.
Bothalia37,l: 23-56 (2007)
Notes on African plants
Various Authors
APOCYNACEAE
ANEW SPECIES OF HUERNIA (ASCLEPIADOIDEAE-CEROPEGIEAE) FROM SOUTHERN ANGOLA
The genus Huernia R.Br. consists of 64 species
according to Leach (1988). However, further research
has suggested that 49 species is a better estimate (Bruyns
2005). The present new species then brings the number
of species in Huernia to 50. Huernia is widely distrib-
uted in sub-Saharan Africa and six species also occur in
the Arabian Peninsula, where the former South Yemen
is the northeastern limit of its distribution. In Angola,
Huernia is represented by H. oculata N.E.Br., H. similis
N.E.Br., H. urceolata L.C. Leach, H. verekeri Stent and
H. volkartii Peitsch. ex Werderm. & Peitsch. Of these,
only H. similis is endemic to Angola, whereas H. oculata
and H. urceolata are found in both Angola and Namibia
and H. verekeri and H. volkartii are more widely distrib-
uted in southern Africa. The new species described here
is also an Angolan endemic and is of very local occur-
rence on the margin of the Namib Desert near Namibe
(Figure 1).
Huernia lopanthera Bruyns, sp. nov., nullo dubio
H. kennedyana Lavranos proxima, praecipue differt cau-
libus gracilioribus, corolla intus uniformiter rubra-brun-
nea cum papillis brevioribus, tubo corollae breviore et
lobis coronae exterioris interiorisque brevioribus.
TYPE. — Angola, NE of Namibe, Jan. 2006, Bruyns
70470 (BOL, holo.;K, iso.).
Dwarf succulent, up to 300 mm diam., forming dense
mats of tightly packed stems, not rhizomatous. Stems
10^0 X 6-12 mm, decumbent, shortly ellipsoidal, grey-
green; tubercles 1-3 mm long, conical, laterally flat-
tened, acute, joined into 6 or 7 low ridges along stem to
give it a tessellate appearance. Inflorescence of 1-3 flow-
ers developing in gradual succession, arising at middle
of or in lower half of stem on short peduncle (< 5 mm
long), with fine filiform bracts 1 mm long; pedicel 7-10
X 1 mm, ascending, holding flower facing horizontally
or slightly downwards; sepals 3^ mm long, 1 mm broad
at base, acuminate. Corolla x 22-27 mm, shallowly
plate-like to flat, scentless; outside obscurely papillate,
pale brownish green, with 3 raised longitudinal veins
running down lobes; inside uniformly deep red-brown
becoming finely speckled with white towards centre,
with many pale yellow papillae from middle of lobes to
around corona, papillae up to 0.5 mm long and widely
spaced on lobes becoming much shorter and densely
crowded around corona; tube absent to very shallow (1-
2 mm deep); lobes 5-7 x 7-10 mm, spreading, deflate,
acuminate, with raised pale yellow papillate margin.
Corona 2.5 x 3 mm, sessile on corolla; outer lobes 0.5
X 1-1.5 mm, spreading onto surface of corolla, ± rect-
angular to slightly notched in centre, pale yellow faintly
suffused with red; inner lobes 1.5 mm long, deep yellow.
adpressed to anthers in lower half then rising slightly to
meet in centre, slightly dorsiventrally flattened, some-
times with swollen gibbosity at base, sometimes with
slightly clavate, finely bristly apex. Follicles and seed
unknown. Figure 2.
The new species differs from all others in Angola
by its 6- or 7-angled, shortly ellipsoidal stems and the
nearly flat flowers. Vegetatively, Huernia lopanthera
is most similar to 77. kennedyana Lavranos, 77. longii
Pillans and 77. pillansii N.E.Br. from the southern edges
of the Great Karoo in South Africa, on account of the
tessellate stems with more than five angles. In 77. longii
and 77. pillansii the corolla has a cupular tube and the
lobes are much longer than broad, whereas in 77. ken-
nedyana the flowers are relatively flat with a shallow
tube and the corolla lobes are roughly as long as broad.
Consequently, among these three species, 77. lopanthera
is most similar to 77. kennedyana. In 77. kennedyana
the stems are almost spherical and, at 10-25 mm thick,
are mostly much thicker than those of 77. lopanthera.
The flower in 77. kennedyana is brilliantly transversely
striped with red-brown to maroon on a cream-coloured
24
Bothalia37,l (2007)
FIGURE 2. Ifiiernia lopanthera, Bruyns 10410. A, portion of plant; B, face view of flower; C, side view of dissected flower; D, papillae inside
flower, larger ones from middle ol lobes, smaller from around corona; E, F, side views of gynostegium; G, pollinarium. Scale bars (at A):
A-C, 5 mm; (at F): D-G; D, 0.5 mm; E, F, 1 mm; G, 0.25 mm. Artist: RV. Bruyns.
Bothalia 37,1 (2007)
25
to dull yellow background (as opposed to uniformly deep
red-brown in H. lopanthera), the corolla tube is 5-6 mm
long (to 2 mm long in the new species, but often absent,
as in Figure 1C) and the inner surface of the corolla has
relatively long papillae, that reach a maximum length
of 3 mm (contrasting with a maximum of 0.5 mm in the
new species).
Although it has been stated (Leach 1969, 1974, 1988;
Meve 1997: 42) that the corona in all species of Huernia
is sessile on the base of the corolla tube, there are some
species for which this is not the case (Bruyns 2005: 132).
In those species of Huernia where the gynostegium is
indeed sessile, the outer corona lobes can be pressed to
the base of the corolla tube and may even be partly fused
to it. The other extreme is presented by H. kennedyana,
where the gynostegium possesses a short stipe (i.e. is
not sessile) so that the outer corona is somewhat raised
above the surface of the corolla tube. In this respect H.
lopanthera differs from H. kennedyana in that it does
not have a stipe beneath the gynostegium and the outer
corona lobes touch the surface of the corolla, though
they are not fused to it.
It has also been stated (Meve 1997: 43) that the gyno-
stegium in Huernia is distinctive for the small tubercle
on the outer corona projecting towards the base of the
guide-rails. This structure is formed by the somewhat
raised and projecting lip of the orifice below the guide-
rails (Bruyns 2005: 133). Again this feature is actually
variably present in Huernia (Bruyns 2005: 69, 133)
and, while it is found in H. kennedyana, it is absent in
H. lopanthera. In H. kennedyana the inner corona lobes
are connivent in the centre and then diverge above this,
whereas in H. lopanthera the inner corona lobes touch in
the centre but do not rise above that and do not diverge
again. The presence of a swollen gibbosity at their base
is surprisingly variable (as is seen in Figure IE, F, drawn
from different plants of the same collection) and the
inner corona lobes are also sometimes swollen and finely
papillate at their tips.
Plants of this new species form dense mats of very
short stems filling crevices between rocks in gently
sloping terrain. They occur amongst a sparse and very
low vegetation, much of which is not higher than 300
mm. This vegetation contains a remarkable number
of succulents. These include various members of the
Portulacaceae, Euphorbia carunculifera L.C. Leach, E.
indurescens L.C. Leach and El subsalsa Hiem, one spe-
cies each of Kalanchoe Adanson and Kleinia Miller,
as well as a wide selection of Apocynaceae, including
Adenium obesum (Forssk.) Roem. & Schultes, Fockea
angustifolia K.Schum., Hoodia currorii (Hook.) Decne.,.
H. mossamedensis L.C. Leach, Huernia oculata Hook.f,
Sarcostemma viminale (L.) R.Br., Stapelia kwebensis
N.E.Br. and Tavaresia angolensis Welw.
REFERENCES
BRUYNS, RV. 2005. Stapeliads of southern Africa. 2 vols. Umdaus
Press, Hatfield, Pretoria.
LEACH, L.C. 1969. Stapelieae from south tropical Africa V. Bothalia
10: 45-54.
LEACH, L.C. 1974. Stapelieae (Asclepiadaceae) from south tropical
Africa 8. Journal of South African Botany 40: 1 5-25.
LEACH, L.C. 1988. A revision of Huernia R.Br. (Asclepiadaceae).
Excelsa Taxonomic Series 4: 1-197.
MEVE, U. 1997. The genus Duvalia (Stapelieae). Springer- Verlag,
Vienna.
P.V BRUYNS*
* Bolus Herbarium, University of Cape Town, 7701 Rondebosch.
MS. received: 2006-06-30.
BORAGINACEAE
LECTOTYPIFICATION OF THE BASIONYM, ECHIUM GLAUCOPHYLLUM
Buck (1837) based his combination Lobostemon glau-
cophyllus (Jacq.) H.Buek on Echium glaucophyllum
Jacq. Although not directly referring to Jacquin, Buek
did so indirectly by citing "E. glaucophyllum Pers.’ refer-
ring to the use of the name by Persoon (1805) who cites
Jacquin’s original publication. Buek also included as a
synonym, E. laevigatum Lam., a superfluous and illegiti-
mate name for E. glaucophyllum. The species in question
was described in Jacquin’s Collectanea (1789a).
Jacquin (1789b) simultaneously and subsequently
compiled leones plantarum rariorum, the publication
dates of which are multifarious (Schubert 1945). The
illustration of E. glaucophyllus appeared in vol. 2, fasci-
cle 3 in 1789 and the accompanying text was issued with
the appearance of fascicle 16 (Jacquin 1795). Many of
the Jacquin names published in Collectanea have been
typified by illustrations in his leones plantarum rario-
rum. Jacquin based many new names on specimens col-
lected from plants cultivated in botanical gardens such
as Schonbrunn (D’Arcy 1970). However, there is a Boos
specimen at W (here chosen as lectotype) which appears
to be one of a few associated with Jacquin that was col-
lected and dried in the field. Franz Boos (1753-1832),
later to become Director of the mentioned garden, proba-
bly collected the specimen while accompanying the offi-
cial Austrian expedition to Mauritius (Neilreich 1855).
Boos arrived at the Cape in May 1786 and remained
there for nine months before proceeding to Mauritius in
February 1787. He called at the Cape again in the sum-
mer of January 1788 on his return to Europe, and arrived
back in Vienna in July (Garside 1942; Gunn & Codd
1981).
Garside (1942: 211, 212) based his statements regard-
ing the collection dates of the Oxalis specimens of Boos
& Scholl, on the analysis by Salter, who had an unri-
valled field knowledge of this genus in South Africa.
Salter assumed that the Oxalis specimens were collected
during Boos’ first expedition from May to July, 1786.
In the protologue, Jacquin mentions "Ad Promon-
torium bonae Spei crelcit; sub dio apud nos floret Julio,
hyemem agens in hybernaculo' , i.e. ‘growing near the
26
Cape of Good Hope, blossoms in July when growing
outside, kept in hibernation’ (under cover) in the winter.
'Julio'’ could mean the month of collection in the Cape.
We interpret the protologue as referring to a plant cul-
tivated in the Schonbninn garden which subsequently
flowered in July.
The herbarium sheet Boos s.u. sub W0001202 consists
of two separate specimens, the one on the left numbered
‘2’ and the one on the right numbered ‘ 1’. On the accom-
panying label is written in an unidentified hand ‘ 1 . Cap.
B. Sp. Boos.’ and ‘2. Hort. Schonbr.’ and on the reverse
similarly, in an unidentified hand ‘1. Cap. B. Spei. Boos.’
and ‘2. Hort. Bot. Schonbr.’ Specimen ‘1’ was therefore
collected in the field by Boos and specimen ‘2’ was
sampled from a plant cultivated later in the Schonbrunn
garden — the one mentioned in the protologue as flower-
ing in July. Accordingly, we choose the specimen on the
right (‘ 1’) as lectotype.
Echium glaucophylhim Jacq., Collectanea: 325,
326. (1789) = Lobostemon glaucophyllus (Jacq.)
H.Buek, Linnaea 11: 139 (1837). Lectotype designated
here: without locality. Boos s.u. sub W0001202 (W!).
This lectotypification maintains application of the
name Lobostemon glaucophyllus which is currently con-
sidered the correct name (Buys 2002).
ACKNOWLEDGEMENTS
We thank the curator of the Natural History Museum,
Vienna (W) for allowing access to the designated type
Bothalia37,l (2007)
specimen and John McNeill and Bruno Wallnofer for
commenting on the manuscript.
REFERENCES
BUEK, H.W. 1837. Echia Capensia. L/wnaea 11; 129-149.
BUYS, M.H. 2002. Lobostemon. In R Goldblatt & J.C. Manning, Cape
plants. A conspectus of the Cape flora of South Africa. Strelitzia
9: 375-377. National Botanical Institute, Pretoria and Missouri
Botanical Garden, St. Louis.
D’ARCY, W.G. 1970. Jacquin names, some notes on their typification.
Taxon 19: 554-560.
GARSIDE, S. 1942. Baron Jacquin and the Schonbrunn gardens.
Journal of South African Botany 8: 201-224.
GUNN, M. & CODD, L.E. 1981. Botanical exploration of southern
Africa. Balkema, Cape Town.
JACQUIN, N.J. 1789a. Collectanea, vol. 2: 325, 326. Wappler,
Vindobonae.
JACQUfN, N.J. 1789b. leones plantarum rariorum, vol. 2, fasc. 3: t.
312. Wappler, Vindobonae.
JACQUfN, N.J. 1795. leones plantarum rariorum, vol. 2, fasc. 16: 10.
Wappler, Vindobonae.
NEILREICH, A. 1855. Geschichte der Botanik in Nieder-Oesterreich.
Verhandlungen des zoologisch-botanischen Vereins in Wien 5:
23-76.
PERSOON, C.H. 1^05. Echium. Synopsis plantarum: 163, 164. Crame-
rum, Paris.
SCHUBERT, B.G. 1945. Publication of Jaquin’s leones plantarum rari-
orum. Contributions of the Gray Herbarium 64: 3-23.
M.H. BUYS* and B. NORDENSTAM**
* Compton Herbarium, South African National Biodiversity Institute,
Private Bag X7, 7735 Claremont and Department of Botany &
Zoology, University of Stellenbosch, Private Bag XI, 7602 Matieland,
South Africa.
** Department of Phanerogamic Botany, Swedish Museum of Natural
History, P.O. Box 50007, SE-104 05 Stockholm, Sweden.
MS. received: 2006-05-31.
ASPHODELACEAE
TRACHYANDRA ARENICOLA AND T. MONTANA, TWO NEW SPECIES FROM SOUTH AFRICA
INTRODUCTION
Trachyandra Kunth is a genus of ± 50 species distrib-
uted throughout sub-Saharan Africa. It is centred in the
southern African winter rainfall region, with three or four
species extending into tropical Africa, one as far north
as Ethiopia (Smith & Van Wyk 1998). Trachyandra is
distinguished from other genera of Asphodelaceae by
its soft-textured leaves and short-lived, white or pink-
ish flowers with a deciduous perianth and ± scabrid fila-
ments (Obermeyer 1962). The seeds appear to be highly
characteristic, developing distinctive idioblasts contain-
ing bundles of needle-like crystals. These are visible as
small, raised excrescences in the tissue of the enveloping
funicular sarcotesta, which is a family characteristic of
Asphodelaceae.
Three sections are currently recognized within the
genus (Obenneyer 1962). In sections Liriothamnus and
Glanditlifera the first leaves of each growth cycle are not
reduced to membranous sheaths and thus grade into the
foliage leaves, the roots are slender and wiry, and the fila-
ments are monomorphic. Species in section Glanditlifera
are invariably sparsely or densely glandular-pubescent
and those in section Liriothamnus either glabrous or
pubescent but not glandular. Section Trachyandra is
readily distinguished from the other sections by having
the first leaves of each growth cycle reduced to sheathing
membranous or papery cataphylls that surround the base
of the annual shoots, by its ± swollen or tuberous roots,
and by often having dimorphic filaments. It is the larg-
est section of the genus, comprising around half of the
species, most of which are endemic to the winter rainfall
parts of the Western Cape and Northern Cape.
Just over 30 species are endemic or near-endemic
(Obenneyer 1962; Manning & Klopper 2006). Two
species have been described since the last revision of
the South African members of the genus (Obenneyer
1962), both from the winter rainfall parts of the Western
Cape and Northern Cape (Manning 1990; Perry 1990).
Bothalia37,l (2007)
27
Another two species are described here. Trachyandra
montana was recently discovered on the middle and
upper slopes of Jonaskop, the highest peak in the west-
ern part of the Riviersonderend Mountain range in the
southwestern Cape. Trachyandra arenicola, from the
arid west coast of Namaqualand and adjacent Western
Cape, was first collected in 1925-1926 and later in 1945
but confused with other species in the genus (Obermeyer
1962). Several recent gatherings made over the past few
decades, however, provide ample evidence of its dis-
tinctness.
Trachyandra montana J.C. Manning & Goldblatt,
sp. nov.
Geophyta decidua, 100-200 mm alta, radicibus gra-
cilibus tuberibus fusiformibus distalibus ferentibus, folds
1 vel 2 ligulatis firmis scabridis 5-10 mm latis, inflor-
escentibus simplice pedunculis dense pubescentibus,
floribus albis nervo medio aurantiacis, tepalis ad basem
medioque pubescentibus, ovario sparse scabrido ovulis
6 in quoque loculo, capsulis dense pubescentibus erectis
6-7 X 4—5 mm.
TYPE. — Western Cape, 3319 (Worcester): Rivier-
sonderend Mtns, Jonaskop, along road to summit at
1 200 m, (-DC), 18 October 2005, J. Manning 2995
(NBG, holo.; MO, iso.).
Deciduous geophyte, 100-200 mm high. Rhizome
short, horizontal; roots slender, lanate, reddish brown,
developing tubers ± 100 mm from base, tubers 30^0
X 5-8 mm. Cataphylls surrounding shoots as well as
leaf and scape bases, papery, brown, 5-10 mm long.
Leaves (1)2, linear-lanceolate, 200-300 x 5-10 mm,
plane, firm-textured, somewhat twisted and flexuous,
striate with raised, thickened veins, scabrid along veins,
bright green flushed pink towards base. Inflorescence
a simple raceme; peduncle flexed 20-40 mm from
base then inclined to erect, terete, 1. 5-2.0 mm diam.
at base, densely pubescent with short, straggling hairs;
raceme lax, few- to many-flowered, 40-80 mm long;
bracts ovate-acuminate, membranous, pubescent at
base and along margins and midrib; pedicels 8-10 mm
long, suberect in bud but arcuate-spreading at anthesis,
becoming erect in fruit and ultimately ± 10 mm long,
pubescent. Flowers rotate, white with orange midribs,
immaculate, opening in afternoon, fragrance unknown;
tepals pubescent at base and along abaxial midrib, outer
elliptical, 10-12 x 2. 0-2. 5 mm, inner obovate, 10-12
X 4—5 mm. Stamens suberect; filaments filiform, taper-
ing below, white, retrorsely scabrid in upper two thirds,
outer ± 7 mm long, inner ± 6 mm long; anthers yellow, ±
2 mm long. Ovary ovoid, ± 2 mm long, sparsely scabrid,
green, with 6 ovules per locule; style erect, filiform, ±
6.5 mm long, white. Capsule subglobose, 6-7 x 4—5
mm, densely retrorsely scabridulous. Seeds tetrahedral,
verrucose, black, ± 3 mm diam. Flowering time: October
and November. Figure 3.
Distribution and habitat: so far known only from
the middle slopes of Jonaskop at the western end of the
Riviersonderend Mountains (Figure 4), Trachyandra
montana is not uncommon on the drier, north-facing
slopes, occurring in stony sandstone in more open, rocky
ground and along the roadside.
Diagnosis and relationships: the papery cataphylls
and fleshy roots place Trachyandra montana in section
Trachyandra, where it is closely allied to T. hirsutiflora
(Adamson) Oberm., another southwestern Cape endemic.
Both are characterized by apomorphic, unbranched,
cylindrical inflorescences with densely hairy scapes
lacking sterile bracts, firm-textured and striate, scabrid
leaves, and pubescent capsules. Trachyandra montana
is distinguished by its distinctive roots, which are ini-
tially slender and only develop fusiform tubers some
distance along their length, by its two (or rarely just
one) plane leaves 5-10 mm wide, surrounded at the
base with chestnut brown cataphylls, its rather lax and
inclined inflorescence, and its relatively small capsules
6-7 mm long. Trachyandra hirsutiflora, in contrast, has
roots that taper from a swollen base, (2)3-5 linear-cana-
liculate leaves (l-)2-5(-8) mm in diameter surrounded
at the base by translucent cataphylls, typically a dense,
stiffly erect inflorescence, and characteristically large
capsules (8-) 10-1 5 mm long. Trachyandra hirsutiflora
ranges through the coastal mountains of the southwest-
ern Cape, from the Piketberg to the Potberg (Figure 4),
but has not been recorded from further inland on the
Riviersonderend Mountain range. It favours sandy, often
seasonally moist flats rather than the drier, rockier situa-
tions in which T. montana occurs, and flowers especially
freely, sometimes prolifically, after fire.
Other material examined
WESTERN CAPE. — 3319 (Worcester): Riviersonderend Mtns,
Jonaskop, along road to summit at 1 200 m, (-DC), 10 January 2006
(fruiting), J. Manning 2996 (NBG, MO).
Trachyandra arenicola J.C. Manning & Goldblatt,
sp. nov.
TYPE. — Northern Cape, 2917 (Springbok): flats
below Anenous Pass, along road to Eksteenfontein,
Farm Grasvlakte, deep red sands, (-DC), 28 July 2006,
J. Manning 3026 (NBG, holo.; K, MO, iso.).
Geophyta decidua, (100-)200-500 mm alta, radi-
cis pluribus leviter incrassatis decrescentibusque, folds
150-200(-500) X 5-10(-30) mm, marginibus scabridulis
retrorsis, ad basem cataphyllis papyraceis, inflorescentia
erecta ramis 1 vel 2, laxe multifloris, pedunculo glabro
vel pubescente, floribus albis maculo flavo ad basem
tepalis, loculis ovulis 10-14 eodem, capsulis erectis 10-
14 mm longis.
Deciduous geophyte, (100-)200-500 mm high.
Rhizome short, vertical; roots several, slightly swol-
len and tapering, lanate, current season’s complement
without lateral rootlets, whitish, 5-6 mm diam. at base,
previous season’s complement developing feeder root-
lets, brownish. Cataphylls ± 10 mm long, surrounding
shoots as well as leaf and scape bases, papery, translu-
cent greyish brown. Leaves (2-)4— 12, lanceolate-fal-
cate, 150-200(-500) X 5-10(-30) mm, plane, soft-tex-
tured and succulent, retrorsely scabridulous on margins,
withered and twisted at tips at flowering time, glaucous
28
Bothalia37,l (2007)
FIGURE 3. — Trachyandra montana. Manning 2995, 2996. A, whole plant; B-F, floral details; B, outer tepal; C, inner tepal; D, outer stamen, E,
inner stamen; F, gynoecium, G, infructescence; H, seed. Scale bars: A, G, 10 mm; B-F, H, 2 mm. Artist: John Manning.
Bothalia 37,1 (2007)
29
FIGURE 4. — Known distribution of Trachyandra montana. O; and T.
hirsutiflora, •.
or dull green. Inflorescence: peduncle erect, sometimes
becoming deflexed with age and in fruit, 3-7 mm diam.
at base, green or flushed reddish, glabrous or pubescent
with soft, straggling hairs; raceme, compound, lax, with
one or two suberect branches, the lower sometimes with
an additional accessory branch, usually several (up to 6)
inflorescences per plant, many -flowered, 100-200(-600)
mm long; bracts ovate-acuminate, 7-9 mm long, mem-
branous, glabrous or pubescent along midrib, ciliate
along margins in upper half; pedicels (10-)12-19 mm
long, erect in bud but arcuate-spreading at anthesis, gla-
brous, sparsely retrorse-scabridulous, strigose, or pubes-
cent with spreading hairs, becoming sinuate- or spread-
ing-erect in fruit but not elongating further. Flowers
rotate, white or flushed pink on reverse with greenish
or greyish midribs, tepals each with greenish or bright
yellow spot near base, opening ± 13:00 and withering
± 18:00, sweetly scented; tepals fused below for 1 mm,
glabrous or pubescent adaxially at base and on midrib,
outer oblanceolate, 10-13 x 2. 3-3.0 mm, inner obovate,
10-13 X 3^ rnm, somewhat clawed in lower 3 mm and
papillate-ciliolate along margins of claw. Stamens sub-
erect; filaments filiform, tapering below, white or flushed
pinkish in lower half, retrorsely scabrid or scabridulous
except in uppermost 0. 5-1.0 mm, outer 7-8 mm long,
inner 8-9 mm long; anthers yellow, ± 1.8 mm long at
anthesis. Ovary ellipsoid, ± 2 mm long, glabrous, green
or greyish, with 10-14 ovules per locule; style erect, fili-
form, white, 10-11 mm long. Capsule ellipsoid, 10-12
X 4—5 mm, glabrous. Seeds tetrahedral, ± 2 mm ‘diam.,
sparsely verrucose, pale reddish brown. Flowering time:
mainly July and August, extending into September under
favourable conditions. Figure 5.
Distribution and ecology. Trachyandra arenicola
is scattered along the edge of the Namaqualand coastal
plain wherever suitable areas of deep, sandy soils occur.
Populations have been recorded from the Anenous Flats
at the southern fringe of the Richtersveld in the north, as
far south as Wallekraal in Northern Cape, and then much
farther to the south around Klawer and Graafwater in
Western Cape (Figure 6). Plants are locally common on
sandy flats or in duneveld, sometimes numbering hun-
dreds of individuals. Like most species in the genus, the
flowers open in the afternoon and wither the same eve-
ning. They are pleasantly scented.
Diagnosis and relationships: in its relatively robust
stature, lanceolate leaves, and sparsely branched inflor-
escence, Trachyandra arenicola most closely resembles
T. ciliata (L.f) Kunth and T falcata (L.f ) Kunth in sec-
tion Trachyandra. The bracts in the immature inflores-
cences of these three species are closely imbricate, and
the young racemes of T. ciliata and T. falcata were viv-
idly likened to ears of com by Obermeyer (1962). These
characteristic young inflorescences make the latter two
species useful as a vegetable under the Afrikaans ver-
nacular name veldkool. On the basis of these several fea-
tures it seems likely that the three species are immedi-
ately related.
Trachyandra arenicola is distinguished from T fal-
cata by its more slender stature, laxer inflorescences
with ± concolorous bracts, and especially by lacking the
amplexicaul peduncular bract diagnostic of T. falcata.
The latter is an altogether more robust species with a
dense inflorescence, characteristically with the outer half
of the bracts suffused dark brown. In addition, the roots
of T. falcata are far more numerous, slender, and not
swollen. The two species are ecologically distinct, with
T. falcata favouring stony, often granitic slopes, and T.
arenicola restricted to sandy flats. From T. ciliata it is
immediately separable by its erect, not trailing or creep-
ing inflorescences (although the older inflorescences
tend to topple over and become decumbent in fmit),
its erect, not pendent fmits, and by the shorter bracts,
7-9 mm long vs 10-16 mm. Trachyandra ciliata, like
T. arenicola, is also commonly encountered on sandy
coastal flats but is much more widely distributed, extend-
ing from southern Namibia through the coastal areas of
Western Cape as far east as Bathurst in the Eastern Cape
(Figure 6).
All three species may be variously pubescent or ±
glabrous, with both forms co-occurring in certain locali-
ties. In Trachyandra arenicola the variation in vesti-
ture is geographical. Populations from Namaqualand in
Northern Cape are glabrous apart from the sparsely sca-
bridulous pedicels, whereas those from the south, around
Klawer and Graafwater, are densely pubescent on the
peduncles and pedicels, and the tepals are also partially
pubescent. The capsules in the southern populations tend
also to be slightly more broadly ellipsoid. Early collec-
tions from the southern populations were identified as T.
falcata by Obermeyer (1962), and a single, incomplete
collection from Namaqualand she assigned to T. ciliata.
Other material examined
NORTHERN CAPE. — 2917 (Springbok): Anenous Flats, Farm
Grasvlakte, red sands, (-DC), 6 October 1991, J. Manning 1032
(NBG); 23 August 1992, P. Goldblatt & J. Manning 9287 (MO,
NBG). 3017 (Hondeklipbaai): Farm Avontuur, 15 km inland from
Hondeklipbaai, deep, loose sand, (-AD), 29 August 1990, P.A. Bean &
M. Viviers 2558 (BOL); 1 km from Wallekraal, red sand flats, (-BC),
without date, PL. Perry 3869 (NBG); 1.5 km along Wallekraal road
30
Bothalia37,l (2007)
FIGURE 5. — Trachyandra arenico-
la. Manning 3026. A, whole
plant; B-G, floral details: B,
outer tepal; C, inner tepal; D,
outer stamen; E, inner stamen;
F, gynoecium; G, capsule; H,
seed. Scale bar: A, G, 10 mm;
B-F, 2 mm; H, 1 mm. Artist:
John Manning.
to Spocgrivicr, red sands, (-BC), 6 October 1991, ./. Manning I039\
Farm Hardekoppic, northwest of Kotzesrust, consolidated sand, (-DC),
180 m, 29 September 1987, C. Reid 1299 (BOL, PRE); 10.5 km west
of Kotzesrust, sandy slope, ( -DD), 28 August 2001, P. Goldhiall &
L. Porter 1 1 778 (MO, NBfJ). Without precise locality: Namaqualand,
1925- 6, G. Meyer s.n. (NBG).
WESTERN CAPE.— 3118 (Vanrhynsdorp): ± 50 km west of N7
highway between Clanwilliam and Vanrhynsdorp, 6.5 km north of
tunioff to Klawer, sandveld, (-DA), 7 August 1985, K. Steiner 906
(NBG); 20 km SSE of Klawer, Sandkraal, (-DC), without date, PL.
Perry 3580 (NBG); Klawer, (-DC), 2 September 1945, W.F. Barker
J63«(NBG).
Bothalia37,l (2007)
31
30°-
32°-
FIGURE 6. — Known distribution of Trachyandra arenicola, A ; T ciliata, O; and T. falcata, •.
ACKNOWLEDGEMENTS
Material was gathered with permits from Western and
Northern Cape Nature Conservation.
REFERENCES
MANNING, J.C. 1990. A new species of Trachyandra section
Liriothamnus (Asphodelaceae) from the Richtersveld. South
African Journal of Botany 56: 1-5.
MANNING, J.C. & KLOPPER. R.R. 2006. Trachyandra. In G.
Germishuizen, N.L. Meyer, Y. Steenkamp & M. Keith, A check-
list of South African plants. Southern African Biodiversity
Network Report No. 41: 912, 913. SABONET, Pretoria.
OBERMEYER, A. A. 1962. A revision of the South African species of
Anthericum, Chlorophytum and Trachyandra. Bothalia 7: 669—
767.
PERRY. PL. 1990. A new species of Trachyandra section Trachyandra
from Western Cape. South African Journal of Botany 56: 257-
260.
SMITH, G.F. & VAN WYK, B-E. 1998. Asphodelaceae. In K. Kubitzki,
The families and genera of vascular plants 3>\ 130-140. Springer,
Hamburg.
J.C. MANNING* and P. GOLDBLATT**
* Compton Herbarium, South African National Biodiversity 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: 2006-05-29.
HYACINTHACEAE
ANEW PYROPHYTIC LACHENALIA SPECIES (MASSONIEAE) FROM WESTERN CAPE, SOUTH AFRICA
The new species described here forms part of a series
of papers towards a revision of the genus (Duncan 1993,
1996, 1997, 1998; Duncan & Edwards 2002, 2006).
Lachenalia lutzeyeri G.D. Duncan, sp. nov.
Bulbus globosus penitus praesens, folia 1 - vel 2
anguste lanceolata marginibus cartilagineis, pedunculus
rigidus, flores oblongo-campanulati nutantes flavo-cre-
mei gibbis viridibus vel brunneis, stamina bene exserta
filamentis leniter declinatis, semina globosa strophiolo
inflate medio-terminali. L. youngii Baker afifinis, sed ab
specie nova bulbis non profundis foliis linearibus mar-
ginibus non cartilagineis pedunculo non rigido floribus
staminibus breve exsertis filamentis rectis differt.
TYPE. — Western Cape, 3419 (Caledon): Witkrans-
berg area of Grootbos Private Nature Reserve, northwest
of Gansbaai, on southwest-facing mountain slope in
Table Mountain Sandstone, (-CB), November 2004, H.
Lutzeyer s.n. (NBG, holo.!; PRE, iso.).
Deciduous, winter-growing geophyte, 200^20 mm
high. Bulb solitary, deep-seated, globose, 20-25 mm
diam., white with membranous, dark brown outer tunics;
roots numerous, mainly contractile, a few fibrous.
Leaves 1 or 2, narrowly lanceolate, 85-290 x 4— 13 mm,
prostrate, spreading or suberect, deeply canaliculate or
sometimes conduplicate, yellowish green or pale to deep
maroon, adaxial surface plain or with large dark green,
brown or maroon flattened pustules, margins cartilagi-
nous, flat or undulate; clasping leaf base white, subter-
ranean, up to 85 mm long, or with short aerial portion
up to 15 mm long. Inflorescence an erect or suberect,
many-flowered, moderately dense raceme 50-120 mm
long, with a sterile apex up to 10 mm long; peduncle
32
Bothalia37,l (2007)
rigid, erect or suberect, 120-200 mm long, green and
unmarked, or yellowish green and heavily marked
with minute maroon speckles or small to large maroon
blotches; rachis green, 50-120 mm long; pedicels suber-
ect during flowering, ascending in fruit, green, 3-4 mm
long; bracts ovate in lower half of inflorescence, becom-
ing lanceolate in upper half, 1-2 x 0. 5-2.0 mm, trans-
lucent white. Flowers oblong-campanulate, nodding
at anthesis, yellowish cream with dark green or brown
gibbosities, with a moderately strong, soapy-sweet
scent, fading to dull maroon and ascending in fruit; peri-
anth tube cup-shaped, pale yellowish cream, 2-3 mm
long; outer tepals ovate, 5-6 x 3^ mm, inner tepals
obovate, 6-7 x 4-5 mm, protruding very slightly beyond
outer tepals, translucent white with a yellowish green
median keel and a dark green or brown marking near
apex. Stamens exserted 5-6 mm beyond perianth; fila-
ments weakly declinate, white, 9-10 mm long; anthers
maroon prior to anthesis; pollen yellow at anthesis.
Ovary obovoid, 2x3 mm, bright green; style straight,
9 mm long, white. Capsule obovoid, 6-7 x 6-7 mm.
Seed globose, 1.1-1. 2 x 1.2-1. 3 mm, shiny black, with
a terminal, inflated, black strophiole 0.8-0. 9 mm long.
Flowering time: early to late November, rarely to late
January, with a peak in mid-November. Figures 7, 8.
Etymology. LachenaUa lutzeyeri is named for Heiner
Lutzeyer of Grootbos Private Nature Reserve near
Gansbaai, who discovered this species and made the first
collection of plants, in recognition of the wonderfiil con-
tribution that he has made to nature conservation in this
part of the southern Cape.
FIGURE 7, — LachenaUa lutzeyeri,
Duncan 5II (NBG): A, inflor-
escence, peduncle and leaves;
B, bulb and clasping leaf base;
C, 1/s flower; D, single flower.
A. B, X 0.8. Scale bar: C, D,
10 mm. Artist: Vicki Thomas.
Bothalia 37,1 (2007)
33
History: Lachenalia hitzeyeri is only known from the
type material, first collected by Heiner Lutzeyer (Figure
8B) in early November 2004, following an exten-
sive wild fire that burnt over the Witkransberg area of
Grootbos Private Nature Reserve, northwest of Gansbaai,
in late March 2004. Large numbers of plants were seen
in flower in November 2004 but during the correspond-
ing period in November and December 2005, the num-
ber of flowering plants had decreased drastically to just
a few individuals. A single flowering specimen was col-
lected on 15 November 2005 for illustration. In February
2006, the L. hitzeyeri type population was burnt again as
a result of a devastating fire that raged across most of
the Grootbos Private Nature Reserve and resulted in the
appearance of large numbers of flowering individuals in
November of that year.
Diagnostic features and affinities: Lachenalia hitzey-
eri is recognized by a moderately dense raceme of
oblong-campanulate, nodding, yellowish cream flowers
with olive green or dark brown gibbosities and well-
exserted stamens with weakly declinate, white fila-
ments. The inflorescence is borne on a rigid peduncle
and the inner tepals protrude 1-2 mm beyond the outer
tepals. The pedicels become ascending and the capsules
become suberect at fruiting, and the globose, shiny black
seeds have a terminal, inflated, medium-long strophiole
0.8-0. 9 mm long. The plants occur as solitary individu-
als within small to large groups, and have deep-seated,
globose bulbs that produce one or two narrowly lanceo-
late, prostrate, spreading or suberect, deeply canaliculate
or sometimes conduplicate leaves with flat or undulate,
cartilaginous margins (Figures 7, 8).
Lachenalia hitzeyeri appears to be most closely allied
to L. youngii Baker, another late-flowering species that
occurs much further east from Mossel Bay in the south-
ern part of the Western Cape to Humansdorp in the
southwestern part of the Eastern Cape. L. youngii has
similar oblong-campanulate, nodding flowers produced
on suberect white pedicels and is also a member of the
group of species having globose shiny black seeds with a
medium-long (0.5-0. 9 mm), terminal, inflated strophiole.
L. youngii differs mainly in having linear leaves with-
out cartilaginous margins, a non-rigid peduncle, shortly
exserted, straight stamens, purplish pink, unscented
flowers, and its shallow-seated bulbs are offset-forming
and not dependent on fire for flowering to oceur.
Distribution and habitat: Lachenalia hitzeyeri is
presently only known from the type locality on the
Witkransberg in the Grootbos Private Nature Reserve,
northwest of Gansbaai in the southern part of the
Western Cape (Figure 9). Possible reasons for it hav-
ing remained undetected until very recently are that
flowering is extremely erratic, due to its dependence
on the occurrence of summer or early autumn fires,
coupled with its very late flowering period and the slim
chance of it being recognized as a distinct taxon due to
its superficial similarity to other members of the genus.
It may well be recorded in other parts of the southern
Cape in due course. In Lachenalia, flowering perfor-
mance is greatly enhanced in a number of species in the
spring and early summer season immediately following
late summer or early autumn fires, such as in L. orchi-
oides (L.) Aiton and L. peersii Marloth ex W.F.Barker.
However, L. hitzeyeri is remarkable in being one of
only three members of this genus known to be entirely
dependent on the effects of fire for flowering to occur.
Until recently, two other pyrophytes endemic to mon-
tane fynbos of the southern Cape, L. montana Schltr. ex
W.F.Barker and L. sargeantii W.F.Barker were the only
FIGURE 8. — A. Lachenalia hitzeyeri
flowering in habitat. Grootbos
Private Nature Reserve; B.
Heiner Lutzeyer with a flow-
ering specimen of Lachenalia
hitzeyeri, 15 November 2005.
Scale bar: A, 10 mm.
34
Bothalia37,l (2007)
FIGURE 9. — Distribution of Lachenalia bitzeyeri.
other species known to flower solely in the wake of wild
fires (Duncan 1988). An example of erratic flowering in
one of these pyrophytic taxa was observed recently in L.
sargeantii. It was discovered in full bloom in November
1970 after a summer bum near Bredasdorp; only a few
individuals flowered there the following year, and it was
only seen in flower again 33 years later, in a new locality
in the southern Cape (Duncan & Edwards 2005; Duncan
et al. 2005). The typical phenological pattern displayed
by these three fire-dependent species is seen as lush
vegetative growth produced in the winter season imme-
diately following summer or autumn fires, followed by
prolific flowering in early summer, and dormancy from
midsummer to midautumn. During the second winter
season following a bum, most of the bulbs remain com-
pletely domiant, with an extremely small number of
individuals (or sometimes none at all) producing leaves
in winter, and flowers in early summer. The bulbs of
these three species all flower at ± the same time of year,
and are deep-seated, especially those of L. Iiitzeyeri that
can be buried up to 85 mm deep. All three species pro-
duce relatively few fibrous roots, but numerous contrac-
tile roots assist in pulling the bulbs deeply into the soil.
Deep-seated bulbs may be an adaptive strategy to escape
damage from severe fires. Rigid peduncles in Lachenalia
are only known in these three species.
At the type locality, Lachenalia lutzeyeh occurs
in a large colony, but as solitary individuals; no plants
seen appear to multiply by offset formation. They grow
in full sun in open aspects or between Table Mountain
Sandstone boulders, on a moderate slope with a south-
westerly aspect, at an altitude of 390 m. Following
a recent severe fire that burnt over most of Grootbos
Private Nature Reserve in Febmary 2006, large num-
bers of L. Iiitzeyeri have been observed growing in a
range of aspects in addition to the southwesterly aspect
of individuals at the type locality, especially on south-
ern and northern slopes. Companion plants of L. Iiitzey-
eri include Mimetes ciicullatus (Proteaceae) and various
genera of the family Restionaceae.
Paratype material examined
WESTERN CAPE. — 3419 (Caledon): Witkransberg in Grootbos
Private Nature Reserve, northwest of Gansbaai, on southwest-facing
mountain slope in Table Mountain Sandstone, (-CB), 15-11-2005,
Duncan 511 (NBG).
ACKNOWLEDGEMENTS
We extend our thanks to Mr Heiner Lutzeyer for
bringing this new taxon to our attention, Ms Vicki
Thomas for executing the line drawing, and Dr E.G.H.
Oliver for translating the diagnosis into Latin.
REFERENCES
DUNCAN, G.D. 1988. The Lachenalia handbook. Annals of
Kirstenbosch Botanic Gardens 17. National Botanic Gardens,
Cape Town.
DUNCAN, G.D. 1993. Lachenalia thomasiae. Flowering Plants of
Africa 52: t. 2062.
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. 1 997. Five new species of Lachenalia (Hyacinthaceae)
from arid areas of South Africa. Bothalia 27: 7-15.
DUNCAN, G.D. 1 998. Five new species of Lachenalia (Hyacinthaceae)
from arid areas of South Africa and Namibia. Bothalia 28: 131-
139.
DUNCAN, G.D. & EDWARDS, T.J. 2002. Anew species ot Lachenalia
from Namaqualand, South Africa (Hyacinthaceae: Massonieae).
Bothalia 32: 190-192.
DUNCAN, G.D. & EDWARDS, T.J. 2005. Lachenalia sargeantii.
Curtis 's Botanical Magazine 22: 1 76-184.
DUNCAN, G.D. & EDWARDS, T.J. 2006. Three new species of
Lachenalia (Hyacinthaceae: Massonieae) from Western and
Northern Cape, South Africa. Bothalia 36: 147-155.
DUNCAN, G.D., MCMASTER, C. & MCMASTER, R. 2005. Out of
the ashes. Veld <& Flora 91 : 66-69.
G.D. DUNCAN* and T.J. EDWARDS**
* South African National Biodiversity Institute, Kirstenbosch, Private
Bag X7, 7735 Claremont, Cape Town.
** Botany Department, University of KwaZulu-Natal, Private Bag
XOl, 3209 Scottsville, Pietermaritzburg. ^
MS. received: 2006-06-06.
FABACEAE
ASPALATHUS THERESAE, A NEW SPECIES FROM WESTERN CAPE, SOUTH AFRICA
INTRODUCTION
A.spalathii.s L., comprising 278 species, is the largest
genus endemic to South Africa (Dahlgren 1988). The
distribution of the genus lies mainly in the Cape Floristic
Region (CFR), where about 98% of the species occur,
and it is the second largest genus in the CFR (Goldblatt
& Manning 2002). Important contributions to the taxon-
omy of Aspalatlms were published in a series of publica-
tions in the 1960s by Dahlgren (I960, 1961, 1963, 1965,
Bothalia37,l (2007)
35
1967, 1968). Since then a single species was described
in the mid 1980s (Dahlgren 1984), but since the treat-
ment for the Flora of southern Africa (Dahlgren 1988)
no new species has been described.
During a field trip in 1997 in the Riviersonderend Moun-
tains, Western Cape, with colleagues Ted and Inge Oliver
and John Manning, I collected various legume plants.
Numerous attempts to identify an Aspalathiis species
failed. The specimen displays a combination of charac-
ters unique in the genus which justify its recognition as a
distinct species following the species concept applied by
Dahlgren (1960, 1963). He recognized species when the
individuals of one or more populations are morphologi-
cally distinct from those of other species by a marked
discontinuity in several characters. The distribution of
the species may or may not overlap with other closely
related species (Dahlgren 1960, 1963). This species
meets these criteria and it is accordingly described here.
Aspalathus theresae C.N.Cupido, sp. nov., A.
aspalathoides (L.) Rothm. affinis a qua differ! carina
rostrata, stylo sigmoideo, vexillo ovato-circulori, alls
lunatis, ovulis 4.
TYPE. — Western Cape, 3419 (Caledon): Rivierson-
derend, Riviersonderend Mtns, Olifantsdoom, northern
head, 1 280 m, (-BB), 7 November 1997, C.N. Cupido
29 (TNBG, holo.; PRE). Figure 10.
Erect, branched shrub up to 700 mm high; branches
densely leafy, villous or woolly. Leaves trifoliolate; leaf-
lets flat, oblong or narrowly elliptic, acute, 5. 5-9.0 x
1.5-2. 8 mm, coriaceous, green, with long villous hairs on
abaxial surface, adaxial surface glabrous. Inflorescence a
terminal head of 5-20 flowers, subtended by uppermost
leaves; pedicel 0. 5-3.0 mm long, covered with long vil-
lous hairs; bracts unifoliolate, linear, 5.8-8. 1 mm long,
covered with long villous hairs; bracteoles unifoliolate,
similar to bracts, 5. 5-5. 8 mm long, covered with long
villous hairs. Calyx campanulate, sericeous; lobes tri-
angular, upper lobes distinctly broader than remaining
lobes, 2. 2-3. 6 mm long. Corolla yellow; standard blade
ovate-circular, 6. 7-9.0 x 6. 8-8.0 mm, sericeous on back,
apex emarginate, base cordate, claw 3. 0-3. 4 mm long;
wing blade lunate, 6. 1-8.0 x 3. 1^.3 mm, sericeous on
most of lower parts, with several rows of minute folds,
claw ± 2.9 mm long; keel blade rostrate, 6.5-8. 1 x 3.5-
4.7 mm, upper margin slightly concave, sericeous on
lower parts, prominent pouch present, claw 2. 8^.0 mm
long. Stamens monadelphous; filaments united basally
for three quarters, forming a split sheath; anthers 5 short,
dorsifixed; 4 long, basifixed; 1 intermediate in size and
attachment. Pistil sessile, ovary and style base sericeous;
ovules 4; style sigmoid, stigma capitate. Pod obliquely
ovate, rostrate, 5. 5-6.0 x 3. 2-3. 5 mm, one-seeded.
Flowering time: November. Figure 11.
Consen’ation status: known from a single small popu-
lation on the Riviersonderend Mountains and is not
under threat. It must be considered as Vulnerable [D2,
World Conservation Union (lUCN) 2001].
Diagnostic features and affinities: no phylogeny for
Aspalathus exists to provide evidence for relationships
within the genus. However, Dahlgren (1988) established
FIGURE 10. — Holotype ol Aspalathus theresae.
informal groups to give recognition to morphological
similarities between supposedly related species. The
placement of A. theresae is uncertain, but a combina-
tion of several characters can be used to suggest possible
affinity. The flat, trifolioliate leaves and capitate inflores-
cence place A. theresae in the group Cephalanthae, one
of 34 groups recognized by Dahlgren (1988). The name
Cephalanthae is derived from the capitate inflorescence
characteristic of species in this group. Species in other
groups, particularly in the Sericeae may have these char-
acters. Within the Cephalanthae, A. theresae superfi-
cially resembles A. aspalathoides (L.) Rothm. The two
species are similar in general appearance, the number
of flowers in the inflorescence and the cordate shape of
the standard base. A. theresae differs from it in having
a rostrate keel, sigmoid style, ovate-circular standard,
lunate wing blades and four ovules. In contrast, the keel
in A. aspalathoides is obtuse, the style upcurved, the
standard elliptic, the wings oblong and the ovules two.
In addition to the differences in morphological features,
A. aspalathoides is not known from the Riviersonderend
Mountains. It occurs mainly on coastal mountains
from the Cape Peninsula to Bredasdorp, as well as
along the Langeberg Mountain range in Swellendam,
Riversdale and Montagu, Rooiberg Mountains south
of Ladismith and Keurkloof northwest of Barrydale.
Within the Cephalanthae, the rostrate keel and sigmoid
style are unique to A. theresae. A rostrate keel and sig-
moid style are characteristic of the groups Rostratae and
36
Bothalia37,l (2007)
Bothalia37,l (2007)
37
Crotalariiformes, but in these groups the inflorescences
are unifloral and situated on reduced lateral shoots or in
terminal groups of two to four flowers. The leaves are
also described as subterete or somewhat angular or flat
(Dahlgren 1988). In the Camosae, A. theresae shares the
rostrate keel, sigmoid style and capitate inflorescence
with the Cape Peninsula endemic species, A. capitata L.
However, this species differs from A. theresae in hav-
ing clusters of linear-pinoid leaves, elliptic to narrowly
obovate bracts, longer pedicels and two ovules.
Distribution and habitat: Aspalathus theresae is only
known from a single collection from the northern head
of Olifantsdoom in the Riviersonderend Mountains
(Figure 12). It grows in sand on an exposed rocky Table
Mountain Sandstone slope at an altitude of 1 280 m.
Etymology: the specific epithet is derived from my
mother’s forename, Theresa, in acknowledgement of her
love for plants, which she also instilled in me.
ACKNOWLEDGEMENTS
I wish to thank Ferozah Conrad for the Latin transla-
tion and Dr John Manning for valuable comments on the
final draft of the manuscript. Deshni Pillay assisted with
determining the conservation status and Tracey Nowell
is thanked for the line drawings.
REFERENCES
DAHLGREN, R. 1960. Revision of the genus Aspalathus I. The species
with flat leaflets. Opera Botanica 4: 1-393.
DAHLGREN, R. 1961. Revision of the genus Aspalathus 11. The spe-
cies with ericoid and pinoid leaflets 1. Opera Botanica 6: 1-69.
DAHLGREN. R. 1963. Studies on Aspalathus and some related genera
in South Africa. Opera Botanica 9: 5-301.
DAHLGREN, R. 1965. Revision of the genus Aspalathus II. The species
with ericoid and pinoid leaflets 4. Opera Botanica 10: 1-23 1 .
DAHLGREN, R. 1967. Some new and rediscovered species of
FIGURE 12. — Known distribution of Aspalathus theresae.
Aspalathus (Leguminosae). Botaniska Notiser 120: 26-40.
DAHLGREN, R. 1968. Revision of the genus Aspalathus II. The species
with ericoid and pinoid leaflets 6. Opera Botanica 2 1 : 1-309.
DAHLGREN, R. 1984. A new species oi Aspalathus (Fabaceae) from
the Prince Albert District. South African Journal of Botany 3:
259-261.
DAHLGREN, R, 1988. Crotalarieae (Aspalathus). Flora of southern
Africa 16, part 3, fasc. 6: 1^23.
GOLDBLATT, P. & MANNING, J.C. 2002. Plant diversity of the Cape
region of southern Africa. Annals of the Missouri Botanical
Garden 89: 281-302.
WORLD CONSERVATION UNION (lUCN) 2001. lUCN Red List
categories and criteria: version 3.1. lUCN Species Survival
Commission. lUCN, Gland, Switzerland and Cambridge, UK.
C.N. CUPIDO*
* Compton Herbarium, South African National Biodiversity Institute,
Private Bag X7, 7735 Claremont, Cape Town. Cupidocn@sanbi.org.
MS. received: 2005-12-19.
LAMIACEAE
RABDOSIELLA LEEMANNII, A NEW SPECIES FROM THE LIMPOPO PROVfNCE OF SOUTH AFRICA
Rabdosiella leemannii N.Hahn, sp. nov., affinis
R.calycinae sed ab ilia habito pererme fruticoso, foliis
minoribus, angustioribusque, ellipticis vel trullatis, (57-)
59-66(-70) X (25-)27-36(-38) mm, perianthio maiore
(6.9-)7.0-9.4(-9.7) mm, tubo (3.6-)3.7-4.9(-5.2) x 2.9-
3.1 mm mediente differt.
TYPE. — Limpopo, 2329 (Polokwane) [Pietersburg] :
Soutpansberg, Lejuma, 23°01'22.584"S, 29°25'57.180"E,
1 638 m, (-AB), 13-04-2005, in flower, N. Hahn 2086
(ZPB, holo.; PRE!, PRU!, iso.).
Phanerophyte, single or multi-stemmed shrub up to 2
m tall, usually not exceeding 1.5 m. Seedlings with dis-
tinctly succulent roots. Main stem up to 50 mm diam.,
showing multiple year rings, not ribbed in fresh mate-
rial; young stems pubescent, pale brown. Bark grey to
grey-brown, glabrous. Leaves decussate, rarely temate;
lamina shortly petiolate; petiole (l.l-)l. 9-3. 4(^.0) mm;
blade ovate to elliptic to trullate, size of mature leaf,
(57-)59-66-(70) x (25-)27-36(-38) mm, upper sur-
face pale green, strigose, lower surface grey, tomentose;
main veins 5-8, venation reticulate; margins crenate to
dentate, teeth 10-15 per side. Inflorescence a terminal
thyrse, 50-100 mm long. Flowers in pedunculate dicha-
sia, 20 mm long; peduncles 1-A mm; pedicel (3.6-)3.7-
4.9(-5.2) mm long. Calyx equally 5-toothed, tubular,
2.2^.9(-5.2) mm long in flowering stage extending to
1 X 6 mm in fruiting stage, green, glandular hairy, lobes
5, (1.1-)1.2-1.4 mm long, tinged purple at apex, at fruit-
ing (4.0-)4.2-5.0(-5.1) mm long. Corolla (6.9-)7.0-9.4
(-9.7) mm long; tube (3.6-)3.7-4.9(-5.2) x 2.9-3. 1 mm.
38
Bothalia37,l (2007)
FIGURE 13. — Rahdosiella leemannii. A, flowering branch, x I; B, infructescence, x 1; C, persistent calyx encasing nutlet, x 5; D, nutlet, x 15; E,
habitat, much reduced. Artist: G. Condy.
Bothalia37,l (2007)
39
FIGURE 14. — Holotype of Rabdosiella leemannii.
white-hairy; upper lobe (1.3-)1.4— 1.9(-2.0) mm, inside
tinged purple-pink, outside densely glandular hairy;
lower lobe navicular, (3.2-)3.1^.3(-^.5) mm, white on
outside, tinged purple-pink on inside. Stamens antrorse,
5.4— 7.5 mm long, attached to tube. Style antrorse, shorter
than stamens, 5.4— 7.2 mm long, bifid. Fruit a small,
brown, rounded nutlet, 0.9 x 0.7 mm. Figures 13, 14.
The genus Rabdosiella Codd was described in 1984
with two species, the widespread grassland suffrutex, R.
calycina (Benth.) Codd from South Africa, and R. ter-
nifolia (D.Don) Codd from India, Burma, Thailand and
China. Rabdosiella leemanii represents the third South
African species, and clearly belongs to Rabdosiella.
Rabdosiella can be distinguished from Plectranthiis
L.Her. by its woody growth, temate leaves and leaf-like
bracts, as well as its distinctly 10-nerved calyx (bearing
subequal teeth) becoming erect with maturity. Although
the delimitation of the genus Plectranthiis is not clear,
it is at present best to uphold Rabdosiella as a s^eparate
genus until a thorough multidisciplinary investigation of
Plectranthiis and its relatives can be undertaken.
Rabdosiella leemannii is a distinct species and differs
from R. calycina in several features such as its habitat,
habit and the vegetative and floral features (Table 1). It
is distinguished by its erect, shrubby nature (Figure 15),
woody stems, smaller leaves and flowers. The leaves of
R. leemannii are only rarely temate, smaller, narrower
(ovate to elliptic or tmllate), never becoming broadly
ovate as those found in R. calycina. The leaves of R.
calycina are often temate. The inflorescences and flow-
ers are also considerably smaller than those of R. calyc-
ina. R. leemannii is endemic to the Soutpansberg and
Blouberg Afromontane mist-belt open bushland (Figure
16) above 1 400 m, where it forms perennial shmbs
with woody stems reaching 50 mm in diameter at the
base and displaying up to 10 clearly defined year rings.
The habitat consists of Soutpansberg Summit Sourveld
and a mosaic of the latter and Soutpansberg Mountain
Bushveld (Mucina et al. 2005). Hahn (2002) has pointed
out the importance of mist precipitation on the endemic
flora of the Soutpansberg. In times of drought, mist pre-
cipitation is the only source of moisture available in
this habitat. However, total precipitation can almost be
doubled if mist precipitation is added to rainfall. On
herbarium sheets its habitat is occasionally stated as
grassland — this is an error as the ‘grassy component’ is
in fact the sedge, Coleochloa setifera (Ridl.) Gilly. The
habitat of R. leemannii is quite sheltered from fire with a
maximum period of five years. In the absence of fire, R.
leemannii forms single-stemmed shrubs, however, plants
exposed to fire do resprout forming multi-stemmed
shrubs. Rabdosiells calycina is clearly a herbaceous suf-
frutex, annually burnt. It is commonly associated with
grassland of the eastern escarpment of South Africa
occurring as far north as the Soutpansberg.
Both Rabdosiella leemannii and R. calycina have
been cultivated at the Kirstenbosch National Botanical
Gardens (E.J. van Jaarsveld pers. comm.). In spite of
being cultivated under similar conditions, each retained
their characteristic growth form. Seed of R. leeman-
nii germinates readily within three weeks, immediately
forming distinctly fleshy roots.
FIGURE 15. — Plant from which type specimens were collected.
40
Bothalia 37,1 (2007)
TABLE 1 . — Comparison of selected characters of Rabdosiella leeman-
The species is named after the great uncle of the first
author, Albert Conrad Leemann (misspelt Leeman in
Gunn & Codd 1981), who in 1933 was the first to collect
a specimen while on an excursion to the Blouberg. His
findings were published in Vegetations bi I der (Leemann
1935). In this article he made mention of the wholesale
destruction of the environment at the foot of the moun-
tain. He stressed the importance of conserving mountains
such as the Blouberg which act as refugia of undisturbed
habitat. These refugia are of immense importance to our
understanding of vegetation changes caused by man.
ACKNOWLEDGEMENTS
A special word of thanks to Ernst van Jaarsveld for
his invaluable comments whilst writing this article and
sharing his knowledge on the cultivation of Rabdosiella
leemannii. We would also like to thank the South African
National Biodiversity Institute for allowing us access
to their herbarium and Mr John Lavranos for the Latin
diagnosis.
Other specimens examined
LIMPOPO. — 2229 (Waterpoort): Soutpansberg, Farm Wellington,
northern side of farm, 1 400 m, (-DC), 25-03-1994, Rossouw 208
(PRE); northern slopes of Hanglip near summit, (-DD), 02-04-1957,
Meeuse 10164 (PRE); Farm Buckworth (Dr Bird’s farm), south-
ern slope north of Bird’s cottage near summit, (-DD), 12-05-1957,
Meeuse 10243 (PRE). 2328 (Baltimore): Blouberg, mountain grass-
land, aspect south, shrub, 23°04'37.922"S, 28°59T0.506"E, 1 990
m, (-BB), 11-05-1999, Hahn 1574 (ZPB); in kloof leading to bea-
con, 5 600 ft [1 847 m], 29-04-1954, (-BB), Codd 8759 (PRE); alt.
6 000 ft [1 968 m], (-BB), 10-03-1933, Leemann 118 (PRE); kloof
near top, 4 ft [1.3 m], (-BB), 26-06-1961, Strey & Schlieben 8517
(PRE); kloof below beacon, (-BB), 26-4-1961, Van der Schyff 5416
(PRE). 2329 (Polokwane): Soutpansberg, Farm Lejuma, Mt Lejuma,
23°01'22.584"S, 29°25'57.180"E, 1 636 m, 22-05-1982, (-AB), Venter
7855 (PRE); Lejuma south of Mt Lejuma, (-AB), 23-05-1982, Venter
8754 (PRE).
REFERENCES
CODD, L.E. 1984. The genus Isodon (Schrad. ex Benth.) Sprach in
Africa and a new genus Rabdosiella Codd (Lamiaceae). Bothalia
15: 7-10.
GUNN, M. & CODD, L.E. 1981. Botanical exploration of southern
Africa. Balkema, Cape Town.
HAHN, N. 2002. Endemic flora of the Soutpansberg. M.Sc. thesis.
University of Natal, Pietermaritzburg. Unpublished.
LEEMANN, A.C. 1935. Der Blaauwberg im Norden von Transvaal.
Vegetationsbilder 24: t. 43—48.
MUCINA, C., RUTHERFORD, M.C. & POWRIE, L. (eds). 2005.
Vegetation map of South Africa, Lesotho and Swaziland, 1 :
1 000 000 scale sheet maps. South African National Biodiversity
Institute, Pretoria.
N. HAHN* and G.J. BREDENKAMP*
FIGURE 16. — Distribution of Rabdosiella leemannii, and R. calyci-
na, O, in South Africa.
* Botany Department, University of Pretoria, 0001 Pretoria.
MS. received: 2006-01-31.
BURSERACEAE
COMMIPHORA KUNENEANA, A NEW SPECIES FROM THE KAOKOVELD, NAMIBIA
Commiphora kuneneana Swanepoel, a new species
from the Kaokoveld Centre of Endemism (Van Wyk
& Smith 2001), is described. During field work for
the Namibian Tree Atlas project in the Kunene River
Valley and Sesfontein areas, the author encountered
Commiphora trees with morphological features that were
characteristic of both C. saxicola Engl, and C. crenato-
serrata Engl. (Van der Walt 1974, 1986). Similar plants
were subsequently found to be widespread in the west-
ern Kaokoveld and are here proposed as representing a
distinct new species.
A study of the Commiphora holdings in PRE and
WIND revealed several collections of the new species,
all filed under either C. saxicola or C. crenato-serrata. In
the Flora of southern A frica treatment of Commiphora,
one of these specimens {De Winter & Leistner 5876) is
mentioned as a possible new species, C. crassifoliolata
Bothalia37,l (2007)
41
FIGURE 17. — Commiphora kime-
neana. A, tree, 6 m tall, with-
out leaves and with flattened
crown; B, tree, 5 m tall, in
leaf; C, shrub-like trees.
Mendes nom. prov. (Van der Walt 1986: 34). Herbarium
specimens of C. kuneneana can be mistaken for C. saxi-
cola and C. crenato-serrata due to similarities in leaf
morphology. In habit, however, C. kuneneana differs
conspicuously from C. saxicola over most of its range,
in that it is a tree, with a distinct, erect trunk (Figure
17A, B). In the field, when without fruit, C. kuneneana
can also be confused with C. crenato-serrata because of
its similarities in habit, bark, exudate and leaves. Apart
from morphological differences, the three taxa also have
42
Bothalia 37,1 (2007)
Illustrations: Van der Walt: t. 31, as C. saxicola (1974: 19). Steyn:
69, top left; 70, bottom left as C. saxicola (2003). Steyn: 53, top left &
right; 54, centre & right as C. crenato-serrata (2003).
Dioecious tree without spines, up to 8 m tall, usually
infundibular in shape, crown rounded or ± flat-topped.
Trunk single, occasionally multi-stemmed, cylindrical, ±
straight, erect, 0.8-3. 5 m long, up to 400 mm in diam.,
rarely a shrub-like tree branching from just above
ground level (Figure 17C). Bark pale grey, reddish grey,
cream-coloured or pale yellowish, smooth, not peeling,
with minute, shallow, longitudinal fissures (Figure 18).
Branches and branchlets with scattered small lenticels,
obtuse, glabrous, youngest ones with glandular hairs and
with long flexuous branched hairs at apex, yellowish,
dark or reddish brown or grey to reddish grey; branchlets
short, scarred. Exudate milky, glutinous, not squirting
when branches or branchlets damaged or cut, aromatic,
producing a soft, caramel-brown or hard olive-green or
pale yellow transparent resin.
FIGURE 18. — Commiphora kuneneana. Bark.
different distribution ranges: C. kuneneana has a more
northerly distribution than C. saxicola and a more west-
erly distribution than C. crenato-serrata. However, in
the northwestern parts of its range in Namibia, in the
Otjihipa Mountains, C. kuneneana is sympatric with C.
crenato-serrata.
Diagnostic morphological characters to differentiate
between C. kuneneana, C. saxicola and C. crenato-ser-
rata are presented. Apart from examining the herbarium
collections of the three species in PRE and WIND, live
material from numerous populations in Namibia were
studied in the field. Morphological characters in the fol-
lowing description were all determined, from mature
leaves, fresh flowering material and ripe fruit.
Commiphora kuneneana Swanepoel, sp. nov., C.
saxicolae Engl, simillima cortice laevi non deglubenti,
foliis pinnatis, laminis viridibus pilis longis glandulari-
busque, saepe subconduplicatis, foliolis sub-orbiculari-
bus, ovatis, obovatis vel ellipticis; inflorescentiis saepe
thyrsoideis, pedunculo glanduloso-piloso, floribus perig-
ynis, disco intus 8-plicato, lobis apice bifidis non hypan-
thio adnato, pseudo-arillo cupulare, putamen 25-60%
tegenti, brachiis commissuralibus 2(0) brevibus, lobis
facialibus 2(1). A C. saxicola statura plerumque arboris,
caule proprio, recto, cylindrico, erecto, foliis plerumque
duplo maioribus, lamina saepe plana, plerumque maiori,
saepe ovata vel lanceolata, raro rhombea, petiolo saepe
cum fasciculis vascularibus medullaribus, petiolulis
saepe longioribus, inflorescentiis saepe paniculoso-thyr-
soideis, pedunculo plerumque longiori (usque ad 400
mm), antheris plerumque longioribus (usque ad 2.1 mm),
pseudo-arillo semper rubro, differt.
TYPE. — Namibia, 1713 (Swartbooisdrif): 3 km
WSW of Epupa Falls on main road to Okangwati, 690
m, (-AA), 14-01-2006, Swanepoel 211, (WIND, holo.!;
PRE, iso.!).
Leaves imparipinnate (Figure 19), rarely paripinnate
or intermediate, (2)3-6(-9)-jugate, rarely trifoliolate,
up to 280 mm long, grouped closely together at end of
branches and branchlets, spirally on shoots, green; leaf-
lets with scattered, short glandular hairs on both sides,
especially on and along midrib, long glandular and
long flexuous hairs, some branched, also usually pres-
ent, otherwise glabrous; lamina flat or subcondupli-
cate, shape varies considerably, even on same tree and
same leaf; terminal leaflets elliptic to broadly elliptic,
ovate, rarely broadly lanceolate, rhombic or suborbicular
(15-)23-38(-68) x (10-) 18-28(^2) mm; lateral leaf-
lets narrowly ovate to broadly ovate, elliptic to broadly
elliptic, lanceolate to broadly lanceolate, rarely obovate
or suborbicular, (12-)19-36(-64) x (6-)18-36(-45)
mm; apex acute, acuminate or obtuse but with minute
tip usually acute, base cuneate, truncate or obtuse and in
lateral leaflets also cordate, base often abruptly attenu-
ate onto petiole, often oblique and then asymmetric with
one side abruptly attenuate onto petiole; margin crenate-
serrate, occasionally dentate near apex, 5-26 teeth per
side, usually entire near base, rarely almost entire; mid-
rib green or yellowish green, conspicuous and promi-
nent abaxially; petiole with scattered, short glandular
hairs, usually with long glandular and long flexuous
simple or branched hairs, otherwise glabrous, (7-)27-
56(-72) mm long, pentagonal, ovate or triangular in
t/s with (8-)10-13(14) vascular bundles, some petioles
with additional one or two meduliary vascular bundles
lacking xylem element, occasionally shallowly sulcate
adaxially, sectional dimensions (0.9-)1.2-1.8(-2.2) x
(1.2-)1.5-1.9(-2.4) mm; rachis and petiolules with short
glandular hairs, often with few additional long flexuous
hairs, some branched; petiolule on terminal leaflets up to
35 mm long, on lateral leaflets up to 23 mm long or leaf-
lets rarely subsessile.
Inflorescence thyrsoid or paniculose-thyrsoid (Figure
20A), with short glandular hairs, often with additional
long, simple or branched hairs near apex; peduncle up to
400 mm long, at apex of branches and on dwarf lateral
branchlets. Flowers subsessile or pedicellate, unisexual.
Bothalia37,l (2007)
43
perigynous, precocious or flowering with leaves. Bracts
linear-triangular, up to 4 mm long, bracteoles narrowly
triangular or narrowly elliptic, up to 1.5 mm long, apex
acute, with glandular hairs abaxially and on margin, often
with long simple or branched hairs. Calyx green, yellow-
ish green or greenish red, with glandular hairs, lobes
triangular or triangular-ovate, apex acute. Petals green-
ish yellow, often reddish along centre, glabrous or with
short glandular hairs abaxially over apical part, recurved
towards apex but minute tip indexed. Disc cylindrical
with 4(5) distinct lobes, adnate to hypanthium but distal
part of lobes free. Male flowers 2. 5-5.4 mm long; pedi-
cel up to 1.3 mm long, with glandular hairs, often with
few long hairs; calyx 1.3-1. 9 mm long, calyx lobes 4(5),
0.2-0. 7 mm long; petals 4(5), oblanceolate to broadly
oblanceolate or narrowly elliptic, 2. 7-5. 2 x 1.2-1. 8 mm;
disc fleshy, inside of disc 8-folded, often with maroon
markings, lobes bifid at apex; stamens 8(10), 4(5) long
stamens with filaments 1.0-2. 8 mm long, inserted on top
of disc lobes, 4(5) short ones with filaments 0.4— 1.7 mm
long, inserted on margin of disc between lobes; anthers
on long stamens 0.7-2. 1 mm long, anthers on short sta-
mens 0.6-1. 6 mm long; filaments subterete, on long sta-
mens slightly flattened and broadened over lower part;
gynoecium rudimentary (Figure 20B-E). Female flowers
2. 8-3. 3 mm long; pedicel up to 0.9 mm long, with glan-
dular hairs, often with few long hairs; calyx 1.1 -1.4 mm
long, lobes 0.3-0. 6 mm long; petals broadly oblanceo-
late or narrowly elliptic, 2. 5-3. 5 x 1.0-1. 3 mm; disc not
very fleshy, lobes bifid at apex; staminodes 8, 4 lojig and
4 short; ovary half inferior, glabrous or with few short
glandular hairs; style from ± short to ± long, sutures
grooved; stigma obscurely 4-lobed; pistil 2. 3-3.0 x 0.6-
0.8 mm (Figure 20F-I).
Fruit a drupe, ellipsoid, oblong-ellipsoid or ovoid,
10-17 X 5-9 X 5-9 mm, often slightly constricted
midway between base and apex, often slightly apicu-
late (Figure 20J); pericarp 2-valved, suture rectilinear
or rarely slightly curved towards sterile locule; exo-
carp glabrous or with scattered, short, glandular hairs,
FIGURE 19. — Commiphora kime-
neana. Leaves. Scale bar: 20
mm.
non-glutinous, reddish green in ripe fruit; mesocarp
not very fleshy; putamen 5.7-10.7 x 3.5-6. 1 x 3.1— 4.4
mm, flattened, asymmetrical-ellipsoid, oblongoid or
oblong-ellipsoid, with one fertile and one sterile loc-
ule, smooth or slightly rugose (Figure 20K-N); fertile
locule in sutural view asymmetrically convex towards
apex, rarely convex or rectilinear and tapering to base
and apex, apex often bent over to sterile locule, convex
in apical view; sterile locule dorsally ridged, variable in
sutural view, convex near base and rectilinear to concave
towards apex, rectilinear and tapering to base and apex,
or slightly humped near base and apex, ± triangular in
apical view; suture rectilinear but convex towards sterile
locule at apex, rarely rectilinear; angle between locules
at apex 50°-105°; pseudo-aril red, fleshy, cupular, cov-
ering 25%-60% of the locules equally, with 2(0) very
short commissural arms up to 1.8 mm long and 2(1)
facial lobes; lobes convex or triangular: on fertile locule,
0.3-1 .4(-2.6) mm long, usually convex, rarely drawn out
into a long fragmented arm, almost reaching apex, often
undeveloped and completely absent; on sterile locule,
0. 5-2.0 mm long, usually triangular and larger; apical
pits small or absent. Flowering time: August to January.
Diagnostic characters and affinities'. Commiphora
kuneneana is probably most closely related to C. saxi-
cola, the species with which it has hitherto most often
been confused, and also to C. crenato-serrata. Salient diag-
nostic morphological characters to differentiate between
the three species are supplied in Table 2.
The shrub-like tree form of Commiphora kuneneana
was only observed in the far western parts of its range,
in the extremely arid area bordering the Namib Desert.
In this area it is not unusual to find both tree and shrub-
like forms growing side by side. Commiphora saxicola is
only rarely a small tree up to 4 m tall with the trunk and
main stem flexuous, semiterete, up to 2.4 m long, often
with thinner lateral branches at the base and an irregular
crown.
44
Bothalia37,l (2007)
FIGURE 20, Commiphora kunenearta. A, male inflorescence. B-E, male flower: C, calyx and corolla partly removed; D, disc as seen from inside;
E, disc as seen from above to depict position of stamens (black) and rudimentary ovary (circle). F-I, female flower: G, calyx and corolla
partly removed; I I, disc as seen from inside; I, disc as seen from above to depict position of stamens (black). J, fruit. K-N, putamen with
p.scudo-aril. K, side with fertile locule; L, side with sterile locule; M, lateral view, fertile locule right, sterile locule left; N, apical view. A— E,
Swanepoel 2J8; F-N, Swanepoel 21 1 . Scale bars: A, 5 mm; B, C, F, G: I mm; J, 3 mm; K-N, 2 mm. Artist: Julia Kreiss.
Bothalia37,l (2007)
45
TABLE 2. — Salient morphological differences between Commiphora kuneneana, C, saxicola and C. crenato-serrata
46
Bothalia 37,1 (2007)
TABLE 2. — Salient morphological differences between Commiphora kimeneana, C. saxicola and C. crenato-serrata (cont.)
Bothalia37,l (2007)
47
12 14 16 18 20 22 24
FIGURE 21. — Known distribution of Commiphora kimeneana.
It is often difficult to determine a specimen’s true
identity using morphological characters alone. However,
a comparative anatomical study of petioles all over the
ranges of the two species revealed that specimens with
more than 11 vascular bundles and/or medullary vascu-
lar bundles in addition, are restricted to the area north of
± 20°S. Hence it is concluded that C. kimeneana occurs
as far south as ± 20°S and C. saxicola to the south of ±
20°S. Although no evidence was found during the pres-
ent study, the two taxa may be sympatric in the area.
In plants of Commiphora kuneneana from the Kunene
River Valley and Otjihipa Mountains, the lamina of the
leaflets tends to be lanceolate more often, the leaf api-
ces to be acuminate and the petiolules to be longer than
elsewhere in its range in Namibia. In these areas the fruit
is also consistently constricted midway between base
and apex. In the absence of fruiting material, confusion
with C. crenato-serrata is likely in these areas, due to
similarities in lamina shape, habit and habitat prefer-
ences. Leaves of C. crenato-serrata, however, usually
are larger, up to 460 mm long, the lamina is also usually
larger, 30-130 x 10-70 mm long, always lanceolate in
shape and the margin has up to 44 teeth per side. The
petiole has 11-17 vascular bundles and 1-6 medullary
vascular bundles, the latter including xylem, are always
present (Van der Walt & Van der Schijff 1973). The
leaves and lamina of C. kuneneana usually are smaller,
variable in shape and the margin usually has fewer teeth
per side. Furthermore, the petiole in C. kuneneana usu-
ally has fewer vascular bundles and only some petioles
have medullary vascular bundles (lacking the xylem
element). The fruit, when apiculate, is only slightly so.
Inflorescences of C. crenato-serrata are always thyrsoid,
the flowers usually larger and the fruit is strongly apicu-
late and not constricted between the base and apex.
Etymology, the specific epithet refers to the Kunene
Region, named after the Kunene River in northwestern
Namibia where the new species is found. As English and
Afrikaans vernacular names, I propose ‘Kunene cork-
wood’ and " Kunene-kanniedood\ respectively. The local
Ovahimba people in the Epupa area use the name omu-
whanga for C. kuneneana. However, in the Otjitanda
area, where C. kuneneana is absent, the same name is
used for C. crenato-serrata. As with many other spe-
cies of Commiphora, the trunk is damaged by a hacking-
knife and the sap so obtained is used as a thirst quencher
by both the local people and domestic animals.
Distribution: Commiphora kuneneana is only known
from the Kaokoveld Centre of Endemism in north-
western Namibia. It occurs from the Kunene River
southwards to ± 20°S, the area between Palmwag and
Sesfontein (Figure 21). It almost certainly also occurs in
the adjacent parts of southern Angola, as it was collected
at several localities on the Namibian side of the Kunene
River Valley. It ranges from common to rare, often
growing amongst other species of Commiphora, such
as C. anacardiifolia, C. crenato-serrata, C. discolor, C.
glaucescens, C. kaokoensis, C. steynii, C. tenuipetiolata,
C. virgata and C. wildii. C. kuneneana is most abun-
dant in the Kunene River Valley to the west of Epupa
Falls and in the mountainous areas between Sesfontein
and Purros. It is rare to the east of Epupa in the Kunene
River Valley.
Habitat and ecology: Commiphora kuneneana occurs
in the Kaokoveld, including the pro-Namib and the
Escarpment area. It is found 35-180 km from the coast
at altitudes of 200-1 800 m, where the mean annual
rainfall is 50-250 mm (Mendelsohn et al. 2002). It is not
very habitat specific and is found on mountain slopes,
plateaus, valley floors and in drainage lines on sandy
flats and rocky areas. It does not appear to be limited to
any specific geological formation or substrate. C. kune-
neana is not threatened as it occurs in remote, unpopu-
lated areas mostly within the limits of conservancies.
Other specimens examined
Basson Bas 1-12-2001 WIND.
Craven 3061, 3068, 3164 WIND. Curtis 417, 559 WIND. Curtis,
Aronson, Le Floe 'h & Le Floe 'h Cur 1662a, b WIND.
De Winter & Leistner 5670, 5876 PRE, WIND.
Giess 8921 PRE, WIND; 9401 WIND.
Hearn 176 WIND.
Merxmiiller & Giess 1430 PRE, WIND. Muller & Loutit 2196 WIND.
Steyn 24 WIND. Sullivan 39 WIND. Swanepoel 2A, B, 3A, B, 4A, 191-
210 WIND; 211 PRE, WIND; 212-220, 223 WIND.
Finley 1615 WIND.
Van der Walt 243 PRE, WIND. Viljoen 306, 452 WIND.
Ward, Ward & Seely 10455 WIND.
ACKNOWLEDGEMENTS
I would like to thank Prof A.E. van Wyk, University
of Pretoria, for advice and support. Dr H.F. Glen,
SANBI, for translating the diagnosis into Latin, Ms
Hester Steyn, SANBI, for preparing the distribution
map and Ms Julia Kreiss for the line drawings. The
curator and staff of the National Herbarium of Namibia
are thanked for their assistance during visits to the her-
barium. The National Herbarium of Namibia and the
South African National Biodiversity Institute are also
thanked for the use of information from their databases:
SPMNDB, Flora DB and PRECIS. The curator. National
Herbarium, Pretoria, is thanked for access to their collec-
48
Bothalia37,l (2007)
tions; the assistance of Dr C.L. Bredenkamp and Ms M.
Jordaan during visits to the herbarium is acknowledged
with thanks. Ms V. Papworth from the National History
Museum, London, is thanked for images of Angolan
material and Mr B. Olivier for flowering material. Mr
K. Verwey of Otjinhungwa is thanked for logistical sup-
port during visits to the Marienfluss. The University of
Pretoria is thanked for financial support. I am especially
grateful to my wife Hannelie for assistance and support
during field trips.
REFERENCES
MENDELSOHN, J., JARVIS, A., ROBERTS, C. & ROBERTSON, T.
2002. Atlas of Namibia. Philip, Cape Town.
STEYN, M. 2003. A field guide, southern Africa Commiphora/ '«
Veldgids, suider-AfHka Commiphora. Published by the author,
Polokwane.
VAN DER WALT, J.J.A. 1974. A preliminary report on the genus
Commiphora in South West Africa. Madoqua 1 : 5-23.
VAN DER WALT, J.J.A. 1986. Burseraceae. Flora of southern Africa
18,3: 5-34.
VAN DER WALT, J.J.A. & VAN DER SCHIJFF, H.P. 1973. The anato-
my of the petiole as an aid to the identification of South African
Commiphora species. Kirkia 9: 95-108.
VAN WYK, A.E. & SMITH, G.F. 200 1 . Regions offloristic endemism in
southern Africa. A review with emphasis on succulents. Umdaus
Press, Hatfield, Pretoria.
W. SWANEPOEL*
*H.G.W.J. Schweickerdt Herbarium, Department of Botany,
University of Pretoria, 0002 Pretoria, South Africa. Postal address:
P.O. Box 21168, Windhoek, Namibia.
MS. received: 2006-05-05.
APOCYNACEAE
TRANSFER OF SCHIZOGLOSSUM UMBELLULIFERUM TO STENOSTELMA, AND ITS NEOTYPIFICATION (ASCLEPIADOIDEAE)
Rudolf Schlechter originally described Schizoglossim
umbelluliferum in 1895 after a collecting trip to South
Africa. The specimen {Schlechter 3687) on which he
based his description was collected between 28 October
and 15 November 1893 (Gunn & Codd 1981) on the
plains below the Magaliesberg, a range of mountains
near Pretoria, South Africa.
Schlechter ’s placing of this species in the genus
Schizoglossum E.Mey. was based on habit and floral
characteristics. As circumscribed at that time in the
protologue, he mentioned the affinities between S.
umbelluliferum and S. orbiculare Schltr. Brown (1907)
agreed with Schlechter ’s generic placement and indi-
cated that, in his opinion, S. umbelluliferum might be
conspecific not only with S. orbiculare but also with S.
crassipes S. Moore. However, he did not see Schlechter ’s
type specimens, and his opinion was based only on the
descriptions of S. umbelluliferum and S. orbiculare.
Unfortunately, Schlechter’s type specimens, previously
housed at the Berlin Herbarium (B), could not be traced
and were probably destroyed during the bombing of
this herbarium during the Second World War (Nicholas
1992). According to Schlechter (1895), he had collected
only one specimen each of S. umbelluliferum and S. orbi-
culare, and it is thus doubtfijl that any other Schlechter
specimens of these species will be found in any other
herbaria.
Some living plants precisely fitting Schlechter’s
description of Schizoglossum umbelluliferum were recently
rediscovered by the senior author in two subpopula-
tions at the foot of the Magaliesberg. This discovery has
enabled a re-assessment of the generic position, neotypi-
fication and re-circumscription of this species.
TAXONOMY
Slenostelma Schltr. is currently represented in southern
Africa by two taxa, namely S. capense Schltr. and S. cor-
niculatum (E.Mey.) Bullock (Victor et al. 2003). Kupicha
(1984), during her study of the genus Schizoglossum,
suggested an expanded concept for Stenostelma and
argued for the inclusion of a number of Schizoglossum spe-
cies, including S. umbelluliferum. However, she never pub-
lished these new combinations. Nicholas (1999) agreed
with Kupicha and expressed the intention to enlarge the
current circumscription of Stenostelma to also include
three species previously placed in Xysmalobium R.Br.
These transfers have not yet been effectively or validly
published but would bring the number of species under
Stenostelma to ten.
The proposed transfer of Schizoglossum umbellu-
liferum to the genus Stenostelma is based on a number of
correlated characteristics which may be considered synapo-
morphies for the genus, namely: a fleshy, tough and fibrous
napiform tuber; globose or subglobose inflorescences;
greenish yellow to cream-coloured flowers; corolla lobes
that are divided almost to the base, usually with a ter-
minal oblique notch; corona lobes free, erect, somewhat
fleshy, sometimes with an inverted v-shaped indentation
with thickened edges, below the apex on the inner sur-
face and a gibbous or keeled outer surface; anther flaps
that usually fully cover the style head; barrel-shaped
(constricted below and above) style head (in longitudinal
section); triangular anther wings with a medial or sub-
tenninal notch in profile from the side; gynostegial head
conical-shaped in outline; apical attachment of the trans-
lator arms to the pear-shaped pollinia; and erect follicles,
narrowly fusiform (not inflated), smooth or with ± six,
slight longitudinal ridges.
Stenostelma umbelluliferum can be easily distin-
guished from the other southern African species cur-
rently recognized in this genus in that it has the smallest
plants (38-200 mm tall above ground) and has globose,
slightly dorsiventrally flattened corona lobes with no
extended horns, teeth or other processes.
Bothalia37,l (2007)
49
FIGURE 22. — Stenostelma umbellulifenim. A, habit with inflorescences, leaves, follicle and tuber; B, inflorescence. C, 1/s flower: Ca, corolla lobe;
Cb, style head; Cc, calyx; Cd, carpel; Ce, hairs. D, pollinarium: Da, corpusculum; Db, translator-arm; Dc, pollinium. E-G, seed: E, dorsal
view; F, ventral view; G, with coma. H, dehiscent follicle. A, B, Chiliza & Bester 2; C-H, Bester 5251. Scale bars: A, G, H, 25 mm; B, 2.2
mm; C, 1 mm; D, 125 pm; E, F, 1.3 mm. Artist: Sibonelo Chiliza.
50
Bothalia 37,1 (2007)
FIGURE 23. — Stenostelma umbelluliferum, Bester 5251. A-C, c/s lamina showing revolute margins: A, basal portion; B, apical portion; C, position
of sections. D-G, leaf: D, E, tip, abaxial and adaxial view; F, G, base, abaxial and adaxial view. Scale bars: A, B, 0.5 mm; D-G, 2.3 itun.
Artist: Sibonelo Chiliza.
Stenostelma umbelluliferum (Schltr.) S.P.Bester
& Nicholas, comb. nov.
Schizoglossum umbelluliferum Schltr., in Botanische Jahrbiicher
20,5, Beibl. 51: 24 (1895). Type: Farm Doompoort, Pretoria, Gauteng,
South Africa, 25°38'36" S, 28°15'01" E, 1 222 m, 16 September 2004,
S.R Bester 5251 (PRE, neo. selected here; K, MO, B, Z, iso.).
Perennial geophytic herb with milky latex; aerial
parts annual, 38-200 x 40-300 mm. Tuber fleshy, napi-
form, broadest above middle, 50-100(-140) x 4-18 mm,
pale cream-coloured, outer layer fibrous (Figure 22).
Underground stem arising from tuber crown, 30-1 10 mm
long, containing numerous adventitious points, usually
unbranched; intemodes 22-25 mm long. Aerial stems
arising from apex of underground stem, 1-6-branched
from soil level with further secondary branching, erect-
spreading to decumbent from base; intemodes 3-12 mm
long, densely foliate. Leaves opposite, simple, erect to
spreading-erect, 9-16 per branch; lamina narrowly lin-
ear to triangularly linear, up to 90 x 5 mm, acute, base
attenuate to cuneate, usually narrowing into a very short
petiole, nearly glabrous, green to tinged purple above,
becoming dark pink to maroon with age, paler green
towards main vein, paler beneath, margins revolute espe-
cially distinct in younger leaves; main vein prominent,
abaxially concave, smooth to scabrid, adaxially convex,
glabrous to pubemlous to scabrid, secondary veins abax-
ially paler (Figure 23). Inflorescence umbellate, axillary,
alternate, up to 22 umbels per plant, up to 7 per branch,
(l-)3-28 flowers per umbel; pedimcles 5-30 mm long,
pubescent, deflexed at apex after flowering; pedicels
2-18 mm long, drooping after flowering, pubemlous,
hairs flat, multicellular; bracts linear to narrowly lan-
ceolate, up to 1.5 X 0.24 mm. Flowers 5-merous. Calyx
free, erect to spreading-erect; lobes lanceolate, 0.8-1 .5 x
0.45-0.65 mm, half as long as corolla, acute, dark green,
outside glabrous, inside glabrous to pubemlous. Corolla
± divided to base, erect to spreading-erect; lobes oblong-
lanceolate, obliquely notched, 1.8-0.24 x 0.10-0.12 mm,
convex in basal half aind concave in apical half, revo-
lute, pinkish inside, greenish outside, both sides darken-
ing to maroon as they mature; apex rounded to obtuse,
both sides glabrous. Staminal corona arising at base of
staminal column; lobes free, erect, ovate to, oblong-
ligulate, 0.5-0. 8 x 0.4-0. 6 mm, obtuse to acute, basally
ovoid, keeled on sides, inner surface flat, outer surface
bulged, much shorter than staminal column, green.
Staminal column cylindrical or barrel-shaped, 0. 7-1.1
mm high. Stamens: anther wings triangular, broadest at
base or middle, 0.2 x 0. 1 mm; anther appendages ovate
to broadly ovate, 0.35-0.4 x 0.2-0.3 mm, apex obtuse to
acute, erect-inflexed over conical style apex. Pollinaria
solitary, pendulous in each anther sac, obliquely pear-
shaped. Ovaries 2, subinferior, many-ovuled, glabrous;
styles fused into a stylar column, terete, style head orbic-
ular to conical-orbicular, exceeding anthers; translators
from lateral surface of style head inverted y-shaped,
60-140 X 40 pm, caudicles filiform, widely diverging,
apically fused to corpusculum, corpusculum oblong-nar-
rowly-ovoid to oblong-ovoid, subacute. Follicles up to 4
per branch, usually 1(2) per umbel, narrowly fusiform,
noninflated, ± equally tapering at both ends, tips slightly
broader, obtuse, up to 85 x 8 mm, minutely papillate or
pubemlous, green striped with cream to green, maturing
to darker green turning to pinkish then maroon, or some-
times green to glaucous with darker lateral striations,
usually longer than 70 mm at maturity, fhiiting stalk
swirled and then upturned. Seeds 25^5 per follicle,
broadly ovate, 3.0-3. 5(-4.0) x 1.4-1. 6 mm, dorsiven-
trally flattened, bifacial, minutely mgulose to favulariate,
above slightly convex, beneath slightly concave, mar-
gin up to 0.3 mm broad, ridges very dark brown, with
light brown wing on outside of darker centre; coma up
to 22 mm long. Flowering time: August-May, peaking
October-January. Fruiting time: September-May.
Bothalia37,l (2007)
51
Distribution and ecology: endemic in South Africa
(North-West, Gauteng, KwaZulu-Natal and Free State)
(Figure 24). Foimd at 1 115-1 280 m in savanna of
Mixed Bushveld (Acocks 1988) on deep black turf soils
mainly associated with or in the vicinity of drainage
lines, in fully exposed situations as well as in light shade
(mainly of species of Acacia sensu lato). Depth of the
tuber below the soil surface varies greatly. The neck of
the deepest seated tuber was found at 20 cm below the
soil surface.
Material examined
NORTH-WEST. — 2527 (Rustenburg): west of settlement of Sonop,
(-DA), 25°39'20"S, 27°38T2"E, 31 January 2006, Bester 6451 (PRE);
± 10 km east of Brits, (-DB), 25°37'47"S, 27°55'5"E, 2 February 2006,
Bester 6470 (PRE).
GAUTENG. — 2528 (Pretoria): Annlin/Sinoville, drainage line running
E-W just south of Wonderboom Airport, (-CA), 25°40'0" S, 28°12'38" E,
24 October 2004, Bester 5255 (PRE, with stem tuber and follicle).
FREE STATE. — 2627 (Potchefstroom): Sasolburg Dist., Uitkomst
Farm 413, 5-6 km directly west of Sasolburg on west-facing hill on banks
of Rietspruit, (-DD), 26°48'20" S, 27°44'47" E, 10 October 1996, Kroon
12084 (PRE, with flowers and follicles).
KWAZULU-NATAL. — 2730 (Vryheid): Golugola Plain, Bloedrivier
Station, near station, (-DC), 17 September 1945, Acocks 11774 (PRE,
with flowers).
A Specimen of Acocks {Acocks 11774) named as Schi-
zoglossum sp. and another identified as S. eustegioides
{Kroon 12084) belong to the current taxon and was re-
named accordingly. Although initially thought to be
FIGURE 24. — Known distribution of Stenostelma umbelluliferum in
South Africa.
restricted in distribution, an extensive search for more
populations, sponsored by the Gauteng Department of
Agriculture, Conservation eind Environment (GDACE),
has lead to the discovery of additional populations east
and west of Brits. A herbarium sheet originating from
the Free State was also foimd at PRE. Given the pres-
ent known distribution of this species and the fact that
the Magaliesberg populations fall mainly within a devel-
oping, high-density residential area, its conservation
status has, after some debate, been classified as Near
Threatened (J.E. Victor & M. Phab 2006 pers. comm.).
ACKNOWLEDGEMENTS
We are grateful to Dr O.A. Leistner who kindly trans-
lated the original Latin and German descriptions. The
South Afncan National Biodiversity Institute, University
of KwaZulu-Natal and Gauteng Department of Agriculture
Conservation and Environment are thanked for making
resources available for this study. Dr H.F. Glen is thanked
for nomenclatural advice.
REFERENCES
ACOCKS, J.P.H. 1988. Veld types of South Afiica, edn 3. Memoirs of
the Botanical Survey of South Africa No. 57. Botanical Research
Institute, Pretoria.
BROWN, N.E. 1907. Schizoglossum umbelluliferum. In W.T. Thiselton-
Dyer, Flora capensis 4,1: 622, 623. Reeve, London.
GUNN, M. & CODD, L.E.W. 1981. Botanical exploration of southern
Africa. Balkema, Cape Town.
KUPICHA, F.K. 1984. Studies on African Asclepiadaceae. /few S«//eri«
38: 599-672.
NICHOLAS, A. 1992. The asclepiadaceous works of Rudolf F.
Schlechter (1872-1925). Willdenowia 22: 215-264.
NICHOLAS, A. 1999. A taxonomic reassessment of the subtribe
Asclepiadinae (Asclepiadaceae) in southern Africa. Ph.D. the-
sis, Department of Botany, Faculty of Science, University of
Durban- Westville, Durban.
SCHLECHTER, R. 1895. Beitrage zur Kenntnis Sudafrikanischer
Asclepiadaceen. In A. Engler, Botanische Jahrbiicher 20,
Beiblatt51: 1-56.
VICTOR, J.E., NICHOLAS, A., BRUYNS, P.V, VENTER, H.T.J.
& GLEN, H.F. 2003. Stenostelma. In G. Germishuizen &
N.L. Meyer, Plants of southern Africa: an annotated checklist.
Strelitzia 14: 172, 173. National Botanical Institute, Pretoria.
S.P. BESTER* and A. NICHOLAS**
* National Herbarium, South African National Biodiversity Institute,
Private Bag XlOl, 0001 Pretoria. E-mail: bester@sanbi.org
** School of Biology and Conservation Sciences, University of Kwa-
Zulu-Natal, Private Bag X54001, 4000 Durban.
E-mail: nicholasa@ukzn.ac.za
MS. received: 2006-06-15.
LEGUMINOSAE
VIGNA VERDCOURTII (PAPILIONOIDEAE), A NEW SPECIES FROM EASTERN AFRICA
The genus Vigna Savi (Papilionoideae) is an impor-
tant group which includes several domesticated species,
five in Asia and two in Africa, the African ones being the
cowpea, Vigna unguiculata (L.) Walp., and the Bambara
groundnut, Vigna subterranea (L.) Verde. Major advances
in the knowledge of the genus are the works of Verdcourt
(1970) and Marechal et al. (1978).
Since 1978, various phylogenetic works, especially
Thulin et al. (2004), indicate a much smaller genus
Vigna, reduced to 50-60 species. Apart from the Asian
species of subgenus Ceratotropis, and despite several
changes made in the nomenclature (Pasquet 2001), most
of the African taxa had already been described by 1978.
Six new species were described after 1978 (Mithen &
52
Bothalia37,l (2007)
Kibblewhite 1989; Pasquet & Marechal 1989; Pienaar
1991, 1993; Du Puy & Labat 2002); however, some of
them are controversial (including Vigna benuensis Pas-
quet & Marechal) and will probably not survive the phy-
logenetic test of DNA sequence analysis.
During the preparation of the account of Figna for
Flora zambesiaca (Pasquet 2001), a few specimens were
difficult to identify. These specimens were close to V.
oblongifolia A. Rich, and pubescent forms of V. luteola
(Jacq.) Benth. (= V. fischeri Harms), but they were char-
FIGURE 25, — Vigna verclcoiirlii. A, flowering and fruiting branches; B, seedling from accession X3080, National Botanic Garden of Belgium,
Meise; C, seed. A, Paget-Wilkes 926 (EA); B, C, Robson & Angus 407 (K). Scale bars: A, B, 10 mm; C, I mm. Artist: Nicholas Muema, East
African Herbarium, Nairobi, Kenya.
Bothalia37,l (2007)
53
acterized by standard dimensions longer than wide, an
unusual character in Vigna, and only encountered in two
unrelated species, i.e. V. schimperi Baker from subgenus
Haydonia (R. Wilczek) Verde, and V. owahuensis Vogel
from subgenus Vigna in the Hawaiian archipelago.
Later, from one matK sequence (Feleke 2007), it
appeared that these specimens clearly belonged to a
fairly rare new species within subgenus Vigna, known at
present from only thirteen specimens.
Vigna verdcourtii Pasquet, sp. nov., (Sect. Vigna-
Papilionoideae), floribus flaveis 12.5-18.0 mm longis,
V schimperi Baker similis sed exino grani pollinis reti-
culato atque ovario ovula 7-10 gerenti differt; fonnis
pubescentibus V liiteolae (Jacq.) Benth. similis sed
vexillo longiore quam latiore differt.
Vigna fischeri sensu Thulin in Opera Botanica 68: 172 (1989), non
Harms.
TYPE. — Ethiopia, Kaffa, Amero, 2 250 m, 1 Decem-
ber 1960, Mooney 8745 (K, holo.; ETH, FT, S, iso.).
Twining herb. Rootstock unknown. Stems covered
with spreading ferruginous hairs. Stipules triangular-
lanceolate, 5-6 X 1.5 mm, slightly bilobed at base, multi-
nerved; lobes rounded, 1.5 mm long. Leax’es trifoliolate;
terminal leaflets lanceolate, rarely ovate, 35-77 x 13-30
mm, acute and mucronulate at apex, rounded-subcordate
at base, sometimes slightly hastate, sparsely to densely
soft-pubescent on both surfaces; petiole 1 5-45 mm long;
rhachis 5-15 mm long. Inflorescence axillary, 2-16-
flowered; peduncles 10-160 x 0.8-2 mm, not winged;
rhachis 8-17 mm long, 1-8-noded; intemode 1-3 mm
long. Flowers 12.5-18 x 10-14 mm, yellow with pur-
ple marks; pedicel 1^ mm long, very slightly expand-
ing during fruiting. Bracteole oblong or lanceolate, 3-6
X 0.5-1. 5 mm, 1-nerved. Calyx pubescent; tube 3 mm
long; lobes 1.5 mm long, the lower twice as long, the
upper pair joined to fonn acute or bifid lip. Standard
longer than wide, obovate, usually 14-16 x 10-13 mm,
with two small oblique appendages. Keel slightly twisted
toward right or not twisted (seen from rhachis top), whit-
ish with a short purple beak. Alternate anthers without
a pair of glands at base. Pollen exine reticulate. Ovary
7-10-ovuled. Pot/ black, slightly compressed, curved or
almost straight, 45^8 x 4. 0-4. 5 mm, slightly constricted
between seeds, covered with ferruginous and white bris-
tly hairs, with a short curved beak. Seed 3-4 x 2.5 mm,
1.5 mm thick, black; hilum 1.5 mm long, eccentric, with
a conspicuous fork-shaped eccentric aril. Figures 25, 26.
Diagnostic features and affinities: Vigna verdcour-
tii is sister to the mainly yellow-flowered group which
includes V. lanceolata Benth., V. pseudovenulosa (Mare-
chal, Mascherpa & Stainier) Pasquet & Maesen, V. sub-
terranea (L.) Verde., V filicaulis Hepper, V. multinervis
Hutch. & Dalziel, V heterophylla A. Rich., V pubigera
Baker, V oblongifolia var. oblongifolia, V luteola, V.
FIGURE 26. — Vigna verdcourtii. A, flower; B, pods; C, seeds; D, seedlings from accession X3080. National Botanic Garden of Belgium, Meise. A,
B, Luke 7032 (EA); C, Robson & Angus 407 (K). Scale bars: A,B,D, 10 mm; C, 1 mm.
54
Bothalia37,l (2007)
FIGURE 27. — Known distribution of Vigna verdcoiirtii.
marina (Bunn.) Merr., V. oblongifolia var. paj'viflora
(Baker) Verde., and V. owahiiensis (Feleke 2007).
The general plant morphology is very similar to both
that of Vigna oblongifolia var. oblongifolia (especially
the pubescenee, the number of ovules, and the pod and
seed morphology), and to that of the pubescent forms
of V. Inteola (= V. fischeri) due to the leaf shape and the
flower size (Figure 25). Previously, Mooney 8745 was
identified as V. fischeri by Thulin (1983, 1989), and
Robson & Angus 407 as V. oblongifolia by the author
(Pasquet 2001). Most of the specimens were labelled V.
fischeri, one V. luteola, and one V. oblongifolia.
The contracted inflorescence and the standard being
longer than wide (Figure 26) suggest Vigna schiniperi,
but the V. verdcoiirtii pollen exine is not smooth, the
stipule is produced below the point of insertion, and
the ovule number (7-10) is much lower than that of V.
schimperi (13-20).
Distribution and ecology: Vigna verdcoiirtii appears
to be restricted to the mountainous areas of tropical East
Africa (Figure 27) occurring at 1 500-2 300 m. This is
higher than the V. luteola altitudinal range, and similar
to that of V. schimperi. Flowever, the ecology is more
related to that of the V. luteola group: streambanks, for-
est edges, fallows and cleared land in forest areas. In
fact, this exactly matches pubescent forms of V. luteola.
The number of specimens available is small, but it
highlights a surprising feature of its distribution. The
taxon is found between latitude 8°N and 12°S, but it has
never been collected between 6°N and 6°S (Figure 27),
notably in Kenya and northern Tanzania, which have
been intensively surveyed in the past and where pubes-
cent fornis of V. luteola are encountered. This may be
due to a photoperiod sensitivity which could prevent
the plant from flowering when day length is not variable
enough. The area of occurrence of V. verdcoiirtii is fully
included within that of V. schimperi, which extends from
13°N to 12°S, with numerous collections from Kenya
and northern Tanzania.
Other material examined
TANZANIA. — Iringa Prov., Mufindi, 6500' [1 980 m], 15 October
1931. Davies 29 (EA); Udzungwa Mountain NP, Ruipa River, I 650 m,
6 October 2000, Luke. Bytebier, Butynski, Ehardt, Perkins & Kimaro
7032 (EA, K); Mufindi, ± 6200' [1 900 m], 25 July 1969, Paget-Wilkes
549 (EA); Mufindi, Lugeme, ± 6200' [1 900 m], 18 September 1971,
Paget-Wilkes 926 (EA, K); Mufindi, Nymalala, ± 6200' [1 900 m], 17
September 1971, Paget-Wilkes 937 (EA, K); Mufindi Dist., Livalonge
Tea Estate, [± 1 700 m], 27 August 1971, Perdue & Kibuwa 11269
(EA, K); Rungwe Dist., Ngozi Poroto Mtns, 2 100 m, 17 October 1956,
Richards 6537 (K); Mbeya Dist., Kikondo, 2 250 m, 20 October 1956,
Richards 6654 (K); Nyassa Hochland, Station Kyimbila, 1913, Stolz
2757(C, HBG, L, LD, S, U).
MALAWI. — Chikangawa, ± 1 800 m, 9 July 1952, Jackson 956
(BR, K); edge of rainforest path leaving Nyika, 7300' [2 225 m], 7 July
\91\,Pawek5033(YA.
ZAMBIA. — Nyika Plateau, below Rest House on path to N Rukuru
waterfall, 2 150 m, 27 October 1958, Robson & Angus 407 (BM, K,
PRE).
REFERENCES
DU PUY, D.J. & LABAT, J.N. 2002. Papilionoideae. In D.J. Du Puy,
J.N. Labat, R. Rabevohitra, J.F. Villiers, J. Bosser & J. Morat,
The leguminosae of Madagascar: 294—714. Royal Botanic
Gardens, Kew.
FELEKE, Y. 2007. Phylogenetic relationships in genus Vigna and cow-
pea revealed by DNA sequences from the chloroplast and nucle-
ar genomes. Ph.D. thesis, Kenyatta University, Nairobi.
MARECHAL, R., MASCHERPA, J.M. & STAINIER, F. 1978. Etude
taxonomique d’un groupe complexe d’especes des genres
Phaseolus et Vigna (Papilionaceae) sur la base de donnees mor-
phologiques et polliniques, traitees par I’analyse informatique.
Boissierali: 1-273.
MITHEN, R. & KIBBLEWHITE, H. 1989. A new species of Vigna
(Leguminosae-Papilionoideae) from southern Africa. Kew
Bulletin 44: 175-177.
PASQUET, R. & MARECHAL, R. 1989. La Vigna beniiensis, une
nouvelle espece de la section Vigna du genre Vigna (Fabaceae).
Canadian Journal of Botany 67: 949-953.
PASQUET, R. 2001 . Vigna Savi. In B. MacKinder, R. Pasquet, R. Polhill
& B. Verdcourt, Flora zambesiaca 3, 5, Phaseoleae: 121-156.
Royal Botanic Gardens, Kew.
PIENAAR, B.J. 1991. A new species of Vigna Savi (Fabaceae) from
southern Africa. South African Journal of Botany 57: 314—3 1 8.
PIENAAR, B.J. 1993. Vigna kokii, a new species from southern Africa.
Bothalialh: 68-70.
THULIN, M. 1983. Leguminosae of Ethiopia. Opera Botanica 68:
1-223.
THULIN, M. 1989. Fabaceae (Leguminosae). In 1. Hedberg & S.
Edwards, Flora of Ethiopia 3: 49-25 1 . Addis Ababa, Asmara &
Uppsala.
THULIN, M., LAVIN, M., PASQUET, R. &'DELGADO-SALINAS, A.
2004. Phylogeny and biogeography of Wajira (Leguminosae): a
monophyletic segregate of Vigna centred in the Hom of Africa
region. Systematic Botany 29: 903-920.
VERDCOURT, B. 1970. Studies in the Leguminosae-Papilionoideae
for the Flora of tropical East Africa: IV. Kew Bulletin 24: 507-
569.
R.S. PASQUET*
* ICIPE, P.O. Box 30772, Nairobi, Kenya. IRD, Departement
Ressources Vivantes, 209-213 Rue La Fayette, 75480 Paris Cedex 10,
France.
MS. received: 2006-08-25.
Bothalia37,l (2007)
55
ASTERACEAE
A KEY TO DICOMA TAXA (DICOMEAE) IN SOUTHERN AFRICA
Recent molecular phylogenetic studies of the fam-
ily Asteraceae suggest that the genus Dicoma Cass,
belongs to the tribe Dicomeae, closely related to the tribe
Cardueae (Panero & Funk 2002). Harvey (1865) revised
the southern African species of Dicoma. In 1923, Wilson
revised the whole genus including seventeen southern
African species. Since the publication of Wilson’s mono-
graph, many new species have been described, and some
taxonomic changes have been made to the genus. Ortiz
(2000) made a phylogenetic analysis of Dicoma and
related genera. Following this analysis he reinstated the
genus Macledium, which was first described by Cassini
(1825). The species that previously belonged to sections
Macledium (Cass.) DC. and Pterocoma DC. of Dicoma
(as they were delimited by De Candolle in 1838 and
Harvey in 1 865) were transferred to the genus Macledium
(Ortiz 2001). Netnou (2001) revised the genus Dicoma in
southern Africa and during the course of this study, vari-
ous herbaria were visited. Due to large numbers of incor-
rectly identified and unidentified species of this genus in
some herbaria, it became apparent that a key to the spe-
cies of Dicoma in southern Africa is much needed.
Key to southern African taxa of Dicoma Cass.
la Heads radiate; rays neuter; disc floret pappus isomorphic of barbellate bristles; rigid shrublets:
2a Leaves many, bright green, narrowly obovate or oblong-spathulate; midrib inconspicuous; heads cylindrical in shape; involucral bracts
often reddish, gland-dotted D. fi-i'ticosa Compton
2b Leaves few. dull grey-green, upper ones minute, basal ones often absent from herbarium specimens, if present broader than upper
ones; midrib conspicuous at least on abaxial surface; heads obconic or campanulate; involucral bracts not gland-dotted:
3a Heads distinctly obconic, cuneate at base; involucral bracts in (5)6(7) rows; pappus 9-11 mm long; leaves linear-ellip-
tic D. obconica S. Ortiz & Pulgar
3b Heads campanulate to subglobose, rounded at base; involucral bracts in 7 or 8(9) rows; pappus < 9 mm long; leaves oblanceolate to
spathulate D. picta (Thunb.) Druce
lb Heads discoid or disciform; disc floret pappus isomorphic or dimorphic; shrubs, prostrate or erect herbs:
4a Heads discoid; disc floret pappus dimorphic or isomorphic; plants erect or prostrate,:
5a Pappus dimorphic, outer rows of barbellate bristles, innermost row of membranous scales becoming barbellate toward apex:
6a Erect annual herb, leaves discolorous. linear, narrowly elliptic or spathulate. margins serrate, base attenuate; involucral bracts pur-
plish. spiny; inner pappus of membranous scales with barbellate tips D. tomentosa Cass.
6b Prostrate perennial herb; leaves concolorous. elliptic or obovate. margins entire, distinctly petiolate; involucral bracts straw-
coloured, not spiny; inner pappus with lower half scale-like, apex barbellate D. cimeneensis Wild
5b Pappus isomorphic, of barbellate bristles, inner row not of membranous scales:
7a Shrubs; leaves spathulate or broadly elliptic to suborbicular, both surfaces greyish white-felted:
8a Leaves broadly elliptic to suborbicular, (10-) 15-25 x (4— )7-10 mm; heads with 2(-5) subtending leaves; heads 13-16 mm long ....
D. kuntmanii S. Ortiz & Netnou
8b Leaves spathulate, 8-12(-15) x 4—6(-8) mm; heads covered with at least 20 subtending leaves; heads less than 12 mm
long D. nachtigallii O.Hoffm.
7b Herbs or shrubs; leaves linear, elliptic, oblanceolate or ovate, upper surface tomentose, glabrous or subglabrous, lower surface
greyish white-felted:
9a Leaves linear, several times longer than wide:
10a Shrub; leaves (5-)20-25 x 2—4 mm. upper surface dull greyish green, tomentose; involucral bracts erect with aristate apices,
often sericeous on either side of midvein D. dinteri S. Moore
10b Erect or prostrate herbs; leaves often longer than 30 mm:
11a Plants erect:
12a Heads solitary; leaves flexuous, upper surface glabrous to glabrescent, often with yellowish glands; heads broadly obconic,
30-32 mm wide; involucral bracts ovate-deltate, glabrous D. swazilandica S. Ortiz, Rodr.Oubina & Pulgar
12b Heads solitary or 3-5 corymbosely arranged at end of branches; leaves not flexuous, upper surface greenish black,
gland-pitted; heads subcylindric to campanulate, relatively small, less than 30 mm wide; involucral bracts lanceolate,
glabrescent D. anomala Sond. subsp. gerrardii (Harv. ex F.C. Wilson) S. Ortiz & Rodr.Oubina
11b Plants prostrate:
1 3a Heads broadly campanulate; involucral bracts finely araneose-lanate or glabrescent, margins scarious, tips often recurved;
leaves not fascicled, upper surface gland-pitted, greenish black D. anomala Sond, subsp. anomala (narrow-leaved form)
13b Heads subcylindric; involucral bracts glabrous, ± entirely scarious, often not recurved; leaves often fascicled, upper sur-
face dull greyish green with black glands D. prostrata Schweick.
9b Leaves elliptic, lanceolate, oblanceolate, ovate or obovate:
14a Prostrate herbs; leaves lanceolate to elliptic; heads sessile up to 40 mm wide; pappus of fine barbellate bristles, 10 mm or
more long D. anomala Sond. subsp. anomala (broad-leaved form)
14b Erect herbs or subshrubs; leaves oblanceolate, obQvate or narrowly ovate; heads pedunculate, up to 25 mm wide; pappus cili-
olate, if barbellate up to 7 mm long:
15a Bushy herbs or subshrubs; leaves oblanceolate to obovate, often fascicled on axils, lower ones usually larger, upper surface
bright green, glabrous at maturity, conspicuously gland-dotted; heads up to 25 mm wide; involucral bracts glabrous, outer
often recurved, with sharp pointed apices; pappus ciliolate up to 9 mm long D. galpinii F.C. Wilson
15b Shrubs; leaves lanceolate or narrowly ovate, not fascicled, upper surface of leaves gland-dotted, visible through tomen-
tose indumenrnm; heads up to 20 mm wide; outer involucral bracts pilose, often recurved, apices acuminate-pun-
gent; pappus barbellate, up to 7 mm long D. montana Schweick.
4b Heads disciform; disc floret pappus dimorphic; prostrate herbs:
16a Disc florets distinctly shorter than pappus; base of inner row of pappus broad, membranous with undulate margins, upper part ciliolate:
17a Leaves concolorous, elliptic, up to 20 mm wide, distinctly petiolate, margins serrulate; disc floret pappus 7-12 mm
long D. schinzii O.Hoffm.
17b Leaves discolorous, oblong-lanceolate, up to 15 mm wide, subsessile, margins subentire; disc floret pappus 21-24(-30) mm
long D. macrocephala DC.
16b Disc florets ± equal to pappus; base of inner pappus of disc florets with basal part scale-like, upper part barbellate:
56
Bothalia37,l (2007)
1 8a Leaves discolorous, ovate to orbicular; inner pappus of disc florets with basal part scale-like for more than length, tip barbel-
late D. arenaria Bremek.
1 8b Leaves concolorous, narrowly oblong-lanceolate; inner pappus of disc florets with basal part scale-like for length, tip plu-
mose D. capensis Less.
ACKNOWLEDGEMENTS
This key is part of an M.Sc. thesis submitted to the
Rand Afrikaans University, now the University of
Johannesburg. We would like to thank the curators of the
following herbaria: BOL, GRA, J, NBG, NH, PRE, PRU
and UNEST for making their collections available for
this study. Financial assistance from the South African
National Biodiversity Institute, National Research
Foundation and University of Johannesburg is acknowl-
edged. Prof. A.E. van Wyk and Dr H.F. Glen are thanked
for their valuable comments on the thesis. We would also
like to thank the staff of the National Herbarium, espe-
cially Drs Elizabeth Retief, Elsie Steyn, Sarie Perold,
and Marinda Koekemoer, and Priscilla Burgoyne, Anne-
Lise Fourie, Paul Herman, Estelle Potgieter and Sandra
Turck for their assistance and support.
REFERENCES
CASSINI, A.H. DE. 1825. Macledimn. In G. Cuvier, Dictiotmaire des
sciences naturelles 34: 39. Paris.
DE CANDOLLE, A.P. 1 838. Prodromus systematis naturalis regni veg-
elabilis 7. Treuttel & Wurtz, Paris.
HARVEY, W.H. 1865. Compositae. In W.H. Harvey & O.W. Sender,
Flora capensis 3: 44—530. Hodges, Smith, Dublin.
NETNOU, N.C. 2001. A revision of the genera Dicoma Cass, and
Macledium Cass. (Asieraceae, Mutisieae) in southern Africa.
M.Sc. thesis, Rand Afrikaans University, Johannesburg.
ORTIz, S. 2000. A phylogenetic analysis of Dicoma Cass, and related
genera (Asteraceae: Cichorioideae: Mutisieae) based on mor-
phological and anatomic characters. Annals of the Missouri
Botanical Garden 87: 459-481.
ORTiZ, S. 2001. The reinstatement of the genus Macledium Cass.
(Asteraceae, Mutisieae): morphological and phylogenetic argu-
ments. Taxon 50: 733-744.
PANERO, J.L. & FUNK, V.A. 2002. Toward a phylogenetic subfamilial
classification for the Compositae (Asteraceae). Proceedings of
the Biological Society of Washington 115: 909-922.
WILSON, F.C. 1923. Revision of the genus Dicoma. Bulletin of
Miscellaneous Information 1923: 377-388. Royal Botanic
Gardens, Kew.
N.C. NETNOU* * and B-E. VAN WYK**
* Department of Biology, University of Limpopo (Medunsa Campus),
P.O, Box 139, 0204 MEDUNSA, South Africa.
** Department of Botany, University of Johannesburg, P.O. Box 524,
2006 Auckland Park, Johannesburg.
MS. received: 2004-04-30.
Bothalia 37,1: 57-73 (2007)
A reconnaissance survey of the woody flora and vegetation of the
Catapii logging concession, Cheringoma District, Mozambique
M. COATES PALGRAVE', A.E. VAN WYK% M. JORDAAN^ J.A. WHITE" and P. SWEET'
Keywords: Catapii, checklist, Cheringoma, ecology, flora, Mozambique, Sena, vegetation, vernacular names
ABSTRACT
A checklist of the trees, shrubs and Hanes of Catapii, Cheringoma District, Mozambique, is presented. Floristically the
study area falls within the Swahilian/Maputaland Regional Transition Zone. In total, 238 woody species and infraspecific
taxa have been recorded, representing 59 families and 167 genera. Most species (64%) occur both to the north and south of
the study area, 26% have their core distribution in the Swahilian Regional Centre of Endemism, 4% have a more southerly
distribution, 14 are endemic to the Swahilian/Maputaland Regional Transition Zone and two near-endemic, extending into the
Zambezian Regional Centre of Endemism only along the Zambezi River Valley as far west as Kariba. The checklist includes
the Sena names for 191 species, 77 of which are recorded for the first time. Comparisons of the Catapii checklist are made
with other checklists.
INTRODUCTION
According to a preliminary checklist of vascular
plants, the flora of Mozambique comprises 3 932 indig-
enous plant taxa and of these 177 are endemic (Da Silva
et al. 2004). The primary purpose of the study was to
establish what trees, shrubs and woody lianes occur
on the Catapii logging concession area. Floristically,
White (1983) recognized the zone occupying the East
African coastal belt, ± 50-200 km wide, from south-
ern Somalia in the north to the mouth of the Limpopo
River in the south as the Zanzibar-Inhambane Regional
Mosaic. Subsequently this phytochorion was split into
two smaller floristic regions by Clarke (1998), namely:
1, the Swahilian Regional Centre of Endemism along
the Kenyan, Tanzanian and northern Mozambique coasts
and marginally extending into southern Somalia; and
2, the Swahilian/Maputaland Regional Transition Zone
extending along the Mozambique coast and into southern
Malawi and eastern Zimbabwe between the Swahilian
Regional Centre of Endemism and White’s (1983)
Maputaland-Pondoland Regional Mosaic. The study
area falls within the Swahilian/Maputaland Regional
Transition Zone.
Wild & Barbosa (1967) broadly classified the vegeta-
tion of the concession area as Dry Deciduous Lowland
Forest (vegetation type 6), Mosaic of Low Altitude Dry
(Mixed) Forest and Miombo (vegetation type 10) and
Discontinuous Dry Savanna Woodland-Tree Savanna
and ‘Tandos’ Grassland (Gorongosa Lowland) (vegeta-
tion type 45) along the lower altitude river valleys. The
Wild & Barbosa (1967) map also indicates the presence
of Open Deciduous Tree Savanna (Lowland) v^ixh. Acacia
nigrescens dominant (vegetation type 52) but this has not
been seen either in or near the study area. White (1983)
' 9 Blue Kerry, 30 Steppes Road, RO. Chisipite, Harare, Zimbabwe, e-
mail: megcp@zol.co.zw'
^ H.G.W. J. Schweickerdt Herbarium, Department of Botany, University
of Pretoria, 0002 Pretoria, e-mail: braam.vanwyk(^up.ac.za
^ South African National Biodiversity Institute, Private Bag XIOI,
0001 Pretoria, e-mail: jordaan(2jsanbi.org
TCT-Dalmann Furniture, Estrada Nacional No. 6, Beira, Mozambi-
que. e-mail:. tctdalmann(^tdm.co.mz
^Companhia de Sena, Caixa Postal 1903, Beira, Mozambique, e-mail:
PatSweet(S;senasugar.com
MS. received: 2005-09-13.
mapped the region as North Zambezian Undifferentiated
Woodland and Wooded Grassland (mapping unit 29c),
which seem to better describe the floristically rich veg-
etation of the study area. In the east, Catapii borders
White’s Zanzibar-Inhambane East African Coastal
Mosaic (mapping unit 16a). Outliers of one of the for-
est types, Zanzibar-Inhambane Undifferentiated Forest,
occur on Catapii.
Although some plant exploration was undertaken in
the Sofala Province of Mozambique between 1940 and
1980, notably by Muller & Pope (1973 pers. comm.) and
Tinley (1977), that part of Mozambique was still remote,
and collecting of herbarium material was largely con-
fined to areas adjacent to roads. In those days, Catapii
was well away from the main road which went between
Inhamitanga and Marromeu. During the civil war in
Mozambique (1975-1994), botanical collecting came to
a halt and has only taken plaee sporadically since then
(Da Silva et al. 2004). Hitherto, most of the botani-
cal inventories undertaken in Mozambique have been
conducted mainly in the south of the country; the cen-
tral and northern parts have been less well documented
(Izidine & Bandeira 2002). The paucity of biodiversity
data for central and northern Mozambique has also been
highlighted by Schipper & Burgess (2004).
The checklist focuses on the woody plants and lianes
and is ongoing. To describe the intricate mosaic of veg-
etation types in the study area, we have adopted a narra-
tive style whieh we hope will be useful for workers on
the ground and as a basis for possible quantitative sur-
veys in future. The common Sena (= Cisena, Chisena)
names — the principal vernacular of the region — are pre-
sented, many for the first time. We also comment on the
impacts of logging on the vegetation and the need for
measures to ensure sustainability and the conservation of
an ecosystem rich in plant diversity.
STUDY AREA
Catapu, the logging concession of TCT-Dalmann
Furniture, Lda., is located 30-40 km south of the
Zambezi River and on the northwestern side of the
Inhamitanga Village in the district of Cheringoma, Sofala
58
Bothalia 37,1 (2007)
Province, central Mozambique (Figure 1 ) at ± S 1 8°00'05"
and E35°08'13", an area covering 24 821 ha. The GPS
reading at the sawmill is S18°02'41"; E35°12'24". The
main road, the EN-1, traverses the concession where it ±
follows the watershed between the Zangue and Tissadze
Rivers. Field work started in 2000 and results up to July
2006 are reported. Hitherto the survey has been concen-
trated within the firebreak around the sawmill area, west
of the EN-1 along the roads known as Via Pungue and
Via Santove and around the pans.
In the west, Catapu is bordered by the Zangue River
and flood plain, up which the Zambezi River floods back
during the wet season. The eastern boundary extends
from the EN- 1 road/railway junction at Gangala in the
north along the obsolete Dondo-Caia railway line to the
Inhamitanga Village. The northern boundary is in the
fonn of a ‘V’ with Via Zangue 2 on the western arm and
the EN-1 on the eastern ann. The southern boundary is
the Inhamitanga road to the EN-1 at Chapa and then the
EN-1 to the Zangue River bridge. The Tissadze River
flows from south to north ± bisecting the eastern half of
the concession and the Chirimadzi River, which is more
a series of pans than a river with the woody vegetation
margin more clearly defined than the waterway, goes
from south to north in the northern third.
The altitude at the Catapu turn-off from the EN-1 is
200 m a.s.l. but drops from ± 100 to 50 m along the river
valleys. Soils are sand with outbreaks of sandstone and
calcareous conglomerates and black cotton or turf soils
around the vleis and in the river floodplains. The sands
are underlain by sublittoral sands which accumulate
water, enabling them to support very tall trees. A bore-
hole was drilled and a sample collected every 3 m to a
depth of 51 m. Every sample was sand except for the
first 3 m in which there was some gravel. Mean annual
rainfall ranges from 700-1 400 mm, although for the
last four years it has barely exceeded 500 mm during
the rainy season (November to March). There are many
pans in the forest/thicket/woodland mosaic, grass-cov-
ered depressions of half a hectare or more which fill with
water during the rains. As a result of below-average rain-
fall, none of these pans has held any water for the last
four or five years. During the other seven months (April
to October) an average of 1 26 mm has been recorded.
Vegetation of Catapu comprises a mosaic of mainly
dry lowland forest, dry deciduous thicket and woodland.
In the Chirimadzi Valley in the northern area of the con-
cession, the vegetation is sparsely wooded grassland.
There is an area of miombo woodland on the southeast
boundary of the concession and open undifferentiated
woodland in the southwestern section.
METHODS AND TERMINOLOGY
Recognizing the rich -plant diversity at Catapu, a sur-
vey of the woody species was started. Specimens have
Bothalia 37,1 (2007)
59
been collected and voucher specimens prepared which
are housed in the recently established field herbarium
to be known as the Cheringoma Herbarium (proposed
acronym CHER). Any species not identified in the field
were taken for identification to the National Herbaria in
Harare (SRGH) and Pretoria (PRE). Regional Eloras,
e.g. Flore de Mozambique (1969-), Flora zambesiaca
(I960-) and Flora of tropical East Africa (1952-) have
also been used to assist in identification. The scientific
names of the plants are based mainly on Coates Palgrave
(2002) and Klopper et al. (2006).
Sena names on the list, (Appendix 1, column 4), were
checked against species in the field to confinn identifi-
cation. Aware that the use of plant vernacular names is
essentially a spoken language, they have been spelt pho-
netically. The names were checked against De Koning
(1993) where a very broad view was taken of the spell-
ing and the regions in which the names were recorded.
Plants recorded at Catapii were assessed phytogeo-
graphically. The checklist of the study area was com-
pared with the preliminary checklist of vascular plants
of Mozambique (Da Silva et al. 2004) and also with
checklists compiled during two recent vegetation and
plant surveys in nearby areas, one of the Cabora Bassa
shoreline (Timberlake 2002) and the other the Zambezi
Delta wetland plant survey (Muller et al. 2000). It was
also compared with the checklists of two South African
forests, Richards Bay forest and the Maputaland for-
est complex (Geldenhuys 1992) as well as with a mid-
Zambezi Valley dry forest biodiversity survey (Hoare et
al. 2002). The Red Data List, Mozambique (Izidine &
Bandeira 2002) was checked for species occurring in the
study area.
Very simplified, the following definitions are used for
the different broad vegetation types:
Forest has a canopy whose crowns interlock at 10 m
or more in height (Hoare et al. 2002). A forest is rich in
species, but due to the availability of soil moisture, it
changes rapidly in composition and physiognomy over
quite short distances arid is difficult to classify (White
1983). Dry deciduous forest is characteristically found
on deep sands which absorb all the incident rainfall or
receive lateral seepage water and remain moist through-
out the greater part of the dry season (White 1983).
Understorey species remain evergreen throughout the
year in dry semi-deciduous forest and are deciduous for
more than one month in dry deciduous forest (Chapman
& White 1970).
Thicket has a canopy between 3-7 m tall formed
by the interlocking branches of small trees and multi-
stemmed shrubs. The presence of emergent trees over
10 m tall protruding above the canopy or shrub layer is
often a feature of thiqkets (Chapman & White 1970).
Woodland comprises open stands of trees at least 5 m
tall, crowns covering at least 20% of the surface, some-
times in lateral contact but crowns not interlocking; grass
cover is usually present (Chapman & White 1970).
Zambezian undifferentiated woodland is defined by
the absence of miombo and mopane dominants rather
than by its own floristic composition. It is composed of
many more tree species than either miombo or mopane
woodland. Although the dominants of miombo wood-
land are normally absent from undifferentiated woodland
some of their associated species are frequently present
(White 1983).
Miombo is a colloquial term used to describe those
central, southern and eastern African woodlands domi-
nated by members of Brachystegia, Julbernardia and/or
Isoberlinia, three closely related genera from the legume
family, Fabaceae, subfamily Caesalpinioideae (Campbell
et al. 1996). It generally occurs on geologically old,
nutrient-poor soils in the uni-modular rainfall zone.
The shrub layer is variable in density and composition.
Ground cover varies from a dense coarse grass growth
to a sparse cover of herbs and grasses (Campbell et al.
1996).
RESULTS
Floristic analysis
A total of 238 woody species and infraspecific taxa
(referred to collectively as species) recorded during
the survey are listed in Appendix 1 . These represent 59
families and 167 genera. Fabaceae is the largest family,
with eight genera and eight species in Caesalpinioideae,
seven genera and 20 species in Mimosoideae and 11
genera and 17 species in Papilionoideae, making a total
of 26 genera and 45 species, followed by Rubiaceae with
12 genera and 14 species. None of the other families has
more than 10 species.
We recorded 191 Sena names (Appendix 1, column 4)
for species in the study area of, which 77 names are not
included in De Koning (1993). The names published in
De Koning (1993) are marked with an asterisk, those not
marked are names not previously published.
Appendix 1, column 5, presents the phytogeographi-
cal distribution of the species in the study area. With
the symbol ‘T’, there are 152 species (64.5%) common
to both the East African coastal flora and the southern
African flora {FSA) region (Germishuizen & Meyer
2003). With the symbol ‘N’ denoting those from the
north there are 61 species (25.5%) extending southwards
from the East African coastal flora. With the symbol ‘S’
there are 10 (5%) species extending northwards from the
southern African flora (FSA) region. There are 14 spe-
cies (5.6%) with the symbol ‘E’, which are endemic to
the Swahilian/Maputaland Regional Transition Zone.
They are: Acacia torrei, Catunaregam swy’nnertonii.
Cola mossambicensis, Combretum kirkii, Cordia stuhl-
mannii, Eiythrina livingstoniana, Glyphaea tomentosa.
Hibiscus mossambicensis, Millettia mossambicensis,
Monodora stenopetala, Oclrna angustata, Pavetta klotzs-
chiana, Tricalysia jasminiflora and Vangueria esculenta.
Ziziphus mauritiana, included in the checklist is an
alien, marked with a (+). Tamarindus indica is treated as
indigenous (Coates Palgrave 2002). There are 40 species
in common with the Maputaland-Pondoland Regional
Mosaic as shown in Appendix 1 , column 6.
Comparison with Da Silva et al. (2004) Appendix 1,
column 7, shows that of the 240 species on the Catapii
60
checklist, 38 species can be added as occurring in
Sofala Province and an additional 22 as occurring both
in Sofala Province and in Mozambique. New records
of particular interest are Dovyalis xanthocarpa, a new
record for Mozambique; new for Sofala and south of the
Zambezi River include Cordia torrei, Grewia forbesii,
Zanthoxyhim holtziamm, also interesting is Combretum
kirkii, with a very limited distribution along the Zambezi
Valley; Elaeodendron transvaalense has not previ-
ously been recorded as far north as this (R. Archer pers.
comm.).
The Zambezi Delta survey (Muller et al. 2000)
included six days spent at Coutada 11, GPS reading
S18°33T1", E36‘’06'02", a hunting concession not very
far from Catapu. It was anticipated that many of the
woody species at Catapu would appear on that list of
plants recorded for the forest and woodland columns in
the survey. But Appendix 1, column 8, shows 86 spe-
cies in common. Lake Cabora Bassa is situated on the
Zambezi River in Tete Province upstream from the study
area. The result of the comparison with that shoreline
survey (Appendix 1, column 9) shows that 152 species
on the Catapu checklist were recorded. Of the Catapu
checklist, 59 species appeared on both the previously
mentioned lists and 60 species did not occur on either
list. Although 162 species (68%) of the plants on the
Catapu checklist occur in the FSA region, only 40 spe-
cies ( 1 7%) occur in the Maputaland/Pondoland Regional
Mosaic (Appendix 1, column 6).
In the mid-Zambezi Valley dry forest biodiversity
survey (Hoare et al. 2002), no checklist was presented,
but a list of 36 species considered to be indicator spe-
cies of dry forest was given. Of those, 20 species occur
in the study area, Kirkia acuminata and Schinziophyton
rautanenii are infrequent and found mainly in woodland
and the shrubs Citropsis daweana and Zanthoxyhim lep-
heurii are only occasional. Cleistochlamys kirkii and
Monodora junodii are classified as shrubs, the fonner
described as being tall trees in the description of the veg-
etation to follow and the latter is distinctly a tree of 3 to
4 m tall. Xylotheca tettensis is plentiful in the study area.
Xylia torreana is considered an indicator of dry forest
and thicket and is very much a constituent of the forest
vegetation in the study area.
Conservation status
Eleven species on the Catapu checklist have been
included in the Red Data List for Mozambique (Izidine
& Bandeira 2002). Only three are considered vulnerable:
Acacia torrei VU D2, Cola mossambicensis VU Ala and
Sterculia appendiculata VU A lad. Izidine & Bandeira
(2002) comment that regeneration in Sterculia appen-
diculata is difficult. In the study area, however, this spe-
cies coppices very readily. On the edge of Via Pungue
where the roots were disturbed by a bulldozer when the
road was being constructed, there has been a prolifera-
tion of saplings growing from root shoots, some now ±
4 m tall. Afzelia quanzensis is given a Threatened status
in Mozambique by Bandeira et al. (1994), Lower-Risk
Near Threatened (Izidine & Bandeira 2002), and as
Vulnerable (Da Silva et al. 2004), citing that large quan-
Bothalia37,l (2007)
tities are being cut for timber, charcoal and fuel wood
production.
Two species. Acacia adenocalyx and Amblygono-
carpus andongensis are incorrectly listed as Mozam-
bique endemics and rated VU D2 in Da Silva et al.
(2004) but not included in Izidine & Bandeira (2002).
Acacia adenocalyx is not an endemic to Mozambique,
as it also occurs in Kenya and Tanzania (Ross 1979)
and in the study area it is widespread and invasive.
Amblygonocarpus andongensis has a wide distribution,
occurring as far afield as Namibia, Botswana, Zambia
and in the savanna regions of tropical Africa (Brenan
1970). It is also fairly widespread in Mozambique.
Vegetation types
The study area comprises a mosaic of three vegeta-
tion types: dry deciduous forest, dry deciduous thicket
and woodland. Areas of lowland deciduous forest, with
the canopy usually ± 20 m high, but varying from 12
to 25 m and not always continuous, are very patchy but
there are good examples along the road known as Via
Pungue, in the area west of the EN-1 and just south of
Mashamba Grande and an area east of the EN-1 just
north of Chapa. Nearly all the canopy species are decid-
uous but there is considerable variation from species
to species and from year to year. The dominant trees
emergent above the canopy are Adansonia digitata,
Berchemia discolor, Bivinia jalbertii, Bombax rhodog-
naphalon, Milicia excelsa, Millettia stuhlmannii, Monts
mesozygia and Sterculia appendiculata. Reaching and
fonning the canopy at 12 to 25 m are Afzelia quanzensis.
Balanites maughamii, Celtis mildbraedii, Cordyla afri-
cana, Fernandoa magnifica, Terminalia sambesiaca and
Xylia torreana. Many of the Terminalia sambesiaca have
died except for some along the EN- 1 highway. Sterculia
appendiculata occurs in dense and open forest.
Below the canopy but reaching up towards it in the
sparse understorey, are numerous Diypetes reticu-
lata. Also frequent are Millettia mossambicensis and
Stiychnos usambarensis. In addition, Hunteria zeylan-
ica, Stiychnos henningsii and S. potatorum are found.
There is a fairly continuous canopy and very sparse
undergrowth with very few to no ferns, herbs or grasses.
Some of the understorey shrubs are Chazaliella abrupta,
Citropsis daweana, Diypetes arguta. Hibiscus mossam-
bicensis, Lasciodiscus pervillei, Sclerochiton kirkii and
Zanthoxyhim leprieiirii.
Lianes in the study area include Abriis precatorius.
Acacia adenocalyx, Adenia gummifera, Artabotiys brach-
ypetalus, Cissus cucumerifolia, C. integrifolia, C. qiiad-
rangularis and C. rotundifolia, Combretum kirkii, C.
microphyllum, C. padoides, Dalbergia arbutifolia,
Dalbergia fischeri, Entada wahlbergii, Grewia caffra,
Hugonia busseana, Landolphia kirkii, Loeseneriella
crenata, Opilia celtidifolia, Reissantia buchananii, R.
indica, Strophanthus kombe, S. petersianus, Synaptolepis
alternifolia, S. kirkii and Tiliacora funifera.
In 1994 there was a devastating fire through parts of
the forest. Acacia adenocalyx proliferated and totally
Bothalia 37,1 (2007)
61
overran large areas. Stands were so thick that nothing
was able to grow underneath and it also climbed over
and smothered surviving trees. This has been cleared in
places and one of the pioneer species that regenerated
in the cleared areas was Fernandoa magnifica. Lianes
and climbers also seem to have proliferated as a result
of fire damage and in the area known as Mashamba
Experimental, a 30 m tall Gyrocarpus americanus has
fallen, apparently having been pulled down by their
weight. In the same section, Cordia stuhlmannii and C.
torrei occur. They seem to have survived the fire or man-
aged to regenerate successfully. Perhaps the latter, as
Cordia torrei has also been observed along the edge of
the road on the EN- 1 .
Continuing west along Via Pungue, the forest thins
towards Mpiao Pan and the vegetation becomes open
wooded grassland. The scattered trees include remarkably
tall Boscia salicifolia, Cleistochlamys kirkii, Combretum
adenogonium, Dalbergia melanoxylon, Philenoptera
violacea, Piliostigma thonningii, Sclerocarya birrea and
Vitex payos. Some of the smaller species are Roiirea ori-
entalis, Senna petersiana and Xylotheca tettensis.
Vegetation east of the EN-1 along the road known as
Via Entrada, is very difficult to classify as it is an intri-
cate mosaic of forest, thicket and woodland. The canopy
varies in height considerably and it is probably best
described as dry thicket with some emergent trees over
10 m high. The taller trees are Afzelia quanzensis, Albizia
harveyi. Balanites maughamii, Berchemia discolor,
Blighia imijugata, Boscia salicifolia, Hymenocardia
idmoides, Millettia stuhlmannii, Monodora jimodii var.
macrantha, Monodora stenopetala, Pteleopsis myrtifo-
lia, Strychnos madagascariensis and Warneckea sansi-
bariciim. Other trees are Carpolobia goetzei. Gardenia
ternifolia, Millettia mossambicensis, M. usaramensis and
Rothmannia fischeri. Some of the shrubs are Allophylus
nibifolius, Bauhinia tomentosa, Coffea racemosa,
Cordia pilosissima, Dovyalis xanthocarpa, Friesodielsia
obovata, Indigofera ormocarpoides, Rourea orientalis
and Tricalysia jasminiflora.
East of the Tissadze River along Via Lavu there is a
very different picture. In this area there is very much a
mosaic of forest, thicket and undifferentiated woodland.
There is a canopy of ± 3 to 4 m with tall emergent trees
well over 10 m high, such as Bombax rhodognaphalon,
Newtonia hildebrandtii and Diospyros mespiliformis, of
which, although few and far between, there is no indi-
cation of previous logging; Adansonia digitata, very
tall and slender, is one of the emergents that also occurs
in forest. Millettia stuhlmannii is easily identified with
characteristic grey-green bark and Sterculia africana
also has a distinctive bark. Forest species forming the
lower canopy are Dalbergia boehmii, Lecaniodiscus
fraxinifolius with characteristic pale bark (can also
become very tall), Drypetes reticulata, Strychnos mada-
gascariensis and Trichilia capitata. On the termite
mounds are Lecaniodiscus fraxinifolius, Spirostachys
africana, Tamarindus indica and the occasional Lannea
schweinfurthii.
Usually considered woodland species, and contrib-
uting to the thicket canopy are Albizia anthelmintica,
Combretum adenogonium, C. hereroense, Terminalia
sericea, Drypetes mossambicensis, Schrebera tricho-
clada, Sclerocarya birrea (which becomes very tall
in semi-closed forest but in the open remains far more
squat) and Strychnos madagascariensis. Spirostachys
africana occurs in clumps, sometimes forming pure
stands of its own little forest, as does Cleistochlamys
kirkii. Tall Acacia nigrescens, Crossopteryx febrifuga
and Kigelia africana are found on the fringes and in
open forest, as are Acacia robusta subsp. usambarensis,
which appear to be dying around the pans, possibly as a
result of past dry years. Combretum imberbe also occurs
around the pans and on the heavier soils.
In the more scrubby forest/woodland mosaic Dio-
spyros loureiriana and D. senensis are found. Balanites
maughamii occurs in deep forest and in scrub forest.
Sometimes in the woodland areas there is a tall grass
cover and at others, shrubs, which include Dovyalis
hispidula, Friesodielsia obovata, Grewia forbesii, G.
inaequilatera, G. lepidopetala, Holarrhena pubescens,
Hoslundia opposita (an up to 2 m shrub in the study
area), Margaritaria discoidea, Markhamia obtusifolia,
M. zanzibarica, Ozoroa obovata subsp. obovata and
Phyllanthus ovalifolius.
Sporadically there is an almost park-like opening
in the forest/thicket with a grass cover and very tall
woodland trees up to 10 m high and very few shrubs.
Trees recorded in those areas include Acacia nigres-
cens, A. robusta subsp. usambarensis, Boscia salicifo-
lia, Combretum hereroense, Dalbergia melanoxylon,
Philenoptera violacea and Terminalia sericea.
North from the Catapu turn-off along the ENl as
the road drops down to Tissadze and Chirimadzi River
Valleys, where the altitude is ± 50-100 m, the trees grad-
ually become less forest and more woodland species.
The river valley is open grassland with solitary trees or
clumps of trees and bushes including Acacia robusta
subsp. usambarensis, A. polyacantha, A. xanthophloea,
Combretum imberbe. Ficus sycomoriis, Hyphaene coria-
cea and Xanthocercis zambesiaca, all indicative of a
high water table.
On the Zangue terrace along Via Santove grows the
little known Mozambique endemic. Acacia torrei on
black soils, and three palm species, Borassus aethiopum,
Hyphaene coriacea and Phoenix reclinata. Other species
are: Acacia galpinii, A. polyacantha, A. sieberiana var.
sieberiana, Albizia versicolor, Croton megalobotiys,
Grewia sulcata, Margaritaria discoidea, Pluchea dio-
scoridis and Sclerocarya birrea.
The area of miombo woodland next to the Tissadze
River bridge on the Inhamitanga road is obviously situ-
ated on nutrient-poor soils. The trees are widely spaced,
there is a poor grass cover and few shrubs. The woody
vegetation seems to be mainly concentrated on termi-
taria, adjacent to small pans or along the bank of the
Tissadze River. These include large Cleistochlamys
kirkii, Dovyalis hispidula, Flueggea virosa, Strychnos
62
Bothalia37,l (2007)
potatorum, a large Tamarindus indica (covered with a
Dalbergia arbutifolia), Trichilia capitata and Ziziphus
mucronata. Of the Miombo species only Brachystegia
spiciformis was recorded. There is no continuous canopy
and other trees scattered around are Acacia nilotica,
A. robusta subsp. usambarensis, Diospyros loureiri-
ana, Philenoptera violacea, Schrebera trichoclada and
Vangueria infausta. Close to the river bank next to a
huge Sclerocarya birrea is an 8 m tall Brackenridgea
zanguebarica.
Logging and conservation
Millettia stuhlmannii known in the timber trade as
panga-panga, jambiri and partridgewood, Afzelia quan-
zensis {chamfuta/chanfuta), Cordyla africana (mutondo)
and other hardwoods have been logged extensively
throughout Mozambique for the last 100 years and con-
tinue to be today. Almost all logging has taken place
on a non-renewable basis (pers. obs.). Judged from
the height of many isolated very tall trees, particularly
solitary baobabs, Adansonia digitata, and Sterculia
appendiculata, trees that are not exploited for their
wood, it is suggested that forest was more prevalent in
central Mozambique in the past than is now apparent.
Furthermore, those that remain are frequently being
replaced by rural settlement and cultivation (pers. obs.;
Izidine & Bandeira 2002; Schipper & Burgess 2004),
with the result that much of the extensive forest that once
covered especially the coastal areas of Mozambique, has
disappeared.
Although Catapu is a timber concession, only certain
species are felled. The three targeted species have the
ability to coppice and silvicultural follow-up operations
are now conducted on all stumps that have been logged.
There is also an active programme of re-planting of
Millettia, Cordyla and Afzelia. The seedlings are raised
in a small nursery adjacent to the sawmill and currently
some 10 000 seedlings are planted out into the logged
areas each year in early summer with the first rains.
This represents roughly four times the number of trees
that are felled in the concession each year. The growth
of the seedlings, those which have escaped the attention
of the porcupines, in some places has been remarkable.
For instance, of 34 plants propagated in ground seedbeds
in Nov. 1997, dug up and replanted in the field in Oct.
1998, 20 survived and measurements taken in October
2005 show an average height of 4.15 m with an aver-
age trunk diameter of 140 mm. Obviously the average of
126 mm of rain which falls in the dry season enables the
deep sands to remain moist throughout the greater part
of the year.
A conservation/sensitive forest area of ± 4 000 hec-
tares within the firebreak around the sawmill and resi-
dential area has been established. This includes many
different vegetation communities and has now been pro-
tected from fire for six years. Although woodland and
miombo vegetation types in the study area tolerate burn-
ing and are even dependent on it for maintaining their
structure, this area of forest has clearly not evolved with
fire as a natural factor. Such intolerance to fire is also
seen in Sand (Licuati) Forest on the coastal plains of
Maputaland in southern Mozambique. Currently fire is
increasingly introduced into forest patches with Ae slash
and bum agricultural activities of an expanding human
population.
DISCUSSION
Checklists of plants have an important role beyond
the call for simple inventories of species. They serve as
foundations for the enumeration of flora and vegetation
in given areas. This paper also serves to demonstrate the
diversity of woody species and vegetation types within
a small area. Comparisons of the Catapu checklist with
checklists of the surveys in nearby regions, confirm the
convergence of several floristic and vegetation elements
resulting in the richness and unique composition of the
vegetation at Catapu.
When a plant has a name it has an identity, and the
inclusion of vernacular names in a checklist has the
potential to increase communication during field work.
The trees at Catapu have been numbered and tagged,
their exact position recorded and voucher specimens
prepared. This has resulted in a living reference collec-
tion, cross-referenced to herbarium specimens, available
on site, an important contribution to any future botanical
study or field work in the area.
The present survey shows that 181 species recorded
in the study area at Catapu were also recorded by one
or other of the two available Zambezi River Valley veg-
etation surveys. A total of 60 species on the Catapu list
were not included in either of the Zambezi River Valley
vegetation surveys. Furthermore, this checklist is essen-
tially a list of woody species and is an ongoing exercise
but when ferns, herbs, geophytes and grasses are taken
into account, the diversity will prove considerable.
There is a so-called forest reserve, the Inhamitanga
Forest Reserve, a 2 km wide belt (1 km on either side
of the road) on the Inhamitanga-Chupanga road, which
starts at Inhamitanga Village on the southeastern bound-
ary of Catapu, where the railway line and road meet,
and runs for ± 10 km. Although it is marked on most of
the 1 : 250 000 maps of the area, very few people know
of its existence and there is no formal protection of the
reserve. It falls within Coutada 12, the hunting conces-
sion on the Catapu eastern boundary, so no timber con-
cessions or simple cutting licenses can be issued in the
area. The damage to the reserve so far has been very
limited but like all Mozambique’s forests they face the
constant threat to slash and bum settlement, the risk of
fire damage and of fuel-wood collection.
Not included in the checklist are at least another 10
or more tree species known to occur in the area but not
yet found on Catapu. Hence the importance of this study
which it is hoped can be extended to cover the rest of
the Cheringoma area, including the Inhamitanga Forest
Reserve and adjacent hunting concessions where the
vegetation is probably protected to some extent, par-
ticularly against the ravages of extensive slash and bum
settlement. An increased conservation initiative in this
Bothalia37,l (2007)
63
area is urgently required, together with further biological
studies. It is essential to know what is there before meas-
ures can be taken to save it.
We believe there is a very strong case for the whole
Cheringoma area to be declared an Important Plant Area
(Smith 2004) as it fulfills the following criteria; the site
contains restricted range species (narrow endemics),
has a botanical diversity with a high number of species
and is a regionally threatened habitat. The area forms
part of the Southern Zanzibar-Inhambane Coastal Forest
Mosaic, an ecoregion of which the conservation status is
‘critical’ (Schipper & Brngess 2004). Furthermore, the
Cheringoma area also falls within the Coastal Forests of
Eastern Afnca Hotspot (Burgess et al. 2004), one of 34
such regions identified as Earth’s biologically richest and
most endangered terrestrial ecoregions.
ACKNOWLEDGEMENTS
TCT-Dalmann Furniture, Lda. is thanked for the oppor-
tunity to conduct this study and for providing support
and generous hospitality during visits made to Catapu.
Mario Jose Inacio has been an invaluable guide. He
knows the trees by the local vernacular names in Sena
and their numbers in English. Antonio Luis Domingos
helped with the phonetic spelling of the Sena names.
Mr R.B. Drummond is thanked for all his help with
the identification of difficult specimens. John Burrows
and the Mpumalanga Plant Specialist Group visited
Catapu and their help with identification is acknowl-
edged with grateful thanks. The staff at SANBI in
Pretoria, including those of the Mary Gunn Library, are
always most helpful and are sincerely thanked. The Head
of the Institute of the National Herbarium and Botanic
Garden in Harare, Zimbabwe, is thanked for allowing
the use of the library and herbarium.
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64
Bothalia 37,1 (2007)
APPENDIX 1 . — A checklist of woody species at Catapu
Column 1 : taxon, author name
Column 2: voucher no., collectors’ abbreviations: Cher, Cheringoma herbarium accession no.; DP, David Peta; JAW, James White; M, Meg Coates
Palgrave; MJI, Mario Jose Inacio; qg, quick guide no.
Column 3: growth form: T, tree; S, shrub over 2 m; L, liane; P, herbaceous perennial.
Column 4: Sena name, * published in De Koning (1993).
Column 5: Distr., phytogeograph ical distribution: N, northerly distribution, occurring mainly north of the study area; S, southerly distribution,
occurring mainly south of the study area; T, distribution transitional, occurring both north and south of the study area; E, species endemic to
the Swahilian/Maputaland Regional Transition Zone; Ziziphus mauritiana, marked +, an alien, is included in the list. Tamarindus indica is
considered to be indigenous for the area (Coates Palgrave 2002).
Column 6: Gel., Geldenhuys (1992): 0, absent; I, present.
Column 7: DaS, Da Silva et al. (2004): 1, present; NS, absent or not recorded for Sofala; NM, absent or not recorded for Mozambique.
Column 8: Mul., Muller et al. (2000): 1, present; 0, absent.
Column 9: Tim., Timberlake (2002): 1, present; 0, absent.
Bothalia37,l (2007)
65
APPENDIX 1. — A checklist of woody species at Catapu (cont.)
Column 1 : taxon, author name
Column 2: voucher no., collectors’ abbreviations: Cher, Cheringoma herbarium accession no.; DP. David Peta; JAW, James White; M, Meg Coates
Palgrave; MJI, Mario Jose Inacio; qg, quick guide no.
Column 3: growth form: T, tree; S, shrub over 2 m; L, liane.
Column 4: Sena name, * published in De Koning (1993).
Column 5: Distr., phytogeographical distribution; N, northerly distribution, occurring mainly north of the study area; S, southerly distribution,
occurring mainly south of the study area; T, distribution transitional, occurring both north and south of the smdy area; E, species endemic to
the SwahiliartTVlaputaland Regional Transition Zone; Ziziphus mauritiana, marked +, an alien, is included in the list. Tamarindus indica is
considered to be indigenous for the area (Coates Palgrave 2002).
Column 6: Gel., Geldenhuys (1992): 0, absent; 1, present.
Column 7: DaS, Da Silva et al. (2004): 1, present; NS, absent or not recorded for Sofala; NM, absent or not recorded for Mozambique.
Column 8: Mul., Muller et al. (2000): 1, present; 0, absent.
Column 9: Tim., Timberlake (2002): 1, present; 0, absent.
66
Bothalia37,l (2007)
APPENDIX 1 . — A checklist of woody species at Catapii (cont.)
Palgrave; MJl, Mario Jose Inacio; qg, quick guide no.
Column 3: growth form: T, tree; S, shrub over 2 m; L, liane.
Column 4: Sena name, * published in De Koning (1993).
Column 5: Distr., phytogeographical distribution: N, northerly distribution, occurring mainly north of the study area; S, southerly distribution,
occurring mainly south of the study area; T, distribution transitional, occurring both north and south of the study area; E, species endemic to
the Swahilian/Maputaland Regional Transition Zone; Ziziphus mauritiana, marked +, an alien, is included in the list. Tamarindus indica is
considered to be indigenous for the area (Coates Palgrave 2002).
Column 6: Gel., Geldenhuys ( 1 992): 0, absent; 1 , present.
Column 7: DaS, Da Silva et al. (2004): 1, present; NS, absent or not recorded for Sofala; NM, absent or not recorded for Mozambique.
Column 8: Mul., Muller et al. (2000): I, present; 0, absent.
Column 9: Tim., Timberlake (2002): 1, present; 0, absent.
Bothalia37,l (2007)
67
APPENDIX 1. — A checklist of woody species at Catapu (cont.)
Column 1 : taxon, author name
Column 2: voucher no., collectors’ abbreviations: Cher, Cheringoma herbarium accession no.; DP, David Peta; JAW, James White; M, Meg Coates
Palgrave; MJI, Mario Jose Inacio; qg, quick guide no.
Column 3: growth form: T, tree; S, shrub over 2 m; L, liane; G, geoxylic suffrutex.
Column 4: Sena name. published in De Koning (1993).
Column 5: Distr., phytogeographical distribution: N, northerly distribution, occurring mainly north of the study area; S, southerly distribution,
occurring mainly south of the study area; T, distribution transitional, occurring both north and south of the study area; E, species endemic to
the Swahilian/Maputaland Regional Transition Zone; Ziziphus mauritiana, marked +, an alien, is included in the list. Tamarindus indica is
considered to be indigenous for the area (Coates Palgrave 2002).
Column 6: Gel., Geldenhuys (1992): 0,. absent; 1, present.
Column 7: DaS, Da Silva et al. (2004): 1, present; NS, absent or not recorded for Sofala; NM, absent or not recorded for Mozambique.
Column 8: Mul., Muller et al. (2000): 1, present; 0, absent.
Column 9: Tim., Timberlake (2002): 1, present; 0, absent.
68
Bothalia37,l (2007)
APPENDIX 1. — A checklist of woody species at Catapii (cont.)
Column 1 : taxon, author name
Column 2; voucher no., collectors’ abbreviations: Cher, Cheringoma herbarium accession no.; DP, David Peta; JAW, James White; M, Meg Coates
Palgrave; MJl, Mario Jose Inacio; qg, quick guide no.
Column 3: growth fonn: T, tree; S, shrub over 2 m; L, liane.
Column 4: Sena name, published in De Koning (1993).
Column 5: Distr., phytogeographical distribution; N, northerly distribution, occurring mainly north of the study area; S, southerly distribution,
occurring mainly south of the study area; T, distribution transitional, occurring both north and south of the study area; E, species endemic to
the Swahilian/Maputaland Regional Transition Zone; Ziziphus maiiritiana, marked +, an alien, is included in the list. Tamarindus indica is
considered to be indigenous for the area (Coates Palgrave 2002).
Column 6: Gel., Geldenhuys (1992): 0, absent; 1, present.
Column 7: DaS, Da Silva el al. (2004): 1 , present; NS, absent or not recorded for Sofala; NM, absent or not recorded for Mozambique.
Column 8: Mul., Muller (2000): I, present; 0, absent.
Column 9: Tim., Timberlake (2002): 1, present; 0, absent.
Bothalia37,l (2007)
69
APPENDIX 1 . — A checklist of woody species at Catapu (cont.)
Column 2: voucher no., collectors’ abbreviations: Cher, Cheringoma herbarium accession no.; DP, David Peta; JAW, James White; M, Meg Coates
Palgrave; MJI, Mario Jose Inacio; qg, quick guide no.
Column 3: growth form: T, tree; S, shrub over 2 m; L, liane.
Column 4: Sena name, * published in De Koning (1993).
Column 5: Distr., phytogeographical distribution: N, northerly distribution, occurring mainly north of the study area; S, southerly distribution,
occurring mainly south of the smdy area; T, distribution transitional, occurring both north and south of the smdy area; E, species endemic to
the Swahilian/Maputaland Regional Transition Zone; Ziziphus mauritiana, marked +, an alien, is included in the list. Tamarindus indica is
considered to be indigenous for the area (Coates Palgrave 2002).
Column 6: Gel., Geldenhuys (1992): 0, absent; 1, present.
Column 7: DaS, Da Silva et al. (2004): 1, present; NS, absent or not recorded for Sofala; NM, absent or not recorded for Mozambique.
Column 8: Mul., Muller et al. (2000): 1, present; 0, absent.
Column 9: Tim., Timberlake (2002): 1, present; 0, absent.
70
Bothalia 37,1 (2007)
APPENDIX 1. — A checklist of woody species at Catapu (cont.)
Column 1 : taxon, author name
Column 2: voucher no., collectors’ abbreviations; Cher, Cheringoma herbarium accession no.; DP, David Peta; JAW, James White; M, Meg Coates
Palgrave; MJl, Mario Jose Inacio; qg, quick guide no.
Column 3: growth form: T, tree; S, shrub over 2 m; L, liane.
Column 4: Sena name, published in De Koning ( 1993).
Column 5: Distr., phytogeographical distribution: N, northerly distribution, occurring mainly north of the study area; S, southerly distribution,
occurring mainly south of the study area; T, distribution transitional, occurring both north and south of the study area; E, species endemic to
the Swahilian/Maputaland Regional Transition Zone; Ziziphus mawiliana, marked +, an alien, is included in the list. Tamahndus indica is
considered to be indigenous for the area (Coates Palgrave 2002).
Column 6: Gel., Geldenhuys (1992): 0, absent; 1, present.
Column 7: DaS, Da Silva el al. (2004): I , present; NS, absent or not recorded for Sofala; NM, absent or not recorded for Mozambique.
Column 8: Mul., Muller el a!. (2000): 1, present; 0, absent.
Column 9: Tim., Timberlake (2002): I, present; 0, absent.
Bothalia37,l (2007)
71
APPENDIX 1 . — A checklist of woody species at Catapu (cont.)
Column 1 : taxon, author name
Column 2: voucher no., collectors’ abbreviations: Cher, Cheringoma herbarium accession no.; DP, David Peta; JAW, James White; M, Meg Coates
Palgrave; MJI, Mario Jose Inacio; qg, quick guide no.
Column 3: growth form: T, tree; S, shrub over 2 m; L, liane.
Column 4: Sena name, * published in De Koning (1993).
Column 5: Distr., phytogeographical distribution: N, northerly distribution, occurring mainly north of the study area; S, southerly distribution,
occurring mainly south of the study area; T, distribution transitional, occurring both north and south of the study area; E, species endemic to
the Swahilian/Maputaland Regional Transition Zone; Ziziphus mauritiana, marked +, an alien, is included in the list. Tamarindus indica is
considered to be indigenous for the area (Coates Palgrave 2002).
Column 6: Gel., Geldenhuys (1992): 0, absent; 1, present.
Column 7: DaS, Da Silva et al. (2004): 1, present; NS, absent or not recorded for Sofala; NM, absent or not recorded for Mozambique.
Column 8: Mul., Muller et al. (2000): 1, present; 0, absent.
Column 9: Tim., Timberlake (2002): 1, present; 0, absent.
72
Bothalia 37,1 (2007)
APPENDIX 1 . — A checklist of woody species at Catapii (cont.)
Column 1 : taxon, author name
Column 2: voucher no., collectors’ abbreviations: Cher, Cheringoma herbarium accession no.; DP, David Peta; JAW, James White; M, Meg Coates
Palgrave; MJI, Mario Jose Inacio; qg, quick guide no.
Column 3: growth form: T, tree; S, shrub over 2 m; L, liane; G, geoxylic suffrutex.
Column 4: Sena name, published in De Koning (1993).
Column 5: Distr., phytogeographical distribution: N, northerly distribution, occurring mainly north of the study area; S, southerly distribution,
occurring mainly south of the study area; T, distribution transitional, occurring both north and south of the study area; E, species endemic to
the Swahilian/Maputaland Regional Transition Zone; Ziziphus manritiana, marked +, an alien, is included in the list. Tamarindtis inJica is
considered to be indigenous for the area (Coates Palgrave 2002).
Column 6: Gel., Geldenhuys (1992): 0, absent; 1, present.
Column 7: DaS, Da Silva et al. (2004): I , present; NS, absent or not recorded for Sofala; NM, absent or not recorded for Mozambique.
Column 8: Mul., Miillcr et al. (2000): I , present; 0, absent.
Column 9: Tim., Timberlake (2002): I , present; 0, absent.
Bothalia37,l (2007)
73
APPENDIX 1 . — A checklist of woody species at Catapii (cont.)
Column 1 : taxon, author name
Column 2: voucher no., collectors’ abbreviations: Cher, Cheringoma herbarium accession no.; DP, David Peta; JAW, James White; M, Meg Coates
Palgrave; MJI, Mario Jose Inacio; qg, quick guide no.
Column 3: growth form: T, tree; S, shrub over 2 m; L, liane.
Colurrm 4: Sena name, * published in De Koning (1993).
Column 5: Distr., phytogeographical distribution: N, northerly distribution, occurring mainly north of the study area; S, southerly distribution,
occurring mainly south of the study area; T, distribution transitional, occurring both north and south of the study area; E, species endemic to
the Swahilian/Maputaland Regional Transition Zone; Ziziphus mauritiana, marked +, an alien, is included in the list. Tamarindus indica is
considered to be indigenous for the area (Coates Palgrave 2002).
Column 6: Gel., Geldenhuys (1992): 0, absent; 1, present.
Column 7: DaS, Da Silva et al. (2004): 1, present; NS, absent or not recorded for Sofala; NM, absent or not recorded for Mozambique.
Column 8: Mul., Muller et al. (2000): 1, present; 0, absent.
Column 9: Tim., Timberlake (2002): 1, present; 0, absent.
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Bothalia37,l: 75-77 (2007)
The concept of ^Musa-pelo and the medicinal use of shrubby legumes
(Fabaceae) in Lesotho
A. MOTEETEE*t and B-E. VAN WYK*
Keywords: adaptogen, bitter tonic, Fabaceae, immune stimulant, sedative, traditional medicine
ABSTRACT
In the Kingdom of Lesotho, 20 plant species are commonly known as 'Musa-pelo. The term literally means ‘the one who
brings back or turns around the heart’, 'Musa-pelo is traditionally used as a sedative and is given as a first aid treatm.ent to
bereaved people who are under severe psychological duress or stress. Of the 20 species known as 'Musa-pelo, 1 7 belong
to nine genera of the Fabaceae (Argyrolobium, Crotalaria, Indigofem, Lessertia, Lotononis, Melolobium, Sutherlandia,
Tephrosia and Trifolium). The three remaining species namely Cleome monophylla, Heliophila carnosa and Cysticapnos
pniinosa, belong to the families Capparaceae, Brassicaceae and Fumariaceae, respectively. In this paper, the concept of
'Musa-pelo in traditional medicine is explored.
INTRODUCTION
The concept of 'Musa-pelo is an important part of tra-
ditional medicine in Lesotho. 'Musa-pelo and its cultural
significance is described and explained here for the first
time in a scientific context. Watt & Breyer-Brandwijk
(1962) did not discuss "Musa-pelo in their comprehen-
sive review of medicinal plants of southern and eastern
Africa. The aim of this paper is to present and discuss
the identity and uses of 20 plant species that are known
as 'Musa-pelo or that have been reported in the literature
as 'Musa-pelo.
METHODS
The study was based mainly on a literature survey
and observations made by the first author on traditional
practices at home and in the community; the first author
has also used some of these remedies. Some traditional
healers were consulted with regards to the use of Cleome
monophylla as 'Musa-pelo — the plant was shown to
them in the field.
DISCUSSION
The concept o/’ Musa-pelo
'Musa-pelo is used to treat fits, circulation problems,
stress-related ailments and chronic illnesses (Maliehe
1997). As a traditional healer, Eric Maliehe (pers.
comm.) pointed out: ‘for ages and ages the Basotho have
been using 'Musa-pelo for trauma because trauma is
considered to affect the heart first’. The term 'Musa-pelo
literally means ‘one who brings back or turns around
the heart’ and according to him, it means ‘bringing back
health to the heart’. It is a term given to several plant
species traditionally used to cure or ease a ‘sore heart’
or dropsy of the heart (Phillips 1917; Jacot Guillarmod
1971). In the first author’s own experience, 'Musa-pelo
is administered as a sedative to bereaved people who are
* Department of Botany and Plant Biotechnology, University of
Johannesburg, P.O. Box 524, Auckland Park 2006, Johannesburg,
t Corresponding author e-mail address: amoteetee@uj.ac.za
MS. received: 2006-08-31.
under severe psychological duress or stress. It is given to
recently widowed women for spasms and heart disease.
Most of the traditional uses point to possible sedative
and/or stress-relieving activity for the plants involved.
Plants known to be used as ’Musa-pelo
Detailed information on the Sesotho names and
the traditional uses (where known) of the 20 species
known to be 'Musa-pelo plants are given in Appendix
1. Authorities for the scientific names are included
in Appendix 1 and will not be given elsewhere. In all,
20 plant species, including 17 belonging to the fam-
ily Fabaceae are called 'Musa-pelo. These include two
species of Argyrolobium, one of Crotalaria, four of
Indigofera, two of Lessertia and one of Sutherlandia.
Three other species Cleome monophylla (Capparaceae),
Cysticapnos pruinosa (Fumariaceae) and Heliophila car-
nosa (Brassicaceae), have also been listed as 'Musa-pelo.
Heliophila carnosa is used for auscultation (Phillips
1917); and is also known in Sesotho as Semameloana,
meaning ‘the small one who listens’. Cysticapnos prui-
nosa is used as a ‘charm to comfort and drug the sorrow-
ing’ (Jacot Guillarmod 1971). Phillips (1971) recorded
'Musa-pelo as a Sesotho name for Cleome monophylla,
but did not provide any details of its usage. After consul-
tation with several traditional healers in Lesotho regard-
ing the use of C. monophylla, none of them could con-
firm that it is used as 'Musa-pelo.
To distinguish between the 20 species with one com-
mon name, variations to the name are applied either to
describe a striking feature of the plants, their size or their
habitat. For example, Melolobium alpinum is 'Musa-
pelo-o-moholo-oa-thaba (the big mountain 'Musa-pelo)
because it is indeed an alpine species. To differentiate
Crotalaria distans, flower colour is added to the name,
i.e. ' Musa-pelo-o-mosehlanyana (the yellowish 'Musa-
pelo). Lotononis sericophylla is characterized by whitish
hairs and is known as ' Musa-pelo-o-mosoeu (the white
'Musa-pelo).
Members of the Fabaceae are well known as rich
sources of nitrogen compounds. Some of the species
in Appendix 1 have been investigated (summarized in
Harbome 1994): Argyrolobium lotoides and Melolobium
76
Bothalia 37,1 (2007)
alpimim contain quinolizidine alkaloids (anagyrine, cyti-
sine, leontidine and camoensine), Lotononis sericophylla
is a source of pyrrolizidine alkaloids (integerrimine and
senecionine), Crotalaria distans has amino acids (2,4-
diaminobutanoid acid and N-y-glutamyltyrosine), and
TrifoUum contains flavonoids (maackiain, medicarpin
and vestiol). Of special interest is the closely related
Sutherlandia and Lessertia, both of which contain high
levels of free amino acids and canavanine (a non-protein
amino acid) in the leaves (Van Wyk & Wink 2004).
Possible mode of action q/’‘Musa-pelo
Sedatives are drugs which depress the central nervous
system. Many plants, including those of ’Musa-pelo, are
also used traditionally as sedatives. The traditional use
of Sutherlandia species to relieve anxiety and stress is
believed to be due to the presence of the amino acid
gamma-aminobutyric acid (GABA), a known inhibi-
tory neurotransmitter with mood-elevating properties
(Van Wyk & Gericke 2000). It is interesting to note
that Smith & Myburgh (2004) have presented evidence
that Sutherlandia alters the corticosterone response in
stressed rats.
Adaptogens are described as agents or biological
substances (of plant origin) that help to increase the
body’s resistance to stress, whether physical, chemi-
cal or biological (Schulz et al. 2001). For a plant to be
adaptogenic, it ‘must be innocuous and cause minimal
disorders in the physiological functions of an organism,
it must have nonspecific action, and it usually has a nor-
malizing action irrespective of the direction of the patho-
logical state’ (Brekhman & Dardymov 1969). Some very
well-known adaptogenic plants include Panax ginseng
(Asian ginseng), P. quinuefolium (American ginseng),
Eleutherococcus senticosus (Siberian ginseng), Rhodiola
rosea (roseroot) and Withania somnifera (often called
Indian ginseng) (Van Wyk & Wink 2004).
Immune stimulants are substances that help increase
the activity of the body’s immune system but without
any antigenic action against specific pathogens, whether
viral or bacterial (Schulz et al. 2001). They are therefore
similar to adaptogens in that their action is nonspecific
and could be multifocal. Two well-known immune stimu-
lant herbs are Echinacea spp. (coneflower) and Viscum
album (mistletoe).
Digestive bitters (amara) are tonic herbs with a bit-
ter taste thought to stimulate the taste buds and promote
(as a reflex via the vagus nerve) the secretion of saliva,
gastric juices and bile (Van Wyk & Wink 2004). The bit-
terness can be due to a number of chemical constituents
including volatile oils, alkaloids and sesquiterpene lac-
tones. Surprisingly, bitters also act on the cardiovascu-
lar system, decreasing the heart rate and cardiac stroke
volume (Schulz et al. 2001). Some commonly used
bitter herbs include Gentiana lutea (gentian), which
occurs in mountains of central and southern Europe,
Centaurium erythraea (century), primarily distributed
in the Mediterranean and Artemisia absinthium (worm-
wood), native to temperate regions of Europe, Asia and
northern Africa.
CONCLUSIONS
It appears to be no coincidence that of the 20 plants
used as ’Musa-pelo, 17 plant species are from the
Fabaceae, a family known for its rich diversity of nitro-
gen compounds (Harbome 1994) — alkaloids, phenolic
amines, amino acids, nonprotein amino acids and other
constituents known to act on the central nervous system.
The chemical composition of most ’Musa-pelo plants
and their pharmacological activities (such as possible
corticomimetic and neuromodulatory effects), are not yet
known, and this opens up a new area of investigation. It
should be pointed out, however, that some plants belong-
ing to certain listed genera (Argyrolobium, Crotalaria,
Indigofera and Lotononis) have been reported to have
indications of toxic potential (Hutchings et al. 1996),
therefore, it is advisable that these remedies should not
be taken without proper knowledge of required dosages.
It is possible that ’Musa-pelo plants have undiscovered
compounds inducing adaptogenic, bitter (amarum) or
neuroreceptor responses which are, individually or col-
lectively, responsible for the sedative or mood-enhancing
effects reported for these plants. Altogether there are 20
species listed as ’Musa-pelo plants, however there is no
evidence that Cleome monophylla is used as ’Musa-pelo
and should perhaps be excluded from the list.
ACKNOWLEDGEMENT
The University of Johannesburg is thanked for finan-
cial assistance.
REFERENCES
BREKHMAN, LI. & DARDYMOV, I.V. 1969. New substances of plant
origin which increase nonspecific resistance. Annual Review of
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BRUMMITT, R.K. & POWELL, C.E. 1992. Authors of plant names.
Royal Botanic Gardens, Kew.
HARBORNE, J.B. 1994. Phytochemistry of the Leguminosae. In F.A.
Bisby, J. Buckingham & J.B. Harbome, Phytochemical diction-
ary of the Leguminosae, vol. 1. Plants and their constituents.
Chapman & Hall, Cambridge.
HUTCHINGS, A., SCOTT, A.H., LEWIS, G. & BALFOUR-CUNNING-
HAM, A. 1996. Zulu medicinal plants. University of Natal Press,
Pietermaritzburg.
HUTCHINGS, A. & VAN STADEN, J. 1994. Plants used for stress-
related ailments in traditional Zulu, Xhosa and Sotho medicine.
Part 1 ; plants used for headaches. Journal ofEthnopharmacology
43: 89-124.
JACOT GUILLARMOD, A. 1971. Flora of Lesotho. Cramer, Lehre.
MALIEHE, E.B. 1 997. Medicinal plants and herbs of Lesotho. Mafeteng
Development Project, Lesotho (in Sesotho).
PHILLIPS, E.P. 1917. A contribution to the 'flora of the Leribe Plateau
and environs. Annals of the South African Museum 16: 1-379.
SCHMITZ, M.O. 1982. Wild flowers of Lesotho. ESS A, Roma.
Lesotho.
SCHULZ, V, HANSEL, R. & TYLER, V.E. 2001. Rational phyto-
therapy, a physician s guide to herbal medicine, edn 4. Springer,
Heidelberg.
SMITH, C. & MYBURGH, K.H. 2004. Treatment with Sutherlandia
frutescens subsp. microphylla alters the corticosterone response
to chronic intermittent immobilization stress in rats. South
African Journal of Science 100: 229-232.
VAN WYK, B-E. & GERICKE, N. 2000. People s plants. A guide to the
useful plants of southern Africa. Briza Publications, Pretoria.
VAN WYK, B-E. & WINK, M. 2004. Medicinal plants of the world.
Briza Publications, Pretoria.
WATT, J.M. & BREYER-BRANDWIJK, M.G. 1962. The medicinal and
poisonous plants of southern and eastern Afiica. Livingstone,
Edinburgh.
Bothalia37,l (2007)
77
APPENDIX 1. — Medicinal plants commonly known in Sesotho as ' Musa-pelo. Note: the word 'Musa-pelo means ‘the one who brings back the
heart (pelo)' and since it is the base name for the Sesotho names of all these species, its meaning will not be repeated in the table. References
are given in brackets, detail? at end of table
1, Jacot Guillarmod ( 1971); 2, Phillips (1917); 3, Maliehe (1997); 4, Own experience of first author; 5, Hutchings & Van Staden (1994); 6, Schmitz
(1982). Author abbreviations according to Brummitt & Powell (1992).
Bothalia 37,1: 79-88 (2007)
Can anthropogenic variables be used as threat proxies for South
African plant richness?
M. KEITHt* * and M. WARREN*
Keywords: endemic species, human population density, land transforaiation, setting conservation priorities
ABSTRACT
Human demographic and socio-economic measures (anthropogenic variables) reflect the detrimental impact of humans
on plant diversity globally. The Pretoria (PRE) Computerised Information System (PRECIS) of the South African National
Biodiversity Institute (SANBI), provided three sets of South African plant richness data, overall (OPR), endemic (EPR), and
threatened (TPR), to investigate the relationships between richness and six anthropogenic variables. Spearman’s Rank order
correlations, Kruskal Wallis Analysis of Variance (ANOVA) and Generalized Linear Models (GLZ) were used. Although all
three plant richness measures were correlated with anthropogenic variables, individual anthropogenic variables contributed
a small fraction to the explained variation in richness. Differences in spatial and temporal scaling of the datasets, or the
response to another causal mechanism, may have contributed to this low explained variation. Because more variation was
accounted for in OPR than EPR or TPR, OPR is a more suitable surrogate measure of plant biodiversity when investigating
the anthropogenic variables used here. Average human density (HD), infrastructure (degree of urbanization and road cover)
(LRU) and percentage land area transformed and degraded (LTD) were identified as useful surrogates of human impacts on
OPR. LTD may be a more inclusive human impact measure when conducting analyses of human impacts using OPR. LTD
includes the effects of urban expansion, road networks and other land transformation impacts, such as agriculture.
INTRODUCTION
Human actions threaten biological diversity at a
global scale. The sources of threat include taxon-specific
threats such as exploitation, introduced taxa and various
forms of ecosystem degradation, including land transfor-
mation and pollution (World Resources Institute 2000).
Strong evidence indicates correlations between rates of
habitat and species disappearance with human demogra-
phy patterns, such as population density and population
growth, and human activities (James 1994; Ceballos &
Ehrlich 2002; Harcourt & Parks 2003; Luck et al. 2003).
As a result, conservation planners have been urged to
integrate biological, socio-economic and human demo-
graphic data in their assessments to effectively determine
real world conservation priorities (Cincotta et al. 2000;
Brooks & Thompson 2001 ).
Human demographic and socio-economic data are
often current and easily available (Harcourt & Parks
2003), and may constitute useful surrogate measures
of the proximate threats to certain life forms and taxa.
A number of human-related variables are correlated
with plant richness measures around the world, both at
regional and global scales. These include human demo-
graphic parameters (e.g. population density, human
population growth rate change, poverty and affluence,
urbanization), land transformation, land fragmentation
and fuel wood consumption (Macdonald 1991; Kerr &
Currie 1995; Cincotta et al. 2000; Ceballos & Ehrlich
2002; Liu et al. 2003).
Of these parameters, human population density has
been considered a reasonably good indicator of threat of
the risk of species extinction (Thompson & Jones 1999;
Harcourt & Parks 2003). Plant population declines are
t Former address: Threatened Species Programme, South African
National Biodiversity Institute, Private Bag XIOI, 0002 Pretoria.
* Department of Zoology and Entomology, University of Pretoria,
0002 Pretoria. mkeith.za(@gmail.com
MS. received: 2005-04-01.
mostly concentrated in areas with either high human
densities or high human impact, such as agriculture
(Burgess et al. 2002; Ceballos & Ehrlich 2002; Araujo
2003). In turn, rapid and continuing population growth,
and the associated human impact on the environment, is
ever increasing (Cincotta et al. 2000; World Resources
Institute 2000; Liu et al. 2003), necessitating urgent
investigation into the relationship between plant richness
and human population increase.
Sub-Saharan countries have some of the highest
population growth rates in the world (United Nations
Development Programme 2001). This, tied with high
human population densities, clearly translates into
considerable landscape transformation (James 1994;
McKinney 2001). Most of Africa’s dense human settle-
ments, intensive agricultural activities and habitat frag-
mentation are concentrated in areas of high animal and
plant endemism (Balmford et al. 2001; Harcourt et al.
2001; Burgess et al. 2002; Chown et al. 2003).
A strong association between population growth and
environmental degradation exists that is mediated, in
part, through income. For example, it has been dem-
onstrated that as human populations grow, agricultural
productivity declines, and this in turn raises rural pov-
erty (Ukpolo 1994). Poverty stricken people are forced
to rely heavily on surrounding resources for survival,
placing increased pressure on vegetation and plant and
animal species in the region (Lucas & Synge 1981;
James 1994). Struggling populations in rural areas may
then move to the cities (Ukpolo 1994; World Resources
Institute 2000) resulting in, for example, urban sprawl
that generally involves complete transformation of rela-
tively large areas (Macdonald 1991; Cincotta et al. 2000;
Liu et al. 2003).
On the other end of the income scale, higher per cap-
ita income may also lead to environmental degradation
(Naidoo & Adamowicz 2001). Areas with a high Gross
National Product (GNP) correspond to areas with a high
80
Bothalia37,l (2007)
proportion of threatened plants in various countries (Kerr
& Currie 1995; Naidoo & Adamowicz 2001). High GNP
usually fuels excessive land conversion and resource
exploitation, increasing the number of threatened plant
and vertebrate taxa (Naidoo & Adamowicz 2001). In
many cases, this association between humans and envi-
ronmental degradation, and humans and species threat,
is mutually reinforced by additional anthropogenic mea-
sures such as household dynamics, urbanization, technol-
ogy and political instability (World Resources Institute
2000; Liu et al. 2003). In this sense, the combined forces
of human population pressure apply tremendous stress to
ecological systems in Africa and other underdeveloped
areas (Ukpolo 1994).
Environmental degradation extends beyond the effects
of human population growth. Land use and land-cover
changes are important elements of the larger problem
of global environmental change. Land use impact and
the loss of species can often be directly related to the
percentage area under urbanization, transformation or
fragmentation (Wood et al. 1994; Pfab & Victor 2002).
Theobald (2003) found that areas with more than 15%
infrastructure development coverage could be deemed
highly fragmented and thus impacting negatively on bio-
diversity, resulting in species loss (Santos et al. 2002;
Tschamtke et al. 2002). Correspondingly, roads have
been claimed to have a disproportionate effect on biotic
diversity (Macdonald 1991; Reyers 2004). Reyers et
al. (2001) illustrated that road construction and mainte-
nance significantly altered surrounding natural habitat
and landscapes. In addition, other forms of landscape
transformation and degradation such as areas utilized for
agriculture and plantations may lower species richness.
Although the effect of high human population density on
species richness may be observable through increased
road infrastructure and urban expansion, agriculturally
degraded or transformed landscapes are likely to occur
in lower human density areas, confounding the relation-
ship between land transformation and human population
density. As a result, land-cover changes may also impact
species richness patterns.
Identifying and managing threatening processes,
threatened areas and taxa at risk of extinction at national
scales, in essence, requires interpreting the impacts
of human activities on biodiversity at smaller spatial
extents (Brooks & Thompson 2001; Gardenfors et al.
2001; Chown et al. 2003; Liu et al. 2003). Incorporating
human demographic and socio-economic variables into
conservation priority setting procedures will allow more
informed conservation decisions to be made (Hannah et
al. 1994; Sisk et al. 1994). An understanding of the rela-
tionships between these variables and plant richness at
national scales is essential to detennining the suitability
of such data as surrogate measures to assist in under-
standing the proximate and ultimate threats to plant
taxa (Kerr & Currie 1995; Brooks &. Thompson 2001).
However, ascertaining the relationships between biodi-
versity and human impacts are not as straightforward as
originally believed. The effects of, and threat posed by,
human demographic and socio-economic activities on
taxon richness at national scale remain unclear because
of the complexity and multiplicity of human activities,
as mentioned above.
The impacts of human activities on biodiversity may
also vary between richness measures (Van Rensburg et
al. 2002). For example, human impacts may be greater,
and therefore more apparent, for threatened and endemic
plant taxa than for total species richness. Higher taxon
richness has been successfully used as a biodiversity sur-
rogate in prioritizing areas of conservation importance
at both national and continental scales (Fjeldsa 1997;
Balmford et al. 2001; Harcourt & Parks 2003; Rouget
et al. 2004). But conservation strategies based solely on
overall plant richness are often of limited use (Jetz &
Rahbek 2002). The large proportion of endemic taxa in
southern Africa has been attributed to the diverse ecolog-
ical conditions, as well as the product of high speciation
within a large number of endemic genera (Cowling &
Hilton-Taylor 1994). It is essential to include an analysis
of threatened and range restricted taxa when classifying
areas of conservation priority (Rouget et al. 2004).
In South Africa, the Western Cape has been favoured
by human settlement for the past 350 years, leading to
substantial land transformation through agricultural
and urban development, and alien plant encroachment
(Deacon 1992; Rebelo 1992a; Richardson et al. 1996).
Subsequently, much of the remainder of southern Africa
has also undergone extensive land transformation over
the past 100 years (Macdonald 1991; Van Rensburg
et al. 2004). People have been attracted to the interior
where mineral and fossil fuel resources abound (Deacon
1992). This has resulted in a multi-faceted combination
of human demands and inherent threats across the south-
ern African landscape (Reyers et al. 2001; Neke & Du
Plessis 2004).
Previous work investigating human impacts in South
Africa has focused on the relationships of taxon (or spe-
cies) richness and human variables for South Africa’s
birds and, recently, frogs (Van Rensburg et al. 2002;
Chown et al. 2003; Van Rensburg et al. 2004; Evans et
al. 2006). Their results show that avian species richness
and human density were positively correlated, apparently
as both responded positively to increasing levels of pri-
mary productivity (Van Rensburg et al. 2002; Chown et
al. 2003; Van Rensburg et al. 2004). Here, we investigate
the relationships between three plant richness measures
that include all endemic and threatened South African
plant taxa and six human demographic and socio-eco-
nomic variables, to better understand potential and real-
istic anthropogenic threats to South African plants.
METHODS
Plant distributions of all South African taxa were
extracted from the Pretoria National Herbarium (PRE)
Computerised Infonnation System (PRECIS) for 1 936
quarter degree squares (QDS) of South Africa. Richness
maps were collated for three sets of plant groups. First,
overall plant richness (OPR) was collated for South
Africa at QDS level. Secondly, a list of ‘endemic’ taxa
believed to occur only in South Africa, were extracted
from the PRECIS database as well as Germishuizen
et al. (2006) to produce a richness map for South
African endemic taxa. Finally, a measure of threat-
ened plant richness per QDS was calculated based on
Bothalia37,l (2007)
81
a list of threatened taxa, which was extracted from the
Threatened Species Programme’s database of December
2003 (Threatened Species Programme unpublished
data). This list of threatened species as used here (pre-
liminary 2003 Red List of South African plants) is cur-
rently in the process of being re-assessed according to
the lUCN (2001) Red List Categories & Criteria, by the
Threatened Species Programme of the South African
National Biodiversity Institute (SANBI). On release,
this SANBI Red List will supersede the current 2003 list
used. All taxa listed either by Hilton-Taylor (1996) as
Endangered (E), or Vulnerable (V) or by lUCN (2001)
categories Critically Endangered (CR), Endangered (EN)
or Vulnerable (VU), were classified as Threatened.
Taxon richness counts across South Africa’s QDSs
were square root-transformed following Rebelo &
Tansley (1993). Endemic and threatened taxon richness
per QDS were also square root-transformed and cor-
rected for total plant richness by dividing by total plant
richness in that QDS (following Rebelo & Tansley 1993)
yielding standardized endemic plant richness (EPR) and
standardized threatened plant richness (TPR) at a QDS
scale across South Africa.
South A frican anthropogenic data
Six human and socio-economic variables (anthro-
pogenic variables from here on) were extracted from
various data sources at a national scale in South Africa.
These include: 1, human population density, 2; human
population growth rate change; 3, a poverty index; 4, an
affluence measure; 5, infrastructure; and 6, the degree
of land transformation and degradation. Human popu-
lation density, human population change, poverty, and
affluence data were derived from magisterial district
data (Central Statistical Service 1995, 1998), whereas all
land-cover and transformation data were collated from
the National Land-Cover (NLC) database (Fairbanks &
Thompson 1996; Fairbanks et al. 2000). To standardize
the scale of this data with the plant distributional data,
data were converted to a spatial scale at the QDS level
(25 X 25 kmQ using ESRI ArcView GIS 3.2.
The 1996 South African population census data
(Central Statistical Service 1998) were used to esti-
mate the weighted average population density per QDS
(human density — HD). Human density was denoted as
the average number of people/km- within each QDS.
The average percentage increase or decrease of human
population per QDS (human growth rate change — HC)
over the period 1996 to 2001 (Central Statistical Service
1998; Rouget et al. 2004) was used as a direct proxy for
the impact of human population growth on the environ-
ment.
A poverty index (economic poverty — EP) was esti-
mated as the proportion of people per municipality earn-
ing less than R200 per month (Central Statistical Service
1998). The United Nations Development Programme
South Africa (2003) report indicated that people earning
less than R354 per month could be regarded as earning
below the poverty line. The census data uses broad cat-
egories of which ‘less than R200/month’ together with
the ‘no income’ category are regarded as earning below
the poverty level. This allowed the computation of a
weighted average of the proportion of people per QDS
earning less than R200/month.
A measure of eeonomic affluence (EA) defined as the
weighted average Gross Geographic Product (GGP) per
capita income per QDS, was based on GGP obtained for
all South African magisterial districts (Central Statistical
Service 1995). GGP represents ‘the remuneration
received by the production factors — land, labour capital
and entrepreneurship for their participation in produc-
tion within a defined area’ (Central Statistical Service
1995). The Central Statistieal Service (1995) provides
1994 estimates of GGP and remuneration of employees
by magisterial district in South African Rand (R). Fine-
scale spatial Gross National Product (GNP) data for
South Africa were not available, forcing the use of GGP
data, which represents the finest-scale data available for
South Africa and was incorporated in the current analy-
sis rather than GNP data used by other authors (Kerr &
Currie 1995; Naidoo & Adamowicz 2001).
To obtain a measure to investigate the effects of
urbanization and related development (e.g. industry) on
plant richness in South Africa, infrastructure coverage
(in the form of the percentage of a QDS covered by road
and urbanized areas) was estimated (land-cover, roads
and urban, LRU). Infrastructure data were extracted
from the NLC database (Fairbanks & Thompson 1996).
The extent of the urban area was extracted from all types
of ‘urban/built-up land’ land-cover type (= land-cover
type 24-30; Fairbanks & Thompson 1996) in the NLC
database. A buffered road network for South Africa was
obtained from Reyers et al. (2001) representing various
buffered road types in South Africa.
The extent of land transformation (land-cover, trans-
formed and degraded (LTD) was obtained from the NLC
database, by calculating and summing the percentage
of each land-cover class in each QDS, based on the six
transformed land-cover classes identified by Fairbanks
& Thompson (1996) and Fairbanks et al. (2000). These
classes were based on seasonally standardized Landsat
TM satellite imagery captured primarily during 1994-
1995 and included anthropogenic effects such as forest
plantations, artificial water bodies, urban/built-up areas,
cultivated lands, degraded land as well as mines/quarries.
LTD may be a more inclusive human impact measure
than HD or LRU alone (when available), as it includes
the effects of human population density on urban expan-
sion, road networks and the effects of other forms of
land transformation that occur in lower human density
areas but that may significantly affect species richness
(e.g. agricultural, industrial and other land transforma-
tions, such as plantations).
The values of the weighted average anthropogenic
variables calculated for each QDS across South Africa
(Table 1) conform reasonably well to human statistics
presented by the Development Bank of South Africa
(DBSA) (2000). This suggests that the rescaled (to
weighted average values for each QDS across South
Africa) human demographic variables used here do not
noticeably differ from the provincial scale data issued by
DBSA.
82
Bothalia37,l (2007)
TABLE 1 . — No. approximate total plant taxa, no. endemic and threatened plant taxa for South Africa and for the nine respective provinces. All
national and provincial taxa counts are approximate counts (rounded off) obtained from PRECIS (see Germishuizen et al. 2006 for up to
date counts)
Endemic taxa in area: approximate no. South African endemic taxa occurring in particular region.
Endemic taxa to area: approximation of taxa endemic to area, reported to occur only in area.
Threatened and endemic taxa: no. threatened taxa [Critically Endangered (CR), Endangered (EN), Vulnerable (VU) (lUCN 2001), Endangered (E),
and Vulnerable (V) pre-1994 criteria] based on preliminary 2003 Red List (Threatened Species Programme 2003 unpubl. data).
Information on average anthropogenic variables for each of areas calculated for all provincial quarter-degree squares (QDS), with abbreviations as
follows: HD, human density (people/km-); HC, human growth rate change (% increase/decrease of people); EP, economic poverty (proportion of
people earning < R200/month); EA, economic affluence [Rands (GGP) per capita x lO*"]; LRU, land use roads and urbanization (% area under urban
or road); LTD, land-cover transformed and degraded (% area transformed or degraded).
Statistical analysis
All three measures of richness (OPR, EPR and TPR)
and the six anthropogenic variables (HD, HC, EP, EA,
LREf, LTD) were log-transformed for statistical analyses.
Kruskal Wallis Analysis of Variance (ANOVA) by Ranks
and Spearman’s R Rank order correlations (Zar 1996)
were used to test for statistical differences and correla-
tions, respectively, between measures.
Generalized Linear Models (GLZ; McCullagh &
Nelder 1989) were used to assess the relationship
between OPR and EPR, and between OPR and TPR,
as well as between each of the three measures of plant
richness and each anthropogenic variable independently.
Anthropogenic variables were not included simulta-
neously in the analysis because of high correlations
between some variables (which may lead to collinearity
in the model), the substantial differences in the manner
in which the variables were measured and subsequently
rescaled, and because we wanted to examine the con-
tribution of each variable to explained deviance in each
richness variable independently, to assess its poten-
tial as a surrogate measure of anthropogenic impact.
Because the measures of plant richness were in the form
of counts, a Poisson distribution with a logarithmic link
function was used in the GLZs (Maggini et al. 2002). A
goodness-of-fit test (a deviance statistic), which yields
the proportion of deviance explained (similar to an R^
value) by the variable in the GLZ (McCullagh & Nelder
1 989) was used to determine which anthropogenic vari-
able contributed the most to explained deviance in the
richness variable used. All statistical analyses were
based on analytical subroutines in STATISTICA version
6.1 (StatSoft 2001).
RESULTS
Based on the 2003 data extracted from the PRECIS
data set, ± 1 1 200 of the 22 000 recognized South
African plant taxa are endemic to South Africa. The
preliminary 2003 Red List regarded about 1 900 taxa as
threatened with extinction (Threatened). The Western
Cape and Eastern Cape proved to have the highest num-
ber of plant taxa, endemic as well as threatened (Table 1 ;
Figure 1), mainly in the Cape Floristic Region.
Considering the anthropogenic measures, Gauteng
displayed the highest human variables except for eco-
nomic poverty (EP) and economic affluence (EA —
GGP/capita). KwaZulu-Natal, was highlighted as the
province with the highest proportion of poverty (EP),
and Northern Cape highlighted with the highest aver-
age GGP/capita (EA), mainly the result of lower human
population density and large GGP contributors (mining
and quarrying, data not shown) in the province (Table 1).
Excluding 'per capita’ from the GGP measure, Gauteng
Province was the largest contributor to South Africa’s
economy.
Human densities (HD) and high human growth rate
change (HC) were evident mostly in the large metro-
politan areas such as City of Cape Town, eThekwini
Municipality (city of Durban) and municipalities in
Gauteng Province (Figure 2A, B). EP in turn was higher
in areas in the North-West, Eastern Cape and northeast-
ern KwaZulu-Natal. EA (affruence/capita) was high
in areas with low human density, e.g. Northern Cape
(Figure 2C, D). Gauteng Province yielded QDSs with
the highest levels of urbanization and road coverage,
with Western Cape, Eastern Cape (Transkei), North-West
and Free State yielding areas of high transformation and/
or degradation (Figure 2E, F).
Relationships between species richness measures
Overall plant richness (OPR), endemic plant richness
(EPR) and threatened plant richness (TPR) were all sig-
nificantly different from one another across South Africa
(Kruskal Wallis 4070; P < 0.001). Nonetheless,
BothaIia37,l (2007)
83
FIGURE 1 . — Patterns of plant richness across South Africa, based on
distribution data from Pretoria National Herbarium Computerised
Information System (PRECIS) presented at quarter-degree square
(QDS, 25 X 25 km). A, overall plant richness (OPR); B, endemic
plant richness (EPR); C, threatened plant richness (TPR). Darker
shades represent higher plant richness for each QDS. Threatened
plant richness was based on the preliminary 2003 Red List of
South African plants. Note: this map should not be used for pro-
vincial conservation purposes until our results are verified by the
updated Red List (shortly to be released by SANBI).
EPR was strongly significantly correlated with OPR
(Spearman R = 0.807; n = 1936; P < 0.05), with EPR
(dependant variable) contributing to 63% (Pearson =
3468.52; d.f. = 1935; P < 0.001) of the total deviance
in OPR. OPR and TPR were also strongly correlated,
but not as strongly as the relationship between OPR and
EPR (Spearman R = 0.679; n = 1936; P < 0.05), with the
regression model explaining 61% (Pearson X- = 724.76;
d.f = 1935; P < 0.01) of the total deviance. The strong
correlation and percentage deviance explained for over-
all plant richness with both EPR and TPR indicates, not
surprisingly, that endemic and threatened South African
taxa are relatively strongly dependent on total taxon
richness. Despite this strong link between richness mea-
sures, ± 40% of the variation in overall plant richness
remains unaccounted for when including either endemic
or threatened taxa. We felt it prudent to investigate the
relationships between each plant richness variable and
the six anthropogenic variables, given that the richness
measures varied significantly from one another across
QDSs and because human impacts on endemic and
threatened taxa may differ from overall plant richness.
Relationships between human demographic and
socio-economic variables
The anthropogenic variables were all strongly signifi-
cantly different from one another (Kruskal Wallis
= 7307.40; P < 0.001). All variables were positively cor-
related with one another except for the measure of afflu-
ence (EA — GGP/capita), which was negatively corre-
lated with all anthropogenic variables calculated within
the current study (Table 2). The poverty index (EP) and
human growth rate change (HC) were the only two vari-
ables that were not significantly correlated with each
other. Human density (HD) was strongly positively cor-
related with all variables, except EA (Table 2), suggest-
ing that this variable may be the single, most inclusive
anthropogenic variable to include in investigations of
human impacts on flora. Human density was particularly
strongly correlated with both land transformation (LTD)
and roads and urbanization (LRU).
The effects of anthropogenic variables on species
richness measures
Statistical analysis indicated varying relationships
between OPR and the six anthropogenic variables. Plant
richness (OPR) was positively correlated with human
density (HD), human population growth rate change
(HC), land-cover roads and urban (LRU) and land-cover
transformed and degraded (LTD). Economic poverty
(EP) and economic affluence (EA) were negatively corre-
lated with plant richness, indicating that the effect of
these variables on OPR was inverse to the effect of the
remaining anthropogenic variables (Table 3). Although
HD and LTD were strongly correlated (Table 2), LTD
was able to contribute 2% more than HD to explained
deviance in OPR (Table 3). No single anthropogenic
variable accounted for more than 10.10% deviance
explained in OPR (Table 3). Therefore, the observed
effect of individual anthropogenic variables on OPR was
small, but significant.
84
Bothalia37,l (2007)
FIGURE 2. — Spatial representation per QDS across South Africa. A, human population density (HD); B, human growth rate change (HC); C,
economic poverty (EP); D, measure of economic affluence (EA) denoted as the weighted average GGP/capita (EA); E, infrastructure of land-
cover road and urban areas (LRU); and F, degree of land-cover transformed and degraded (LTD). Darker shades represent higher impacts.
Endemic plant richness (EPR) and the anthropogenic
variables were all weakly correlated with one another
(Table 3). Only the poverty (EP) variable, which was
weakly negatively correlated with EPR, explained a
significant proportion of the deviance (10.50%) in EPR
(Table 3). Other than EP, the anthropogenic variables
used here do not appear to contribute sufficiently to
understanding the effects of human impacts on endemic
plant species.
Threatened plant richness (TPR) and the human vari-
ables were all weakly significantly correlated (Table 3).
Bothalia37,l (2007)
85
TABLE 2. — Spearman Rank order correlation (R-values) for the six
anthropogenic measures
Values in bold: statistical significance of P < 0.05; ns, not significantly
correlated; HD, human density; HC, human growth rate change; EP,
economic poverty; EA, economic affluence; LRU, percentage land-
cover under roads and/or urban; LTD, percentage land-cover trans-
formed and degraded.
Poverty (EP) and economic affluence (EA) were again
negatively correlated with the plant richness variable
(TPR). Only these variables contributed significantly to
explained deviance in TPR, although the contribution
was minimal (Table 3).
DISCUSSION
Strong evidence links rates of habitat and species loss
with human demographic patterns (Cincotta et al. 2000;
Ceballos & Ehrlich 2002; Luck et al. 2003). Given South
Africa’s low economic growth during the past decade
(2.7%), increased population size (2.2% per annum) and
increasing demand on urbanized areas (DBSA 2000;
Statistics South Africa 2002; Chown et al. 2003), anthro-
pogenic effects on species should be readily apparent.
None of the anthropogenic variables measured here
contributed more than 10% towards explaining varia-
tion in any of the three species richness measures used.
Nonetheless, the contribution of single, anthropogenic
variables on overall species richness was both signifi-
cant and observ'able. It is important to remember that
anthropogenic variables entered in combination, may
contribute substantially more than individual variables.
This analysis is, however, beyond the scope of the cur-
rent paper.
Various studies have found a significant, albeit weak,
relationship between human population size, human
population growth, poverty, per capita income, urbaniza-
tion and species per country (Ehrlich & Holden 1971;
Kerr & Currie 1995; Cincotta et al. 2000). Balmford et
al. (2001) reported a marked congruence of high spe-
cies richness and human population density across the
African continent. Chown et al. (2003) state that a strong
significant relationship exists between South African
birds and human population density (R^ = 0.67) at a
quarter-degree scale that is driven, in part, by available
energy (Evans et al. 2006). Moreover, strong evidence
supports remarkably strong correlations between plant
richness and potential evapotranspiration, annual pre-
cipitation, as well as geographic variation in plant taxo-
nomic richness (Rutherford & Westfall 1986; Currie &
Paquin 1987; O’Brien et al. 1998, 2000; Van Rensburg
et al. 2002). Yet none of the human predictors under
consideration here had any such strong relationship with
any of the richness variables, even though weak signifi-
cant correlation was present.
The discrepancy between temporal scales of the data
may have contributed to the weak relationships that were
detected between richness and anthropogenic variables
in this study. The time scale of the plant richness distri-
bution data range from specimens collected in the early
1700s until the present time. Most collections occurred
in and prior to 1970 with only 7% of all the specimen
collections occurring since 1990 (PRECIS unpublished
data). Furthermore, plant distribution data are rarely rep-
resentative and accurate, and in most cases old and out
of date (Rebelo 1992a, b; Freitag & Van Jaarsveld 1997;
Maddock & Du Plessis 1999; Maddock & Samways
2000; Rouget et al. 2004). Conversely, the human and
socio-economic variable data used in the current study
generally date from 1994 to present.
Other than time scale differences in data collection,
another major factor influencing statistical results and
analysis could be the different spatial scales of the biotic
and anthropogenic databases. Most of the anthropogenic
data were measured at magisterial district scale and con-
sequently were transformed by up- or down-scaling to
QDS scale and converted to weighted averages. A QDS
is often too large to reflect finer-scale topographical and
vegetation differences and will most likely not reflect
many of the finer interactions between human predic-
tors and plant richness measures (Rebelo & Tansley
1993; Van Rensburg et al. 2004). However, this effect
TABLE 3. — Spearman Rank (R values) and P value, as well as Generalized Linear Model with Poisson error distribution and Log Link function
(McCullagh & Nelder 1 989) parameters for relationships between overall plant richness (OPR), endemic plant richness (EPR) and threatened
plant richness (TPR) with each anthropogenic variable
HD, human density; HC, human growth rate change; EP, economic poverty; EA, economic affluence; LRU, percentage land-cover under roads
and/or urban; LTD, percentage land-cover transformed and degraded.
Statistical significance: 0.001; **,/*< 0.01; *, P<0.05;ns, not statistically significant.
86
Bothalia 37,1 (2007)
is unlikely to be substantial as the summarized statis-
tics for each province, using the data rescaled to QDS
(see Table 1), are remarkably similar to the results of the
Development Bank of South Africa issued at provincial
scale (DBSA 2000). Notwithstanding these temporal and
spatial scaling issues, all the databases are still useful to
investigate the presence of broader spatial scale relation-
ships and trends, albeit if finer scale interactions may be
weakened.
Finally, other more important causal mechanisms may
dominate plant richness patterns, e.g. climatic variables,
topographic variables and P-diversity (Bailey et al.
2002). Indeed, Evans et al. (2006) suggest that available
energy may be driving the observed patterns in bird spe-
cies richness and human population change.
Even though conservationists are urged to include
human variables when setting conservation priorities,
human population variables are imperfect indicators of
risk to biodiversity (Thompson & Jones 1999; Cincotta
et al. 2000). Indeed, the results of this study show that
the strength of the relationships between human vari-
ables were weak, albeit significant, and the strength of
the relationship did not improve when endemic or threat-
ened taxa were used in the analyses instead of overall
plant species richness (OPR).
Thompson & Jones (1999) found that human popula-
tion density accounted for almost 35% of the variability
in rare and threatened plant loss in a study conducted in
Britain. Our study shows that human population density
did explain a significant proportion of the variation in
OPR but that its contribution to explained variation in
endemic (EPR) or threatened (TPR) plant species rich-
ness was not significant. Therefore, although there is a
strong link between the three richness measures used
here, OPR appears to be the most suited richness surro-
gate when using the anthropogenic variables included in
this study. Nonetheless, it remains important to consider
how human impacts may affect endemic and threatened
plant richness patterns. This study shows that the only
suitable anthropogenic variable to use for EPR and TPR
is economic poverty. This was the only anthropogenic
measure to show a significant relationship with these
richness measures. In addition, TPR data used here are
based on preliminary 2003 data, which will be super-
seded by the Red List due in 2007 (Threatened Species
Programme, SANBI). This will allow for more up-to-
date richness data, as well as updated threat data and
subsequent threat analysis. The results from the present
study should be verified when the updated SANBI Red
List becomes available.
Although clear evidence for different measures of
human impact affecting plant extinction have been
highlighted, it is difficult to assign a risk value to the
impacts of these measures (Kerr & Currie 1995; Czech
& Krausman 1997; Chertow 2001; Ceballos & Ehrlich
2002). Anthropogenic activities form a complex web of
threats that is influenced by various socio-economic and
political factors, e.g. national policies, economic condi-
tions and a host of other factors varying among nations
(Macdonald 1991; Kerr & Currie 1995; DBSA 2000;
McKinney 2001; O’Neill et al. 2001; Liu et al. 2003).
Testing certain human threat predictors (as used in the
current study) should not be taken to mean that other
anthropogenic variables are insignificant, as many addi-
tional human predictors are also extremely important at
local scales (Macdonald 1991; Rouget et al. 2004).
For the current study, human population density (HD),
land-cover that is transformed and degraded (LTD) and
land-cover roads and urban areas (LRU), appear to be
useful measures of anthropogenic impacts on OPR. The
pooled strength of the relationship between these vari-
ables and OPR may not increase as HD is strongly cor-
related with LTD and LRU. LTD is, however, a more
inclusive human impact measure than HD or LRU when
conducting analyses of human impacts using OPR. The
analyses show that slightly more variation in OPR is
explained by this variable than for HD. Also, the effects
of large human populations on urban expansion and road
networks are included in HD and LRU. However, other
forms of land transformation that occur in lower human
density areas that may significantly affect species rich-
ness, such as, areas under agricultural or industrial use
or plantations, are not included in HD and LRU. These
impacts are included in LTD. Plant population decreases
are concentrated in areas with high human densities or
high human impact, such as agricultural areas (Burgess
et al. 2002; Ceballos & Ehrlich 2002; Araujo 2003), also
suggesting that the use of LTD may yield more informa-
tion on human impacts than HD and LRU alone. It has
been shown that human density is clearly a proximate
threat, whereas agriculture, urbanization, land trans-
formation, and roads are ultimate threats (Thompson
& Jones 1999). Further analysis is required to ascertain
whether other human measures are proximate or ultimate
threats.
The differing anthropogenic threats should be
assessed individually for each province prior to any
risk assessment. Plant richness and endemism are not
evenly distributed across South Africa, and most of these
endemic taxa are confined to the predominantly winter
rainfall Fynbos and Succulent Karoo Biomes (Cowling
& Hilton-Taylor 1994; Figure lA, B). Also, as shown
here, anthropogenic variables vary substantially between
provinces. For example, Gauteng is an important eco-
nomic region in terms of business and industrial devel-
opment, mining and agriculture, and is undergoing rapid
expansion of urban areas, constituting the most serious
threat to plant populations in this province (Phab &
Victor 2002).
It is self-evident of the conservation movement today,
that conservation targets are set by incorporating a wide
variety of suitable data, ranging from taxa/species infor-
mation, land types and habitat types (Pressey et al. 2003;
Rouget et al. 2004). Using anthropogenic variables to
improve conservation priority, setting procedures for
plants is somewhat confounded by the low correlation
between anthropogenic and richness variables found in
the current study. Studies that do use anthropogenic vari-
ables as threat predictors for taxa should not be taken
up haphazardly (Cincotta et al. 2000). To further assist
in identifying South African plant taxa threatened by
human activity, additional research is clearly required.
Analysis of different human and socio-economic vari-
ables may well yield substantially different results.
Undertaking a more detailed study on the effects of cli-
Bothalia37,l (2007)
87
mate, water energy dynamics, and topography on vari-
ous species richness measures (not only woody tree and
shrub richness; see O’Brien et al. 1998), could perhaps
provide insight into processes driving species distribu-
tion, and subsequently assist in identifying threats to
these species. Furthermore, it would perhaps be more
relevant to investigate human threats and its impact on
broader units (e.g. vegetation type, habitat units) for
which the QDS data is adequately suited. The analysis
performed here provides insight into the relationships (or
lack thereof) between human and socio-economic vari-
ables as used in the current study, and provides conser-
vation planners with a better understanding of potential
anthropogenic variables that could be regarded as threat
proxies for plant taxa.
ACKNOWLEDGEMENTS
This work would not have been possible without the
assistance of the Royal Norwegian Embassy (NORAD),
via the Department of Environment Affairs and Tourism
(DEAT). We would like to thank Statistics South Africa
for data and the South African National Biodiversity
Institute for use of data from the National Herbarium
(PRE) Computerised Information System (PRECIS). H.
Snyman, J. Victor, W. Foden and D. Pillay are thanked
for their assistance with data acquisition and review of
the manuscript. We thank the reviewers for comments on
the earlier version of the manuscript.
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Bothalia37,l: 89-102 (2007)
Threatened Limestone Fynbos plant communities of Andrew’s Field
and Tsaba-Tsaba Nature Reserve, Western Cape
M.M. ZIETSMAN* and G.J. BREDENKAMP*
Keywords: conservation area, endemic plants, Limestone Fynbos, phytosociology, plant communities, TWINSPAN, Western Cape
ABSTRACT
The vegetation of inland plains and hills of the Andrew’s Field and Tsaba-Tsaba Nature Reserve, Bredasdorp District,
Western Cape was classified using TWINSPAN and Braun-Blanquet procedures. The resulting four plant communities and nine
subcommunities were described and interpreted ecologically. The vegetation was sampled using 97 randomly stratified plots. The
floristic composition, Braun-Blanquet cover-abundance of each species, and various environmental variables were recorded in each
sample plot. The relationship between the vegetation units and the associated environmental gradients was confirmed by ordination,
using the DECORANA computer program, applied to the floristic data set. The conservation priority of each vegetation unit was
determined by taking the occurrence of Red Data List species, limestone endemic species and Cape Floristic Region endemic species
into consideration. The distribution of the plant communities can mainly be ascribed to differences in the clay/sand content of the soil
and the degree of exposure of the vegetation to the dominating winds (southeastern and northwestern) of the area.
INTRODUCTION
The Andrew’s Field and Tsaba-Tsaba Nature Reserve
is situated in the Bredasdorp/Riversdale Centre of
Endemism (Cowling 1992). This centre refers to a well-
defined group of plants confined to the limestones of the
Bredasdorp Formation (Moll et al. 1984) and associ-
ated colluvial deposits (Heydenrych 1994). Limestone
Fynbos is one of the most threatened vegetation types
in the Cape Floristic Region (Hilton-Taylor & Le Roux
1989). Factors threatening Limestone Fynbos vegetation
include alien plants (considered to be the biggest threat
to the natural environment), land clearing, resort devel-
opment, incorrect fire management and over-harvesting
of flowers (Heydenrych 1994).
Cowling et al. (1988) and Rebelo et al. (1991) indi-
cated that very little is known about the vegetation ecol-
ogy of Limestone Fynbos. Both these authors attempted
vegetation classifications based on a combination of
structural characters (Campbell 1986) and dominant spe-
cies. Vegetation classification and description using total
floristics were avoided due to the highly variable floristic
strucmre and the very high number of species in Fynbos
communities (Cowling et al. 1988). Detailed vegetation
studies based on total floristics are therefore very limited
or do not exist at all in Limestone Fynbos. Furthermore,
21% of all Limestone Fynbos species are not conserved
in state-owned nature reserves (Heydenrych 1994). The
conservation importance of Limestone Fynbos, which
dominates the inland plains and hills, makes the effec-
tive management of the area all the more important, and
justifies a detailed vegetation study based on total floris-
tics.
STUDY AREA
Location
The study area is situated in the Bredasdorp District,
Western Cape, between Struisbaai North in the south
* African Vegetation and Plant Diversity Research Centre, Department
of Botany, University of Pretoria, 0002 Pretoria.
MS. received 2005-08-19.
and the De Mond State Forest in the north (Figure 1).
The area is bordered in the west by the Bredasdorp/
Struisbaai road, and in the south by the sea. According to
the geographical division of Heydom & Tinley (1980),
the study area is situated on the south coast, which is a
transitional zone, between tropical and temperate waters.
Offshore is the major retroflex area of Agulhas Current
waters, recurving eastwards and landwards (Heydom &
Tinley 1980).
The study area is ± 979 ha in extent and consists of
Andrew’s Field (± 129 ha) and Tsaba-Tsaba Nature Re-
serve (± 850 ha), which are adjacent areas on the same
farm (Portion 7 and the remainder of Portion 8 of the
Farm Zoetendalsvlei No. 280). Approximately two thirds
of the study area consists of inland plains and hills of
Limestone Fynbos; the rest of the area consists of coastal
thickets. According to Low & Rebelo (1998), two veld
types occur on the inland plains and hills: Limestone
Fynbos and South- and Southwest Coast Renosterveld,
whereas Mustard et al. (1997) also mentions Dune
Asteraceous Fynbos for this coastal area. According to
Mucina et al. (2005), the area consists of Overberg Dune
Strandveld and De Hoop Limestone Fynbos.
Geology
Two formations of the Bredasdorp Group namely the
Strandveld Formation and the Waenhuiskrans Formation
are found in the study area. The Bredasdorp Group is
described by Malan et al. (1994) as Cenozoic sediments
of marine and marine related origin, stretching up to 22
km inland from the current coastline. The rocks are dis-
cordantly laid down on marine-paved rocks of the Table
Mountain, Bokkeveld and Uitenhage Groups (Malan et
al. 1994).
The Strandveld Formation, found along the coast,
consists of white to pale grey dune sands with a high
percentage of shell fragments. Partial cementing of
sands with a high calcium carbonate content took place.
The lithology of this formation can be described as white
dune sand with finely divided shell and alluvial stones
(Malan e/ a/. 1994).
90
Bothalia37,l (2007)
The Waenhuiskrans Formation forms outcrops next
to the current coastline. The Waenhuiskrans stratiotype
is 12.4 m deep, overlain with 1 m thick calcretes locally,
and consists of medium granulated cross-layered cal-
carenite with well-rounded quartz and a few glauconite
grains. Large-scale aeolic cross layers are character-
istic of the unit. The lithology of this formation can be
described as partially calcified dune sand (Malan et al.
1994).
A small portion of the study area’s mother mate-
rial consists of sedimentary rocks (pale grey to pale red
sandy soil), underlain by the Waenhuiskrans Formation
(partly calcified dune sand with calcrete lenses) (Malan
et al. 1994).
Physiography
The study area is situated on the Agulhas Plain, which
is a coastal lowland. Most of the study area falls below
the 10 m contour with a ridge running roughly north-
FIGURE 1. — The location of the
study area.
south reaching an altitude of 31 m in places (Jeffery
1996). The area mostly consists of gradual, sloped lime-
stone hills and flat to gradual, sloped plains.
Soil
The soil is shallow and sandy, overlying limestone, or
deeper and sandy, overlying clay, silt and gravel. Four
different soil forms have been distinguished in the area.
Coega (Orthic A on hard bank carbonate horizon), fam-
ily Marydale (lime containing A-horizon); Immerpan
(Melanie A on hard bank carbonate horizon), family
Kalkpan (lime containing A-horizon); Brandvlei (Orthic
A on soft carbonate horizon), family Koike (signs of wet-
ness in the carbonate horizon); and Namib (Orthic A on
regie sand), family Beachwood (containing lime within
1 500 mm from the soil surface) (MacVicar 1991).
The Coega soil form is found on the limestone hills
and shallow-soil limestone plain. The Immerpan soil
form is found on the deeper soil of a proteoid-dominated
Bothalia37,l (2007)
91
limestone plain. The Brandvlei soil form is found in the
marsh area, as well as on the renosterveld plain, whereas
the Namib soil form is found on the dune plain and the
deep sand found on proteoid-dominated foothills and
plains.
Climate
The main factors affecting the climate are the con-
trasting sea surface temperatures of the two major ocean
currents and the inshore circulation (Tinley 1985). The
south coast has a warm temperate climate with all sea-
sons and bimodal equinoctial rainfall (Strydom 1992).
The study area is situated in the Southern Overberg,
which is a transition between the winter rainfall region
and the nonseasonal rainfall region in the east (Mustard
et al. 1997). According to Heydom & Tinley (1980)
the study area is situated in a low rainfall region, on
an arid inland tongue reaching the coast. The low rain-
fall regime appears to be due to the occurrence of cold
inshore waters, which inhibit shoreline rains. The average
annual precipitation of the area is 444 mm. Precipitation
mostly takes place in the form of rain, but also occurs in
the form of fog. The maximum precipitation is in June,
and the minimum precipitation during February and
December. A climate diagram (Figure 2) was compiled
from data obtained from the Agulhas Weather Station.
The average annual minimum and maximum temper-
atures for the area are 6°C and 13.3°C, respectively. The
maximum and minimum temperatures for the area were
36.1°C in February and 3.9°C in June respectively.
The wind along the south coast is bi-directional;
southeasterly winds alternate with northwesterly and
southwesterly winds (Heydom & Tinley 1980).
METHODS
Releves were compiled in 97 plots, placed in a strati-
fied random way in relatively homogeneous areas, repre-
sentative of particular plant communities. A plot size of
10x10 mwas used, which is considered as large enough
to ensure that all species of regular occurrence in the
stand are present in the sample plot (Rebelo et al. 1991 ).
Sample plots were placed to ensure that each plot ade-
quately represented the stmcture of the particular vegeta-
tion (Werger 1974). A list of plant species found in each
sample plot was compiled. The cover-abundance of each
species in the sample plot was assessed, using the Braun-
Blanquet cover-abundance scale (Mueller-Dombois &
Ellenberg 1974; Werger 1974), with the modification
proposed by Barkman et al. (1964) and Westhoff & Van
der Maarel (1978). Vegetation stmcture, based on h'eight
and plant cover classes, is described according to the
stmctural classification of Edwards (1983).
The Edwards (1983) height classification is as fol-
lows;
FIGURE 2. — A climate diagram for Andrew’s Field and Tsaba-Tsaba
Nature Reserve. A, altitude; B, mean annual temperature; C,
mean annual rainfall; D, mean daily minimum (coldest month);
E, mean daily maximum (hottest month).
The following habitat characteristics were recorded
in each sample plot: altitude, topographical position,
geomorphology, exposure to sun and wind, slope angle,
slope direction, geology, soil, percentage rock cover and
biotic influence.
All data were incorporated into a vegetation database
created in TURBOVEG (Hennekens & Schaminee 2001).
To obtain a first approximation of the plant communities
of the area, releves were extracted from TURBOVEG
into MEGATAB (Hermekens 1996) and then classified
using the TWINSPAN algorithm (Hill 1979) contained
within MEGATAB. These results were then refined by
application of the classical Braun-Blanquet methodology
(Behr & Bredenkamp 1988), using MEGATAB as the
table editor. The results obtained from the classification
are presented in a phytosociological table (Appendix 1).
Plant names conform to Germishuizen & Meyer (2003).
RESULTS
Classification
Six main habitat types are found in the study area,
namely wetlands, limestone hills, shallow limestone
plains, deep-soil limestone plains, undulating dune plains
with neutral sands and Renosterveld.
The plant communities are classified as follows:
1. Phragmites australis-Jimcus kraussii Tall to High
Closed Reed Wetland Community
2. Chondwpetalum microcarpum-Metalasia pungens
Low Closed Ericoid and Restioid Fynbos Community
2.1 Adromischus caryophyllaceiis-Chondropetalum micro-
carpnm Low Closed Limestone Fynbos Subcommunity
2.2 Passerina corymbosa— Chondwpetalum micwcar-
pum Low to Short Closed Fynbos Subcommunity
2.3 Empodium gloriosim-Chondwpetalum micwcarpum
Low to Short Closed Limestone Fynbos Subcommunity
2.4 Eriocephalus kingesii— Chondwpetalum micwcar-
pum Low to Short Closed Fynbos Subcommunity
3. Leucadendron meridianum— Protea obtusifolia Low
to High Limestone Fynbos Community
3.1 Amphithalea alba-Leucadendron meridianum Tall
Closed Limestone Fynbos Subcommunity
3.2 Erica abietina— Leucadendron meridianum Tall
Closed Limestone Fynbos Subcommunity
3.3 Ehrharta calycina-Leucadendron meridianum Low
to Short Limestone Fynbos Subcommunity
92
Bothalia37,l (2007)
3.4 Metalasia muricata-Leucadendron meridiamim Tall
Closed Limestone Fynbos Subcommunity
3.5 Protea susannae-Leucadendron conifenm High Closed
Fynbos Subcommunity
4. Oedera uniflora-Elytropappus rhinocerotis Low Closed
Renosterveld Community
A diagrammatic presentation of the hierarchical clas-
sification and associated environmental interpretation of
the plant communities is given in Figure 3. The commu-
nity numbers in Figure 3 correspond with the plant com-
munity numbers used in the text.
The distribution of the communities is shown in a
vegetation map (Figure 4).
Description of the communities
1. Phragrnites australis-Juncus kraussii Tall to High
Closed Reed Wetland Community
Species per releve: 4.
Diagnostic species: the reed Phragrnites australis, the
rush Juncus kraussii as well as the shrub Plecostachys
serpyllifoUa (Species Group A, Appendix 1).
Dominant plants: Phragrnites australis and Juncus
kraussii.
Structure: high, closed grass Phragrnites australis in
the wetland, with high Phragrnites australis, tall Juncus
kraussii and some short shrubs in the area surround-
ing the dam. The surrounding marshy area contains
tall, closed Juncus kraussii, with some short shrubs in
between.
The community is situated near the comer of the
northern and western borders of the Reserve. It is found
in the flat wetland/marsh area close to a dam, at an alti-
tude of 3 m. Small, irregular, white-grey limestone
stones, originating from the Waenhuiskrans Formation
of the Bredasdorp Group, and orange-brown ferricrete
pebbles associated with Table Mountain Rocks (Malan
et al. 1994), cover 10% of the soil surface. Soils are of
the Coega Form.
2. Chondropetalum microcarpum-Metalasia pungens
Low Closed Ericoid and Restioid Fynbos Community
Diagnostic species: the shmbby Metalasia pungens,
Indigofera meyeriana, Acmadenia obtusata, Agathosma
collina, Muraltia satureoides var. satureoides, the forb
Lobelia setacea, the restio Calopsis viminea, and the
grasses Cymbopogon pospischilii and Setaria sphacelata
var. sphacelata (Species Group B, Appendix 1).
Dominant plants: the restios Ischyrolepis eleocharis
and Chondropetalum microcarpum (Species Group N).
Prominent species: the shrubs Metalasia pungens,
Acmadenia obtusata, Agathosma collina (Species Group
B), Diosma guthriei (Species Group C), Disparago ano-
mala, Phylica ericoides (Species Group M), Morelia
quercifolia (Species Group N), Passerina paleacea
(Species Group P) and Passerina galpinii (Species
Group Q), and the grass Stipagrostis zeyheri subsp. seri-
cans (Species Group M).
Structure: varies from predominantly low, closed eri-
coid and asteraceous shmbs, with low restioids in abun-
dance and with short shmbs sparsely scattered in the
area.
This widespread and diverse community, consisting of
four subcommunities, is found at the western and north-
ern borders of Tsaba-Tsaba Nature Reserve and also in
a mosaic of scattered patches in the central parts of the
study area. It occurs at an altitude of 3-23 m, at a gra-
dient of 0°^5°, on various habitat types, including the
shallow-soil limestone plains, undulating dune plains,
sandy footslopes and southerly slopes of limestone hills.
Small, irregular, white-grey pebbles, stones and rocks
from the Waenhuiskrans Formation cover 0^0% on
the limestone plains and 2-60% on the limestone hills.
Small limestone pebbles originating from the Strandveld
Formation cover 0-5% on the dune plains.
Soils are of the Coega and Brandvlei Forms on the
limestone plain, the Namib Form on the dune plain and
the Coega and Immerpan Forms on the limestone hills.
This restio-dominated fynbos community is floristi-
cally different from the Restioid Fynbos recognized by
Cowing et al. (1988) or Rebelo et al. (1991). It should,
however, be mentioned that Rebelo et al. (1991) com-
bined stmctural and floristic characters in their analysis,
and only dominant species were surveyed. This may
account for some of the floristic differences between
the plant communities from the current study and those
from the Riversdale Plain. Many floristically different
plant communities are, however, expected to be found in
Restioid Fynbos from the limestone areas, as four sub-
communities are recognized in this study:
2.1 Adromischus caryophyllaceus-Chondropetalum micro-
carpum Low Closed Limestone Fynbos Subcommuni-
ty
Species per releve: 28.
Diagnostic species: the shrubby Diosma guthriei, the
succulent forb Adromischus caryophyllaceus and the
sedge Ficinia truncata (Species Group C, Appendix 1).
Dominant plants: the shrubs Metalasia pungens and
Acmadenia obtusata, (Species Group B), and the restios
Ischyrolepis eleocharis and Chondropetalum microcar-
pum (Species Group N).
Prominent species: the shrubs Disparago anomala,
Phylica ericoides (Species Group M) and Passerina gal-
pinii (Species Group Q) and the grass Stipagrostis zey-
heri subsp. sericans (Species Group M).
Structure: low, closed ericoid and asteraceous shrubs,
with low restioids in abundance; short shrubs are also
sparsely distributed.
The subcommunity is found scattered throughout the
study area and is situated at an altitude of 3-10 m on a
gradient of 0°-8°, on the shallow-soil limestone plain,
where pebbles, stones and rocks cover 0-30% of the
soil. Soils are of the Coega and Brandvlei Forms.
2.2 Passerina corymbosa-Chondropetalum microcar-
pum Low to Short Closed Fynbos Subcommunity
Species per releve: 35.
Diagnostic species: the shrubby Passerina corymbosa
and Agathosma dielsiana and the restio Ischyrolepis tri-
flora (Species Group D, Appendix 1).
Dominant plants: the shrubs Metalasia pungens,
Acmadenia obtusata (Species Group B), Agathosma
collina (Species Group B), Morelia quercifolia
Bothalia37,l (2007)
93
FIGURE 3. — The hierarchical classification and associated environmental characteristics of plant communities of the inland plains and hills.
94
Bothalia37,l (2007)
Phragmites austrelis-Juncus kraussii tall to high
closM Reed Wetland community
I Adromischus caryophyllacaus-Chondropetalum
I microcarpiim low closed Limestone Fynoos
subcommunity
I Passerina con
I microcarpum fc
' subcommunity
] Empodium gloriosum-Chondropetalum
I microcarpum low to short, closed Limestone
■ Fynbos subcommunity
t Eriocephalus kingesii-Chondropetalum
5 microcarpum low to shod, closed fynbos
subcommunily
Amphithalea alba-Leucadendron meridianum
I tall closed Limestone Fynbos subcommunity
I Erica abieiina-Leucadendron mendianum tail
! closed Limestone Fynbos subcommunity
Ehrharla calycina-Leucadendron meridiar)um
low to short Limestone Fynbos subcommunity
I Melalasia muncata-Leucadendron mendianum
[ tall closed Limestone Fynbos subcommurtrty
I Protea susannae-Leucadendron coniferum
( high closed fynbos subcommunity
I Oedere uniflora-EMropappus rhinocerolis low
I closed Renosterveld community
FIGURE 4. — Vegetation map of the
Andrew’s Field and Tsaba-
Tsaba Nature Reserve.
(Species Group N) and Passerina paleacea (Species
Group P), and the restios Ischyrolepis eleocharis and
Chondropetaliim microcarpum (Species Group N).
Prominent species: the shrubs Phylica ericoides,
Disparage anomala, (Species Group M), Otholobium
bracteolatum (Species Group P) and Passerina galpinii
(Species Group Q), the restio Thamnochortus insignis
(Species Group Q) and the grass Stipagrostis zeyheri
subsp. sericans (Species Group M).
Structure: mostly low to short, closed ericoid and
asteraceous shrubs; taller shrubs scattered throughout the
area; some tall and short restios in abundance.
The subcommunity is found scattered over the entire
study area at an altitude of 0-12 m, at a gradient of 0°-
26°, on the dune plains where pebbles cover 0-5% and
the sandy, shallow-soil limestone plains with pebbles
covering 7^0%. Soils are of the Namib Fonn on the
dune plains and the Brandvlei Form on the limestone
plain.
2.3 Empodium gloriosum-Chondropetalum microcarpum
Low to Short Closed Limestone Fynbos Subcommu-
nity
Species per releve: 26.
Diagnostic species: the forb Empodium gloriosum.
(Species Group E, Appendix I). Although this species
group is poorly defined, with low constancy of the diag-
nostic species, the subcommunity is further confirmed by
the presence of Species Group B and absence of Species
Groups C, D, and E.
Dominant plants: the shrub Phylica ericoides (Species
Group M) and restioids Ischyrolepis eleocharis and
Chondropetaliim microcarpum (Species Group N).
Prominent species: the shrubs Disparage anomala
(Species Group M), Passerina paleacea (Species Group
P) the restoid Thamnochortus insignis (Species Group
Q) and the grass Stipagrostis zeyheri subsp. sericans
(Species Group M).
Structure: mostly low to short, closed ericoid and
asteraceous shrubs; some taller shrubs scattered through-
out the area; some tall and short restios in abundance.
This subcommunity is situated near the western bor-
der and in the eastern middle part of the Reserve. It
occurs at an altitude of 3-23 m, at h gradient of 0°^5°,
on the southeastern- and southwestern-facing slopes of
limestone hills. Pebbles, stones and small rocks cover
2-60% of the soil. Soils are of the Coega and Immerpan
Fonns.
2.4 Eriocephalus kingesii-Chondropetalum microcarpum
Low to Short Closed Fynbos Subcommunity
Species per releve: 22.
Diagnostic species: the shrub Eriocephalus kingesii
(Species Group F, Appendix 1).
Dominant plants: the shrubs Acmadeuia ohtusata and
Agathosma collina (Species Group B), Phylica ericoides
(Species Group M), Passerina paleacea (Species Group P),
Bothalia 37,1 (2007)
95
the restios Ischyrolepis eleocharis and Chondropetalum
microcarpum (Species Group N).
Prominent species: the shrubs Metalasia pungens
(Species Group B), Disparago anomala (Species
Group M), Morelia quercifolia (Species Group N)
and Passerina galpinii (Species Group Q), the restios
Calopsis viminea (Species Group B) and Thamnochortus
ins ignis (Species Group Q) and the grass Stipagrostis
zeyheri subsp. sericans (Species Group M).
Structure: mostly low to short, closed ericoid and
asteraceous shrubs; taller shrubs scattered throughout the
area with tall and short restios in abundance.
The subcommunity occurs scattered over the entire
study area on footslopes and slopes of limestone hills
and on dune plains at an altitude of 3-2 1 m, with gra-
dients of 0°-16°. Pebbles cover 1-35% of the soil. Soils
are of the Namib Form on the dune plain and Immerpan
and Coega Forms on the limestone hills.
3 . Leucadendron meridianum-Protea obtusifolia Low to
High Limestone Fynbos Community
Diagnostic species: the shrub Helichryswn patulum
and the trees Leucadendron meridianum and Protea
obtusifolia (Species Group G, Appendix 1).
Dominant plants: the shrubs Rims glaiica, Euclea rac-
emosa and the restio Thamnochortus insignis (Species
Group Q).
Prominent species: the shrubs/trees Amphithalea
alba, Leucospermum trimcatum, Erica longifolia, E.
propinqua, Stoebe cinerea and Aspalathus globulosa
(Species Group I), Protea snsannae (Species Group L),
Metalasia muricata (Species Group N), Passerina pale-
acea (Species Group P), and Passerina galpinii (Species
Group Q), the forbs Dimorphotheca nudicaulis (Species
Group I), Dorotheanthus bellidiformis, Gladiolus
gracilis, Dianthus albens (Species Group J), the grass
Eragrostis sarmentosa (Species Group K) and the restios
Ischyrolepis eleocharis and Chondropetalum microcar-
pum (Species Group N).
Structure: a mixture of low, open ericoid-dominated
fynbos shrubland, with scattered groups of tall to high,
closed proteoid-dominated shrubs. High restios are found
in the community, usually more abundant in the closed
proteoid-dominated shrubland areas of the community.
Low restios are also found quite abundantly throughout
the community.
The community occurs from the comer formed by
the northern and western borders of the Reserve up to
the middle of the study area, mnning along the western
border, and on the eastern section of the northern bor-
der, at an altitude of 1-31 m, at a gradient of CT“-16°,
on the shallow and deeps soils of the limestone plains,
along the limestone hills, on the northern and northeast-
ern midslopes of limestone ridges, on the deep sands of
footslopes as well as in the dune troughs between dunes.
Large, irregularly shaped, white-grey rocks, with
some smaller stones and pebbles, as well as rock sheets,
cover 3-65% of the soil on the ridge. Small, irregu-
lar white-grey pebbles and stones cover 0-7% of the
limestone plains. The sand in the dune troughs have no
rocks. The limestone originated from the Waenhuiskrans
Formation of the Bredasdorp Group. Soils are of the
Brandvlei Form on the shallow-soil limestone plain, the
Immerpan Form on the deep-soil limestone plain, the
Immerpan and Coega Forms on the limestone hills, and
the Namib Form in the dune through areas.
The tall-, high-, short-, and low shmb strata have
an average canopy cover of 2 1 %, 1 6%, 1 1 % and 1 1 %
respectively, and an average height of 2.5 m, 1.5 m, 0.75
m and 0.4 m respectively. The forb stratum has an aver-
age canopy cover of 4% and an average height of 0.3
m. The high-, tall-, short-, and low restioid strata have
an average canopy cover of 0.5%, 6%, 2%, and 14%
respectively, and an average height of 2.5 m, 1.5 m,
0.75 m and 0.4 m respectively. The sedge stratum has an
average canopy cover of 0.7% and an average height of
0.15 m. The short- and low grass strata have an average
canopy cover of 1% and 3% respectively, and an aver-
age height of 0.5 m and 0.3 m respectively. Rebelo et al.
(1991) emphasized the conspicuous high cover of prote-
oids and restioids in this type of vegetation.
Similar communities were described from the
Riversdale Plain by Cowling et al. (1988) and Rebelo et
al. (1991) as Leucadendron meridianum/Protea obtusi-
folia Mesic Proteoid Fynbos.
Five subcommunities could be distinguished. The
diagnostic species groups of particularly subcommuni-
ties 3.2, 3.3 and 3.4 are rather weakly defined, though
TWfNSPAN delimited these subcommunities on the
basis of total floristic composition, and the diagnostic
species, though present with rather low constancy, are to
a large extent restricted to the particular subcommunity.
3.1 Amphithalea alba-Leucadendron meridianum Tall
Closed Limestone Fynbos Subcommunity
Species per releve: 42.
Diagnostic species: the shrubs Amphithalea alba,
Leucospermum trimcatum, Oedera capensis and Erica
spectabilis (Species Group I, Appendix 1).
Dominant plants: the shrubs Leucadendron meridi-
anum and Protea obtusifolia (Species Group G) and
the restios Ischyrolepis eleocharis and Chondropetalum
microcarpum (Species Group N).
Prominent species: the shrubs Helichrysiim patulum
(Species Group G), Metalasia calcicola (Species Group
H), Phylica ericoides (Species Group M), Disparago
anomala (Species Group M), Rhus glauca and Euclea
racemosa (Species Group Q).
Structure: low, open ericoid-dominated fynbos shrub-
land, with scattered groups of tall, closed proteoid-domi-
nated shrubs.
The subcommunity is situated near the comer of the
northern and western border of the Reserve, adjacent to
the marsh area. The subcommunity is also found at the
northwestern inland border, as well as near the middle of
the western border of the Reserve, at an altitude of 3-3 1
m, at a gradient of 0°-16°, on shallow-soil limestone
plains and along the limestone hill summit.
Large irregularly shaped, white-grey rocks, smaller
stones and pebbles cover 3-5% of the soil on the lime-
stone hills, while pebbles are present on the limestone
96
Bothalia37,l (2007)
plain. The limestone originated from the Waenhuiskrans
Formation of the Bredasdorp Group. Soils are of the
Brandvlei Form on the limestone plain and the Coega
and Immerpan Forms on the limestone hills.
3.2 Erica abietina-Leiicadendron meridiatmm Tall Closed
Limestone Fynbos Subcommunity
Species per releve: 34.
Diagnostic species: the shrubs Erica abietina var.
abietina, Heterolepis peduncularis and Stoebe cinerea
(Species Group J, Appendix 1).
Dominant plants: the shrubs Leucadendron meridi-
anum and Protea obtusifoHa (Species Group G) and the
restioids Ischyrolepis eleocharis and Chondropetalum
microcarpum (Species Group N).
Prominent species: the shrubs Helichrysum patidum
(Species Group G), Metalasia calcicola (Species Group
H), Phylica ericoides, Disparage anomala (Species
Group M), Metalasia miiricata (Species Group N,
Appendix 1), Passerina paleacea (Species Group P),
Rhus glauca and Euclea racemosa (Species Group Q) and
the restio Thamnochortus insignis (Species Group Q).
Structure: tall, closed, proteoid-dominated shrubland
fynbos.
The subcommunity is found along the northern bor-
der of the Reserve, near the marsh area and at the
northwestern border, as well as near the middle of the
Reserve, at an altitude of 3-3 1 m, at a gradient of 0°-
16°, on the summit of limestone hills, deep soils of the
limestone plains and the deep sand on the footslopes of
the limestone hills. Small, irregular, white-grey pebbles
and stones on the limestone plains, and large, irregular,
white-grey rocks, with some pebbles and stones, as well
as sheets of rock on the limestone hills cover 0-65% of
the soil. Soils are of the Immerpan Form on the lime-
stone plains, the Immerpan and Coega Forms on the
limestone hills, and the Namib Form in the sand area.
3.3 Ehrharta calycina-Leucadendron meridianum Low
to Short Limestone Fynbos Subcommunity
Species per releve: 33.
Diagnostic species: the shrub Cassine peragua and
the grasses Ehrharta calycina, Bromiis diandrus and
Eragrostis sarmentosa (Species Group K, Appendix 1).
Dominant plants: the shrubs Rhus glauca, Euclea
racemosa (Species Group Q) and Passerina paleacea
(Species Group P), and the restio Thamnochortus insig-
nis (Species Group Q).
Prominent species: the shrubs Leucadendron meridia-
num, Protea obtusifolia (Species Group G), Helichrysum
patulum (Species Group G), Metalasia muricata,
Morelia quercifolia (Species Group N), Otholobium
hracteolatum (Species Group P) and Passerina galpinii
(Species Group Q) and the restios Ischyrolepis eleocha-
ris and Chondropetalum microcarpum (Species Group
N) and the grass Stipagrostis zeyheri subsp. sericans
(Species Group M).
Structure: low to short, open ericoid-dominated fyn-
bos shrubland, with groups of tall to high, closed prote-
oid-dominated shrubs; high and low restios are abundant.
The subcommunity occurs in patches from the middle
section of the northern border to the central part of the
Reserve; it also occurs in the centre of the whole of the
study area, near the western and northern borders, at an
altitude of 3-21 m, at a gradient of 0°-16° on the north-
ern and northeastern midslopes and upper slopes of lime-
stone hills and also on the sandy, deep-soil limestone
plain. Irregular white-grey limestone pebbles, stones and
small rocks and rock sheets cover 1-10 % of the soil on
the limestone plain. Large, irregular, white-grey rocks,
with pebbles, stones and rock sheets, cover up to 60% on
the limestone hills. Soils are of the Immerpan Form on
the limestone plain, and the Immerpan and Coega Forms
on the limestone hills.
3.4 Metalasia muricata-Leucadendron meridianum Tall
Closed Limestone Fynbos Subcommunity
Species per releve: 13.
Diagnostic species: Species Group G and the absence
of Species Groups H-L (Appendix 1).
Dominant plants: the shmb Metalasia muricata and the
restio Chondropetalum microcarpum (Species Group N).
Prominent species: the shrubs Leucandendron meridi-
anum, Protea obtusifolia, Helichrysum patulum (Species
Group G) and Passerina paleacea (Species Group P).
Structure: tall, closed, proteoid-dominated shrubland.
The subcommunity is found in patches near the cen-
tral part of the whole study area and in patches near
the southeastern border of the Reserve, at an altitude
of 1-14 m, at a gradient of 0°-8°, on the sandy sum-
mits of limestone hills, and on the deep-soil limestone
plain and on dune troughs. Small, irregular, white-grey
pebbles and stones cover 3% of the soil on the limestone
hill, but no rocks occur on the limestone plain and in the
dune troughs. Soils are of the Namib Form in the dune
area, the Immerpan Form on the limestone plain, and the
Immerpan and Coega Forms on the limestone hills.
The absence of the widespread species of Species
Group M is a further diagnostic feature. An obvious
explanation for the lack of diagnostic species is the very
low species richness of this subcommunity. The species
present in this subcommunity are from Species Group G
(diagnostic species of the Leucadendron meridianum-
Protea obtusifolia low to high Limestone Fynbos com-
munity), and other widespread species (Species Groups N,
P and Q).
3.5 Protea susannae-Leucadendron coniferum High
Closed Fynbos Subcommunity
Species per releve: 24.
Diagnostic and dominant species: the trees Protea
susannae and Leucadendron coniferum (Species Group
L, Appendix 1 ).
Dominant plants: the trees Protea susannae and Leuca-
dendron coniferum (Species Group L, Appendix 1).
Prominent species: the shrubs Heliclnysum patulum
(Species Group G) and Passerina paleacea (Species
Group P) and the restios Ischyrolepis eleocharis and
Chondropetalum microcarpum (Species Group N).
Structure: high, closed, proteoid-dominated shrubland.
The subcommunity is situated in patches along the
northern border of the Reserve, adjacent to the marsh
area, in the comer of the northern and inland northwest-
Bothalia37,l (2007)
97
em border and in patches near the centre of the whole
study area, at an altitude of 3-25 m, at a gradient of 3°-
8°, on the southern and southwestern deep sands on the
footslopes of limestone hills and on the deep-soil lime-
stone plain. There are no rocks in this subcommunity.
Soils are of the Namib Form on the sand areas and the
Immerpan Form on the deep-soil limestone plain.
A further characteristic feature of this subcommu-
nity is the absence of the widespread species of Species
Groups M and N.
Rebelo et al. (1991) also identified a community
with Protea sussanae from the Riverdale Plain, while
Cowling et al. (1988) described a Protea sussanae/
Leucadendron coniferum Mesic Proteoid Fynbos from
the Agulhas Plain.
4. Oedera imiflora-Elytropappus rhinocerotis Low Closed
Renosterveld Community
Species per releve: 25.
Diagnostic and dominant species: the shrubs Elytro-
pappus rhinocerotis, Oedera uniflora and Euchaetis
meridionalis, the grass Andropogon appendiculatus
and the forbs Berkheya coriacea and Tritonia deusta
(Species Group O, Appendix 1).
Prominent species: the shrubs Passerina galpinii and
Helichrysum teretifolium (Species Group Q).
Structure: low, closed renosterveld shrubland.
The community is found in patches near the middle of
the western border of the study area, as well as adjacent
to the wetland area, at an altitude of 3-18 m, at a gradi-
ent of 0°-13°, on the northern and northwestern slopes
of limestone hills, and on the Renosterveld plain. Small,
irregular, white-grey pebbles and stones cover 1-10%.
The tree stratum has an average canopy cover of 6%
and an average height of 2.5 m. The high shrub stra-
tum has an average canopy cover of 6% and an average
height of 1.5 m, while the medium shrub stratum has an
average canopy cover of 7%, and an average height of
0.75 m and the low shrub stratum has an average canopy
cover of 32% and an average height of 0.4 m. The high,
medium and low restio strata have an average canopy
cover of 2%, 0.6% and 4% respectively, and an aver-
age height of 1.5 m, 0.75 m, and 0.20 m respectively.
The medium grass stratum has an average canopy cover
of 17% and an average height of 0.6 m while the low
grass stratum has an average canopy cover of 2% and an
average height of 0.25 m. The sedges have an average
canopy cover of 0.5% and an average height of 0.10 m.
The forb stratum has an average canopy cover of 3% and
an average height of 0.15 m.
The absence of widespread Limestone Fynbos species
listed in Species Groups M, N and P (Appendix 1) is a
feature of this Renosterveld community. This community
is considered to be rare and threatened by encroachment
from alien Acacia species (Zietsman & Bredenkamp
2006). A similar community was recognized by Cowling
et al. (1988) from the Agulhas Plain west of the study
area and Rebelo et al. (1991) from the Riversdale Plain,
east of the study area.
DISCUSSION
A large variety of plant species grouped into various
plant communities occur on the Reserve. This study indi-
cated that a total floristic approach can be successfully
applied in the species-rich Limestone Fynbos communi-
ties. It is also shown that floristically derived plant com-
munities can be mapped at the scale of nature reserves,
and that this can be applied in conservation management
plans. It is, however, recognized that a total floristic
approach for regional studies remains difficult within the
Limestone Fynbos area (Van der Merwe 1977; Cowling
et al. 1988; Rebelo et al. 1991). The use of TWINSPAN
and the application of Braun-Blanquet procedures for
refinement were successful to delimit the communities
and classify them in a hierarchical system. Four plant
communities, with nine subcommunities, were identified.
The plant communities could all be related to specific
environmental conditions and are therefore floristically
and ecologically distinguishable and interpretable. These
plant communities are interpreted as separate ecosystems
(Bredenkamp & Brown 2001), they therefore represent
different management units, and can be incorporated in a
veld management program.
The communities are well defined by diagnostic spe-
cies, though diagnostic species for the subcommunities
often have low constancy values. However, these sub-
communities are retained because they were delimited
by TWINSPAN, based on total floristic composition, and
the diagnostic species are largely restricted to the particu-
lar subcommunities.
CONSERVATION
All the plant communities of the inland plains and
hills are considered to have a high conservation value.
Community 1, the Phragmites australis-Juncus kraussii
tall to high closed Reed Wetland is a sensitive wetland
system that could easily be disturbed and should there-
fore be protected (Doust & Doust 1995; Van Wyk et
al. 2000). The conservation of wetlands is furthermore
enforced by law (Environment Conservation Act, 1989:
Act No. 73 of 1989 and National Water Act, 1998: Act
No. 36 of 1998).
Among the terrestrial plant communities identified in
the study area, nine species encountered are listed in the
Red Data List of southern African plants (Hilton-Taylor
1996) and a further four are listed in the later Southern
African plant Red Data List (Golding 2002), giving a
total of 13 Red Data plant species. Furthermore, 23 lime-
stone endemic species were present. From a PRECIS
list (SANBI 2006), 38 from 238 species collected from
the 3420CC grid, were Red Data species. The Agulhas
plain, on which the study area is situated, is considered,
from a botanical viewpoint, as an area of high irreplace-
ability and high vulnerability, being rich coastal lowland
with remnant patches of coastal renosterveld and low-
land fynbos, which are considered among the highest
priorities for conservation in South Africa and globally
(Schwegler 2003).
Heydenrych (1994) listed 110 plant species endemic
to limestone outcrops. The Agulhas Plain Centre, one
98
of the six phytogeographic centres of the Cape flora,
contains a total of 1 374 species (Goldblatt & Manning
2000). Of the 285 species recorded during the plant
surveys in the Reserve, 138 species are endemic to the
Cape, giving a figure of 48% endemism for the study
area.
It can clearly be seen that the study area is one of
exceptional species richness and conservation impor-
tance. Communities 2, 3 (Limestone Fynbos) and 4
(Renosterveld) should be protected due to the endemic
and Red Data plant species present. Furthermore, large
areas of renosterveld have been destroyed, mainly
because of agricultural land clearing (Low & Rebelo
1998).
CONCLUSIONS
Andrew’s Field and Tsaba-Tsaba Nature Reserve are
of great conservation significance. This area comprises
an important part of a unique natural floral heritage
found at the southern tip of Africa, with several endemic
and Red Data species, and it should be protected and
conserved to maintain its biodiversity for future genera-
tions.
The resulting classification could provide a useful
tool, not only for the management of the plant communi-
ties of Andrew’s Field and Tsaba-Tsaba Nature Reserve,
but also for similar vegetation areas, e.g. the Agulhas
National Park, found in the surrounding region.
ACKNOWLEDGEMENT
Mrs F. Siebert is thanked for her assistance during
data processing.
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APPENDIX I . Phytosociological table of inland plains and hills of Andrew’s Field and Tsaba-Tsaba Nature Reserve
Bothalia37,l (2007)
99
1
APPENDIX 1 . — Phytosociological table of inland plains and hills of Andrew’s Field and Tsaba-Tsaba Nature Reserve (cont.)
100
Bothalia 37,1 (2007)
APPENDIX I , — Phytosociological table of inland plains and hills of Andrew’s Field and Tsaba-Tsaba Nature Reserve (cont.)
Bothalia37,l (2007)
101
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APPENDIX 1 . — Phytosociological table of inland plains and hills of Andrew’s Field and Tsaba-Tsaba Nature Reserve (cont.)
102
Bothalia37,l (2007)
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Bothalia37,l: 103-108 (2007)
Miscellaneous notes
AMARYLLIDACEAE
CHROMOSOME STUDIES ON AFRICAN PLANTS. 20. KARYOTYPES OF SOME CYRTANTHUS SPECIES
INTRODUCTION
The genus Cyrtanthus Aiton (Amaryllidaceae) is
endemic to sub-Saharan Africa. The centre of distribution
is the southeastern Cape with smaller centres in other parts
of the Western and Eastern Cape, Gauteng, Mpumalanga
and KwaZulu-Natal Provinces (Du Plessis & Duncan
1989; Meerow & Snijman 1998; Snijman & Archer 2003).
Cyrtanthus is the largest amaryllid genus in southern
Africa and currently consists of 56 species, one subspecies
and seven varieties (Snijman & Archer 2003, 2006).
There are two major trends characterizing amaryllida-
ceous karyotype evolution (Meerow 1995). Certain gen-
era, e.g., Crinum L. and Hippeastrum Herb., show great
karyotypic stability, with a low frequency of polyploidy.
Similar chromosome morphology among the species of
these genera is characteristic. Their polyploids have a ten-
dency to be autoploid in origin. A genus may also exhibit
great variation in both chromosome number and morphol-
ogy, e.g. Hymenocallis Salisb. and Lycoris Herb. Both
allopolyploidy and Robertsonian changes have been impli-
cated as important factors in speciation in such genera.
Chromosomes in Cyrtanthus are large, as in the rest
of the Amaryllidaceae (Wilsenach 1963). The genus has
base numbers of x = 6, 8 and 11 (Meerow 1995). Most
Cyrtanthus species studied (Figures 1, 2) have a somatic
FIGURE 1. — Mitotic chromosomes in various Cyrtanthus species with 2n = 16. A, C. bicolor. Spies 7185; B, C. brachyscyphus, Spies 7186; C, C.
breviflorus. Spies 7189; D, C. elatus. Spies 7193; E, C. epiphyticus. Spies 7200; F, C. eucallus. Spies 7262; G, C.falcatus, Spies 7208; H, C.
herrei, Spies 7267; I, C. loddigesianus. Spies 7203.
104
Bothalia37,l (2007)
FIGURE 2. — Mitotic chromosomes in various Cyrtanthus species. A, C. mackenii var. cooperi. Spies 7211; B, C, C. mackenii van mackenii (dip-
loid); D, tetraploid. Spies 7268, 7274, 7370; E, F, C. macowanii. Spies 7201; G, H, C. montamis. Spies 7275; 1, C. obliqutis. Spies 7276; J,
C. ohrienii. Spies 7376; K, C. sanguineus. Spies 7216; L, C. Oickii var. viridilobus. Spies 73 77.
chromosome number of 2n = 16 (Taylor 1925; Gouws
1949; Tjio & Levan 1950; Flory 1955; Ising 1962,
1966, 1970; Wilsenach 1963; Bose 1965; Nandi 1973;
Venkateswarlu & Lakshmi 1976; Laksmi 1980) or 2n =
32 (Ising 1966), but 2n = 14 and 22 were recorded by
Bose (1965) and Sato (1938, 1942), respectively for
C. ohliquus, and Mookerjea (1955) and Bose (1965)
recorded 2n = 1 8 for C. sanguinem. Variations in karyo-
types have been described (Wilsenach 1963; Ising 1966).
The aim of this study was to determine the chromo-
some numbers of more species and to compare their
karyotypes.
MATERIALS AND METHODS
Bulbs were collected in the field and grown in green-
houses at ARC-Roodeplaat, Pretoria. Herbarium speci-
mens are stored in the Geo Potts Herbarium (BLFU),
Bloemfontein (Appendix 1).
Root tips were collected between 08:00 and 10:00
in glass bottles filled with tap water (4°C) and refriger-
ated for 24 hours. Subsequent to the cold pre-treatment,
the water was replaced with Pienaar’s fixative (Pienaar
1955). The root tips were macerated in IN HCl for 7
minutes at 60"C and stained with 2% propionic-carmine,
BothaIia37,l (2007)
105
FIGURE 3. — Ideograms of some
Cyrtanthus species. A, C.
bicolor, B, C. brachyscyphus\
C, C. breviflonis; D, C. elatus\
E, C. epiphyticus; F, C. eucal-
lus; G, C. falcatus', H, C. her-
rei.
to which a trace of ferric-acetate was added. Permanent
slides were analysed under a Nikon Microphot FXA
microscope, chromosome counts were made and cells
with well-spread chromosomes were photographed with
a Cool Pix digital camera.
All chromosomes in a minimum of 20 well-spread
metaphase cells were measured to determine the karyo-
types. The following parameters were used for each
metaphase plate: 1, chromosome length of each indivi-
dual chromosome (CL); 2, length of the short arm (p); 3,
total chromosome length (TCL); 4, relative chromosome
length (CLyXCL); 5, relative short arm length (pyTCL).
The arithmetic averages of the relative chromosome
lengths and relative short arm lengths were used to draw
the ideograms and to indicate the standard deviation
from the mean for each chromosome pair.
RESULTS AND DISCUSSION
Sixty-two specimens, representing 19 species were
studied. A somatic chromosome number of 2n = 2x =
16 was observed in all specimens (Figures 1, 2), except
two C. mackenii var. mackenii specimens. Spies 7274
(Figure 2D) and 7372, which were both tetraploid with
2n = 4x = 32 (Appendix 1). This is the first report on
a natural occurring tetraploid in C. mackenii and these
somatic chromosome numbers indicate a basic chromo-
some number of x = 8 (Figures 1, 2).
Reported chromosome numbers for Cyrtanthus indi-
cate 2n = 22 as the previous highest somatic chromo-
some number (Sato 1938, 1942). This report further
supports the chromosome numbers observed by Taylor
(1925), Gouws (1949), Tjio & Levan (1950), Flory
(1955), Ising (1962, 1966, 1970), Wilsenach (1963),
Bose (1965), Nandi (1973), Venkateswarlu & Lakshmi
(1976) and Laksmi (1980). The apparent different basic
chromosome numbers described in the literature, such
as X = 6 and 1 1 in Cyrtanthus, are most probably the
results of incorrect identifications or incorrect interpre-
tation of metaphase plates. In the Sato report (1938),
the material was embedded in paraffin and cut in cross
sections. This method does not enable researchers to
examine single layers of large cells in their totality and
thus may result in miscounts of chromosomes (Darling
& La Cour 1942). No vouchers were cited in this report
and therefore they cannot be consulted for accuracy of
chromosome acounts. The Bose report (1965) is also a
miscount, given ample evidence from recent counts that
C. sanguineus has 2n = 16. Ising (1970) already com-
mented on this issue, concluding that the secondary con-
strictions occurring sometimes, could also be a reason
106
Bothalia 37,1 (2007)
FIGURE 4. — Ideograms of some
Cyrtanthus species. A, C.
loddigesianus; B, C. mackenii
van mackenii', C, C. mackenii
van cooperi', D, C. montanus',
E, C. obliquus', F, C. obrienii',
G, C. sanguineus', H, C. tuckii
van viridilobus.
for incorrect counts. The real base number is x = 8, as
supported by this study. In this study, the somatic chro-
mosome number of 2n = 32 indicates a doubling of the
somatic chromosome number of 2n = 16.
Previous studies indicated that ideograms vary among
the different Cyrtanthus species and may be used in the
identification of certain species (Ising 1970). This vari-
ability is not supported by this study. Chromosome con-
traction varies greatly between different chromosomes
and in a single root tip, different ideograms can be
obtained from different cells, varying slightly depending
upon the time the cell entered mitosis. The ideograms
of 16 diploid taxa are shown in Figures 3 and 4. Each
taxon has eight meta- to submetacentric chromosomes,
arranged in the ideograms from the longest to the short-
est. There is a significant difference in length between
the longest and shortest chromosomes in each of these
taxa. However, when a large number of metaphase cells
are measured, and the average length of a chromosome
pair in all cells are used, the variation in ideograms is
minimal and cannot be used for identification purposes.
This study clearly indicated that Cyrtanthus has a
chromosome complement of 2n = 2x = 16. Although
ideograms provided additional information, they proved
to be relatively unreliable for identification purposes.
ACKNOWLEDGEMENTS
The University of the Free State and the Roodeplaat
Vegetable and Ornamental Plant Institute of the
Agricultural Research Council are thanked for finan-
cial assistance during this study. The latter Institute and
Johan Wentzel are also thanked for collecting and pro-
viding the bulbs used in this study.
REFERENCES
BOSE, S. 1965. Cytological studies in the genus Cyrtanthus. Bulletin of
the Botanical Society of Bengal 19:
DARLINGTON, C.D. & LA COUR, L.F. 1942. The handling of chro-
mosomes. Allen & Unwin, London.
DU PLESSIS, N. & DUNCAN, G. 1989. Bulbous plants of southern
Africa. Tafelberg, Cape Town.
FLORY, W.S. 1955. Chromosome complements of some Cyrtanthus
species. Virginia Journal of Science 6: 250, 251.
GOUWS, J.B. 1 949. Karyology of some South African Amaryllidaceae.
Plant Life (Herbertia) 5: 54-81.
ISING, G. 1962. Chromosome balance in Cyrtanthus. Plant Life
(Herbertia) 18: 95-128.
ISING, G. 1966. Cytogenetic studies in Cyrtanthus. I. Segregation in an
allotetraploid. Hereditas 56: 27-53.
ISING, G. 1970. Evolution of karyotypes in Cyrtanthus. Hereditas 65:
1-28.
LAKSMI, G. 1980. Cytotaxonomical studies in eight genera of
Amaryllidaceae. Cytologia 45: 663-673.
MEEROW, A.W. 1995. Towards a phylogeny of Amaryllidaceae.
In P.J. Rudall, P.J. Cribb, D.F. Cutler & C.J. Humphries,
Monocotyledons: systematics and evolution: 169-179. Royal
Botanic Gardens, Kew.
Bothalia37,l (2007)
107
MEEROW, A.W. & SNIJMAN, D.A. 1998. Amaryllidaceae. In K.
Kubitzki, The families and genera of vascular plants, vol. Ill:
83-110. Springer- Verlag, Berlin.
MOOKERJEA, G. 1955. Cytology of amaryllids as an aid to the under-
standing of evolution. Caryologia 7: 1-71.
NANDI, S. 1973. Chromosome studies in several genera of
Amaryllidaceae with special reference to the status of the tribe
Zephyrantheae. Journal of Cytology and Genetics 7: 24—35.
PIENAAR, R. DE V. 1955. Combination and variation of techniques
for improved chromosome studies in the Gramineae. Journal of
South African Botany 2 1 : 1-9.
SATO, D. 1938. Karyotype alteration and phylogeny. IV. Karyotypes
in Amaryllidaceae with special reference to SAT-chromosome.
Cytologia 9: 203-242.
SATO, D. 1942. Karyotype alteration and phylogeny in Liliaceae and
allied families. Japanese Journal of Botany 12: 57-161.
SNIJMAN, D.A. & ARCHER, R.H. 2003. Amaryllidaceae. In G.
Germishuizen & N.L. Meyer, Plants of southern Africa: an anno-
tated checklist. Strelitzia 14: 957-967.
SNIJMAN, D.A. & ARCHER. R.H. 2006. Amaryllidaceae. In G.
Germishuizen, N.L. Meyer, Y. Steenkamp & M. Keith, A check-
list of South African plants. Southern African Botanical Diversity
Network Report No. 41: 858-869. SABONET, Pretoria.
TAYLOR, W.R. 1925. The chromosome morphology of Veltheimia, Allium
and Cyrtanthus. American Journal of Botany 12: 104—115.
TJIO, J. & LEVAN, A. 1950. The use of oxyquinoline in chromosome
analysis. Analysis Estacidn Experimental de Aula Dei 2: 21-64.
VENKATESWARLU, J. & LAKSHMI, N. 1976. Karyotype of
Cyrtanthus mackenii Hook. f. Current Science 45: 148, 149.
WILSENACH, R. 1 963. A cytotaxonomic study of the genus Cyrtanthus.
Cytologia 2i\ 170-180.
A. STRYDOM*, R. KLEYNHANS** and J.J. SPIES*
* Department of Genetics (62), University of the Free State, P.O. Box
339, 9300 Bloemfontein.
** Agricultural Research Council, Roodeplaat, Private Bag X293,
0001 Pretoria.
MS. received: 2006-05-29.
APPENDIX I. — Specimens of Cyrtanthus taxa studied with their voucher numbers, localities or author and somatic chromosome numbers
108 Bothalia 37,1 (2007)
APPENDIX 1 . — Specimens of Cyrtanthus taxa studied with their voucher numbers, localities or author and somatic chromosome numbers (cont.)
Bothalia37,l: 109-118 (2007)
The handling of the proposal to conserve the name Acacia at the 17th
International Botanical Congress — an attempt at minority rule
G. MOORE*
Keywords: Acacia, conservation, International Botanical Congress, International Code of Botanical Nomenclature, nomenclature, Nomenclature
Section, Racosperma C.Mart., Senegalia Raf., Vachellia W.F. Wight & Am.
ABSTRACT
The handling of controversial Proposal 1 5 84 to conserve the name Acacia with a conserved type for the Australian acacias
during the Nomenclature Section meeting at the 1 7th International Botanical Congress (Vieima) in 2005 is reviewed. Through
a simple majority vote, this Section adopted mles requiring a 60% majority of votes to approve any proposal to modify the
International Code of Botanical Nomenclature and a simple majority to approve all other motions; motions not receiving the
required majority were to be rejected. However, for the motion addressing Proposal 1584,45.1% voted to conserve the type of
the name Acacia for Australian acacias, and 54.9% voted to retain the current African type for the name Acacia. Even though
this motion failed to get a 60% majority either way as required by the Section’s own mles. Section officials have concluded
that the name Acacia is to be conserved for Australian acacias. Treating a motion as approved, even though it received only
minority support, also violates the fundamental principle of standard parliamentary procedure — the right of the majority to
approve proposals. For Acacia to be formally conserved, the Nomenclature Section needed to approve a motion addressing
Proposal 1584 with a majority vote, and this never happened in Vienna. Recommendations are made on how this process
might be improved.
Introduction
The Nomenclature Section of the 17th International
Botanical Congress met during July of 2005. Division
III, Provision 1 of the International Code of Botanical
Nomenclature (Code) states that the Code ‘may be
modified only by action of a plenary session of an
International Botanical Congress on a resolution moved
by the Nomenclature Section of that Congress’ (McNeill
et al. 2006: 117). The Section at Vieima acted on numer-
ous proposals to amend the Code, including proposals to
conserve and reject names, and the decisions made by
the Section have been incorporated into the latest printed
version of the Code (McNeill et al. 2006).
One of the proposals that the Section addressed was
Proposal 1584 (Orchard & Maslin 2003) to conserve the
name Acacia Mill, with a conserved type. This proposal
was made so as to maintain the name Acacia for the bulk
of Australian species that would otherwise have to go by
the name Racosperma C.Mart. under a pending new tax-
onomy (Pedley 2003).
Unlike proposals to amend the rules of the Code,
proposals to conserve or reject names are usually first
reviewed by two committees, the relevant nomencla-
ture committee (Committee for Spermatophyta in the
case of Proposal 1584) and the General Committee,
before a decision is taken by the Nomenclature Section
(Articles .14. 12, 56.2 in McNeill et al. 2006: 3-1, 99).
Proposal 1584 was met with strong resistance but did
receive positive recommendations from the Committee
for Spermatophyta and General Committee, although
the vote was divided in both committees. This set the
stage for action on Proposal 1584 by the Nomenclature
Section and Congress. At the Nomenclature Section
meeting, when the motion dealing with Proposal 1584
was put up for a vote. Section officials required that 60%
* Brooklyn Botanic Garden, 1000 Washington Avenue, Brooklyn, New
York 11225 USA. E-mail: gerrymoore(gibbg.org
MS. received: 2006-10-07.
of the votes cast on this matter be opposed to Proposal
1584 in order for that proposal to be rejected. The vote
on this motion received only 45.1% to conserve the
name Acacia for Australian acacias.
Despite the minority support of 45.1% for Proposal
1584 by the Nomenclature Section, Section officials
treat Proposal 1584 as having been approved, lead-
ing Rijckevorsel (2006) to ask, ‘Is it possible that the
Nomenclature Section made a decision to modify the
Code, i.e. to include the name Acacia with a conserved
type by voting 54.9% against?’ In order to address
Rijckevorsel’s question, a detailed examination of the
history of the process of the conservation and rejec-
tion of names is provided. A review is also provided on
what occurred at the Nomenclature Section meeting in
Vienna. Given that the Section at Vienna, through voting,
adopted rules of procedure that required a 60% majority
vote for any motion to pass, an answer of ‘no’ is given to
Rijckevorsel’s query.
Review of procedure for conservation of names
Prior to the Paris Congress of 1954, proposals to con-
serve names, like proposals to amend the rules of the
International Code of Botanical Nomenclature, were pre-
sented directly to the Nomenclature Section. For exam-
ple, at the Stockholm Congress of 1950, over 39 sets of
250-1- individual proposals to conserve names of fami-
lies or genera were made by over two dozen botanists
(Lanjouw 1950). This led Acting Rapporteur Lanjouw
(1950: 221) to note that all these proposals could not
be individually dealt with at the Nomenclature Section
meeting, and he recommended they be handled by vari-
ous special (nomenclature) committees.
While there was general agreement by the Nomen-
clature Section with Lanjouw’s suggestion, there was
considerable discussion regarding what the relative
authorities should be among the special (nomenclature)
committees. General Committee and Nomenclature Sec-
110
Bothalia 37,1 (2007)
tion. A relevant portion of this discussion is given below
(Lanjouwl953; 538, 539):
‘Dr Rogers asked what was the power of the Special Committee[s]
with regard to nomina conservanda proposita.
Prof. Merrill said they had power to conserve.
Prof. Lanjoltw thought they would refer to the General Committee.
Dr Sprague said in the last Congress these committees had been given
power and they had worked in some cases exceedingly well, espe-
cially where the committee had an energetic secretary. He suggested
power be given to the committee and then it should secure the sanc-
tion of the General Committee for conservation.
Prof. Baehni asked how we were to know what names were con-
served.
Prof. Lanjoltw asked if he meant between Congresses.
Prof. Baehni; Yes.
Prof. Lanjoltw: By means of our new association [International
Association for Plant Taxonomy] we will publish new lists as soon as
they are available.
Dr Blake inquired whether the list of nomina generica conservanda
would be published in a preliminary way so that the names would be
subject to criticism before final adoption.
Dr Ramsbottom replied that the names would be accepted informally
in the interim, the list to be tentative until formally approved by the
next Congress.
Dr Rogers said he agreed to that.
Prof. Merrill put the motion before the meeting that Committees have
power to select names for conservation but the list will be tentative
until the next Congress. [Italics in original.]
This was carriedhy a large majority. [Italics in original.]’
The current procedure for the conservation and rejec-
tion of names has remained largely unchanged since
its establishment with the founding of the International
Association for Plant Taxonomy in 1950 and the addi-
tion of Division III to the Code at the Paris Congress of
1954. The following overview is from Rickett & Smith
(1958: 151; see also Figure 1):
‘[C]onservation can be effected only by action of a plenary session
of an International Botanical Congress, as specified in Division III,
Provision I of the Code....
(1) the author sends his proposal to the rapporteur, and it is published
in Taxon',
(2) by this publication the proposal is brought to the attention of the
General Committee;
(3) the proposal, with advance agreement by the General Committee, is
automatically referred to the appropriate study committee;
(4) after study of the case, this special [nomenclature] committee pub-
lishes its report in Taxon',
(5) the General Committee studies this report and any collateral infor-
mation sent to it, reaches a decision on each proposal by mail ballot,
and prepares a recommendation for the next International Congress;
(6) the next Congress at a plenary session affirms or rejects the recom-
mendation of the General Committee;
(7) the nomen conservandwn proposition, if all decisions up to this
point are affirmative, is listed in the next edition of the Code and
becomes a part of that document.’
My only quibble with Rickett & Smith’s over-
view is that (6) actually involves two steps: first the
Nomenclature Section approves or rejects the proposal
to conserve or reject, taking into consideration the
committees’ recommendations, and then a plenary ses-
sion ratifies (or not) the Nomenclature Section’s deci-
sion. To summarize, the committees recommend, the
Nomenclature Section decides and the International
Botanical Congress, through a plenary session, ratifies.
The new printed edition of the Code that incorporates the
decisions taken by the previous Nomenclature Section
is also subject to approval by the next Nomenclature
Section (see Greater el al. 2000: 15). Under the current
Code (McNeill el al. 2006), besides the conservation of
names of families and genera, it is also possible to con-
serve the names of species and reject names at any rank.
Proposals to conserve or reject names are clearly pro-
posals to amend the Code, since conserved or rejected
names must appear in one of the Code’s Appendices (II,
III, IV, V); hence Rickett & Smith’s (1958) citation of
Division III, Provision I of the Code that governs the
procedure for its modification. Stafleu & Lanjouw (1954:
8): ‘Dr Lanjouw explained that at present the Appendices
are parts of the Code and that some of them even contain
rules’.
Currently, the nomenclature committees (Commitees
for Vascular Plants*, Bryophyta, Fungi, Algae, Fossil
Plants) and the General Committee operate under a pro-
cedure whereby it takes a three-fifths (60%) majority to
either recommend acceptance or rejection of a proposal
(Nicolson 1996). Proposals that fail to receive a 60%
vote either way remain unresolved. Previously, commit-
tees have operated under rules requiring a simple major-
ity (Voss 1979: 179) or a two-thirds majority (Greater et
al. 1989: 215) to recommend acceptance of a proposal,
and proposals that were not approved were automatically
considered as rejected. Recently, proposals to conserve
or reject names have not been included in the synopses
of proposals prepared by the rapporteurs (Greater &
McNeill 1993; Greater & Flawksworth 1999; McNeill &
Turland 2005), although they were included in the past
(Stafleu & Voss 1969; Voss & Greater 1981; Greater &
McNeill 1987).
The Nomenclature Sections’ voting majority required
to approve proposals to amend the Code has varied from
Congress to Congress, with a simple majority (Lanjouw
1953: 488, 489; 1959: 29), two-thirds majority (Stafleu
1954: 192) and three-fifths majority (Stafleu 1966: 8)
being used. The Nomenclature Section at Seattle in 1969
required a 60% majority to approve proposals to amend
the articles (rules) of the Code and a simple majority to
approve proposals to amend other portions of the Code
(Stafleu & Voss 1972: 4, 5). Nomenclature Sections since
Leningrad in 1975 have required a 60% majority for all
proposals to modify the Code; in the case of competing
proposals, a vote of 60% has been required to authorize
the change, and a simple majority to choose between the
competing proposals (Voss 1979: 132; Greater & Voss
1982: 9; Greater et al. 1989: 14; 1994b: 12; 2000: 14,
15; McNeill et al. 2005: 1058).
Occasionally there has been debate at Nomenclature
Sections regarding what type of majority vote (e.g. sim-
ple, three-fifths, two-thirds) was required to approve a
proposal to amend the Code. During the Section meeting
at Leningrad in 1975, Greater moved for a 60% major-
ity for proposals to amend the Code (Voss 1979: 132),
but at Yokohama in 1993 (Greater et al. 1994b: 12), he
moved for a simple majority; Brummitt objected both
times, in Leningrad (Voss 1979: 132) arguing for a sim-
ple majority, stating ‘a 59% vs 41% vote would be frus-
trating’, but in Tokyo (Greater et al. 1994b: 12) defend-
ing the 60% majority, noting that the 60% requirement
’'‘Prior to the Vienna Congress there was not a Committee for Vascular
Plants but rather separate committees for Spemiatophyta and Pterido-
phyta.
Bothalia37,l (2007)
111
At the next Nomenclature Section meeting, the Code as a printed document is approved.
T
If the proposal to conserve or reject is approved by the Nomenclature Section and ratified by a plenary session of the International Botanical
Congress, the name appears in one of the Code’s appendices.
A plenary session of the International Botanical Congress ratifies (or not) the decisions of the Nomenclature Section.
t
The Nomenclature Section at the next International Botanical Congress decides whether or not to approve the proposal, taking into consider-
ation the recommendation of the General Committee.
t
The General Committee issues recommendation. If the General Committee recommends the proposal’s acceptance, the Code authorizes (Art
14. 14) ‘retention (or rejection) of that name. ..subject to the decision of a later International Botanical Congress’.
T
The Nomenclature Committee issues recommendation, publishes a report in Taxon.
t
The General Committee refers proposal to the appropriate permanent Nomenclature Committee. At this point, the Code recommends (Rees.
14A.1 ) that authors ‘should follow existing usage of names as far as possible’ pending the recommendation of the General Committee.
T
Proposal is published in Taxon
FIGURE 1. — Steps a proposal to conserve or reject a name must go through under the current procedure authorized by the International Code of
Botanical Nomenclature (McNeill et al. 2006) and administered by the International Association for Plant Taxonomy.
‘has always been applied in these sessions and that it had
worked very well’.
The primary justification for the 60% requirement to
approve any proposal to amend the Code is to ensure
that changes to it are generally accepted (Greuter et al.
2000: 14). The 60% requirement is similar to the proce-
dure in Robert et al. (2000), which recommends a two-
thirds majority to amend existing rules. Sturgis (2001:
132), however, suggests that requiring greater (e.g.
three-fifths, two-thirds) than a simple majority is unwise
because of the ‘power it gives to a minority’ to reject
proposals thereby overriding ‘the majority’s wishes.’
Neither Robert et al. (2000) nor Sturgis (2001) permits
the approval of a motion when it receives a minority
vote.
Actions on proposals to conserve or reject names by
Nomenclature Sections: Paris Congress (1954) — St
Louis Congress (1999)
Since the establishment of the current nomenclature
committee structure in 1950, proposals to conserve or
reject names have always been subject to the approval
of the Nomenclature Section. At earlier Nomenclature
Section meetings, proposals to conserve names were
presented as proposals to amend the relevant appendices
in the Code where the entries would appear if approved
(Rousseau et al. 1960: 30-33; Stafleu 1966: 61-65; Voss
1979: 173) or as a batch of nomina conservanda propos-
ita (Stafleu & Voss 1972: 125). At the Section meeting
at Sydney in 1981, proposals to conserve or reject were
addressed when the General Committee reported; two
votes were taken, one accepting the General Committee
report and a second ‘to accept the proposals to conserve
and reject names as approved by the General Committee’
(Greuter & Voss 1982: 105). At the Nomenclature
Section meetings at Berlin in 1987, Yokohama in 1993
and St Louis in 1999, a single vote was taken approving
the General Report and the proposals to conserve and
reject as recommended by the General Committee (Greuter
etal. 1989:215; 1994b: 252, 253; 2000: 237, 238).
The changes in how proposals to conserve or reject
names have been presented to Nomenclature Sections
(i.e. proposals to amend appendices, nomina conser-
vanda proposita, General Committee reports) should not
be taken as evidence that the Section’s authority on this
matter has declined, while that of the committees has
expanded. The full authority given to the Nomenclature
Section and the International Botanical Congress to
modify the Code under Division III, Provision I has not
changed since it was introduced with the Paris Code in
1956 (Lanjouw et al. 1956).
While there has always been considerable debate by
Nomenclature Sections regarding proposals to amend the
rules of the Code, there has seldom been discussion on
the proposals to conserve or reject individual names (see
Greuter et al. 1989: 215, 216 for an example involving
the conservation of the name Lycopersicon esculentum
Mill.), and these proposals have always been approved
112
Bothalia37,l (2007)
by the Section. There has also never been any serious
discussion of decisons taken by Nomenclature Section
meetings during any of the plenary sessions responsible
for ratifying the proposals passed at the Section meet-
ings, although a question was raised at the plenary ses-
sion at Vienna regarding the proposal to conserve the
name Acacia with a conserved type. The plenary sessions
have always ratified the decisions of the Nomenclature
Sections through a single resolution.
Actions on proposals to conserve or reject names at
Vienna (2005)
Orchard & Maslin (2003) proposed (Proposal 1584)
to conserve the name Acacia Mill., replacing the name’s
current type, A. scorpioides (L.) W.F.Wight (= A.
nilotica (L.) Delile), with A. penninervis Sieber ex
DC. The aim of the proposal is to preserve the name
Acacia for the vast majority of species known from
Australia (± 1 000 spp.) that would otherwise have to
go by the name Racosperma C.Mart. under a proposed
new classification (e.g. Pedley 2003). Under the new
classification, and provided the proposal was accepted,
the generic names of many African and American spe-
cies traditionally in Acacia would change to Vachellia
W.F.Wight & Am., whereas other species would change
to Senegalia Raf, regardless of the proposal’s accep-
tance.
The proposal was reviewed by the Committee for
Spermatophyta, with many submissions by both those
in favour and those opposed, and it was approved by
a 9-6 (60%) vote in early 2004 (Bmmmitt 2004). (For
the sake of transparency, I mention that at the time I
was a member of this committee and I voted to reject
this proposal, although, working in a floristic region
with no species of Acacia, I did not have a strong opin-
ion on the case.) This proposal was strongly supported
by the committee’s Secretary, Richard Bmmmitt (Smith
et al. 2006), who along with Tony Orchard, one of the
authors of Proposal 1584, subsequently posted dozens
of letters from Australian citizens in support of Proposal
1584 on several notice boards outside the lecture hall
in Vienna where the Section meetings were taking
place; correspondence submitted to the Committee for
Spermatophyta from opponents of Proposal 1584, such
as letters signed by twelve legume taxonomists and 24
African botanists, was not posted. Prior to the Vienna
Congress there was much debate within the botanical
community with articles for (Maslin 2004a, b; Maslin
& Orchard 2004; Orchard & Maslin 2005) and against
(Walker & Simpson 2003; Pedley 2004; Luckow et al.
2005) the proposal being published.
The General Committee had still not issued its rec-
ommendation on Proposal 1584 when the Nomenclature
Section convened in July 2005, over one year after the
Committee for Spermatophyta had made its recom-
mendation. During the Section meeting, the General
Committee finally voted to approve the recommendation
of the Committee for Spermatophyta by a vote of 14-6-2
(63.6%; percentage figure assumes three positions on the
General Committee were vacant or had been eliminated
at the time of the vote, since the Nomenclature Section
at St. Louis elected 25 members to serve; see Nicolson
1999). In the 1 July 2006 General Committee circular to
members, the Secretary of the General Committee, Fred
Barrie, concluded ‘the best approach for the GC would
be to approve this report, thus providing the foundation
for the inevitable debate in Vienna’. This advice may
have influenced committee members to cast a positive
vote, even though debate on Proposal 1584 could have
occurred at the Nomenclature Section meeting in Vienna
regardless of the outcome of the General Committee’s
vote (Smith et al. 2006).
Action on the General Committee’s recommendations
occurred on the last day of the Nomenclature Section
meeting (16 July). After the nomenclature committees’
reports were received, the General Committee reported.
Instead of a single report, the Section received two
reports from the General Committee. The first covered
all recommendations, except the one for Proposal 1584
to conserve the name Acacia with a conserved type. This
first report, and therefore the recommendations regarding
proposals to conserve or reject contained therein, was
approved by a show of hands vote. The Nomenclature
Section then received a second report from the General
Committee that contained only one recommendation,
that to approve Proposal 1584. Section officials stated
that there would have to be a 60% vote in order to reject
any recommendation made by the General Committee.
There is some confusion on the wording of the motion
regarding the General Committee report on Acacia. The
official report (McNeill et al. 2005: 1059) indicates that it
was a vote on a motion from the floor to reject the report,
whereas the unedited transcript (copy of the relevant por-
tion provided by John McNeill) states that the vote was
on a motion to adopt the report (and therefore the recom-
mendation on Proposal 1584). Regardless of the wording
of the motion, the Nomenclature Section, after a lengthy
and reasoned debate, took a card vote with 203 (45.1%)
in favour of adopting the General Committee’s recom-
mendation on Proposal 1584 and 247 (54.9%) voted for
rejecting the recommendation. Therefore, the General
Committee’s recommendation on Proposal 1584 received
only minority support from the Section.
Article 14.14 — the justification for the procedure used
for Proposal 1584 at Vienna
Article 14.14 of the Code is central to those who
defend the procedure used at Vienna for Proposal 1584.
The essential language of this article dates back to a pro-
posal (Ramsbottom et al. 1929: 11; Hall 1926) to the 5th
International Botanical Congress at Cambridge in 1930.
Despite Rapporteur Briquet’s (1930: 14) initial nega-
tive reaction to the proposal [‘Le rapporteur eprouve
un sentiment de malaise a voir consacrer par un article
special (20bis) des Regies un regime de nomenclature
provisoire.’], it was passed at the Cambridge Congress
(Briquet 1931) and appeared in the Cambridge Code
(Briquet 1935) as Article 22: ‘When a name proposed
for conservation has been provisionally approved by
the Executive Committee [a precursor to the General
Committee], botanists are authorized to retain it pend-
ing the decision of the next International Botanical
Congress’.
Bothalia 37,1 (2007)
113
Rickett & Smith’s (1958) successful proposal to
amend this language — deleting the word ‘provisionally’
and adding the phrase ‘after study by the Committee for
the taxonomic group concerned’— did not change its
meaning but simply clarified that the rule did not take
effect until after the General Committee approved the
recommendation of the special [nomenclature] commit-
tee rather than when the proposal was first published in
Taxon. Starting with the Sydney Code (Voss et al. 1983),
language was added to cover the rejection of names, giv-
ing Article 14.14 its current language (McNeill et al.
2006: 31, 32): ‘When a proposal for the conservation
of a name, or its rejection under Article 56, has been
approved by the General Committee after study by the
Committee for the taxonomic group concerned, retention
(or rejection) of that name is authorized subject to the
decision of a later International Botanical Congress’.
Prior to the Tokyo Code (Greuter et al. 1994a),
Article 14.14 was used as justification for provision-
ally listing in the Code, names recommended by the
General Committee for conservation or rejection but
not ratified by an International Botanical Congress, with
a statement that such names were subject to a decision
by a later Congress. In the Seattle (Stafleu et al. 1972),
Leningrad (Stafleu et al. 1978), Sydney (Voss et al.
1983) and Berlin Codes (Greuter et al. 1988), names
approved by the General Committee but not acted on
by the Nomenclature Section or International Botanical
Congress were asterisked, indicating ‘conservation
approved by the General Committee; use authorized
under Article 15 [currently Article 14.14] pending final
decision by the next Congress’. Earlier Codes (Lanjouw
et al. 1956, 1966) also listed provisional entries that
still required action by a later International Botanical
Congress.
The justification for the 60% vote to reject the
General Committee’s recommendation to conserve the
name Acacia Mill, with a conserved type was, that once
the General Committee recommended the name Acacia
for conservation, it had de facto nomen consetx'andiim
status under Article 14.14 (McNeill 2006; Stafleu 1964:
6). Therefore, opposition to the General Committee’s
recommendation was viewed as analogous to a proposal
to amend the Code since the name Acacia could appear
as a provisional entry in a future printed edition of the
Code if no action were taken at a Congress (McNeill
2006).
Was the procedure used at Vienna appropriate?
The intent of Article 14.14 was not to give primary
authority of conservation/rejection of names to the
General Committee, as the previously cited discussion
from the Nomenclature Section at Stockholm in 1950
(Lanjouw 1953: 538, 539) makes clear. The intent was
simply to prevent the possible creation of ‘superfluous
names’ (Ramsbottom et al. 1929: 38) by giving authors
guidance during the interim period when a General
Committee recommends a proposal for approval and
action at the next Congress (there was a 20 year inter-
val between the Cambridge Congress of 1930 and
the preceding Brussells Congress in 1910 where the
Ramsbottom et al. proposal was passed). For example,
if the General Committee recommended conservation of
a name shortly after a Congress had adjourned, Article
14.14 would allow users to retain this name until the
next Congress.
However, in the case of Proposal 1584, the interim
period between the General Committee finalizing its rec-
ommendation and the vote of the Nomenclature Section
was not years but a mere few days! The continent of
Africa did not suddenly abandon the name Acacia and
adopt the names Vachellia and Senegalia during those
few days.
More importantly, such provisionally conserved
names have de facto conserved status but not de jure
conserved status. It is obvious that not everything in the
printed book that goes by the name International Code
of Botanical Nomenclature is part of the de jure Code
as approved by the Nomenclature Section and Congress
(Rijckevorsel 2006). Examples of such items include
tables of contents, prefaces, and indices. Likewise,
things can be part of the de jure Code although they
do not appear in the printed version of the Code, such
as what appears on the errata slip accompanying some
printed copies of the Tokyo Code (Greuter et al. 1994a).
Furthermore, even when a new version of the Code is
published, it itself is provisional until it is approved by
a subsequent Nomenclature Section and International
Botanical Congress (e.g. Greuter et al. 2000: 15;
McNeill et al. 2005: 1058). Ramsbottom, the lead author
of the proposal (Ramsbottom et al. 1929) that resulted
in the basic language now in Article 14.14, was very
clear that names recommended for conservation by the
General Committee were to be ‘accepted informally
in the interim, the list to be tentative until formally
approved by the next Congress’ (Lanjouw 1953: 539).
Most critically, the procedure used when voting on the
General Committee recommendation on Proposal 1584
at the Nomenclature Section meeting in Vienna violated
the most fundamental principle of parliamentary pro-
cedure— the right of the majority to approve decisions,
as noted in the following passage from Sturgis (2001:
130): ‘The most fundamental rule governing voting is
that at least a majority vote is required to take an action.
... Jefferson said, “Until a majority has spoken, nothing
has changed’’. It is obvious that to permit fewer than a
majority to decide for any group would subject the many
to the rule of the few, and this would be contrary to the
most basic democratic principle. Democratic peoples
universally accept decisions by majority vote’.
While it is standard parliamentary practice (Robert et
al. 2000) to permit in some cases a supermajority vote
to approve a motion (such as the rule adopted by the
Nomenclature Section in Vienna to require a 60% vote
to amend the Code) and therefore pennit a minority vote
to result in the rejection of a proposal, it is against stan-
dard parliamentary practice (Robert et al. 2000; Sturgis
2001; Sylvester 2004) to allow a motion to be consid-
ered approved when the motion receives only minor-
ity support, such as the motion to approve the General
Committee’s recommendation on Proposal 1584.
Declaring a proposal approved when the vote on it regis-
tered a majority disapproving, also belies basic common
sense.
114
Bothalia37,l (2007)
The procedural switcheroo — forcing the 60% require-
ment on those opposed to changing the Code — used for
the General Committee’s recommendations on proposals
to conserve or reject names, also created much confu-
sion at the Nomenclature Section meeting. This confu-
sion is not surprising when one considers that, like all
preceding Nomenclature Section meetings, the Section
at Vienna required a positive majority or supermajor-
ity to approve all motions. However, when the General
Committee’s recommendation on Proposal 1584 was up
for a vote, they were told the vote would require 60%
to reject. What was the Section being asked to reject?
A proposal from the floor not to accept the General
Committee’s recommendation on Proposal 1584? Or
the General Committee recommendation itself? These
two different interpretations would require different
votes from supporters and opponents of Proposal 1584.
Nothing perhaps exemplifies this confusion better than
the fact that the official report (McNeill et al. 2005:
1059) and the unedited official transcript give different
interpretations as to what was meant by a ‘yes’ vote, the
official report indicating that it was to support a motion
from the floor to reject the General Committee’s recom-
mendation on Proposal 1584 (the vote being reported as
247:203, 54.9%) and the official transcript indicating a
‘yes’ vote supported adoption of the recommendation
(the vote here being reported as 203:247).
Further evidence of confusion at the Nomenclature
Section meeting at Vienna was that when the vote on
Proposal 1584 was being readied ‘Africa’ and ‘Australia’
were written on the whiteboard with ‘60%’ written next
to Australia seemingly indicating that 60% would have
to vote to move the type to the name of the Australian A.
penninervis. The ‘60%’ was later erased and written next
to ‘Africa’. Robert et al. (2000: 100): ‘It is preferable
to avoid a motion containing a negative statement even
in cases where it would have meaning, since members
may become confused as to the effect of voting for or
against such a motion. ... In this connection, it should be
noted that voting down a motion or resolution that would
express a particular opinion is not the same as adopting
a motion expressing the opposite opinion, since — if the
motion is voted down — no opinion has been expressed’.
Such procedures may have been especially confusing for
those at the Section meeting whose first language was
not English. (Requests to officials of the Nomenclature
Section to receive a copy of the audio recording of the
proceedings of the Nomenclature Section meeting so as
to better understand the precise wording of the motion
concerning Acacia were not honoured.)
It is also important to remember that, unlike the
nomenclature committees which rarely meet, the Nomen-
clature Section meeting is the only time botanists truly
deliberate on the Code (Robert et al 2000: xx): ‘[T]he
opportunity for simultaneous aural communication
among all participants is central to the deliberative char-
acter of a meeting... [AJlthough e-mail or faxes may
provide a suitable substitute for postal mail (the primary
methods of communication used by the nomenclature
committees)... they are not suited for the conduct of the
deliberative process....’ Furthennore, the Nomenclature
Section meeting is more likely to better represent all
users of plant names due to its openness and, therefore.
greater number of participants. All of this argues for the
vote of the Nomenclature Section to prevail when it con-
flicts with the vote of the General Committee. No excep-
tions.
Was there a valid outcome at Vienna on Proposal
1584?
There are no bylaws or standing rules that govern the
procedure to be used at a Nomenclature Section meet-
ing, Division III, Provision I of the Code stating merely
that the Code can be modified ‘only by action of a ple-
nary session of an International Botanical Congress on
a resolution moved by the Nomenclature Section of that
Congress’. Therefore, each Nomenclature Section is free
to establish its own rules of procedure.
On the first day, the Nomenclature Section at Vieima,
like all Sections since the Leningrad Congress in 1975,
approved through a simple majority vote, rules requir-
ing a three-fifths (60%) majority to pass any proposal to
amend the Code and a simple majority to pass any other
motion; motions not receiving the required majority
were to be rejected. The Section never voted to suspend
these rules, nor was a rule permitting a motion to be car-
ried with less than a majority vote, ever voted on and
approved (such a rule would be highly unorthodox with
respect to standard parliamentary practice).
The statement from the Nomenclature Section offi-
cials that the vote on the General Committee recom-
mendation on Acacia would not require a majority vote
for approval was in direct conflict with the rules of pro-
cedure the Section had previously established and was
therefore operating under invalid circumstances — Sturgis
(2001: 130): ‘Any requirements permitting decisions by
less than a majority vote.. .are not valid unless they are
included in the law, the rules of parliamentary law, or the
bylaws’ (italics in original). Robert et al. (2000: 332):
‘[MJotions are out of order if they conflict with a motion
that has been adopted by the society and has neither been
rescinded, nor reconsidered and rejected after adoption.
Such conflicting motions, if adopted, are null and void
unless adopted by the vote required to rescind or amend
the motion previously adopted’.
Therefore, in no way can the vote on the General
Committee’s recommendation on Proposal 1584 be taken
as an approval of that recommendation, since the motion
received only minority (45.1%) support. Nothing done at
the plenary session (held on 23 July 2005) resolved this
matter, since it simply approved the decisions taken by
the Nomenclature Section (Stuessy 2006: 246).
Some will say that the arguments in this paper have
come too late and that the time to object was at the
Nomenclature Section meeting. However, most bota-
nists are not experts on parliamentary procedure, and
should not be expected to have an objection ‘at the
ready’ when something like the unusual procedure used
for Proposal 1584 occurs. Furthennore, Section officials
never indicated that they were establishing a new proce-
dure by requiring only a 40% positive vote to approve
the General Committee recommendation on Acacia, but
rather simply presented this approach as though it were
Bothalia37,l (2007)
115
established procedure (Rijckevorsel 2006). And it is
never too late to point out that the procedure used for the
motion on the General Committee recommendation on
Acacia was invalid.
Conclusions
The procedure used at the Nomenclature Section
meeting at Vienna on the General Committee recom-
mendation on Acacia was not only inappropriate,
unprecedented, and confusing, but conflicted with the
Section’s established rules of procedure (Rijckevorsel
2006). Therefore, the vote by the Section on the General
Committee’s recommendation on Proposal 1584 cannot
be taken as an approval of the proposal. Indeed, based on
the Section’s established rules of procedure, the vote on
the General Committee’s recommendation on Proposal
1584 resulted in its rejection. Nonetheless, Acacia is now
listed in the Vierma Code in Appendix III as a conserved
name with a conserved type (McNeill et al. 2006: 286).
This matter can be raised again at the 18th Inter-
national Botanical Congress at Melbourne in 2011 when
the printed version of the Vienna Code comes up for
ratification. Such challenges to the printed version of the
Code, while rare, are not unprecedented (Stafleu 1966:
6).
Such a challenge to what occurred at Vienna is needed
because of the unfortunate precedent this case will set if
not challenged. Up until the Acacia entry in the Vienna
Code, nothing substantive or nonprovisional had ever
been added to or deleted from the Code without obtain-
ing approval from a Nomenclature Section through
a majority vote. This is a history that spans over 100
years.
To quote from Greuter & McNeill (1994: xiv): ‘...
plant nomenclature is not governed by a bureaucracy of
committees but, in an open and democratic manner, by
the community of its users represented by the enrolled
members of International Botanical Congresses. The
user-driven process by which plant nomenclature is reg-
ulated is of utmost importance for a Code, which, hav-
ing no ‘teeth’ in the way of penalties for infringements,
entirely depends on user consensus for its universal
application and implementation’.
However, if committee recommendations are allowed
to be treated as approved even though a majority of the
votes cast at the Nomenclature Section meeting was
opposed to them, has not plant nomenclature indeed
become a bureaucracy of committees? Incredibly, some
who opposed Proposal 1584 were further told tha't even
if the Section had rejected the General Committee’s rec-
ommendation on Acacia this ‘still would not necessar-
ily have resulted in its absence from the Vienna Code,
in which it could have been included with an asterisk,
pending further discussion at the 2011 International
Botanical Congress’ (Smith et al. 2006: 224, 225).
Apparently, this bureaucracy of committees also ignores
Section decisions that are not to its liking.
The primary reason for Nomenclature Sections requir-
ing a 60% supermajority to amend the Code is to ensure
that there is widespread acceptance of these changes
(Greuter et al. 2000: 14). Treating motions as approved
even though they fail to get majority support is a recipe
for the opposite.
It has been argued that the special procedure used
for the Acacia case was beeause ‘overturning a deci-
sion of the General Committee is clearly a momentous
step and, like an amendment to the Code, is not to be
taken lightly’ (McNeill 2006). However, until the Acacia
debacle, there had never been serious opposition by
a Nomenclature Section to a recommendation by the
General Committee, clear evidence that Section mem-
bers do not take it lightly. The precedent set in oppos-
ing this General Committee recommendation does not
justify the precedent-setting procedure used for han-
dling this recommendation, because there was a long
history of requiring majority votes to approve General
Committee recommendations on proposals to conserve
or reject names. While a proposal’s controversial status
may justify it being handled differently with respect to
a separate consideration and a longer debate, it does not
justify a change in the voting requirement.
It should also be lost on no one that much of the
opposition to the proposal to conserve Acacia largely
came from parts of the world, Africa and South America,
that are under-represented in plant taxonomy in general
and in plant nomenclature specifically. For example, of
the nine officers of the Nomenclature Section at Vienna
(McNeill et al. 2005: 1058), five were from Europe,
three from North America, one from Australia and
none from Africa or South America. At the end of the
Nomenclature Section meeting at the 16th International
Botanical Congress at St. Louis, C. Kabuye from Kenya
noted (Greuter et al. 2000: 239), ‘Several African bota-
nists would have been interested in being present at the
sessions, to learn more about nomenclature through dis-
cussion and debate, but they could not afford to come
on their own’. Nonetheless, some complained about all
of the proxy institutional votes from Africa and South
America that were carried to Vienna, even though this
was wholly within the rules, and, at best, probably
served only to counteract the lack of individual voting
members at the Section from these areas.
Looking to the future, 1 propose the following recom-
mendations that might improve this process:
Recommendations for future Nomenclature Sections
1 . Nomenclature Sections should determine what the vot-
ing procedure will be regarding the approval of General
Committee recommendations to conserve or reject
names. All recommendations of the General Committee
should be subject to the same vote requirement — a
straight up or down vote to approve; all proposals not
receiving a majority vote should be rejected.
Based on the published proceedings, the Nomen-
clature Section at the Seattle Congress in 1969 (Stafleu
& Voss 1972: 4, 5) is apparently the only recent (post
1950) Section to adopt a voting procedure that made a
clear distinction between the rules of the Code and the
other portions of the Code, requiring a 60% majority to
116
Bothalia 37,1 (2007)
amend the rules and a simple majority for proposals to
amend portions other than the articles (Rolla Tryon at
the Edinburgh Congress unsuccessfully proposed that
this procedure be included in the Code; see Stafleu 1966:
8). Subsequent Sections have not been clear as to what
the procedure was when voting on General Committee
recommendations — was a 60% or simple majority
required? In the absence of an explicit rule for voting
on these recommendations, I argue that the procedure
used for amending the Code would apply, since such
proposals, when approved, effect changes to the Code.
Therefore, the proposals to conserve or reject names at
the Nomenclature Section meeting (presented as General
Committee recommendations) at Vienna should have
been subject to a 60% vote to approve since they would
effect changes to the Code.
The second sentence of this recommendation prevents
the voting inconsistency and confusion that occurred at
the Nomenclature Section meeting in Vienna regarding
Proposal 1584. It also ensures that majority rule prevails.
2. A specific recommendation/proposal within a package
of recommendations/proposals must be considered sepa-
rately, if one person in the assembly so moves.
Robert et al. (2000: 265) and Sturgis (2001: 96-100)
state that when ‘a series of independent resolutions
relating to completely different subjects is offered by a
single main motion... any resolution in the series must
be taken up and voted on separately at the request of a
single member’. This allows efficiency when there is
no disagreement, and debate when there is. If a General
Committee recommendation rejects a proposal to con-
serve or reject a name, the original proposal is rejected
and the name should not appear in the Code. If a General
Committee recommendation to conserve or reject is
rejected by the Section, this action does not approve the
original proposal (Robert et al. 2000: 100). A separate
motion would have to be made on the original proposal
and it would be subject to approval by the Section.
3. Rules for voting and membership for the General
Committee should be re-evaluated.
Currently, all members can vote on all propos-
als before the General Committee. All Secretaries of
the nomenclature committees (committees for Algae,
Bryophyta, Fossil Plants, Fungi, and Vascular Plants)
are also ex-officio members of the General Committee.
In addition, some ad hoc members of the General
Committee are also members of a nomenclature com-
mittee. These dual members, therefore, can vote twice
on the same proposal. Procedures should be enacted pre-
venting double voting by these members. The purpose
of the General Committee is to evaluate the recommen-
dations made by the nomenclature committees, and this
should be done with as much independence as possible.
This double voting is tantamount to a judge simultane-
ously sitting on a lower court and a supreme court and
ruling twice on the same case, and it conveys an image
that the General Committee, at least in part, is simply a
rubber stamp of the nomenclature committees. Perhaps
those who are already members of a nomenclature com-
mittee should not serve on the General Committee.
Secretaries of the other committees need to be members
to explain specific cases before the General Committee,
but perhaps they should not vote on cases in which they
have already voted as a member of the other committee.
It is up to the General Committee to determine how it
will evaluate the nomenclature committee recommenda-
tions. For example, should the General Committee con-
sider the merits of the case or should it limit its evalua-
tion to technical aspects, such as making sure a nomen-
clature committee interpreted the rules of the Code cor-
rectly? However, members of the General Committee
should never vote to recommend a proposal simply so it
can be sent to the Nomenclature Section for action. The
Nomenclature Section is omnipotent with respect to the
Code and is free to consider any topic with or without
the General Committee’s recommendation.
4. After the General Committee receives recommen-
dations on proposals to conserve or reject names by a
nomenclature committee, the General Committee should
act on these recommendations within a reasonable time
frame. General Committee recommendations should be
published in advance of the Nomenclature Section meet-
ing.
Stafleu (1964: 6) suggested that the General
Committee should act within three months of publica-
tion of the nomenclature committee recommendation.
This seems like a reasonable time frame. A General
Committee report needs to be published before the
Congress (no such report was published prior to the
Vienna Congress), so it is clear which proposals will
be up for approval at the Nomenclature Section meet-
ing. While it may be impossible to publish the recom-
mendations made on all proposals, ‘last minute’ General
Committee recommendations that go unpublished prior
to the Nomenclature Section meeting in which they are
acted on, should be kept to a minimum.
5. Consideration should be given to expanding the con-
tent in Division III of the Code to clarify the charge and
relative roles of the General Committee and the nomen-
clature committees that report to it.
While the Code spells out the charge of the Editorial
Committee, it says little regarding the roles of the
nomenclature committees. Language should be added
here explaining the general roles of these committees.
Any language would have to be as general as possible,
so as not to hamstring future congresses’ ability to assign
these committees with new duties.
Acknowledgements
The following are thanked for reviewing drafts of
the manuscript: M. Alford, M. Luckow, L. Pedley, C.
Rushworth, B. Schrire, P. van Rijckevorsel, G. Smith, A.
van Wyk and three anonymous reviewers. B. Momberg
and G. Gennishuizen are thanked for assistance with
editing the manuscript. J. McNeill is thanked for his
extensive discussions with me on this matter even
though we disagree. Parliamentarians, N. Sylvester and
L. Wynn, are thanked for providing valuable assistance
regarding procedure and reviewing the manuscript.
Bothalia37,l (2007)
117
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Bothalia37,l: 119-127(2007)
OBITUARY
ELSIE ELIZABETH ESTERHUYSEN (1912-2006)
On Tuesday, 9th June 2006, some 200 persons gath-
ered in the conference room at the Kirstenbosch Research
Centre of the South African National Biodiversity
Institute in the Kirstenbosch National Botanical Garden,
Cape Town, to pay homage to and celebrate the life of
the late Elsie Esterhuysen. The memorial gathering was
hosted and conducted by her nephew, Allan Lighten
and his wife, Megan (nee Esterhuysen), and comprised
three speeches of tribute from the botanical fraternity
and one from the Mountain Club of South Africa on
her mountaineering prowess, followed by refreshments,
with many personal reminiscences among those attend-
ing. The following are the contributions from the three
botanists outlining her life and work and paying tribute
to her wonderful contributions to botany, with personal
reminiscences and anecdotes.
John Rourke
‘Today we are here to celebrate the life of a legend,
for Elsie was a legend, certainly within the local botani-
cal community (Figure 1). She was bom 94 years ago
in Observatory, Cape Town, on 11 April 1912. Elsie
attended the Wynberg Girls High School and after
matriculating, proceeded to the University of Cape Town
where she graduated with an M.A. degree in botany in
1933, the subject of her thesis being the anatomy of the
resurrection plant, Myrothamnus flabellifoUus Welw. But
anatomy and physiology held little attraction for her —
right from those early days it was clear that field work
was to be her abiding interest. After a brief period of
employment as a clerk in the Education Department, her
chance came with the award of the Solly Scholarship at
Kirstenbosch in 1935. There she made a detailed study
of the fynbos regeneration after the felling of a 50 year-
old plantation of Pimis pinaster at the northern end of
the Kirstenbosch estate — a mere few hundred metres
from where we are gathered this afternoon. The results
of this study form the basis of the few research papers
she ever published.
Elsie had long wanted to join Dr LB. Pole Evans’ Bota-
nical Survey group in the Department of Agriculture, but
the Pretoria bureaucrats turned her down merely because
she was a woman. In those days the very prospect of a
female doing botanical survey work in the remoter parts
of the South African bush was unthinkable! One of her
classmates at university, John Acocks, was accepted
by Pole Evans and went on to produce his famous Veld
Types of South Africa. With the Great Depression at its
peak, jobs were hard to come by and so in 1936 Elsie
gratefully accepted a position as assistant to the redoubt-
able Maria Wilman, Director of the McGregor Museum,
Kimberley, who was then undertaking a survey of the
flora of Griqualand West. Miss Wilman insisted that
Elsie should learn to drive in order to reach the more
distant parts of the survey area. All went well until one
day Elsie stalled the car on a level crossing. Despite the
fact that there was no belching locomotive thundering
down the line towards her, this incident proved to be a
life-altering experience for her. She never drove again!
Instead she took to pedal power and the bicycle, perfect-
ing her cycling technique on a variety of models over
the years. Who will ever forget the spectacle of Elsie’s
FIGURE lA-C. — Elsie Esterhuysen (dates unknown; photographs courtesy of Bolus Herbarium).
120
Bothalia37,l (2007)
daily homeward journey from the Bolus Herbarium on
her trusty bike, head forwards, hair streaming behind
her, careering down the viciously steep Stanley Road
at breakneck speed, then whipping into Lovers Walk,
followed immediately by another swooping twist into
Grotto Road — all with the panache of a Tour de France
professional — before coming to a heart-stopping halt in
Main Road, Rondebosch.
After two years in Kimberley, Elsie returned to
Cape Town in 1938. It was here that her real career
began when she joined the Bolus Herbarium (Figure
2) under another formidable woman. Dr Louisa Bolus.
Her former professors, Robert Adamson and Harold
Compton, knowing of her interest in mountaineering,
suggested that she explore and document the high-alti-
tude flora of the Cape Mountains as herbarium records
from such areas were at that stage sparse or even lack-
ing completely. This idea appealed immensely to Elsie
and became the focus of her life’s work for the next 60
years, during which time she amassed over 37 000 her-
barium collections. What makes this achievement even
more remarkable is that having no transport of her own,
she relied entirely on Mountain Club friends to provide
lifts to the various sites she investigated, be they in the
Western Cape mountains or the KwaZulu-Natal (KZN)
Drakensberg.
It’s an astonishing fact that for the first 18 years of
her employment she received no proper salary and was
FIGURE 2. — Elsie Estcrhuysen, taken in the Bolus Herbarium in 1962
(photograph by ,I.R .lessop).
paid out of petty cash at a rate not much better than a
labourer. When Prof. Ted Schelpe arrived as curator of
the Bolus Herbarium in 1956, he had to do battle with
the bureaucrats in the University of Cape Town’s admin-
istrative office to get a permanent establishment post
created for her. It was the first time she had any financial
security. Such was the extent of her loyalty and dedica-
tion.
She did not collect randomly; Elsie was above all
an intelligent collector, seeking range extensions, local
variants, or even new species, filling voids in the Bolus
Herbarium’s records, often returning months later to col-
lect seeds or fruits that were of diagnostic importance.
To do this effectively pre-supposed an encyclopedic
knowledge of the published literature on the Cape flora
and this she possessed to a remarkable degree. Despite
being responsible for this avalanche of botanical dis-
coveries, sustained over several decades, Elsie could not
be persuaded to publish the results of her work under
her own name. Instead she delighted in helping others;
indeed Elsie seemed to take vicarious pleasure in seeing
her own discoveries written up in a host of monographs
by several generations of young taxonomists. Was it lazi-
ness, a fear of criticism or modesty that caused this curi-
ous reluctance to publish? Knowing Elsie, I feel sure it
was her innate modesty. Always self deprecating, one of
her favourite comments was T’m only filling in gaps’.
As postgraduate botany students at the University
of Cape Town working in the Bolus Herbarium during
the 1960s, many of us had the privilege of engaging in
regular lunch-time discussions with her on a great many
aspects of the Cape flora, while eating our sandwiches
perched on the rather decrepit old sofa that graced the
tearoom of the Bolus Herbarium. Though never part of
the formal teaching staff, her influence was incalculable.
But it was not only local botanists whom she helped.
Eminent specialists from abroad all beat a path to her
door to seek her expertise and assistance in the field —
from Switzerland, Sweden, Denmark, Germany and the
USA, among others. It was a well-known fact interna-
tionally that if you were coming to South Africa to work
on the taxonomy of any plant group in the Cape flora,
the first person you had to see was Elsie Esterhuysen.
Although she published very little herself, the South
African Association of Botanists recognized the huge
contribution she had made in assisting others by award-
ing her their Senior Medal in 1987. Her old university
also acknowledged her achievements. On the occasion
of the University of Cape Town’s confennent of an hon-
orary Master of Science degree on Elsie in 1989 (Figure
3), the university orator referred to her as a ‘botanical
hunter gatherer’. The metaphor was very apt. Like a
hunter gatherer, she was in perfect harmony with nature,
never being wasteful, never taking more than was needed
at the time, never defacing or contaminating the environ-
ment. She had a deep and profound respect for nature. It
was almost like a religion to her.
Elsie was a person of huge wannth and generosity.
She cared deeply for the oppressed, the downtrodden
and the underdog. Tramps and vagrants who knocked at
her door were often sent away in astonishment with a fist
Bothalia37,l (2007)
121
FIGURE 3. — Elsie Esterhuysen
with the Chancellor of the
University of Cape Town,
Harry Oppenheimer, after
her graduation in 1989 (pho-
tograph courtesy of Bolus
Herbarium).
full of cash in their hands (but sadly, in her late eight-
ies, she was attacked by one of them when he became
greedy). We all know how fimgal she was. Who else
could survive for days in the mountains on half a raw
cabbage, a few carrots, some powdered milk and weak
tea? But on occasions she could let her hair down. I well
remember offering to get her a drink at a book launch
held at Kirstenbosch. A double brandy and ginger ale
was requested and consumed with gusto! She was one of
the most broad-minded and tolerant people I have ever
met, yet there was one human folly of which she was
extremely intolerant and that was smoking. Cigarette
smoke was anathema to Elsie. How she coped with the
chain-smoking of her boss I do not know. There was a
memorable incident when an eminent but chain-smok-
ing professor of biochemistry from Berlin ignored her
pleas to desist from smoking on a field trip. Elsie calmly
reached up, tore the cigarette from his lips, hurled it to
the ground and crushed it underfoot! Needless to say, he
did not light up again in her presence.
Today Elsie’s botanical legacy lies within the fold-
ers of the Bolus Herbarium — a superb botanical archive,
some of it already cited in major taxonomic publica-
tions by a host of local and international specialists. But
many of her collections remain to be worked on like a
huge untapped data base waiting to be sifted through
and mined by future botanists in decades to come. And
so Elsie’s name will live on as later generations examine
and dissect the specimens she gathered over a lifetime
of prodigious activity. They will marvel at her achieve-
ments rather as we today marvel at the work of 1 8th and
19th century explorers such as Thunberg and Burchell.
Elsie was unique in the truest sense of that overworked
word. We will not see her like again.’
Peter Linder, Zurich, Switzerland (read by E.G.H.
Oliver)
‘When I first started working in the Bolus Herbarium
in 1973, as an undergraduate student wanting to know
the names of the plants on the Piketberg, it was a curi-
ous place. There was a wonderful chaos of specimens,
stacked on every available surface. On two floors was a
superb library of the most magnificent books. Down in
the tearoom was Ted Schelpe, twirling his moustache
and smoking endless Mills cigarettes, a source of many
ideas and inspiration. Tony Hall was in an office on the
top floor, from where he organized a large number of
conservation initiatives. Elsie was on the middle floor,
and she was what I thought a botanist was supposed to
be. She was in the mountains every weekend, and came
back with big black plastic bags full of plants, that she
sorted and passed to Gert Syster to press. She was the
one person who could put names on plants that defeated
my attempts. And she had little time for academic nice-
ties— for her the important things were plants in the
mountains, their welfare, their relationships. She was
immersed in plants and mountains.
However, I only started to know her somewhat bet-
ter when I worked on restios — although she knew the
orchids, these were not a particular theme to her. But
the restios had fascinated her since before the war. After
Pillans had died (in 1964), she had taken on the mantle
of the restio specialist. The really close collaboration
only started after I had left for Kew as South African
Botanical Liaison Officer — we had an intense corre-
spondence about the species limits and the new generic
limits I was proposing. When I left she had said that
she does not write letters — well, I have a huge pile of
her correspondence: detailed comments about each spe-
cies, discussions about the best Latin for the diagnoses,
discussions about the characters for the genera and the
groupings that would be best to use. Her letters were a
delight and a constant inspiration for me. She had built
up a formidable knowledge about each species, knew its
ranges, knew when it flov/ered. She would wait, some-
times for many years, until a mountain club trip took her
back to where they occurred to collect seed, to extend
the distribution ranges, to document the flowering times.
Whereas Ted Schelpe taught me how to write descrip-
122
Bothalia37,l (2007)
tions and deal with nomenclature, Elsie taught me that
each species has an essence, a character — that it liked
some habitats but not others and that it flowered at a
particular time. She was curious about the plants, not
because they informed her about some theory or other,
but she was interested in the plants themselves — she
cared about them.
Elsie lived in a suburban house in Rondebosch, on
a bleached and leached sandy soil. For many years she
let most of the house, and lived in the servant’s quarters.
But the garden at the back was hers, and her cats’. It was
full of remarkable bulbs that she had brought back from
all over the Cape, and was delightfully overgrown. It
was a pleasure to sit in the shade in the unmowed lawn,
amongst a remarkable collection of species, drinking tea
and talking about mountains and plants. Rumour had it
that she also bathed in a tub in the back garden.
She also went to the mountains in the same way — not
to climb them per se, but to be in them. She did not need
clothing and equipment to keep nature out. She was sim-
ply at home in nature. The only natural element that she
could not relax in was the sun.
The older of us probably all still remember Elsie in
her working bay in the Bolus, immediately to the left
on the first floor. She had a large table laden with speci-
mens, comments on fading paper written in a fast, almost
abbreviated handwriting, and tattered paper-covered
revisions and books lying about. Under her table was a
wooden footrest, in the tearoom her unusually low chair.
She talked about her cats, and had a dry sense of humour.
When Schelpe tested a prospective Ph.D. student by ask-
ing her what the differences between Selaginellaceae
and Scrophulariaceae were, and then gleefully explained
when the poor student did not know, Elsie remarked
sotto voce that she, also, did not know what the differ-
ence was. Then there was the poor Austrian botanist that
could not keep up with her in the mountains, when, after
all, he came from the Alps. Or when I was rather nerv-
ous scrambling over an exposed cliff at the top of Ferny
Gully and she said — well, you do have children to worry
about.
1 asked her once how she worked; my curiosity was
aroused when 1 started a revision of Pentaschistis and
found all specimens neatly identified in the herbarium,
complete with the new species sorted out correctly at
the end of the genus. She told me that she would col-
lect all plants that she could not identify. This way she
found genera in which the species taxonomy was inad-
equate. Then she would focus on a genus for several
years, curating the old collections and attempting to
name her new collections. When she found what she
thought might be new species, she would start searching
for more localities, in this way mapping the distribution
ranges, flowering times, and ecology of each new spe-
cies. By the time she was finished, the taxonomy was
sorted out, the new species well documented, and all
mounted and laid in the herbarium. In the case of the res-
tios this even meant that there were little capsules with
seeds, carefully collected in the field. It was left for the
‘academics’ to write the descriptions and the keys. This
was the situation in Restionaceae and Pentaschistis, and
I would love to know whether it was also the situation in
other genera, such as Agathosma and Phylica. And was
she intrigued, or amused, by the antics of the academics?
Or did she simply prefer fieldwork to writing, and fight-
ing with reviewers?
She was, however, at times concerned about the
recognition that she got for her work. She felt that the
importance of her work was not always recognized by
the University authorities. It was therefore really good
that Hugh Amore took the initiative to convince the
University to grant her an M.Sc. honoris causa. Hugh
was at the time the Registrar of the University, a difficult
and demanding task, he had enough to worry about. So
much more to his honour, and to the fame of Elsie, that
he took it upon himself to organize and co-ordinate this
degree (see additions at end).
Several people had an important influence on Elsie. I
do not know where her love for the plants came from —
her origins were always a mystery to me. I never heard
her talk Afrikaans, yet Esterhuysen is a good Afrikaans
surname. Her focus on the montane plants was appar-
ently started by a comment from Adamson, who wanted
her to document the mountain summit floras. And the
late interest in lowland floras I suspect came from
Richard Cowling. Elsie once commented to me that she
should have concentrated more on the lowland flora, as
it was much more threatened than the montane flora.
And how many people did Elsie influence? That is
hard to know. She had a subtle, but very powerful, influ-
FIGURE 4. — Elsie Esterhuysen and Prof. Peter Linder after the gradua-
tion in 1989 (photograph G, Scott, courtesy of Bolus Herbarium).
Bothalia 37,1 (2007)
123
ence on me (Figure 4). And I suspect that many of her
fellow mountaineers, and also many of the botanists that
passed through the Bolus, were deeply affected by this
tiny, remarkable woman, who was so much at one with
nature.’
Ted Oliver
‘We are here this afternoon for a very special pur-
pose— to pay tribute to an extraordinary person who in
some way, small or large, came into our lives. Death,
memorial services, funerals are usually solemn occa-
sions filled with sadness, sometimes tears. But don’t let
us go that route here. Let us rather celebrate a wonderful
life that has finally come to an end. Yes, it is sad that we
have lost Elsie, but then at 94 she did live well beyond
her three score years and ten. In the end, through loss
of memory, she was a burden unto herself and her death
was indeed a happy release for her on 3 1 May 2006.
I first got to know Elsie in 1958, and like Peter, as
an undergraduate wanting to know something about the
Cape flora. At that time I also met her because she lived
in a flat above a double garage along the road from my
family home in Rondebosch (the house of Brian and
Joan Quail of the Mountain Club). Just behind our house
was a large residence for old people. The Abbotsleigh,
where Elsie’s mother had moved in. And so Elsie often
came past our house to visit her mom.
In the Bolus Herbarium I was keen to learn about eri-
cas which I had seen on our family holidays near Betty’s
Bay — who better than Elsie to help me. She had concen-
trated on Erica collecting at the insistence of Mrs Bolus
who had helped her uncle Harry Bolus write them up
for Flora capensis (1896-1905). She was very generous
with her encouragement and guidance for this youngster
who was now encroaching on her territory, but never
once relayed any thoughts of this to me. Thanks very
much Elsie.
The first of Elsie’s few publications was the descrip-
tion of a new species. Erica viscidiflora in 1936, and then
E. cremnophila in 1940 with Capt. Salter. Then came Dr
Hans Dulfer, in Vierma, with his revision of Erica in the
early 1960s. Elsie sent him a few batches of some of her
problem material to show him some of the difficulties
in the genus, whereupon he promptly thanked her, and
described them all as new species. This put the wind up
Elsie’s sails and she decided to send no more material
and promptly wrote a paper on her views on certain spe-
cies just before his revision came out. This was her only
major publication as sole author. Two things must have
galled her (from the grunts one picked up) — Dulfer put
the holotypes of her material in Vienna and he sank her
E. viscidifloral One felt from her veiled comments that
this must have stifled any further ideas of publications.
Little did she know that her viscidiflora is a good species
that my late wife and I have reinstated. Later, of course,
through Peter’s research, she was encouraged to publish
in his papers (Esterhuysen in Linder 1985, 1990) all her
new species of Restionaceae that she had so assiduously
been collecting over the years — some 47 species.
We have heard that Elsie concentrated her efforts on
collecting, but not only in the Cape Florsitic Region
(CFR), she did much in the KZN Drakensberg mainly
on the annual July camps of the Mountain Club (Figure
5). One of her early mentors was Dr Henri Georges
Fourcade with whom she worked and collected for his
FIGURE 5. — Elsie Esterhuysen at the
Mountain Club’s Drakensberg
July Camp of 1987 where she
is being made ‘a free woman’
on receiving the Freedom of
July Camps from the chair-
man of the local branch. Dr
Graham Smith (photograph
by B. Cooke).
124
Bothalia37,l (2007)
FIGURE 6. — Elsie Esterhuysen with
Ted Oliver and the type plants
of Erica atrovinosa in the
Hex River Mtns in 1964; she
discovered it in 1942 (photo-
graph M.F. Thompson).
work on the flora of the Knysna/Uniondale area. As
a collector she was certainly the most prolific for the
CFR and ranks among the top three in South Africa with
Compton and Acocks, all with just short of 40 000 col-
lections.
But how did she manage this? John has mentioned her
lack of enthusiasm for cars, although she did once much
later say that if Mrs Bolus in her late 80s still drove her
tiny black Austin, perhaps she should take up driving
again! So how did she get to all those remote places. The
answer — the Mountain Club as John mentioned. Because
the MCSA members did not roam around the flats, this is
why I suspect she tackled them only later in life as Peter
mentioned. She joined the club in 1935!
But her mode of transport was the bicycle (we have
her latest model here today). She rode to UCT up that
dreadful steep road every day for a lifetime, come sun-
shine or rain, heat or cold. Now one knows why she was
so fit and could outstrip any poor unsuspecting younger
botanist in the mountains! Every day she would come
up and park her bicycle behind the Bolus Herbarium
building and then often jump through the window in the
preparation section rather than walk all the way around
to the front door. One day I happened to be looking out
of the window and saw Elsie leaping onto her bicy-
cle to go home, and lo and behold, a bit further down
the University Ave between the Physics Dept and the
Science Lecture Theatre, she actually fell off it in front
of a group of students! The next day she told me she had
never been so embarrassed.
Peter noted her quiet presence in the comer of the tea-
room on that extraordinarily low chair. There she would
sit with her oversized enamelled mug full to the brim
with skimmed milk and a touch of tea. One hardly saw
her with anything else, perhaps an orange or an apple,
even on trips into the mountains. It is amazing how she
managed to climb so many mountains and live so long
on powdered skim milk. But she did have one serious
health problem — her very thin skin which was always
getting scratched and infected in older life.
Conversation at tea time was very varied with only
the occasional comment from Elsie who spent most of
the time endlessly stirring and sipping her milky tea. But
things changed for her when something political was
mentioned (yes Elsie was quite a liberal) or if someone
mentioned the word CAT. Elsie loved her cats — some-
times too much. She was known to buy choice meat for
the cats rather than herself; and to give up her bed and
sleep on the floor rather than disturb the sleeping cats on
her bed at bedtime!
I had wonderful trips with Elsie in those early days
(Figures 6, 7) and later in the 1980s with Dolf Schu-
mann— Hex River Mountains, Kogelberg and Swartberg.
She always slept apart from us, never on a mattress, pre-
ferring to make a nice nest out of restios and slangbos,
or, in the Drakensberg, lots of grass (Figure 8). Peter
FIGURE 7. — Elsie Esterhuysen with the orchid, Disa longicormi, on
Table Mountain (date unknown, photograph by M. Burger).
Bothalia37,l (2007)
125
FIGURE 8. — Elsie Esterhuysen in the Drakensberg with a load of grass
bedding for the night in July 1982 (photograph courtesy of Bolus
Herbarium
has alluded to her behaviour in nature — with mumbled
apologies she was not averse to shedding everything
and hopping into a little stream to cool off! One of
her greatest experiences was coming along with Dolf
Schumann and myself in 1990 from Cape Town on a
helicopter trip which Dolf organized to photograph the
rare Erica alfredii in the Riviersonderend Mountains. On
the return flight I persuaded the pilot to go low along a
very devious route through the mountains via Jonaskop,
Kaaimansgat, Goudini Sneeukop and Victoria Peak. She
was amazed at seeing from the air all those mountains
that she knew personally on the ground.
I shall cherish the moments I had with Elsie in the
herbarium and in the mountains, I am only sorry that I
was away from the Bolus working at Kew, Pretoria and
Stellenbosch. Elsie was not a religious person but she
must definitely have communed with God and nature
in her own way in her countless days and nights in the
mountains.
Let us give thanks for Elsie’s life — for her great con-
tribution to the study of our unique flora, through her
superb collections that will last for as long as man con-
tinues to study plants from the Cape, and, hopefully,
maintain herbaria. She will be remembered in two gen-
era and about 56 species with Erica having three of them
(Table 1).
When any of you next go up into the mountains, know
that Elsie’s spirit will be around somewhere in her para-
dise. Perhaps you will feel her presence in those large
restiad clumps.
Thank you Elsie — you were a great person, mentor
and friend. ’
Note added in the preparation of this obituary
Elsie’s parents were Johannes Petrus le Roux
Esterhuysen (Afrikaans speaking) and her mother,
Florence Ethel Larkin (English speaking) — hence her
mother tongue. Her father attended Victoria College,
Stellenbosch, where he studied law and then worked in
the master’s office of the Supreme Court in Cape Town.
He was transferred from Cape Town to Windhoek where
Elsie visited her parents by train during holidays. He
retired early due to Florence not liking the Windhoek
climate, and bought a smallholding at De Hoek, outside
Joubertina in the southern Cape, where his son, Cyril,
was the local attorney. Elsie visited the family there dur-
ing her early botanical years at the Bolus Herbarium,
hence her contact and friendship with Dr Fourcade who
lived across the Tzitzikama Mountains at Witelsbosch.
A newspaper cutting found among her personal
effects after her death showed a side of Elsie that none of
us knew existed. It was from the local Kimberley news-
paper (undated) and reported on a Reading on life and
works of Franz Schubert at a meeting of the Kimberley
Philharmonic Society. The lecture was given by Miss
E. Esterhuysen. It was preceded by another describ-
ing BurchelTs travels in South Africa during Schubert’s
time and was followed by ‘a delightful short programme
of the composer’s music played as piano solos by Miss
Esterhuysen’.
Elsie was also an accomplished artist as evidenced
by the few charming watercolours of flowers that were
among her effects. She illustrated with black and white
line drawings the species of Erica which she described
with Salter.
Elsie celebrated her 80th birthday by climbing the
2 000 m high Sneeuberg in the Cederberg Wilderness
area with friends from the Mountain Club.
Also in her effects was a handwritten letter to the
principal of the university about her honorary degree and
this shows something of her character:
‘Dear Prof Saunders
Thank you for your kind letter of 9th Jan. I accept the award of the
honorary degree and appreciate the honour.
I wonder, though, whether the staff of the Botany Dept were con-
sulted. If the matter were not confidential I should like to consult them
myself because it does not seem to me that my work warrants the hon.
degree. I hope Council were not misled into supposing that the discov-
ery of undescribed species was a great achievement because it wasn’t.
Apart from that I worked as a technician keeping the herbarium in
order and up to date.
Yours sincerely
Elsie Esterhuysen’
126
Bothalia37,l (2007)
TABLE 1 . — Plants named after Elsie Esterhuysen. Accepted names in roman t5^pe and synonyms in italics
Direct names
Bryophyta
Lophocolea eslerhuysenii(ae) S.W.Amell (1963) [= L. bewsii (Sim)
Grolle]
Plagiochila esterhuysenii(ae) S.W.Amell (1963) [= P. squamulosa Mitt.
var. crispulo-caudata (Gottsche) Vander Berghen]
Pteridophyta
Dryopteris esterhuyseniae Schelpe & N.C. Anthony ( 1 982) [= Dryopteris
dracomontana Schelpe & N.C. Anthony]
Apiaceae
Chamarea esterhuyseniae B.L.Burtt (1991)
Asphodelaceae
Bulbine esterhuyseniae Baijnath (1987)
Trachyandra esterhuysenae Oberm. (1962)
Asteraceae
Anderbergia elsiae B.Nord. (1996)
Athanasia elsiae Kdllersjo (1991)
Chrysocoma esterhuyseniae Ehr.Bayer (1981)
Corymbium elsiae Weitz (1989)
Felicia esterhuyseniae Grau (1973)
Hydroidea elsiae (Hilliard) P.O.Karis (1990) [= Atrichantha elsiae
Hilliard (1981)]
Troglophyton elsiae Hilliard (19^3)
Brassicaceae
Heliophila esterhuyseniae Marais (1966)
Bruniaceae
Lonchostoma esterhuyseniae Strid (1968)
Tittmannia esterhuyseniae Powrie (1969)
Chenopodiaceae
Salsola esterhuyseniae Botsch. (1973)
Cyperaceae
Ficinia esterhuyseniae Muasya (2005)
Droseraceae
Drosera esterhuyseniae (T.M.Salter) PDebbert (1992) [= D. curviscapa
T.M. Salter var. esterhuyseniae T.M.Salter (1939)], [D. aliciae
Raym.-Hamet]
Ericaceae
Erica
elsieana (E.G.H.Oliv.) E.G.H.Oliv. (1987) [= Philippia elsieana
E.G.H.OIiv. (1984)]
esterhuyseniae Compton ( 1941 )
esteriana E.G.H. Oliv. ( 1 987) [= Philippia esterhuyseniae E.G.H.Oliv.
(1984)]
Fabaceae
Amphithalea esterhuyseniae (Granby) A.L.Sclmtte (1998) [= Coelidiwn
esterhuyseniae R.Granby ( 1 980)]
Aspalathus esterhuyseniae R.Dahlgren (1960)
Lotononis esterhuyseniana B-E.van Wyk ( 1 990)
Hyacinthaceae
Elsiea F.M. Height. (1944) see Bothalia 12: 323 [= Omithogalum
paludosum Baker ( 1 874)]
Lachenalia esterhuyseniae W.F.Barker (1978)
Omithogalum esterhuyseniae Oberm. (1978)
Iridaceae
Geissorhiza
elsiae Goldblatt (1985)
esterhuyseniae Goldblatt (\9S5)
Hesperantha elsiae Goldblatt ( 1 984)
Ixia esterhuyseniae M.P.de Vos (1988)
Moraea elsiae Goldblatt ( 1 976)
Romulea atrandra G.J. Lewis var. esterhuyseniae M.P.de Vos (1972)
Watsonia elsiae Goldblatt ( 1 989)
Linaceae
Linum esterhuyseniae C.M.Rogers (1982)
Lobeliaceae
Lobelia esterhuyseniae E.Wimm. (1968) [= L. comptonii E.Wimm
(1968)]
Mesembryanthemaceae
Aridaria esterhuyseniae L.Bolus (1939) [= A. noctiflora (L.) Schwantes
var. straminea (Haw.) Gerbaulet ( 1 996)]
Delosperma esterhuyseniae L.Bolus (1959)
Erepsia esterhuyseniae L.Bolus (1939)
Esterhuysenia L.Bolus (1967)
Gibbaeum esterhuyseniae L.Bolus (1958)
Lampranthus esterhuyseniae L.Bolus (1958)
Ruschia esterhuyseniae L.Bolus (1950)
Orchidaceae
Disa esterhuyseniae Schelpe ex H.P.Linder (1981)
Polygalaceae
Muraltia
elsieaePmVa (1981)
esterhuyseniae Levyns (1949), in obs., sine descr. lat. (IK)
Proteaceae
Paranomus esterhuyseniae Levyns (1970)
Restionaceae
Askidiosperma esterhuyseniae (Pillans) H.P.Linder (1985) [=
Chondropetalum esterhuyseniae Pillans (1945)]
Calopsis esterhuyseniae (Pillans) H.P.Linder (1985) [= Leptocarpus
esterhuyseniae Pillans (1945)]
Elegia esterhuyseniae Pillans (1945)
Ischyrolepis esterhuyseniae (Pillans) H.P.Linder (1985) [= Restio
esterhuyseniae Pillans (1945)]
Ceratocaryum fimbriatum (Kunth) H.P.Linder [= Willdenowia ester-
huyseniae Pillans (1945)]
Rosaceae
ClifFortia esterhuyseniae Weim. (1948)
Rubiaceae
Anthospermum esterhuysenianum Pujf{\9%6)
Rutaceae
Agathosma esterhuyseniae Pillans (1950)
Euchaetis
elsieae /. Williams ( 1 979)
esterhuyseniae /. Williams (1979)
Scrophulariaceae
Diascia esterhuyseniae K.E. Steiner ( 1 999)
Selago
elsiae Hilliard ( 1 999)
esterhuyseniae Hilliard (1998)
Trieenea elsiae Hilliard (\9S9)
Thymelaeaceae
Passerina esterhuyseniae Bredenk. &A.E.van Wyk (2003)
Names in allusion
Asteraceae
Monticapra (Karis) Koekemoer (2001). .[= Disparage section
Monticapra Karis (1993)]: mons, mentis, a mountain, and capra, a
she-goat; hence like a mountain she-goat (see below)
Ericaceae
Erica oreotragus E.G.H.Oliv. & I.M.Oliv. (2001) [= E. esterhuyseniae
Compton var. trimera Compton (1941)]: Oreotragus oreotragus,
the klipspringer antelope; referring to Elsie’s adept way of hopping
around the mountains like a free klipspringer in search of choice
plants! (see above)
Scrophulariaceae
Trieenea Hilliard (1989): Tri-eenea, three e’s for E.E. Esterhuysen
(Trieenea elsiae).
Bothalia37,l (2007)
127
PUBLICATIONS BY ELSIE ESTERHUYSEN
ESTERHUYSEN, E.E. 1936a, Plantae novae africanae. Series VI.
Erica viscidiflora Esterhuysen. Journal of South African Botany
2: 63, 64.
- 1936b. Regeneration after clearing at Kirstenbosch. Journal of South
African Botany 2\ 177-185.
- 1943. Coleonema nubigemim Esterh. Journal of South African Botany
9: 137.
- 1950. Rutaceae: Coleonema, Diosma, Adenandra, Macrostylus &
Barosma. In R.S. Adamson & T.M. Salter, Flora of the Cape
Peninsula. Juta, Cape Town.
- 1963. Notes on South Afiican species of Erica. Journal of South
African Botany 29: 51—58.
- 1985. New species in Askidiosperma, Calopsis, Chondropetalum,
Elegia, Hypodiscus, Ischyrolepis, Nevillea, Platycaidos, Restio,
Staberoha and Thamnochortus. In H.P, Linder. Conspectus of
the African species of Restionaceae. Bothalia 15: 396, 420, 421,
423, 428. 432, 434, 459^62, 465, 466, 472, All, 489.
- 1990. New species in Anthochortus, Calopsis and Ischyrolepis. In
H.P. Linder, New species of African Restionaceae. South African
Journal of Botany 56: 454, 456.
ESTERHUYSEN, E.E. & SALTER, T.M. 1940. Plantae novae afri-
canae. Series XIII. Erica cremnophila Esterh. & T.M. Salter.
Journal of South African Botany 6: 1,2.
- 1941. Plantae novae africanae. Series XVI. Erica calycina L. var.
longibracteata Esterh. & T.M. Salter. Journal of South African
Botany 1: 82, 83.
- 1950. Erica calycina var. longebracteata Esterh. & T.M. Salter. In R.S.
Adamson & T.M. Salter, Flora of the Cape Peninsula: 651 . Juta,
Cape Town.
E.G.H. OLIVER* (Compiler), J.P. ROURKE** and H.P. LINDER***
* Formerly: Compton Herbarium. South African National Biodiversity
Institute. Present address: Dept of Botany & Zoology, University of
Stellenbosch, Private Bag XI, 7602 Matieland, South Africa. Email:
eoliver@sun.ac.za.
** Compton Herbarium, South African National Biodiversity Institute,
Private Bag X7, 7735 Cape Town,
*** Institute for Systematic Botany, University of Zurich, Switzerland.
Bothalia37,l: 129, 130 (2007)
Book Review
CHECK-LIST DES PLANTES VASCULAIRES DU GABON/
CHECKLISTOFGABONESEVASCULARPLANTS(&rz>to5oto«/ca
Belgica 35) by M.S.M. SOSEF, J.J. WIERINGA, C.C.H. JONGKIND,
G. ACHOUNDONG, Y. AZIZET ISSEMBE, D. BEDIGIAN, R.G.
VAN DEN BERG, F.J. BRETELER, M. CHEEK, J. DEGREEF,
R.B. FADEN, P. GOLDBLATT, L.J.G. VAN DER MAESEN, L.
NGOK BANAK, R. NIANGADOUMA, T. NZABI, B. NZIENGUI,
Z.S. ROGERS, T. STEVART, J.L.C.H. VAN VALKENBURG, G.
WALTERS & J.J.F.E. DE WILDE. 2006. National Botanic Gardens
of Belgium. Meise. Pp. 438. Soft cover: ISBN 90-72619-69-2; ISSN
0779-2387, price € 53.00.
More information is available on: http://www.br.fgov.be
Without up-to-date information on biodiversity, no area can be
properly managed or conserved. Therefore, the compilation of a species
inventory or checklist should be the first step towards conservation.
However, despite this obvious value, very few countries maintain com-
plete checklists of their biodiversity, especially in Africa. Less than half
of the sub-Saharan African countries have country-level checklists or
inventories of their plants, most of them produced more than ten years
ago. Gabon is one of the few African countries that can boast a recent
vascular plant checklist. It is the result of years of botanical exploration
and research in Gabon. The publication was made possible by the col-
laborative effort of eight institutions and 22 authors.
The bilingual (French and English) introduction is very concise
(ten pages and nine figures), but provides valuable information about
the checklist and the Plants of Gabon Project. It briefly introduces
the reader to Gabon, a relatively small country on the equatorial west
coast of Africa. The country has an extremely rich flora and its lowland
rainforests (covering around 80% of the country) are claimed to be
amongst the richest in Africa. Owing to the selective nature of logging
in Gabon, which concentrates on one near-endemic but a common
pioneer species that regenerates rapidly, the Gabonese forests have
escaped the immense destruction of rainforests typically found in other
tropical parts of Africa and the world. The country also has a network of
13 National Parks, covering 10% of the country’s surface. The impor-
tance of inventories or checklists in the management and conservation
of these parks is highlighted. Furthermore, the general introduction is
well illustrated with three maps showing the main rivers, regions and
altitude zones, the main vegetation types and the network of National
Parks of the country. The first map and the National Parks map, as well
as a further map showing the provinces of Gabon, are repeated on the
inside back cover of the publication, facilitating easy referencing.
The rest of the introductory chapter is divided into sections deal-
ing with checklist preparation, plant collecting in Gabon and analyses
of the data. The methods used to create and hone the specimen-based
database from which the checklist was extracted, as well as those used
to improve the resulting draft checklist, are briefly explained under the
first subheading. The database contains over 65 000 specimen records
for the more than 4 700 vascular plants known in the country, as well
as over 800 literature references associated with the Gabonese flora.
Most (98%) of these specimen records include accurate map refer-
ences. Images and other additional data are currently being added to
the database.
The database is available online at http://herbaria.plants.ox.ac.uk/
bol/home. This Plants of Gabon website is very informative and useful.
It provides relevant information on the specimens in the database. In
addition, maps can be produced and, as mentioned above, inTages will
become available. The website also links to various other important
taxonomic and biodiversity databases (including IPNI, TROPICOS,
GBIF and others) where queries are run automatically on the taxon
selected in the Gabonese database. Another important online database
on Africa's flora to which the Plants of Gabon database should link is
the African Flowering Plants Database of the African Plant Checklist
and Database Project (http://www.ville-ge.ch/cjb/bd/affica). The reason
why it is not linked might simply be that the African Flowering Plants
Database website was not yet fully functional when the Plants of Gabon
website was launched.
The database is undoubtedly the most comprehensive of its type
for Gabon. Nevertheless, the claim by the authors that it is unique in
Africa, is not quite true. The information contained in the Plants of
Gabon database is very similar to that in the PRECIS database, kept
by the South African National Biodiversity Institute (SANBI) for the
Flora of southern Africa (FSA) region. The website and the accuracy of
the Plants of Gabon data do make the database exceptional in Africa.
However, SANBI is in the process of developing a web interface for
PRECIS, but, being a much larger database (containing data on 24 000
taxa linked to around 900 000 specimens and almost 30 000 literature
references), the process of quality control and enhancing the PRECIS
data for online availability takes much longer.
Under the Plant collecting in Gabon subheading, the most important
collectors and collecting efforts in Gabon are briefly mentioned. The
authors are apparently preparing a full account on all botanists that col-
lected in Gabon, which will be published elsewhere.
The section of the introductory chapter dealing with the analyses
of the data is again subdivided into various subsections. It offers valu-
able information about the Gabonese flora and the state of botanical
knowledge in the country. According to these analyses, the number of
collections from Gabon (around 72 000 herbarium specimens in total)
reveals that the country is still regarded as under-collected or poorly
known, with a mere 0.24 plant specimens per km-. For a botanical
country checklist to be of an acceptable level, a minimum of one speci-
men per km^ is required. This is also reflected by the fact that almost
every botanical expedition to Gabon reveals novel taxa that have not
yet been recorded for the country or that are entirely new to science.
Fortunately the analysis on collecting history indicates that collecting
activities in Gabon have increased substantially over the past 30 years
and are still increasing. This is very well illustrated with a histogram
showing the number of plant collections per decade from 1846-1849
up to 2004. A map of the number of plant collections per quarter-degree
square demonstrates the distribution of collecting efforts in Gabon. This
map clearly shows a few areas that are botanically well known or fairly
well known. Not surprisingly, the region with the most specimens is
that around the capital, Libreville. Also very obvious on this map is the
fact that most areas in Gabon have very few collections, whereas others
have never been included in plant collecting surveys at all. An interest-
ing analysis is that of collecting throughout the seasons. This was done
to determine whether certain species could have been overlooked, since
most collecting took place outside their flowering or fruiting season.
The accompanying histogram, illustrating the number of collections
per month of the year, shows that collecting is fairly evenly distributed
throughout the year. The least specimens were collected from June
to August, which coincides with the dry season when less flowering
material is available, and which is also the holiday season. The greatest
number of specimens were collected in November, but the authors have
no concrete explanation for this phenomenon.
Since no exhaustive search for type collections was performed
during this project, the figure of 1 318 type specimens indicated in the
database is lower than the actual number of types based on Gabonese
material. The total number of taxa, as well as species, genera and fami-
lies currently listed for Gabon is presented in a very useful table. This
table not only shows the total number, but also the separate numbers
for the classes Lycopsida, Pteropsida, Pinopsida and Magnoliopsida.
The checklist currently contains 4 710 species of vascular plants in
Gabon, including 82 introduced species. This number will fluctuate
with new records and new species being discovered in the country, as
well as with advances in taxonomic knowledge and improved identifi-
cation of specimens. The authors estimate the total number of species
present in Gabon to be between 7 000 and 7 500. The map illustrating
the recorded species richness across Gabon per quarter-degree square,
shows great similarity to the previous map indicating number of plant
collections. This is not unexpected, since information on species den-
sity is largely dependent on the number of collections, even more so in
an area that has not yet been thoroughly explored.
There are 508 strict endemics (10.8% endemism) in Gabon and
a further 97 near-endemics. This figure will also fluctuate with new
records and expanded knowledge. The map showing the distribution
of endemic and near-endemic taxa in Gabon clearly shows the regions
richest in endemics. This map also shows similarities to the previous
maps indicating number of collections and species density. These
endemic-rich areas are of the utmost importance in conservation efforts
and it is good to see that some, although not all, are currently included
in Gabon’s National Parks network.
130
Bothalia 37,1 (2007)
Under the last subheading, the number of collections per species
is discussed. As can be seen from the accompanying histogram, over
half of the species are known from only one to five specimens. This
further highlights the fact that Gabon is severely under-collected and
that the discovery of many more species can be expected as collecting
is expanded in future.
The introductory chapter is concluded by a list of all the literature
cited.
On the first two pages of the 404-page checklist, the format is ex-
plained in detail. The checklist records taxa in the following sequence:
Lycopsida, Pteropsida, Pinopsida and Magnoliopsida. Under each
class, families are listed alphabetically, and then genera and species
also alphabetically under each family. The most important synonyms of
an accepted name are listed. The endemic, near-endemic or introduced
status of relevant taxa is indicated. Cultivated taxa are not included in
the checklist. At least one abbreviated literature reference is provided.
This usually contains a good description of the taxon, or otherwise it
is a publication that was used to verify the correctness of the name. An
abbreviated list of Gabonese provinces in which the listed specimens
were collected is given. The map on the inside back cover showing the
provinces of Gabon also appears on the second page of this chapter,
where the details of the checklist are explained. The altitudinal range
of the taxon in Gabon is indicated, as well as any available notes relat-
ing to nomenclature, distribution or natural occurrence of the taxon.
Each taxon account concludes with an alphabetical listing of specimens
according to collector’s name and number. Other information provided
for each specimen includes the acronym of the herbarium or herbaria
where duplicates of the specimen can be found, the name of the most
recent identifier and the year of identification, if available. Type speci-
mens and doubtful identifications are also indicated.
A reference chapter follows, providing the full version of the refer-
ences abbreviated in the checklist. The publication ends with an index
to genera, which facilitates the location of genera in the checklist.
The authors and institutions involved are congratulated on this pro-
duction of the Plants of Gabon Project. As the authors of the publication
anticipated, this checklist, as well as the database, will undoubtedly
contribute to the continued conservation and wise utilization of Gabon’s
immense botanical wealth. More checklists of this quality and similar
online databases are urgently needed to improve our knowledge, under-
standing and ability to manage and conserve Africa’s unique flora.
R.R. KLOPPER*
* Research and Scientific Services Directorate, South African National
Biodiversity Institute, Private Bag XI 01, 0001 Pretoria.
Bothalia37,l (2007)
131
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2006-12-12
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Latest Publications
FLOWERING PLANTS OF AFRICA
national
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S A N B I
Voi. 60. Plates 2221-2240 (June 2007]
Twenty full-colour plates and descriptions of plants appear in this biennial series, which has
become a collector's item of the South African flora. This issue includes, among others, a new
species of Hypoxis and a new subspecies of Ledebouria ovatifolia, an Ornithogalum from sheer
ciiff faces in the Eastern Cape, a beautiful yellow arum lily, two gingers called Siphonochilus kirkii
and Costus afar, the parasite Viscum crassulae and the mangrove Bruguiera gymnorrhiza. Also
included are two pelargoniums and Stenostelma umbelluliferum, which was re-discovered after
more than 1 00 years. The botanical art is mainly the work of SANBI resident artist, Gillian Condy;
other artists contributing to this issue are Tamlin Blake, Angela Beaumont, Auriol Batten, Elsa
Pooley and Sibonelo Chiliza. There is a guide for authors and artists and an index to species. 250
X 190 mm. pp. 144.
Soft cover: ISBN 978-1-919976-34-1.
Price: SADC R1 90.00/other countries $48.00.
Flowering Plants of Africa
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STRELITZIA 19
The vegetation of South Africa, Lesotho and Swaziland
L. Mucina S. M.C. Rutherford Ceds] [20061
An up-to-date and comprehensive overview of the vegetation of South Africa and the two small
neighbouring countries of Lesotho and Swaziland. This account is based on a vegetation survey
using appropriate tools of contemporary vegetation mapping and vegetation description. The aim
was to draw a new vegetation map that depicts the complexity and macro-scale ecology and
reflects the level of Cand identifies and reveals gaps ini current knowledge of the vegetation of
the region. This is an extensive account of the vegetation of a complex and biologically intriguing
part of the world, offering not only insights into structure and dynamics of the vegetation cover,
but containing a wealth of base-line data for further vegetation-ecological, biogeographical, and
conservation-oriented studies. Included towards the back of the book is an atlas as a systematic
series of A4 maps depicting the various vegetation types. The electronic version on the CD
inside the front cover allows the user to zoom in at any scale to discern detail. The Map and
the descriptive account of the vegetation .of South Africa, Lesotho and Swaziland target not
only scientific academia and the secondary and tertiary education sectors, but offers a powerful
decision-making tool for conservationists, land and resource planners, and politicians as well as
the interested public at large. 297 x 210 mm. pp. 816.
Hard cover: ISBN-13: 978-1-919976-21-1 CISBN-10: 1-919976-21-31.
Price SADC R390. 00/other countries $100.00.
STRELITZIA 18
A revision of the southern African genus Babiana, Iridaceae: Crocoideae
R Goldblatt S. J.C. Manning (May 20071
Babiana, as now constituted, is largely a genus of the winter rainfall zone of western South Africa
and southwestern Namibia. Just two species occur in the southern African summer rainfall zone,
B. hypogaea and B. bainesii, the latter widespread and extending from the Upper Karoo through
Botswana and Namibia to Zimbabwe and southern Zambia, The authors, Peter Goldblatt and
John Manning, recognize 88 species, a substantial increase over the 61 included in Babiana by
G.J. Lewis in her 1959 monograph. This revision also presents a new infrageneric classification
of Babiana, which is divided into three sections: the more generalized section Teretifolieae C40
speciesi, which has inner bracts forked apically or to about the middle; the derived section
Babiana C37 speciesi, which has inner bracts divided to the base; and section Anthoiyzoides (1 1
speciesi, with bilabiate flowers with clawed tepals and a short perianth tube, and inner bracts
divided to about the middle. Each species is described and some species are accompanied by a
line drawing by John Manning, 15 in total, with 48 maps and 45 colour photographs. Excluded
names, a list of references and an index to species and synonyms are provided at the back of the
book, 297 X 210 mm. pp. xii + 98.
$oft cover: ISBN 978-1-919976-32-7,
Price SADC countries R1 30.00/other countries $33.00.
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BOTHALIA SPECIALS
Contents to vols 1-20
by H.F. Glen, B.A. Momberg & E. Potgieter (1991)
A brief history of Bothalia; a list of all papers published; a list of all authors, co-authors, keywords and titles; and tables with
publication dates, major subjects covered and some information on authors.
Price: SADC countries, R15.00 / Other countries US$4.00
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by B.A. Momberg & J.M. Mulvenna (1996)
List of papers alphabetically arranged according to senior author and dates and including all co-authors in alphabetical listing.
Subject index compiled from keywords and titles, with reference to individual articles.
Price: SADC countries, R15.00 / Other countries USS4.00
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by B.A. Momberg (2000)
List of papers alphabetically arranged according to senior author and dates and including all co-authors in alphabetical listing.
Subject index compiled from keywords and titles, with reference to individual articles.
Price: SADC countries, R18.00 / Other countries USS5.00
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BOTHALIA
Volume 37,1
CONTENTS
May 2007
1. Notes on new and misunderstood taxa of Cyrtanthus (Amaiyllidaceae: Cyrtantheae) from the Western
Cape, Eastern Cape and KwaZulu-Natal, South Africa. D.A. SNIJMAN 1
2. New species and notes on the genus Cliffortia (Rosacea&). C.M. WHITEHOUSE and A.C. FELLING-
HAM 9
3. Notes on African plants:
Apocynaceae. Transfer of Schizoglossum umbelluliferum to Stenostelma, and its neotypification
(Asclepiadoideae). S.R HESTER and A. NICHOLAS 48
Apocynaceae. A new species of Huernia (Asclepiadoideae-Ceropegieae) from southern Angola.
P.V. BRUYNS 23
Asphodelaceae. Trachyandra arenicola and T. montana, two new species from South Africa. J.C.
MANNING and P. GOLDBLATT 26
Asteraceae. A key to Dicoma taxa (Dicomeae) in southern Africa. N.C. NETNOU and B-E. VAN
WYK 55
Boraginaceae. Lectotypification of the basionym, Echium glaucophyllum. M.H. BUYS and B.
NORDENSTAM 25
Burseraceae. Commiphora kuneneana, a new species from the Kaokoveld, Namibia. W. SWANE-
POEL 40
Fabaceae. Aspalathus theresae, a new species from Western Cape, South Africa. C.N. CUPIDO . . 34
Hyacinthaceae. A new pyrophytic Lachenalia species (Massonieae) from Western Cape, South
Africa. G.D. DUNCAN and T.J. EDWARDS 31
Lamiaceae. Rabdosiella leemannii, a new species from the Limpopo Province of South Africa. N.
HAHN and G.J. BREDENKAMP 37
Leguminosae. Vigna verdcourtii (Papilionoideae), a new species from eastern Africa. R.S. PAS-
QUET 51
4. A reconnaissance survey of the woody flora and vegetation of the Catapii logging concession, Cheringoma
District, Mozamb-que. M. COATES PALGRAVE, A.E. VAN WYK, M. JORDAAN, J.A. WHITE
and P. SWEET 57
5. The concept of ’Musa-pelo and the medicinal use of shrubby legumes (Fabaceae) in Lesotho. A.
MOTEETEE and B-E. VAN WYK 75
6. (ian anthropogenic variables be used as threat proxies for South African plant richness? M. KEITH and
M. WARREN 79
7. Threatened Limestone Fynbos plant communities of Andrew’s Field and Tsaba-Tsaba Nature Reserve,
Western Cape. M.M. ZIETSMAN and G.J. BREDENKAMP 89
8. Miscellaneous notes:
Amaryllidaceae. Chromosome studies on African plants. 20. Karyotypes of some Cyrtanthus
species. A. STRYDOM, R. KLEYNHANS and J.J. SPIES ’. . 103
9. The handling of the proposal to conserve the name Acacia at the 1 7th International Botanical Congress —
an attempt at minority rule. G. MQORE 109
10. Obituary: Elsie Elizabeth Esterhuysen (1912-2006). E.G.H. OLIVER (Compiler), J.P ROURKE and
H.P LINDER 119
11 Book review 129
1 2 Notice on Change of policy for reprints of Bothalia articles 131
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ABSTRACTS • ISI: Current Contents. Scisearch, Research Alert • Kew Record of Taxonomic Literature • Taxon', reviews and notices.
ISSN 006 8241
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