Bothalia
A JOURNAL OF BOTANICAL RESEARCH
Vol. 38,1
May 2008
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BOTHALIA
Bothalia is named in honour of General Louis Botha, first Premier and Minister of Agriculture of the Union of
South Africa. This house journal of the 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 subjects are published annually.
Three booklets of the contents (a) to Vols 1-20, (b) to Vols 21-25, (c) to Vols 26-30, and (d) to Vols 31-37 (2001-
2007) are available.
STRELITZIA
A series of occasional publications on southern African flora and vegetation, replacing Memoirs of the Botanical
Survey of South Africa and Annals of Kirstenbosch Botanic Gardens.
MEMOIRS OF THE BOTANICAL SURVEY OF SOUTH AFRICA
The memoirs are individual treatises usually of an ecological nature, but sometimes dealing with taxonomy or
economic botany. Published: Nos 1-63 (many out of print). Discontinued after No. 63.
ANNALS OF KIRSTENBOSCH BOTANIC GARDENS
A series devoted to the publication of monographs and major works on southern African flora. Published:
Vols 14-19 (earlier volumes published as supplementary volumes to the Journal of South African Botany).
Discontinued after Vol. 19.
FLOWERING PLANTS OF AFRICA (FPA)
This serial presents colour plates of African plants with accompanying text. The plates are prepared mainly by
the artists at the 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 Letty (over 700 plates), Claire Linder-Smith and Ellaphie Ward-Hilhorst.
The Editor is pleased to receive living plants of general interest or of economic value for illustration.
From Vol. 55, twenty plates are published at irregular intervals.
An index to Vols 1-49 is available.
FLORA OF SOUTHERN AFRICA (FSA)
A taxonomic treatise on the flora of the Republic of South Africa, Lesotho, Swaziland, Namibia and Botswana,
the FSA contains descriptions of families, genera, species, infraspecific taxa, keys to genera and species, syn-
onymy, 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 information is pre-
sented in the form of tables and photographic plates depicting fossil populations. Now available:
Molteno Formation (Triassic) Vol. 1. Introduction. Dicroidium, 1983, by J.M. & H.M. Anderson.
Molteno Formation (Triassic) Vol. 2. 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.).
Heyday of the gymnosperms: systematics and biodiversity of the Late Triassic Molteno fructifications,
2003, by J.M. Anderson & H.M. Anderson.
Brief history of the gymnosperms: classification, biodiversity, phytogeography and ecology, 2007, by
J.M. Anderson, H.M. Anderson & C.J. Cleal.
SANBI BIODIVERISITY SERIES
A series of occasional reports on projects, technologies, workshops, symposia and other activities initated by or
executed in partnership with SANBI.
BOTHALIA
A JOURNAL OF BOTANICAL RESEARCH
Volume 38,1
Scientific Editor: G. Germishuizen
Technical Editor: B.A. Momberg
national
biodiversity
institute
S A N B I
2 Cussonia Avenue, Brummeria, Pretoria
Private Bag XI 01, Pretoria 0001
ISSN 0006 8241
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.
Balkwill, Prof. K. University of the Witwatersrand, Johannesburg, RSA.
Bohs, Dr L. University of Utah, Salt Lake City, USA.
Burrows, J.E. P.O. Box 710, 1120 Lydenburg, RSA.
Carter Holmes, Mrs S. Royal Botanic Gardens, Kew, UK.
Child, Dr A. 31 York Rd, Driffield, East Yorkshire, UK.
Demissew, Prof. S. National Herbarium, Addis Ababa University, Ethiopia.
Dreyer, Dr L. University of Stellenbosch, RSA.
Geldenhuys, Prof. C.J. Forestwood cc, P.O. Box 228, La Montagne, Pretoria, RSA.
Goyder, Dr D.J. Royal Botanic Gardens, Kew, UK.
Herman, P.P.J. South African National Biodiversity Institute, Pretoria, RSA.
Knapp, Dr S. The Natural History Museum, London, UK.
Koekemoer, Dr M. South African National Biodiversity Institute, Pretoria, RSA.
Lavranos, J. Apartado Postal 243, 8100 Louie, Portugal.
Leistner, Dr O.A. South African National Biodiversity Institute, Pretoria, RSA.
Manning, Dr J.C. South African National Biodiversity Institute, Cape Town, RSA.
McDonald, Dr D.J. Bergwind Botanical Surveys & Tours CC, 14A Thomson Rd, Claremont, Cape Town, RSA.
McNeill, Dr J. Royal Botanic Garden, Edinburgh, Scotland, UK.
Meve, Dr U. Department of Plant Systematics, University of Bayreuth, Germany.
Meyer, Mrs N.L. 598 Vacy Lyle St., Elardus Park, 0181 Pretoria, RSA.
Moffett, Dr R.O. P.O. Box 326, Kestell, RSA.
Nordenstam, Prof. R.B. Naturhistoriska Riksmuseet, Stockholm, Sweden.
Oberlander, K.C. University of Stellenbosch, RSA.
Oliver, Dr E.G.H. University of Stellenbosch, RSA.
Retief, Dr E. South African National Biodiversity Institute, Pretoria, RSA.
Shackleton, Dr C.M. Rhodes University, Grahamstown, RSA.
Snijman, Dr D. South African National Biodiversity Institute, Cape Town, RSA.
Stedje, Prof. Dr B. Botanical Garden & Museum, Oslo, Norway.
Thulin, Dr M.L. Department of Systematic Botany, University of Uppsala, Uppsala, Sweden.
Van Jaarsveld, E.J. South African National Biodiversity Institute, Cape Town, RSA.
Van Wyk, Prof. A.E. University of Pretoria, RSA.
Walters, Dr B.B. Mount Allison University, Sackville, Canada.
Werger, Prof. M.J.A. Department Plant Ecology and Evolutionary Biology, Utrecht, The Netherlands.
Williamson, Dr G. 26 Starke Rd, Bergvliet, Cape Town, RSA.
CONTENTS
Bothalia 38,1
1 . Systematics of the southern African genus Ixia (Iridaceae). 1 . The I. rapunculoides complex. P. GOLD-
BLATT and J.C. MANNING 1
2. Three new species of Asparagus (Asparagaceae) from South Africa, with notes on other taxa. S.M.
BURROWS and J.E. BURROWS 23
3. A new species of Euclea (Ebenaceae) from ultramafic soils in Sekhukhuneland, South Africa, with notes
on its ecology. E. RETIEF, S.J. SIEBERT and A.E. VAN WYK 31
4. The genus Solarium (Solanaceae) in southern Africa: subgenus Leptostemonum, section Giganteiformia.
W.G. WELMAN 39
5. Two new species of Babiana (Iridaceae: Crocoideae) from western South Africa, new names for B.
longiflora and B. thunbergii, and comments on the original publication of the genus. P. GOLDBLATT,
J.C. MANNING and R. GEREAU 49
6. The genus Wellstedia (Boraginaceae: Wellstedioideae) in southern Africa. E. RETIEF and A.E. VAN
WYK 57
7. Notes on African plants:
Apocynaceae. A new species of Huernia (Asclepiadoideae-Ceropegieae) from Angola. P.V.
BRUYNS 83
Asphodelaceae. Notes on the nomenclature and typification of Aloe natalensis (Alooideae). N.R.
CROUCH, G.F. SMITH and R.R. KLOPPER 70
Asphodelaceae: Alooideae. Bulbine triebneri, an earlier name for Bulbine alba , as well as additional
and new localities in Eastern and Northern Cape, South Africa. R.R. KLOPPER, A.W.
KLOPPER, H. BAIJNATH and G.F. SMITH 67
Asphodelaceae: Alooideae. New evidence in support of a disjunct distribution of Aloe karasbergensis.
R.R. KLOPPER, P.J. DU PREEZ and G.F. SMITH 82
Asteraceae-Gnaphalieae. Metalasia helmei, a new member of a small clade from the Western
Cape? P.O. KARIS and N. HELME 65
Asteraceae. Tripteris calcicola, a new calciphilous species from Western Cape, South Africa. J.C.
MANNING and P. GOLDBLATT 85
Colchicaceae. Further new combinations in Colchicum. C. ARCHER 83
Errata in Bothalia 37,2 (2007): MOFFETT, R.O. 2007. Name changes in the Old World Rhus and
recognition of Searsia (Anacardiaceae) 71
Hyacinthaceae. Drimiopsis linioseta, a new species from the Sekukhuneland Centre of Endemism,
South Africa. A.J. HANKEY, M.H. BUYS and P.D. LEBATHA 72
Iridaceae. Romulea lutea and R. tubulosa (Crocoideae), two new species from Namaqualand, South
Africa. J.C. MANNING and P. GOLDBLATT 78
Oxalidaceae. A new species of Oxalis from the Hantam-Roggeveld Plateau, Northern Cape, South
Africa. J.C. MANNING and P. GOLDBLATT 75
Rosaceae. Validation of three Cliffortia taxa in Bothalia 37, 1 (2007). C.M. WHITEHOUSE 75
8. Vegetation and vegetation-environment relationships at Grootbos Nature Reserve, Western Cape, South
Africa. M. MERGILI and S. PRIVETT 89
9. Resource demand estimates for sustainable forest management: Mngazana Mangrove Forest, South
Africa. C.H. TRAYNOR and T.R. HILL 103
10. Obituary: Eily Edith Agnes Gledhill (nee Archibald) (1914-2007). E. BRINK and G. GERMIS-
HUIZEN Ill
New combinations, names, species, statuses, subspecies and varieties in Bothalia 38,1 (2008)
Asparagus elephantinus S.M.Burrows, sp. nov., 23
Asparagus hirsutus S.M.Burrows, sp. nov., 25
Asparagus sylvicola S.M.Burrows, sp. nov., 26
Babiana hirsuta (Lam.) Goldblatt & J.C. Manning, comb, nov., 53
Babiana symmetrantha Goldblatt & J.C. Manning, sp. nov., 49
Babiana tubaeformis Goldblatt & J.C. Manning, nom. nov., 53
Babiana virescens Goldblatt & J.C. Manning, sp. nov., 51
Colchicum leistneri (U.Miill.-Doblies & D.Mull.-Doblies) C.Archer, comb, nov., 83
Colchicum palaestinum (Baker) C.Archer, comb, nov., 83
Colchicum schimperianum (Hochst.) C.Archer, comb, nov., 83
Drimiopsis linioseta A.J.Hankey & P.D.Lebatha, sp. nov., 72
Euclea sekhukhuniensis Retief, Siebert & A.E.van Wyk, sp. nov., 32
Huernia calosticta Bruyns, sp. nov., 83
Ixia contorta Goldblatt & J.C. Manning, sp. nov., 21
Ixia divaricata Goldblatt & J.C. Manning, nom. nov., 18
Ixia flaccida (G.J. Lewis) Goldblatt & J.C. Manning, comb, et stat. nov., 17
Ixia lacerata Goldblatt & J.C. Manning, sp. nov., 10
Ixia oxalidiflora Goldblatt & J.C. Manning, sp. nov., 17
Ixia rivulicola Goldblatt & J.C. Manning, sp. nov., 9
Ixia robusta (G.J. Lewis) Goldblatt & J.C. Manning, comb, et stat. nov., 9
Ixia sobolifera Goldblatt & J.C. Manning, sp. nov., 11
Ixia sobolifera Goldblatt & J.C. Manning subsp. albiflora Goldblatt & J.C. Manning, subsp. nov., 14
Ixia sobolifera Goldblatt & J.C. Manning subsp. carnea Goldblatt & J.C. Manning, subsp. nov., 13
Metalasia helmei P.O.Karis, sp. nov., 65
Oxalis odorata J.C. Manning & Goldblatt, sp. nov., 75
Romulea lutea J.C. Manning & Goldblatt, sp. nov., 78
Romulea tubulosa J.C. Manning & Goldblatt, sp. nov., 80
Tripteris calcicola J.C. Manning & Goldblatt, sp. nov., 86
Wellstedia dinteri subsp. gracilior ( D.R.Hunt ) Retief & A.E.van Wyk, comb, et stat. nov., 62
IV
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2
Bothalia 38,1 (2008)
culoides by Lewis (1962) on the basis of its included fila-
ments and partly included anthers.
Vegetatively, all members of the complex have slen-
der, erect flowering stems bearing few to many, short,
± wiry, often thread-like lateral branchlets and typi-
cally, but not exclusively, just three (rarely two or four
to six) foliage leaves. The lower two (rarely one or up to
four) leaves have well-developed blades, and the upper
one (or two) completely sheath the stem and reach to
between the middle of the stem and the base of the spike.
The main spike bears relatively few flowers and the lat-
eral spikes have fewer, sometimes only a single flower.
Plants referred to I. rapunculoides var. robusta normally
have four leaves but sometimes up to six, with three or
four basal and one or two sheathing the stem, and two
other varieties rarely have three expanded foliage leaves.
None of these vegetative features are, however, unique
to the complex, but are shared by several other species
of subgenus Morphixia (Goldblatt & Manning 1999),
notably Ixia latifolia D.Delaroche and I. marginifolia
G.J. Lewis.
Lewis’s taxonomic treatment seems to reflect a per-
ception that the morphological variation within the com-
plex was less significant than that encountered between
species elsewhere in the genus, presumably because
the distinctive flowers of the complex were given
undue weight in a genus that is relatively conservative
in floral morphology. Some of the variation, especially
floral scent, corm features, and the nature of the bracts
subtending the branchlets was simply overlooked. The
morphological variation in the complex is, nevertheless,
extensive and includes: 1, corm tunics that range from
coarsely to finely fibrous, or softly papery; 2, cormlets
that are either sessile at the base of the main corm or
borne on flattened, ribbon-like stolons; 3, flower colour
that is often blue to blue-mauve, but also purple, pink or
white; 4, leaf shape, which may be narrow, linear, and
erect, or sword-shaped and erect, or arching outward
and falcate; 5, degree of thickening of the margins of the
basal leaves, which is least developed in 7. rapunculoi-
des var. fiaccida and most pronounced in var. robusta
and var. rapunculoides ; 6, shape and degree of develop-
ment of the bracts and prophylls subtending the thread-
like, lateral branches (here called branchlets when short,
or branches when well-developed); 7, length of the peri-
anth tube, mostly 5. 5-8.0 mm in var. rapunculoides , and
1 2—1 6(— 1 8) mm in var. namaquana-, 8, length of the fila-
ments and anthers, features linked to tube length, and
together range from 4. 3-6. 5 mm in var. rapunculoides to
a maximum of 9.5 mm in var. namaquana and 10.5 mm
in var. subpendula.
The morphological variation often correlates with
ecological and geographical differences but the pat-
terns have until now been inadequately understood.
Particularly noteworthy, and from a systematic point of
view unsatisfactory, is the local sympatry of some vari-
eties or the parapatric and edaphic or geographic sepa-
ration of others, in all cases without the occurrence of
plants of intermediate morphology.
For example, on dolerite-derived clay soils on the
Bokkeveld Plateau and on the Hantamsberg at Calvinia
in Northern Cape, both Ixia rapunculoides var. rapun-
culoides and plants that conform to var. fiaccida (sensu
Lewis 1962), flower contemporaneously within a few
metres of one another with no intermediates, indicating
complete genetic isolation. Sympatry and synchrony of
flowering of two distinct races without the presence of
intermediate individuals is usually accepted as an indi-
cation of genetic isolation and therefore evidence for
their recognition as separate species. Similarly, Lewis
(1962) noted that at one of its two localities then known,
var. robusta grew on rocky dolerite slopes a few metres
from typical var. rapunculoides , growing on flat, stony
loam near a seasonal watercourse. We have confirmed
Lewis’s observation and amplify it, having seen var.
robusta growing side by side with var. rapunculoides,
both taxa maintaining their typical morphology. A com-
parable situation occurs on the Bokkeveld Plateau near
Nieuwoudtville, where typical var. rapunculoides grows
on tillite-derived clay soils and var. namaquana on adja-
cent sandy soils.
These examples are particularly surprising because
the flowers of all these taxa are apparently adapted for
pollination by the same insects, a range of medium-
and large-bodied bees, mostly Apidae: Anthophorinae,
that forage for nectar and acquire loads of pollen while
probing the perianth tube (Goldblatt et al. 2000 and con-
firmed here).
Field observations suggest to us that varietal sepa-
ration within Ixia rapunculoides of var. namaquana ,
var. fiaccida and var. robusta is unsatisfactory because
the biological situation is not concordant with this tax-
onomy. The recognition of varieties of a single species
is notoriously unevenly applied (Stuessy 1990), but is
most often used to denote trivial regional variation, and
sometimes only small genetic differences such as flower
colour or leaf shape. In our estimation it should never be
associated with genetic isolation and effective crossing
barriers at the microgeographic level. Raising the rank
at which members of the complex are separated to that
of subspecies has similar problems. The rank of subspe-
cies is most often used for sets of populations that are
isolated geographically and have relatively modest, and
often overlapping, morphological differences (Davis &
Heywood 1973). As with varietal rank, subspecific rank
is inconsistently applied. The examples in I. rapuncu-
loides listed above thus accord poorly with treatment at
either varietal or subspecific rank.
MATERIALS AND METHODS
We have attempted to resolve this unsatisfactory taxo-
nomic situation outlined above where so-called varie-
ties of a species co-occur and flower synchronously yet
maintain their morphological differences by examination
of the morphology and ecology of plant populations in
the field. These studies were complemented by a study
of herbarium collections of the genus at BOL, K, MO,
NBG and PRE, the herbaria with the most complete col-
lections of species from the southern African winter rain-
fall zone. Then, using a morphological species concept,
we have compared all significant taxonomic characters
of the different sets of populations with their ecology
and geography to establish a revised taxonomy of the
complex.
Bothalia 38,1 (2008)
3
RESULTS
Our results, obtained from measuring both living
plants in the field and carefully preserved specimens
with laid out flowers are presented in tabular form
(Table 1). We do not use Lewis’s (1962) or De Vos’s
(1999) measures for any taxa because we apply names
in different ways. We have also been careful to include
measurements of only well-pressed herbarium speci-
mens because floral parts are particularly susceptible to
distortion and can shrink by as much as 20% as they dry,
less so when rapidly pressed on glue-covered paper or
between the pages of a heavy book.
Members of the lxia rapunculoides complex are not
reported in the literature to have scented flowers (Lewis
1962; De Vos 1999; Manning et al. 2002). Nevertheless,
we have found that plants of most populations have
distinctive floral odours. Plants from the eastern part
of the range of the complex with nodding white flow-
ers have a particularly intensely sweet, carnation-like
odour; typical var. rapunculoides has a faint rose scent;
var. namaquana usually has a violet scent or are occa-
sionally unscented; and var. robusta has a faint, unpleas-
ant chemical odour. A population of white-flowered
plants allied to var. rapunculoides from the Roggeveld
Escarpment has a strong fruity odour recalling a combi-
nation of banana and grenadilla, very similar to Virgilia
oroboides (Fabaceae).
Another feature ignored in the past, is the nature of
the bracts and prophylls subtending the branchlets.
These range from minute and truncate (var. rapunculoi-
des, var. robusta ) to forked with attenuate tips (Western
Karoo plants assigned to var .flaccida), to filiform, elon-
gate and recurved (plants from the Klein Roggeveld
referred by Lewis (1962) to var. rapunculoides or var.
namaquana).
Comas of members of the complex have until now
been described as ‘having fairly coarse, often subligne-
ous, reticulate fibres’ (Lewis 1962) and neither Lewis
nor De Vos (1999) mention the presence of cormlets
(cormels) or underground stolons. Texture and compo-
sition of the tunics, however, varies considerably and
more so than is ever found within a single species of any
allied genus of Iridaceae. Hard, coarsely fibrous tunics,
matching Lewis’s description, are characteristic of var.
rapunculoides, var. robusta and var. namaquana and
these taxa bear ± sessile cormlets at the base of the main
corm. Soft, short-lived tunics of ± membranous layers
characterize some plants included in var. flaccida and
the southern populations of pink-flowered plants referred
to var. namaquana, and in addition these plants produce
TABLE 1 . — Characteristics of members of lxia rapunculoides complex. All varieties recognized by short stamens with filaments and at least bases
of anthers included in floral tube (var. subpendula is exceptional in this respect as anthers are usually fully exserted). Data are taken from
new collections and well-pressed herbarium specimens
fil, filaments; anth, anthers.
Bothalia 38,1 (2008)
5
quana but represent a novel species, which we describe
below as I. oxalidiflora.
4. Var. flaccida: the type of this variety, from the
Olifants River Valley near Clanwilliam in Western
Cape, is distinguished by the fairly thin-textured, nar-
rowly lanceolate leaves less than 10 mm wide and ±
half as long as the stem. The white to palest blue flow-
ers are carried ± horizontally, have a perianth tube 7-9
mm long, while the stamens are inserted ± 2 mm from
the base of the tube, the filaments are 2. 5-3.0 mm long
and the anthers ±2.5 mm long. Plants have three or
four leaves, the uppermost one (or sometimes the upper
two when four leaves are present) is typically entirely
sheathing. The corms have tunics composed of fine or
medium-textured, netted fibres and bear cormlets at the
base. Plants grow in light sandy soil, among sandstone
rocks, on south-trending slopes, in light shade, in thicket
communities. They most closely match var. rapunculoi-
des in tube and stamen length but apart from differences
in leaf shape and texture, they have narrower tepals and
corm tunics of finer fibres. Dimensions given by Lewis
(1962) and De Vos (1999) for the variety have a wider
range because they included additional sets of popula-
tions within the taxon.
The first of these populations comprises slender, pink-
flowered plants from the Bokkeveld Plateau, the second
slate-blue-flowered plants from the Klein Roggeveld,
and the third represents white-flowered plants from the
central Little Karoo. These variants are recognized by
the taller than usual flowering stem, up to 750 mm long
when growing under conditions of adequate rainfall,
a half or fully nodding flower, and perianth with a pale
yellow cup. The leaves of these plants are narrow, lin-
ear to narrowly sword-shaped, slightly twisted and often
trailing above. Most significantly, the corms differ from
all other populations of I. rapunculoides sensu Lewis
including typical war. flaccida, in having soft, soft-papery
corm tunics and broad, flat stolons 30-100 mm long ter-
minating in a large cormlet.
At two sites where we have seen the pink-flowered
variant, at Glenlyon near Nieuwoudtville and on the
Hantamsberg at Calvinia, plants were sympatric with
typical var. rapunculoides. At a third site, near Grasberg
Farm, northwest of Nieuwoudtville, pink-flowered plants
were sympatric with var. namaquana. The taller pink-
flowered plants grew in grass tufts or in bush clumps,
whereas var. rapunculoides and var. namaquana grew
on open ground. We conclude that these pink-flowered
plants represent an unnamed taxon. Apart from the nod-
ding flower, pink perianth and narrow tepals, flowers of
this plant have a tube ± 9 mm long, filaments ± 3 mm
long, and anthers ± 4 mm long. The style divides at the
anther apices in fully open flowers and the style branches
are short, slightly exceeding 1 mm long, thus visible
above the anthers. Blue-flowered var. rapunculoides
growing nearby has a perianth tube 7-8 mm long, fila-
ments ± 2 mm long, and anthers ± 3 .mm long, whereas
var. namaquana has a tube (1 2-) 14- 16 mm long, anthers
± 4 mm long and a shorter style.
We have also re-collected and examined the grey-
blue- and the white-flowered plants referred by Lewis to
var. flaccida at sites in the Klein Roggeveld and Little
Karoo. Plants from the latter area grow on south-trend-
ing shale slopes in light loamy ground near Oudtshoom
and in the foothills of the Kammanassie Mountains (e.g.
Goldblatt & Porter 12291 , MO, NBG; Vlok & Schutte
494, MO, NBG), and at a few other sites. These plants
resemble most closely the pink-flowered plants from the
Bokkeveld Plateau in their tall stature, and narrow, ± lin-
ear leaves and, most significantly in their soft, somewhat
papery corm tunics and long, slender stolons bearing a
terminal cormlet. The Klein Roggeveld plants, which
also have corms with soft tunics and stolons, favour
clay or sandy soils in more mesic sites in this semi-arid
area, often south-facing slopes. Because these three sets
of populations of stoloniferous plants differ in flower
colour, branching pattern, and in the strength and quality
of their scents, as well as in small differences in size of
floral parts, they are probably best regarded as subspe-
cies of a single species, which we call I. sobolifera. We
propose the names subsp. albiflora for the Little Karoo
populations, subsp. carnea for those from the Bokkveld
Plateau, while the Klein Roggeveld populations con-
stitute subsp. sobolifera. The disjunction between the
northern subsp. carnea and Klein Roggeveld subsp.
sobolifera may prove to be apparent rather than real as
more collecting on the Roggeveld Escarpment is done.
Likewise the disjunction between the ranges of subsp.
sobolifera and the central Little Karoo subsp. albiflora
will likely be bridged when more exploration at suitable
sites between their ranges is undertaken.
5. Var. subpendula: plants assigned to this taxon have
a discrete ecogeographic range, encompassing the Cold
Bokkeveld and surrounding Grootwinterhoek and Hex
River Mtns where they grow in seasonally waterlogged
sandstone-derived soils. They also comprise a coherent
morphological unit, having moderate-sized, white to pale
pink flowers with a perianth tube 8-11 mm long, sub-
equal tepals 11-14 x 6-7 mm, anthers that are 4.0-5. 5
mm long and usually fully exserted from the floral tube
(sometimes the bases are included), and style branches
1-2 mm long. The narrow leaves are typically 3-5 mm
wide (but sometimes up to 10 mm). The remarkable long,
straight lateral branches diverge at more than 45° from
the main axis and bear flowers in the distal half. These
populations seem out of place in the Ixia rapunculoides
complex, particularly in the large, excluded anthers and
relatively longer style branches, and the flowers match
most closely those of the related I. capillaris which is
distinguished not only by the shortly exserted anthers
but by the narrow, linear leaves, and particularly by the
slender lateral branchlets, which are fairly short, rarely
exceeding 1 0 mm, and usually bearing only one or two,
rarely three, flowers. We suspect that var. subpendula
is not immediately allied to I. rapunculoides. While its
flowers resemble most closely those of I. capillaris, the
unusual branching pattern recalls the pink- or red-flow-
ered/. latifolia.
Whereas Ixia capillaris and I. latifolia favour clay
soils and are fairly widespread in the southwestern Cape,
I. rapunculoides var. subpendula is restricted to season-
ally waterlogged, sandy, stony flats and rocky sites. We
recognize var. subpendula as a separate species, which
we call I. divaricata for the unusual, stiff, straight lat-
eral branches that diverge at a sharp angle from the main
6
Bothalia 38,1 (2008)
axis. Use of the epithet subpendula seems inappropri-
ate, for the lateral branches are rarely subpendulous, and
usually held above the horizontal.
6. Var. rigida: the type of var. rigida from the Hex
River Valley, does not differ in any significant respect
from var. subpendula and we therefore include it in that
taxon. Other collections from the Cedarberg and Cold
Bokkeveld that were assigned to var. rigida by Lewis
(1962), however, appear to represent a different spe-
cies. Despite Lewis’s comment that the androecium and
gynoecium are like those of var. rapunculoides, we find
the stamens of these plants rather different: the filaments
are ± 3 mm long (versus ± 2-3 mm in var. rapunculoi-
des), and the anthers are 3^1 mm long, again longer than
is usual for var. rapunculoides. The flowers also differ
from typical var. rapunculoides (Table 1) in their upright
orientation, and the slightly longer perianth tube, 9-12
mm long (versus 5.5— 8(— 10) mm in var. rapunculoides).
We see no particularly close relationship of these plants
to typical 7. rapunculoides or any other member of the
complex and treat them as a separate species, 7. contorta.
Additional variants: there are two more sets of popu-
lations allied to Ixia rapunculoides. The first of these
comprises white-flowered plants from the Roggeveld
Escarpment, unknown to Lewis when she revised Ixia in
1962. Flowering in October (when all other members of
the 7. rapunculoides complex in the Western Karoo and
Roggeveld are in fruit), these plants are confined to the
edges of streams that usually have running water until
October and remain moist at least until December. Plants
are tall, up to 1.2 m high, have membranous corm tunics
that do not accumulate with age, and bear small corm-
lets at the base of the main corm. The relatively large,
nodding flowers have a tube 8-9 mm long, tepals 17-18
x 5. 0-7. 5 mm, and anthers 4. 5-5. 5 mm long, the lower
halves included in the perianth tube. The flowers have a
strong fruity odour, reminiscent of banana and grenadilla
(passion fruit), unique in Ixia, but closely matched by
the scent produced by Virgilia oroboides.
We suggest that the most appropriate treatment for
this plant is recognition at species rank and propose
the name Ixia rivulicola. While its features recall other
varieties of the 7. rapunculoides complex, notably var.
rapunculoides and var. flaccida, it is distinct in flower
colour and fragrance, leaf number and shape, in the
poorly developed corm tunics, and in the unique riparian
habitat.
The second set of populations comprises short plants
from the Klein Roggeveld with pale bluish grey flow-
ers. Assigned previously either to var. rapunculoides
or to var. namaquana (Lewis 1962; De Vos 1999) the
two lower leaves of the plant have expanded blades
and a third sheathing leaf, flowers with a narrowly fun-
nel-shaped perianth tube 10-14 mm long, branchlets
subtended by long, recurved, thread-like bracts and pro-
phylls that are bifurcate, and unusual floral bracts with
dry, often tom tips, the outer usually with five veins and
the inner with three or four. The conns bear basal corm-
lets and have tunics composed of coarse fibres typical of
Ixia rapunculoides. We treat these plants as the new spe-
cies, 7. lacerata.
Key to Ixia rapunculoides and its close allies in subgenus Morphixia
Note: care must be taken in measuring floral parts of preserved specimens: depending on the method of drying, the
perianth can shrink as much as 20%. Presence of stolons is difficult to establish as they are often left in the ground
unless corms are removed with particular care. Filaments are measured from point of insertion on the perianth tube
(although they are decurrent) to the base of the anther. The perianth tube is measured from the top of the ovary to the
point at which the tepals separate from the upper portion of the tube.
la Foliage leaves linear-filiform, less than 2 mm wide when alive; midvein lying closer to abaxial margin but not evident when alive unless
held to the light 7. capillaris complex
lb Foliage leaves linear to lanceolate or falcate but never leathery and filiform, usually more than 2 mm wide in mature plants; midvein
evident when alive, central or slightly displaced toward abaxial margin:
2a Filaments reaching at least to top of perianth tube or exserted, thus anthers always completely exserted 7 latifotia and close allies
including 7 divaricata (= 7 rapunculoides var. subpendula), I. marginifolia, and most of the remaining members of subgenus Morphixia
2b Filaments included and anthers partly or sometimes also fully included within perianth tube (7 rapunculoides complex);
3a Flowers upright; perianth tube 8-13 mm long;
4a Flowers purple with pale yellow cup; main spike and branchlets twisted and flexuose, inclined to nearly horizontal; plants up to
150 mm high I. contorta
4b Flowers white, pink or rarely purple with white or pale yellow cup; main spike long, straight and erect, lateral spikes ascending to
spreading, with flowers crowded in distal half; plants mostly 300-600 mm high (this taxon does not correctly belong to the com-
plex as filaments are normally exserted 1-2 mm but occasionally reach to just short of mouth of tube) 7. divaricata
3b Flowers ascending, horizontal or nodding; perianth tube 5-22 mm long:
5a Perianth tube ( 1 2— )1 3— 20 mm long; anthers partially or fully included in perianth tube:
6a Anthers half exserted from perianth tube; perianth tube ( 1 2—) 1 3— 1 6(— 18) mm long; flowers ± horizontally oriented; corm mostly
12-14 mm diam., with tunics of medium to coarse fibres; cormlets borne at base of main corm 7 namaquana
6b Anthers fully included in perianth tube; perianth tube 16-22 mm long; flowers mostly ascending; corm mostly 8-10 mm diam.,
with tunics of fine fibres; cormlets borne at ends of flattened stolons 7. oxalidiflora
5b Perianth tube 5— 12(— 1 4) mm long; anthers partly included in perianth tube:
7a Leaf blades of lower two or three leaves sword-shaped to falcate (rarely sublinear), mostly 8-15 mm wide (rarely 3-7 mm wide),
usually less than one third as long as stem (longer if growing through bush); flowers ascending to horizontal; filaments 2-4
mm long; anthers 2. 3-4.0 mm long: ■r
8a Bracts and prophylls subtending branchlets bifurcate and attenuate, often directed downward distally; leaves 2(1) with well-deve-
loped blades; perianth tube 10— 12(— 14) mm long; outer bracts with (4)5 major veins, 5-toothed, teeth attenuate 7. lacerata
8b Bracts and prophylls subtending branchlets short, obtuse to truncate, less than 2 mm long; leaves 2 or 3(4) with well-developed
blades; perianth tube 5.5-8.0(-10.2) mm long; outer bracts with 3 major veins, shortly 3-toothed to bluntly 3-lobed:
9a Flowers blue-mauve, occasionally pale pink, with yellow to white throat; expanded foliage leaves 2(3); blades usually falcate;
Bothalia 38,1 (2008)
7
base of stem rarely with fibrous collar; perianth tube (5.5— )8.0— 1 0 mm long; anthers 2. 3-3. 5 mm long I. rapunculoides
9b Flowers pale pink with white throat; expanded foliage leaves 3(4); blades usually sword-shaped; base of stem covered by col-
lar of fibres; perianth tube 7.5-10.2 mm long; anthers 3. 5^4.0 mm I. robusta
7b Leaf blades linear or narrowly sword-shaped, erect, weakly twisted or trailing distally, mostly 1.5-3. 5 mm wide; flowers ascend-
ing to horizontal or nodding; filaments 3—4 mm long, inserted at least 4 mm from base of tube; anthers 2. 5-5. 5 mm long:
10a Plants of streambanks and marshes, often growing in water; perianth tube 8-9 mm long; flowers white with yellow
throat I. rivulicola
10b Plants of hillsides and flats; perianth tube 5-10 mm long; flowers white, pale blue, pale pink or slate-blue with white or yel-
low throat, horizontal or nodding:
11a Corm tunics of fine to medium-textured, netted fibres; corms bearing sessile cormlets at base (rarely on slender stolons);
perianth whitish or flushed palest blue with white throat; style dividing opposite middle of anthers, style branches extend-
ing between them I. flaccida
lib Corm tunics of soft, ± papery (not fibrous) layers, not accumulating significantly; corms bearing flattened, ribbon-like
stolons; perianth pink, bluish grey, slate blue or white; style dividing opposite anther tips, rarely beyond, style branches
extending above them I. sobolifera
Key to subspecies of Ixia sobolifera
la Perianth pink with yellow throat; perianth tube 9-10 mm long; anthers 3—4 mm long, yellow with dark longitudinal lines on anther
lobes subsp. carnea
lb Perianth white with pale yellow throat or slate-blue with white throat; perianth tube 5-7 mm long; anthers 2-3 mm long, uniformly yellow:
2a Perianth white with yellow throat; perianth tube 6-7 mm long subsp. albiflora
2b Perianth slate-blue with white or pale yellow throat; perianth tube 5-6 mm long subsp. sobolifera
TAXONOMY
1. Ixia rapunculoides Delile in Redoute, Les
Liliacees 8: t. 431 (1816). Type: South Africa, with-
out precise locality, collector unknown, illustration in
Redoute, Les Liliacees 8: t. 431 (1816). Epitype: South
Africa, Northern Cape, Klipkoppies, red clay, 9 August
1961, G.J. Lewis 5853 (NBG, epi., here designated; K,
PRE, iso.).
Plants mostly 150-400 mm high. Corm subglobose,
mostly 14—18 mm diam., with tunics of firm, wiry fibres
accumulating in dense mass, bearing small cormlets
at base. Leaves (2)3(4), lower 2 lanceolate to falcate,
mostly (3-) 7-1 2 mm wide, ± one quarter to half as long
as stem, margins thickened, plane or lightly crisped,
uppermost leaf sheathing stem below spike, rarely unifa-
cial in upper 10-30 mm. Stem erect, nodding above, with
several to many (up to 10) short, thread-like, ascending
to horizontal branchlets subtended by obtuse bracts and
prophylls ± l(-2) mm long. Main spike half-nodding,
mostly 3-5-flowered, lateral spikes (1)2-5 -flowered;
bracts translucent sometimes tinged brown especially in
fruit, outer with three dark veins, mostly 5-6 mm long,
3-toothed or bluntly 3-lobed, inner about as long, with
two dark veins and forked apically. Flowers ± hori-
zontally oriented to half nodding, pale blue (mauve or
blue-grey) with pale yellow cup, or pink with white cup,
faintly rose-scented or evidently unscented; perianth
tube 5.5-8.0(-10.0) mm long, widely funnel-shaped,
flaring in upper 3-5 mm; tepals subequal, ovate, 1 1—1 6 x
4. 5-6.0 mm, proximal ± 3 mm forming part of floral cup,
spreading at right angles to tube distally. Stamens paral-
lel; filaments 2-3 mm long, inserted 2-4 mm above base
of tube; anthers 2. 3-3. 5 mm long, upper half exserted
from tube but included in floral cup, pale yellow. Style
dividing shortly below anther tips, branches 0.7-1. 2 mm
long, not reaching anther tips. Flowering time : August to
mid-September, sometimes July. Figure 1A-C.
Distribution : widespread across the Western Karoo,
from the Langberg west of Loeriesfontein and the
Hantamsberg in the north across the Bokkeveld Plateau
and Roggeveld Escarpment as far south as Sutherland
(Figure 2).
Diagnosis and variation : Ixia rapunculoides, as cir-
cumscribed here, is fairly uniform across its range: it is
recognized by the horizontally oriented, mostly blue
flowers with a yellow cup (locally pale pink with a white
cup), short perianth tube mostly 5. 5-8.0 mm long, short
filaments 2-3 mm long and anthers 2. 3-3. 5 mm long
and half included in the perianth tube. In the vicinity of
the Hantamsberg, the species co-occurs with I. robusta,
another member of the complex, which was distinguished
at the varietal level by Lewis (1962) by the presence of a
short collar of fibres sheathing the base of the stem, and
three or sometimes four, rather than the usual two leaves
with expanded blades. These plants are restricted to dol-
erite outcrops along the eastern and northern end of the
Hantamsberg and wherever we have seen them, typical I.
rapunculoides grew nearby or among the I. robusta plants
but was always shorter in stature and with grey-blue
flowers, in marked contrast to the tall, pink-flowered I.
robusta. The occasional presence of three, rather than the
usual two expanded leaves, must be regarded as no more
than part of the pattern of variation and not as evidence
for intergradation with I. robusta. Leaf number occasion-
ally varies in typical I. rapunculoides, thus some speci-
mens of Lewis 5808 have three broad basal leaves (in two
specimens the uppermost of these partly sheaths the stem)
very like those of I. robusta-, and all four specimens of De
Vos 2561 have four leaves, the penultimate one partly to
almost entirely sheathing the stem. Some plants from the
northeastern end of the Hantamsberg (Moordenaarspoort
and the Farm Vanrhynshoek) stand out in the species in
their unusually narrow leaves (± 3 mm wide), narrow
tepals and single flowered branchlets (e.g. Snijman 2068 )
but at present seem merely to be depauperate rather than
representing a separate race or genotype.
In order to fix the application of the name Ixia rapun-
culoides, the type of which is a painting in Redoute’s
Les Liliacees that is not, as far as we know, associated
with preserved specimens and possibly of doubtful iden-
tity, we have designated an epitype. Plants from the
Bothalia 38,1 (2008)
FIGURE 1. — A-C, Ixia rapunculoides, Goldblatt & Porter 12730 (MO, NBG): A, whole plant; B, 1/s flower; C, bracts: outer (left), inner (right).
D-F, Ixia robusta, Goldblatt & Porter 12738 (MO, NBG): D, whole plant; E, 1/s flower; F, bracts: outer (left), inner (right). Scale bars: 10
mm. Artist: J.C. Manning.
Bokkeveld Escarpment near Nieuwoudtville match the
illustration best and of the collections available from
there, Lewis 5853 is representative of the species and
distributed in several herbaria.
Flowers of Ixia rapunculoides have been shown to be
pollinated by large-bodied Anthophorine bees, includ-
ing Anthophora diversipes and Pachymelus peringueyi
(Goldblatt et al. 2000). We have confirmed this initial
report at a second population of the species, south of
Nieuwoudtville, where only female Anthophora diver-
sipes individuals were captured visiting the flowers. The
reward for visiting bees is nectar, moderate amounts of
which are secreted from septal nectaries and retained in
the base of the perianth tube.
Selected specimens
(Collections marked with an asterisk* have a collar of
fibres around the stem base; those marked with a + denote
gatherings with at least one plant with four leaves.)
NORTHERN CAPE. — 3018 (Kamiesberg): summit of Langberg,
among dolerite boulders, 3 563' [± 1 050 m], (-DB), 5 September
2006, Goldblatt & Porter 12769 (MO, NBG, PRE). 3019 (Loeries-
fontein): Farm Taaiboskloof, NW of Loeriesfontein, clay flats, (-CA),
2 September 1982, Snijman 465 (NBG). 3119 (Calvinia): near Nieu-
woudtville, (-AC), September 1930, L. Bolus s.n. (BOL19590)\
Farm Rietfontein, dolerite hill next to road, (-BC), 5 September
2006, *Goldblatt & Porter 12774 (MO, NBG); 51 km on road from
Nieuwoudtville to Calvinia, (-BC), 1 September 2006, Goldblatt &
Porter 12730 (MO, NBG, PRE); Agter Hantam, near Groot Toring,
(-BC), 1 September 2006, *Goldblatt & Porter 12754 (MO, NBG); 5
Bothalia 38,1 (2008)
9
miles [8 km] S of Calvinia, (-BD), 21 July 1961, Lewis 5791 (BOL,
K, NBG, PRE); Hantamsberg slopes, (-BD), 4 September 2002,
Goldblatt & Porter 12162 (MO); Farm Vanrhynshoek, east base
of Hantamsberg, (-BD), 5 Sept. 2006, Snijman 2068 (NBG); Farm
Wilgebos, S of Calvinia, 10 August 1961, Lewis 5879 (K, NBG);
Kareeboomfontein, SW of Calvinia, (-DA), Hanekom 2364 (K,
MO. PRE); Bloukransberge, Farm Kneu east of escarpment, (-DB),
21 August 1975, Thompson 2501 (NBG, PRE). 3120 (Williston):
near Elandsfontein [between Middelpos and Calvinia], (-AD), 10
October 1928, + Hutchinson 731 (K); Perdekloof SE of Calvinia to
Middelpos, (-DD), Goldblatt & Porter 12786 (MO, NBG, PRE). 3220
(Sutherland); Farm Kompromis, between Middelpos and Sutherland,
(-AB), 5 September 1986, +De Vos 2561 (NBG); Roggeveld, Bo-
Visrivier road opposite Noudrif Farm, (-AB), 8 September 2006,
*+Goldblatt & Porter 12800 (MO, NBG); Roggeveld Escarpment,
Farm Kruisrivier, 3 km past Ouberg Pass turnoff, stony red-brown
clay, 1 270 m, (-AD), August 1986, *+Cloete & Haselau 81 (NBG);
Houdenbek, NW of Sutherland, 1 311 m. (-AD), 2 September 1973,
*+Oliver 4409 (NBG, PRE); Roggeveld, Sneeukrans, Uitkyk, 1 600 m,
(-AD), October 1920, Marloth 9876 (NBG, PRE).
2. Ixia robusta (G.J. Lewis) Goldblatt & J.C. Man-
ning, comb, et stat. nov.
I. rapunculoides var. robusta G.J. Lewis in Journal of
South African Botany 27: 74 (1962). Type: South Africa,
[Northern Cape], Moordenaarspoort, 26 miles NE of
Calvinia, 26 September 1952, G.J. Lewis 2503 (BOL!,
PRE!, SAM!).
Plants mostly 0. 5-1.0 m high. Conn subglobose,
mostly 14—18 mm diam., tunics of firm, wiry fibres accu-
mulating in dense mass, bearing small cormlets at base.
Leaves usually 4(5 or 6), lower 3 or 4 lanceolate to sub-
falcate, mostly 8-14 mm wide, ± one quarter to one third
as long as stem, margins thickened, usually plane, upper-
most 1 or 2 leaves sheathing stem below spike. Stem
erect, sheathed below ground by collar of fibres, with
up to 10 short, slender, ascending branchlets subtended
by truncate-apiculate bracts and prophylls 0.5-1.0(-2.0)
mm long. Main spike suberect, mostly 4— 6-flowered. lat-
eral spikes ascending, mostly 2-6-flowered; bracts trans-
lucent becoming tinged brown at tips in fruit, outer with
three dark veins, (6 — )8— 1 0 mm long, 3-toothed or bluntly
3-lobed, subequal or inner slightly longer, with two dark
veins and forked apically. Flowers ± horizontally ori-
ented, pale pink with white cup, faintly acnd-metallic-
scented or apparently unscented; perianth tube 7.5-10.2
mm long, widely funnel-shaped, flaring in upper 4-5
mm; tepals subequal, narrowly ovate to subspathulate,
(1 5-) 17-20 x 7-9 mm, proximal ± 3 mm forming part
of floral cup, spreading at right angles to tube distally.
Stamens parallel; filaments ± 4 mm long, inserted ±3.5
mm above base of tube; anthers 3.5-M.O mm long, half
exserted from tube but included in floral cup, yellow.
Style dividing opposite middle third of anthers, branches
1. 5-2.0 mm long, extending between anthers. Flowering
time\ late August to late September. Figure 1D-F.
Distribution-, locally in the western Karoo east and
north of the Hantamsberg, on flats and lower slopes in
heavy, dolerite clay, among dolerite boulders (Figure 3).
Diagnosis and variation'. Ixia robusta is recognized
by its tall stature, pink flowers with a white cup, and at
least three (and often four) lanceolate basal leaves that
are thick and leathery to almost succulent in texture.
The corms, which are difficult to remove from the rocky
ground, are large, and covered with wiry tunics, and
the underground part of the stem is enclosed by a collar
of fibres, a feature often not preserved when the deep-
seated corms are removed from the rocky ground.
Wherever we have seen Ixia robusta , typical I. rapun-
culoides has been growing nearby and sometimes among
plants of I. robusta , never with any sign of intergrada-
tion. Ixia robusta is always taller, has 3 or 4 expanded
leaf blades, and branches with more and larger, pink
flowers. Associated with the larger flower, the anthers are
± 4 mm long, the perianth tube is 7.5-10.2 mm long and
the style branches are 1. 5-2.0 mm long, contrasting with
anthers 2. 3-3. 5 mm long and style branches 0.7-1 .0 mm
long in I. rapunculoides. Lewis’s comment that what she
called var. robusta and var. rapunculoides favoured dif-
ferent habitats, the latter in flatter, more sandy situations,
is not fully borne out by our observations. We could
usually find I. rapunculoides growing among I. robusta
plants, although I. rapunculoides also grew in flat, less
rocky places where I. robusta did not occur. The mor-
phological differences between Lewis’s var. rapunculoi-
des and var. robusta , combined with their sympatry, syn-
chronous flowering and the absence of morphological
intermediates, provides convincing evidence that they
are separate species.
Although Ixia rapunculoides usually has blue-
mauve to grey-blue flowers, plants from west of the
Hamtamsberg and near Loeriesfontein have a pale pink
perianth, identically coloured to I. robusta and care must
be taken not to confuse I. robusta with pink-flowered /.
rapunculoides : the latter always has short, falcate leaves,
usually only two with expanded blades, and lacks a collar
of fibres around the stem base. Flower colour alone is not
a consistent difference between the two species but where
they co-occur, flower colour and scent always differ.
Additional specimens
NORTHERN CAPE. — 3119 (Calvinia); Calvinia District, near
Elandskop, Agter Hantam, 765 m, (-BB), 17 September 1956, Acocks
19038 (BOL, K, PRE); Agter Hantam, between Moordenaarspoort and
Klipwerf, (-BB), 1 September 2006, Goldblatt & Porter 12746 (MO,
NBG); Moordenaarspoort, northeast of the Hantamsberg, among doler-
ite boulders, (-BD), 1 September 2006, Goldblatt & Porter 12738 (K,
MO, NBG, PRE, S), 12741 (K, MO).
3. Ixia rivulicola Goldblatt & J.C. Manning, sp. nov.
Plantae 450-800 mm altae, cormo tunicis submem-
branosis basi cormo sessile ferenti, foliis (3)4 vel 5, infe-
rioribus 2-4 anguste ensiformibus ad linearibus 3-7 mm
10
Bothalia 38,1 (2008)
FIGURE 3. — Known distribution of Ixia robusta, O; 7. rivulicola, •;
and I. lacerata, A.
latis, spica 2 vel 3-flora, floribus albidis cupula pallide
flava intense odoratis, tubo perianthii campanulato 8-9
mm longo, tepalis subaequalibus 17-18 x 5. 0-7. 5 mm,
filamentis 3^1 mm longis, antheris 4. 5-5. 5 mm longis,
stylo tertiam partem superam antherarum adversus divi-
dend, ramis ± 0.8 mm longis.
TYPE. — Northern Cape, 3220 (Sutherland): Rogge-
veld Escarpment, drift across Bo- Vis River at Noudrif
Farm, in muddy ground, (-AB), 11 October 2004,
Goldblatt & Porter 12666 (NBG, holo.; K, MO, PRE,
iso.).
Plants 450-800 mm high. Corm subglobose, tunics of
submembranous fibres soon disintegrating, bearing 1-
few sessile cormlets at base. Leaves 3(4 or 5), lower 2^1
narrowly sword-shaped to linear, 3-7 mm wide, mar-
gins moderately thickened, hyaline when dry, straight,
usually ± one third to half as long as stem, uppermost
leaf sheathing stem, penultimate leaf often sheathing
in lower half. Stem with 3-5 straight lateral branchlets
mostly held at ± 30° to main axis, filiform, bearing flow-
ers in upper half. Main spike mostly 2- or 3-flowered;
lateral spikes 1- or 2-flowered; bracts translucent, outer
with three dark veins, mostly ± 8 mm long, inner with
two dark veins and forked apically. Flowers ascending
to nearly horizontal, white with pale yellow cup, with
strong fruity scent; perianth tube funnel-shaped, 8-9 mm
long; tepals subequal, 17-18 x 5. 0-7. 5 mm, proximal
3^4 mm forming part of floral cup, spreading at right
angles to tube distally. Stamens parallel; filaments 3^4
mm long, included, inserted ± 3 mm above base of tube;
anthers 4. 5-5. 5 mm long, lower half included in tube.
Style dividing opposite upper third of anthers, branches
± 0.8 mm long, extending between anthers. Capsules
and seeds unknown. Flowering time : mainly October.
Figure 4A-C.
Distribution'. Ixia rivulicola is apparently restricted
to the upper reaches of the Bo-Visrivier (Upper Fish
River) on the Roggeveld Escarpment (Figure 3). First
collected in 1981 by Harry Hall, this plant was not
known to Lewis (1962) at the time that she completed
her revision of Ixia, nor was it dealt with by De Vos
(1999). Revisiting Hall’s locality in the Roggeveld in
October 2004 we found plants growing in shallow water
close to the banks of the Visrivier near the entrance to
Voelfontein Farm. Further downstream at the drift across
the river at Noudrif Farm an extensive population was
flowering particularly well where the reeds and cattails
in the riverbed had been burned the past summer. The
plants are tallest when growing in water among reeds
but shorter when the surrounding vegetation is removed.
The essentially aquatic habitat of this species is unique
in the genus.
Diagnosis and variation : Ixia rivulicola is not sim-
ply a tall, white-flowered variant of 1. rapunculoides, for
the flowers are larger, with a tube 8-9 mm long, tepals
17-18 mm long, and anthers 4. 5-5. 5 mm long, com-
pared with the shorter tube, 5.5-8.0(-10.0) mm long in
I. rapunculoides, tepals 11-16 mm long, and anthers
2. 3-3. 5 mm long. The ascending to horizontally oriented
flowers are white with a pale yellow cup and are particu-
larly sweetly scented, the odour like that of Virgilia oro-
boides (Fabaceae), a combination of grenadilla fruit and
violets. Typical blue-flowered I. rapunculoides grows on
the slopes above the Visrivier and blooms in August and
early September, and is in fruit by the time I. rivulicola
comes into flower at the end of September or in early
October. Ixia rivulicola responds well to cultivation, and
survives outdoors in several degrees of freezing. Plants
reproduce liberally by vegetative reproduction, unlike
typical I. rapunculoides, although both produce cormlets
at the base of the main corm.
Like other members of the Ixia rapunculoides com-
plex, the flowers of I. rivulicola are pollinated by bees.
At the type locality they were actively visited by a range
of medium-sized and large-bodied bees, including Apis
mellifera, Amegilla spilostoma (both Apidae), Melitta
capensis (Melittidae), Megachile sp. (Megachilidae),
Plesanthidium calvini and Patel/apis sp. (both
Halictidae). Male and female bees were captured as well
as Apis workers, and all were found to carry Ixia- type
pollen on their bodies.
Additional specimens
NORTHERN CAPE. — 3220 (Sutherland): Roggeveld Escarpment,
Farm Voelfontein, in vlei in several inches of water, (-AB), 21 October
1981, Hall 5165 (MO, NBG).
4. Ixia lacerata Goldblatt & J.C. Manning, sp. nov.
Plantae 150-350 mm altae, cormo tunicis fibrosis,
foliis 3, inferioribus 2 linearibus 5-10 mm latis, floribus
± horizontaliter extensis pallide caeruleis in cupula pal-
lide flavis, tubo perianthii 10— 1 2(— 14) mm longis anguste
campanulato, tepalis subaequalibus ovatis ( 1 0—) 1 2—14 x
5-6 mm, filamentis ± 3 mm longis ± 6 mm supra basin
tubi insertis, antheris ± 3 mm longis partim supra ex
tubo exsertis, stylo mediam antherarum adversus divi-
dend, ramis ± 1 mm longis.
TYPE. — Western Cape, 3320 (Montagu): Klein Rogge-
veld, about 24 km north of N 1 on road to Sutherland,
dolerite slope, (-BA), 26 August 2006, Goldblatt &
Porter 12702 (NBG, holo.; K, MO, PRE, iso.).
Plants 150-350 mm high. Corm subglobose, 14—20
mm diam., with tunics of firm-textured, coarse fibres,
bearing small cormlets at base. Leaves 3, lower 2 lan-
ceolate to falcate, 5-10 mm wide, plane, sometimes
slightly twisted distally, ± one third to one quarter as
Bothalia 38,1 (2008)
11
long as stem, margins slightly to moderately thickened,
uppermost leaf sheathing stem in lower half to two
thirds. Stem erect, wiry and sometimes somewhat coiled
below spike, simple or with 1-3 short lateral branchlets,
branchlets thread-like, short and twisted, subtended by
filiform, forked or lacerate, distally spreading, silvery
bracts and prophylls up to 5 mm long. Main spike (2)3-
6-flowered, suberect, lateral spikes (l-)3- or 4-flow-
ered; bracts translucent below, becoming rust-coloured
in upper third, 6-7 mm long, outer mostly with 5 main
veins and 5 -toothed at apex, teeth attenuate, sometimes
slightly lacerate, inner ± as long as outer, with 2 main
and 2 smaller veins, with 2—4 attenuate teeth. Flowers
± horizontally oriented or suberect, pale blue-mauve,
often whitish at base, with pale yellow cup, weakly
rose-scented; perianth tube 1 0— 1 2(— 14) mm long; tepals
subequal, ( 1 0 — ) 1 2— 1 4 x 5-6 mm. Stamens parallel; fila-
ments ± 3 mm long, inserted ± 6 mm from base of tube;
anthers ± 3 mm long, yellow. Style dividing opposite
middle of anthers, branches ± 1 mm long. Capsules glo-
bose, ± 4 mm long. Seeds subglobose, shiny, ± 1.5 mm
at longest axis. Flowering time : mainly August to early
September. Figure 4D-F.
Distribution-. Ixia lacerata is restricted to slopes and
flats in the Klein Roggeveld, which lies south of the main
Roggeveld Escarpment, and the Koedoes Mountains
to the west (Figure 3). Plants mostly grow on shales or
mudstones of the Ecca System, and only occasionally on
rocky, dolerite slopes in heavy clay, the habitat in which
typical I. rapunculoides most frequently occurs.
Diagnosis and variation : until now the handful of
specimens resembling typical Ixia rapunculoides from
the Klein Roggeveld and Koedoes Mountains have been
referred either to this taxon or to I. namaquana. Careful
examination of flowers and bracts, however, shows that
these plants differ consistently in several respects. The
perianth tube is 10-14 mm long (vs 5.5— 8(— 10) mm in
var. rapunculoides and ( 1 2 — ) 1 3— 1 6(— 18) mm in I. nama-
quana)-, the outer bracts usually have five major veins
and the inner, two main and often two smaller veins,
with the veins terminating in attenuate teeth; and the
margins are dry and often somewhat tom. The bracts
in particular are highly diagnostic, contrasting with I.
rapunculoides and most other species of Ixia subgenus
Morphixia, which have 1- or 3 -veined outer bracts and
2-veined inner bracts, the veins terminating in short,
acute or blunt teeth. In addition to the floral differences,
the flowering stem has only 1-3 branchlets, each bear-
ing 1-3(4) flowers, and the branchlets are subtended by
forked, thread-like bracts and prophylls that are spread-
ing or directed downward distally, unlike the short, trun-
cate and inconspicuous bracts in most other members of
the I. rapunculoides complex, including I. namaquana
and I. rapunculoides itself.
Ixia lacerata is poorly represented in herbaria and
appears to have first been collected by the traveller-
explorer William Burchell, who found the species on the
Windheuwel in the Koedoes Mountains in July, 1811.
Surprisingly, Burchell’s collection attracted no botanical
attention, although at the time the only Ixia known with
short stamens and included filaments was I. rapuncu-
loides, described in 1816 and not then known from any
wild locality.
Our observations on the pollination of Ixia lacerata
made in September 2006 at two sites showed that the
most common insect visitor was an unnamed long-pro-
boscid fly, Prosoeca sp. (Nemestrinidae). This fly, also
the main pollinator of the long-tubed Romulea syrin-
godeoflora M.P.de Vos (Goldblatt & Manning 2007),
has a proboscis 10.5-11.5 mm long, a close fit for the
perianth tube of I. lacerata, the base of which contains
concentrated nectar, of 38—49% sucrose equivalents.
Because of the short stamens in the species, contact
between the anthers and the fly’s body would be limited
to the base of the proboscis. A less frequent visitor was
the bee Anthophora diver sipes (Apidae: Anthophorinae).
Anthophorine bees are common visitors to flowers of
members of the I. rapunculoides complex (Goldblatt et
al. 2000) and are assumed to be their most important
pollinators. The significance of the visits by Prosoeca
species remains to be determined.
Selected specimens
NORTHERN CAPE.— 3220 (Sutherland): Houthoek, Sutherland.
(-CA), 13 August 1968, Hanekom 1069 (K, PRE); Roggeveld
Escarpment, Farm Kraairivier, 5 km from Ceres turnoff on Sutherland-
Matjiesfontein road, yellow clay-gravel, (-CB), 25 August 1986,
Cloete & Haselau 8 (NBG); Windheuwel, Koedoes Mountains, (-CC—
CD), 22 July 1811, Burchell 1285 (K); 82 km S of Sutherland, rocky
ridge, (-DC), 31 August 1993, Goldblatt & Manning 9658 (NBG,
MO); valley ± 9.5 km below Komsberg Pass, Klein Roggeveld. (-DB),
8 September 2006, Goldblatt & Porter 12802 (MO, NBG, PRE); Klein
Roggeveld, Farm De Hoop S of Komsberg Pass, (-DC), 30 September
2004 (fruiting), Goldblatt & Porter 12659 (MO, NBG).
WESTERN CAPE.— 3220 (Sutherland): 19 miles [± 30 km] N of
Matjiesfontein, (-DC), 15 September 1955, Acocks 18440 (K, PRE); 1
km along turnoff to Komsberg Pass, (-DC), 26 August 2006, Goldblatt
& Porter 12708 (MO, NBG); north of Matjiesfontein on road to
Sutherland, Farm Nuwerus, (-DC), 14 September 2004, Snijman 1929
(NBG).
5. Ixia sobolifera Goldblatt & J.C. Manning, sp.
nov.
Plantae ( 150-)250-750 mm altae, cormo stolonifero
tunicis ± papyraceis, foliis 3, inferioribus 2 laminis lin-
earibus vel anguste ensiformibus (1.5-)2.0-5.0(-8.0)
mm latis, floribus nutantibus cameis vel ardesiaco-cae-
ruleis cupula alba vel pallide flava usitate fragrantibus,
tubo perianthii 5-10 mm longo infundibulari, tepalis
subaequalibus ovatis 10-17 x 4. 5-7. 5 mm, filamen-
tis 2-3 mm longis 3^4 mm supra basem tubi insertis,
antheris 3.0— 4.5 mm longis partis supra ex tubo exsertis,
stylo apicem antherarum adversus dividenti, ramis styli
1 .0-1.5 mm longis.
TYPE. — Northern Cape, 3320 (Montagu): north
of Matjiesfontein on road to Sutherland, (-BA), 9
September 2006, Goldblatt & Porter 12809 (NBG,
holo.; MO, PRE, iso.).
Plants ( 1 50— )250— 750 mm high. Conn subglobose,
8-12 mm diam., with soft, ± papery tunics, producing
flattened, corm-bearing stolons from base. Leaves 3,
lower 2 linear, often trailing above, ( 1 .5— )2.0— 5 .0(— 8.0)
mm wide, ± half as long as stem, margins plane, barely
thickened, edges flat or concave, uppermost leaf sheath-
ing stem below spike. Stem erect, with 1-5 twisted, fili-
form branchlets subtended by forked, attenuate bracts
and prophylls 2-5 mm long. Main spike nodding, 2-5(-
7)-flowered, lateral spikes nodding, (1)2— 4(5)-flowered;
12
Bothalia 38,1 (2008)
FIGURE 4. — A-C, lxia rivulicola, Goldblatt <£ Porter 12665 (MO, NBG): A, whole plant; B, 1/s flower; C, bracts: outer (left), inner (right). D F,
Ixia lacerata, Goldblatt & Porter 12702 (MO, NBG): D, whole plant; E, 1/s flower; F, bracts: outer (left), inner (right). G-I, lxia sobolifera
subsp. carrtea, Goldblatt & Porter 11810 (MO, NBG): G, whole plant; H, 1/s flower; I, bracts: outer (left), inner (right). Scale bars: 10 mm.
Artist: J.C. Manning.
bracts translucent, or becoming dry and brown distally
with age, outer with three dark veins, 6-10 mm long,
inner similar but with two dark veins and forked api-
cally. Flowers half to fully nodding, pink or slate blue
with white cup, or white with pale yellow cup, faintly
rose- to strongly carnation-scented; perianth tube 5-10
mm long, funnel-shaped, flaring in upper ± 5 mm; tepals
subequal, ovate or narrowly ovate, 10-17 x 4. 5-7.0 mm,
proximal ± 3 mm forming part of floral cup, spreading at
right angles to tube distally. Stamens parallel; filaments
2-3 mm long, inserted 3-4 mm above base of tube;
anthers 3. 0-4. 5 mm long, included or half exserted from
tube but included in floral cup, uniformly pale yellow or
marked with dark, longitudinal streaks. Style dividing
opposite or slightly beyond anther tips, branches 1.0-
1.5 mm long, usually exceeding anthers when mature.
Flowering time : August to mid-September.
Distribution : Ixia sobolifera extends from the Lang-
berg and Kubiskou ranges northwest of Loeriesfontein
in Northern Cape though the western Karoo and inte-
rior Western Cape to the Little Karoo (Figure 5). More
detailed distribution and habitat information is provided
under the subspecies.
Diagnosis and variation : included by Lewis (1962)
in Ixia rapunculoides var. flaccida, I. sobolifera is rec-
14
Bothalia 38,1 (2008)
on shale slopes but northwest of Nieuwoudtville, near
the Farm Biekoes, plants grow on sandy tillite soils
among sandstone rocks. The species is fairly common in
the rocky dolerite hills of the Nieuwoudtville Wildflower
Reserve and in similar habitats to the south on the farms
Glenlyon and Oorlogskloof.
Diagnosis and variation : apart from the distinctive
pink perianth, for which the taxon is named, an odd fea-
ture of subsp. carnea is the presence of dark longitudinal
lines on the anthers that contrast with the otherwise yel-
low colour. Plants are sympatric and bloom at the same
time as the longer-tubed I. namaquana northwest of
Nieuwoudtville and with 7. rapunculoides in the dolerite
hills east of Nieuwoudtville and on the Hantamsberg.
Additional specimens
NORTHERN CAPE.— 3018 (Kamiesberg): Langberg W of Loe-
riesfontein, wet gully, on shale, (-DB), 5 September 2006, Goldblatt
& Porter 12764 (MO, NBG, PRE). 3019 (Loeriesfontein): upper
slopes and summit of Kubiskou, in shale or dolerite, (-CD), 14
September 2006, Goldblatt & Porter 12820 (MO, NBG, PRE). 3119
(Calvinia): Klipkoppies, among rocks, (-AC), 9 September 1961,
Lewis 5862 (NBG); 4 miles [6.4 km] N of Nieuwoudtville, (-AC),
September 1930, L. Bolus s.n. ( BOL19589 ); between Grasberg and
Nieuwoudtville, (-AC), 8 August 1961, Lewis 5840 (NBG), [co-
blooming with I. namaquana ], 1 1 September 2004, Goldblatt & Porter
12406 (MO, NBG); dolerite koppies, Nieuwoudtville Wildflower
Reserve, (-AC), 12 September 2004, Goldblatt & Porter 12422 (MO,
NBG); Hantamsberg, lower S-facing slopes, (-BC), 4 September 1991,
Goldblatt 9169 (MO, NBG); Hantamsberg slopes, after fire, in tufts
of grass and among shrubs [sympatric and blooming with I. rapuncu-
loides], (-BD), 4 September 2002, Goldblatt & Porter 12161 (MO,
NBG).
5c. subsp. albiflora Goldblatt & J.C. Manning,
subsp. nov.
Plantae 400-650 mm altae, laminis foliorum anguste
ensiformibus vel linearibus 4-8 mm latis, caule 2 vel 3 (-5)
ramoso, spica 4— vel 5(-7) flora, bracteis usitate 6-8 mm
longis, floribus albis cupula pallide flava intense fragran-
tibus, tubo perianthii 6-7 mm longo, tepalis 12-17 x
5. 0-7. 5 mm, antheris ± 3 mm longis flavis.
TYPE. — Western Cape, 3322 (Oudtshoom): Farm
Buffelskloof, 35 km from Dysseldorp on Laudina road,
south-facing shale hillside, (-DA), 12 September 2003,
Goldblatt & Porter 12291 (NBG, holo.; K, MO, PRE,
iso.).
Plants 400-650 mm high. Leaves linear to narrowly
sword-shaped, 4—8 mm wide. Stem erect, with 2 or 3(-5)
suberect to spreading, filiform branchlets subtended by
silvery-translucent bracts mostly 3^1 mm long. Main
spike 4- or 5(-7)-flowered, lateral spikes (1)2- or 3(-5)-
flowered; bracts membranous and translucent, mostly
6-8 mm long. Flowers creamy white with pale yellow
cup, sometimes flushed grey-mauve outside, nodding,
intensely carnation-scented; perianth tube 6-7 mm long,
tepals 12-17 x 5. 0-7. 5 mm. Filaments ± 3 mm long;
anthers ± 3 mm, uniformly pale yellow; style reaching
anther tips. Flowering time : August to mid-September.
Distribution', mainly restricted to the central Little
Karoo, subsp. albiflora is known from just a handful
of sites, mostly south and east of Oudtshoom, with one
record from the northern foothills of the Swartberg,
east of Prince Albert (Figure 5). We also provisionally
include here a collection from Montagu ( Page s.n.),
which is isolated some 150 km west of the remaining
stations. The plants have white flowers, typical of albi-
flora, with a perianth tube ±7.5 mm long and anthers ±
4 mm long, but the corm has finely fibrous tunics resem-
bling those of I. flaccida and shows no evidence of sto-
lons, critical in recognizing 7. sobolifera. The bracts are
± 7 mm long, also consistent with 7. sobolifera rather
than 7. flaccida which has slightly shorter bracts, typi-
cally ± 6 mm long. The locality of the Page collection,
the Little Karoo, accords better with subsp. albiflora
than 7. flaccida, which occurs to the west in the Olifants
River Valley. Subsp. albiflora favours moist, south-fac-
ing slopes and grows in pockets of loamy soil on shale
in renosterveld. The habitat for the collection from
Montagu is from rocky slopes and cliffs, but the rock
type is not recorded.
Diagnosis and variation : subsp. albiflora is distin-
guished from its sister subspecies mainly by the white,
strongly carnation-scented flowers with a yellow cup,
and also by the 4- or 5(-7)-flowered main spike, peri-
anth tube 6-7 mm long (perianth tube 9-10 mm long
in subsp. carnea and 5-6 mm in subsp. sobolifera ), and
often broader leaves, 4-8 mm wide.
Lewis (1962) included the two collections of Ixia
sobolifera subsp. albiflora known to her (Page s.n. and
Thorne s.n.) in 7. rapunculoides var. flaccida (now 7.
flaccida), which occurs in the Olifants River Valley, well
to the west. As we have discussed above, this is distinct
from the stoloniferous 7. sobolifera, which also has larger
flowers with a longer perianth tube and longer anthers.
Additional specimens
WESTERN CAPE. — 3320 (Montagu): Montagu Baths, common
on rocky slopes and cliffs in gorge, (-CC), August 1920, Page s.n.
(BOL). 3321 (Ladismith): SW of Oudtshoom near Perdebont, (-DB),
12 August 2004, Vlok & Schutte 494 (MO, NBG). 3322 (Oudtshoom):
lower north slopes of the Swartberg, Farm Frisgewaagd, burned S-fac-
ing slope, 900 m, in loamy soil, (-AD), 12 September 1986, Vlok 1586
(MO, NBG, PRE); Oudtshoom, Grootkop, 440 m, deep loamy soil
on S-facing slope, rare, (-CA), 18 August 2005, Vlok & Schutte 506
(NBG); Oudtshoom, Kleinfontein, (-CA), August 1931, Thorne s.n.
{SAM51702)\ Farm Buffelskloof, 35 km from Dysseldorp on Laudina
road, south-facing shale hillside, (-DA), 12 September 2004, Goldblatt
& Porter 12484A (MO, NBG).
6. Ixia namaquana L. Bolus, South African Garde-
ning and Country Life 21: 368 (1931). 7. rapunculoides
var. namaquana (L. Bolus) G.J. Lewis: 76 (1962). Type:
South Africa, [Northern Cape], Klipfontein, September
18 83 , 77. Bolus as Herbario Normale Austro-Africana
698 (BOL!, lecto., designated by De Vos (1999: 15); B,
G, K!, PRE!, SAM!, Z, isolecto.).
Plants mostly 300-500 mm high. Corm 12-15 mm
diam., with tunics of medium-textured to coarse fibres.
Leaves 3, lower 2 sword-shaped to falcate, mostly 8-18
mm wide (particularly broad in the type), margins thick-
ened and hyaline, sometimes minutely crisped, usually ±
one third as long as stem but sometimes much shorter,
uppermost leaf sheathing stem. Stem with several short,
filiform, twisted lateral branchlets, subtended by short
bracts ± 2 mm long. Main spike erect, 2- or 3-flowered,
lateral spikes 1- or 2-flowered; bracts translucent, outer
with 3 dark veins, ± 10 mm long, with 3 short, sub-
Bothalia 38,1 (2008)
15
equal teeth, inner with 2 dark veins and forked at apex.
Flowers held ± horizontally, whitish to pale mauve, lilac,
pale blue or pink with yellow cup, rim of cup marked
with a band of short vertical lines, often flushed lilac
to mauve outside, violet-scented or unscented; perianth
tube 12— 1 6(— 1 8) mm long, narrowly funnel-shaped;
tepals subequal, ovate, ( 1 2—) 1 4 — 1 7 x 5-7 mm (inner),
6-8 mm (outer), proximal 4—6 mm forming part of floral
cup, spreading at right angles to tube in distal 10-12
mm. Stamens parallel; filaments erect, 4-5 mm long,
inserted ± 6-10 mm above base of tube; anthers 3.5— 4.5
mm long, upper half exserted from tube but included in
floral cup. Style mostly dividing opposite upper third
of anthers, branches ± 1 mm long, extending between
anthers (in the type, dividing at anther tips with branches
arching above anthers and 1.5 mm long). Capsules
ovoid, ± 9 mm long. Seeds unknown. Flowering time :
mainly August to mid-September, rarely October. Figure
6A-D.
Distribution : Ixia namaquana has a scattered dis-
tribution in Northern and Western Cape, and has been
recorded from the Richtersveld and the high ground
around Steinkopf in northern Namaqualand, from the
southern Kamiesberg in central Namaqualand, and
from the Bokkeveld Mtns and northern Cedarberg to
the south (Figure 7). The disjunct distribution between
the Steinkopf area of Namaqualand and the southern
Kamiesberg is a pattern shared by Brunsvigia pulchra
(Amaryllidaceae) and is probably related to the some-
what lower altitude and associated reduction in precipita-
tion of the country between these two high-lying points.
Another species pair with this distribution is Hessea
pilosula (Amaryllidaceae) from the Steinkopf area and
its sister species H. incana from the Kamiesberg. There
are no significant differences between the northern
Namaqualand populations of I. namaquana and those
from the south except for the bright pink flower colour
(rather than pale mauve to blue) in the type form, which
occurs on clay soils. In the Kamiesberg, I. namaquana
occurs on granitic gravel but in the Bokkeveld Mtns,
Gifberg and Cedarberg it grows in sands derived from
Cape Sandstone formations. Plants that we assign to /.
namaquana from the Wiedouw River ( Lewis 1980),
south of Vanrhynsdorp, probably grow on loam among
limestone outcrops, the main formation there but we
have not seen the species at this site ourselves.
Diagnosis and variation'. Ixia namaquana is readily
distinguished from other members of the I. rapunculoi-
des complex by the longer perianth tube, mostly 13-16
mm, short branchlets bearing 1 or 2 flowers each, and
corm with tunics of coarse, netted fibres. The flowers
are relatively large with tepals mostly 14—16 mm long
and anthers 3.5r4.5 mm long. The proximal part of the
tepals is directed forward, forming part of the floral
cup, which fully encloses the stamens. The flowers are
usually sweetly scented but are apparently unscented
in plants from Vanrhyn’s Pass and Nieuwoudtville in
the Bokkeveld Mtns. Pink-flowered plants with a peri-
anth tube 16-22 mm long from the Hex River Pass and
nearby that were identified as I. namaquana by De Vos
(1999) and others, are here referred to the new I. oxa-
lidiflora. This species can be distinguished from I. nama-
quana by the longer perianth tube, suberect rather than
horizontally spreading flowers, fully included anthers,
and perhaps most significant, by the corms with papery
to finely fibrous tunics and bearing ribbon-like stolons.
Plants from a seasonally wet site near Leliefontein in
the Kamiesberg, referred to Ixia namaquana by De Vos
in various herbaria, differ in their later flowering time,
October and November, and in having a white peri-
anth with tepals pale blue at the tips, filaments shortly
exserted, a perianth tube ± 7 mm long, and narrow
leaves. They are obviously misplaced in Ixia namaquana
and apparently represent a late-flowering ecotype of I.
latifolia var. ramulosa from moist habitats.
First collected by Harry Bolus in 1883 at Klipfontein
near Steinkopf in northern Namaqualand, Ixia nama-
quana was described by his niece, H.M.L. Bolus in
1931. The species was based on the Bolus collection and
one more ( Herre s.n.) from nearby, both having the large,
pink flowers that characterize the northern populations of
the species. The presence of pale lilac- to almost white-
flowered plants otherwise resembling /. namaquana from
the Bokkeveld Mountains, well to the south, was only
established later and Lewis (1962) included these as well
as the northern Namaqualand plants in I. rapunculoi-
des var. namaquana. In De Vos’s (1999) account of var.
namaquana, its range is extended to include the northern
Cedarberg ( Compton 24239), and the Laingsburg and
Worcester Districts, but no species are cited from the lat-
ter two areas, and we do not recognize the taxon from
south of the Cedarberg. The presence of I. namaquana
in the Kamiesberg of central Namaqualand was first
recognized by De Vos (1999); some plants from there
were referred by Lewis to the shorter-tubed var. rapun-
culoides. The Kamiesberg plants more closely resemble
those from the Bokkeveld Mountains rather than those
from northern Namaqualand. The type of I. namaquana
has particularly large flowers, with the tube up to 18 mm
long, and the style divides opposite the anther tips with
the branches extending above them. Specimens of other
northern Namaqualand collections are less well pressed
and it is impossible to say if the features of the type
plants are shared with all other specimens from the area.
The northern populations need more careful study.
Selected specimens
NORTHERN CAPE.— 2917 (Springbok): Richtersveld, Kalk-
fontein, (-AA), without date, Meyer s.n. (NBG1 78595); Klipfontein,
(-BA), without date, Marloth 12678 (PRE), 29 August 1935,
Compton 5411 (BOL, K, NBG); Kasteelpoort, Steinkopf, (-BA), 7
September 1929 (NBG); Steinkopf, western mountains, (-BA), 9
September 1929, Herre s.n. (STEU11846 in BOL, K, PRE); Steinkopf,
(-BA), 24 August 1959, Lewis 5494 (NBG); 4 km W of Steinkopf,
(-BA), September 1993, Williamson 5301 (NBG). 3018 (Kamiesberg):
Leliefontein, (-AC), 2 October 1947, Rodin 1474 (PRE); Kamiesberg,
Rondefontein, slopes south of Nartjiesdam, (-CA), 1 September 1975,
Oliver 5972 (NBG, PRE). 3119 (Calvinia): between top of Vanrhyn’s
Pass and Nieuwoudtville, (-CA), 23 August 1950, Lewis 2287 (SAM);
Nieuwoudtville, Glenridge, (-AC), 18 August 1960, Lewis 5725 (BOL,
K, NBG); Grasberg, NW of Nieuwoudtville, (-AC), 16 September
1961. Barker 9553 (BOL, MO, NBG, PRE); Grasberg road, NW of
Nieuwoudtville, Farm Biekoes, (-AC), 11 September 2004, Goldblatt
& Porter 12407 A (MO); Lokenburg, (-CA), 29 August 1941,
Esterhuysen 5755 (BOL, PRE); Oorlogskloof Nature Reserve (grid
A6), 680 m, sandy loam, (-CA), 4 August 1988, Pretorius 75 (MO,
NBG, PRE); top of Botterkloof Pass, (-CC), 21 July 1961 .Lewis 5789
(BOL, K, NBG, PRE).
WESTERN CAPE.— 3119 (Calvinia): Kobee Valley, (-CA),
1 September 2001, Goldblatt & Porter 11805 (MO, NBG). 3118
-• • ■ -■ lv- •"
r
Bothalia 38,1 (2008)
17
7. Ixia flaccida (G.J. Lewis) Goldblatt & J.C. Man-
ning, comb, et stat. nov.
I. rapunculoides var. flaccida G.J.Lewis in Journal of South
African Botany 27: 74 (1962). Type: South Africa, [Western Cape],
Boschkloof, 7 Aug 1896, R. Schlechter 8432 (BM, BOL!, S, K!, L,
MO!, PRE (two sheets)!, S, Z, syn.).
Plants 200-350 mm high. Corm subglobose, tunics of
medium-textured, netted fibres, bearing 1-few sessile,
basal cormlets and rarely a short stolon. Leaves 3(4),
lower 2(3) sword-shaped to linear, 4-9 mm wide, mar-
gins hardly thickened, straight, usually about half as long
as stem, uppermost leaf sheathing stem. Stem erect, slen-
der-filiform, with 4-6 loosely twisted, filiform branch-
lets up to 35 mm long, spreading at right angles to main
axis, subtended by minute, acute bracts and prophylls ±
1 mm long. Main spike 2-4-flowered, lateral spikes 1-3-
fiowered; bracts translucent light brown, outer with three
dark veins, mostly 5-6 mm long, inner as long or slightly
longer than outer, with two dark veins and forked at
apex. Flowers horizontally oriented, white or pale blue
with white to greenish cup, apparently unscented; peri-
anth tube funnel-shaped, 7-9 mm long; tepals subequal,
± 14.0 x 3. 5^1. 5 mm, proximal 3^1 mm forming part
of floral cup, spreading at right angles to cup distally.
Stamens parallel; filaments 2. 5-3.0 mm long, included
in tube, inserted 3-4 mm above base of tube; anthers ±
2.5 mm long, lower half included in tube. Style divid-
ing at mid-anther level, branches ± 1 mm long, extend-
ing between anthers. Capsules and seeds unknown.
Flowering time : late July to early September.
Distribution-, restricted to the Olifants River Valley
and nearby, Ixia flaccida grows on seasonally moist,
south-facing, sandy, or sometimes light, loamy clay
slopes, among rocks usually in the shade of shrubs and
small trees (Figure 8).
Diagnosis and variation : the flowers of Ixia flaccida
are relatively small, with tepals ± 14 x 3. 4-4. 5 mm, peri-
anth tube 7-9 mm long, and anthers ± 2.5 mm long. The
tunics of medium-textured to fine fibres and the ± linear
or narrowly sword-shaped, soft-textured leaves, as well
as the white or pale blue flowers, readily distinguish the
species from typical I. rapunculoides, which has blue
to blue-mauve or sometimes pink flowers with a yellow
cup.
In addition to the Olifants River Valley collections
that closely match the type of I. rapunculoides var. flac-
16° 18° 20° 22° 24° 26° 28°^
16° 18° 20° 22° 24° 26° 28°
FIGURE 8. — Known distribution of Ixia flaccida , •; I. oxalidiflora , ■.
cida, Lewis (1962) included pink-flowered plants from
the Bokkeveld Plateau and white-flowered plants from
the Little Karoo in the taxon. As outlined above, we
regard the Bokkeveld plants as I. sobolifera subsp. cor-
nea and the Little Karoo plants as I. sobolifera subsp.
albiflora. Both differ from var. flaccida in their corms,
which have soft-textured, short-lived tunics and produce
long, ribbon-like stolons. They also differ in some flo-
ral features, notably the longer bracts, 6-10 mm long (vs
5-6 mm long in I. flaccida) and, in subsp. carnea, a lon-
ger perianth tube 9-10 mm long and longer anthers 3-4
mm long.
The earliest record we have found of the species is
the collection made by P.A. Mader near Clanwilliam, ±
1874. A later collection made by Rudolf Schlechter in
1894 is from an unlikely locality, near Porterville, that
requires confirmation. These collections and the few
made later were referred to Ixia rapunculoides until
1962 when Lewis (1962) chose a second Schlechter col-
lection, made in 1896, from Boskloof [Boschkloof] east
of Clanwilliam, as the type of her new I. rapunculoides
war. flaccida.
Selected specimens
WESTERN CAPE.— 3218 (Clanwilliam): Clanwilliam, (-BB),
without date, Mader 193 (K), August 1905, H. Bolus 10619 (BOL);
near Clanwilliam Dam, (-BB), July 1948, Lewis 1981 (SAM); 9 miles
[21 km] S of Clanwilliam, damp slopes above road, (-BB), 26 August
1957, Lewis 5210 (NBG); N of Citrusdal at Marcuskraal turnoff, sandy,
S-facing bank, (-BD), 31 August 2005, Goldblatt 12678 (MO, NBG).
3219 (Wuppertal): Cedarberg, Langrug Farm, moist sand in shade,
(-AC), 21 August 1982, Viviers 508 (PRE). Doubtful locality: 3218
(Clanwilliam): sandy places near Porterville, 240 m, (-DD), 20 August
1894, Schlechter 4913 (MO, PRE).
8. Ixia oxalidiflora Goldblatt & J.C. Manning, sp.
nov.
Plantae 200-450 mm altae, cormo 6 — 8(— 1 0) mm diam.
stolonifero, tunicis fibrosis tenuibus, foliis 3, inferiori-
bus 2 anguste ensiformibus 6 — 1 2(— 1 5) mm latis, spica
2^1-flora, floribus pallide purpureo-cameis cupula alba,
inodoris, tubo perianthii 16-22 mm longo, tepalis sub-
aequalibus 14-18 x 6-7 mm, filamentis ± 4 mm longis,
antheris ±3.5 mm longis in tubo inclusis, stylo bases
antherarum adversus dividenti, ramis ± 1 mm longis.
TYPE. — Western Cape, 3319 (Worcester): Hex River
Pass, south-facing clay slopes, (-BC), 2 September 1992,
Goldblatt & Manning 9397 (NBG, holo.; MO, iso.).
FIGURE 7. — Known distribution of Ixia namaquana.
18
Bothalia 38,1 (2008)
Plants 200-450 mm high. Corm 6 — 8(— 1 0) mm diam.,
with broad, compressed stolons, tunics of fine netted
fibres, soon decaying. Leaves 3, lower 2 narrowly sword-
shaped to linear, fairly soft-textured, 6— 12(— 15) mm
wide, attenuate, uppermost leaf sheathing stem, margins
straight and not thickened. Stem usually with 1 or 2 fili-
form, twisted lateral branches up to 12 mm long. Main
spike 2-4-flowered, lateral spikes 1 or 2-flowered; bracts
translucent, outer with 3 dark veins, 10-12 mm long,
with 3 subequal acuminate tips, inner with 2 dark veins
and forked at apex. Flowers ascending to ± upright, pur-
ple-pink with ± white cup, unscented; perianth tube lb-
22 mm long, funnel-shaped; tepals subequal, ovate, 14-
18 x 6-7 mm, spreading at right angles to tube in distal
10-12 mm. Stamens parallel; filaments erect, ± 4 mm
long, inserted 8-10 mm above base of tube; anthers ± 3.5
mm long, included in tube, tips reaching up to 0. 5-2.0
mm below mouth of tube. Style dividing opposite middle
of anthers, branches ± 1 mm long, extending between
anthers. Capsules and seeds unknown. Flowering time :
mid-August to mid- September. Figure 6E-G.
Distribution : Ixia oxalidiflora is currently known from
two sites, the top of Hex River Pass, and near Tweedside
Station, ± 60 km to the east (Figure 8). It favours south-
trending slopes and clay or loamy soils, where it typi-
cally grows wedged in crevices of shale outcrops. The
purple-pink flowers with a white cup (the latter some-
times described as pale yellow) closely resemble in
shape and colouring those of co-occurring Oxalis hetero-
phylla (Oxalidaceae) which blooms at the same time.
Diagnosis and variation: the earliest record of the spe-
cies that we have traced is one made by F.A. Rogers in
1915. Early collections of Ixia oxalidiflora were referred
to I. rapunculoides var. namaquana without comment by
Lewis (1962) and De Vos (1999), although these speci-
mens equally resemble similarly large-flowered I. pau-
ciflora G.J.Lewis in general aspect. Careful examination
of the flowers, however, shows that they differ markedly
from both taxa in having the stamens fully included in
the perianth tube, with the anther tips reaching 0.2-0. 5
mm below the tube apex, and a perianth tube 16-22 mm
long. In I. namaquana, only the filaments and bases of
the anthers are included in the tube, which is shorter,
mostly 13-16 mm long, and in I. pauciflora, the tips of
the filaments are typically exserted from the tube. The
corms, which have soft-textured tunics and long stolons,
resemble neither species, and the lanceolate, soft-tex-
tured leaves are quite different to the linear, very nar-
row, firm leaves of I. pauciflora or the broader, leathery
leaves of I. namaquana. Ixia oxalidiflora is thus rec-
ognized by the combination of fully included stamens,
a perianth tube 16-22 mm long, soft-textured leaves,
6— 1 2(— 1 5) mm wide, and the small corms that bear long,
flat stolons. The flowers are also held upright, a feature
only apparent when plants are seen alive, thus unlike the
horizontal flowers of I. namaquana and I. pauciflora.
Selected specimens
WESTERN CAPE — 3319 (Worcester): Hex River Pass, south-fac-
ing shale slopes, (-BC), 13 September 2005 (late flowering), Manning
& Goldblatt 2992 (MO, NBG), 1 September 1963, Hardy 48 (BOL,
K, PRE); below summit of Hex River Pass, SE slopes on shale, 625
m, (-BC), 16 September 1974, Mauve & I. Oliver 152 (NBG, PRE);
between Osplaats and Tunnel Sidings, (-BC), August 1915, Rogers
16743 (BOL). 3320 (Montagu): hill approx. 2 km W of Tweedside
Station, in loam among rocks, 1 200 m, (-AB), 12 August 1988, Vlok
1989 (NBG, PRE); south-facing slopes of Memorial hill above cem-
etery, in light clay, (-AB), 31 Aug. 2007, Goldblatt & Porter 12924
(K, MO, NBG, PRE).
9. Ixia divaricata Goldblatt & J.C. Manning,
nom. nov., pro I. latifolia var. angustifolia G.J.Lewis in
Journal of South African Botany 28: 86 (1962). Type:
South Africa, [Western Cape] Witzenberg, middle east
face opposite Farm Rosendal, 26 November 1941, N.S.
Pillans 9790 (BOL, holo.!).
/. scariosa var. longifolia Baker: 165 (1892), in part. Types: South
Africa, [Western Cape], near Ceres, October 1873, H. Bolus 2621 (K!,
BOL!, SAM!); Avontuur, H. Bolus 2487 (BOL!, lecto., designated by
De Vos (1999), K!, = I. orientalis L. Bolus).
1. rapunculoides var. rigida G.J.Lewis: 77 (1962), syn. nov. Type:
South Africa, [Western Cape], Hex River Valley, 1 October 1893, P.
MacOwan in Herbario Normale Austro- Africana 1653 (BOL! [as Hex
River Pass, Bolus s.n. in Guthrie 3077], G, K! [as near Hex River East,
1500' (± 460 m)], MO! [as Hex River Valley, De Dooms, Oct. 1893, H.
Bolus s.n.], SAM!, Z).
I. rapunculoides var. subpendula G.J.Lewis: 77 (1962), syn. nov.
Type: South Africa, [Western Cape], between Groot River and Elands
Kloof, Oct. 1939, C.L. Leipoldt 3026 (BOL, holo.!).
Plants 300-500 mm high. Corm subglobose, tunics
of medium-textured, netted fibres, bearing 1-few ses-
sile cormlets at base. Leaves 3-5, lower 2-4 narrowly
sword-shaped to linear, (1.5-)3.0-5.0 mm wide, margins
moderately thickened, hyaline when dry, straight, usu-
ally about one third to half as long as stem, uppermost 1
or 2 leaves sheathing stem. Stem with (l-)3 straight lat-
eral branches mostly held at 30-45(-50)° to main axis,
filiform, bearing flowers in upper half. Main spike 4-7-
flowered, lateral spikes (1— )3— 5 -flowered; bracts trans-
lucent, outer with 3 dark veins, mostly 6-7 mm long,
inner with 2 dark veins and forked apically. Flowers
upright, whitish or pale to deep pink or light purple with
pale yellow cup, unscented or with faint sweet scent;
perianth tube funnel-shaped, 8-11 mm long; tepals sub-
equal, 11-14 x 5. 5-7.0 mm, proximal 3-4 mm forming
part of floral cup, subpatent. Stamens parallel; filaments
4-5 mm long, distal 1-2 mm exserted from tube, rarely
reaching only to top of tube, inserted ± 5 mm above
tube base; anthers 4. 0-5. 5 mm long, fully exserted from
tube, tips exserted from floral cup. Style dividing oppo-
site middle of anthers, branches 1-2 mm long, extend-
ing between anthers. Flowering time : from late August
at low elevations to October, November and December
at higher elevations. Figure 9.
Distribution: Ixia divaricata is scattered through the
mountains of the southwestern Cape, from Elandskloof,
east of Citrusdal in the southwestern Cedarberg, through
the Cold Bokkeveld to Ceres and Tulbagh (Figure 10).
Plants favour seasonally wet, stony sandstone flats and
rocky sites.
Diagnosis and variation: Ixia divaricata is unmistak-
able in its unusual, divaricate branching pattern, with dis-
tinctive, stiff, straight lateral branches held at angles of
30-50° to the main axis and bearing mostly 3-5 flowers
crowded in the distal half. The white or pink, or really
purple flowers are held erect, with the tepals loosely
spreading but not fully patent. Why this plant was treated
as var. subpendula of I. rapunculoides is puzzling for not
only is it distinctive in its branching pattern but the flow-
Bothalia 38,1 (2008)
19
FIGURE 9. — Ixia divaricata, Gold-
blatt & Porter 12610 (MO,
NBG). A, whole plant; B, 1/s
flower; C, bracts: outer (left),
inner (right). Scale bar: 10
mm. Artist: J.C. Manning.
ers are upright, not horizontal, and the anthers, 4. 0-5. 5
mm long, are usually fully exserted from the tube or
rarely have their bases included. Specimens with anthers
exserted 1-2 mm were often included in I. latifolia by
De Vos (1999) but true I. latifolia has larger, deep pink
flowers with a perianth tube 14-17 mm long, filaments ±
10 mm long, and broad basal leaves, mostly 10-18 mm
wide. We regard var. subpendula as a separate species,
which we name I. divaricata for the divaricate branch-
ing pattern. Although valid synonyms at varietal rank
20
Bothalia 38,1 (2008)
16° 18° 20° 22° 24° 26° 28°
I I I I l I I
FIGURE 10. — Known distribution of Ixia divaricata.
exist for the species (see above), we have chosen the
new name divaricata at species rank. The name I. angus-
tifolia is preoccupied in the genus by I. angustifolia
(Andrews) Klatt (1895), a later synonym of I. monadel-
pha (Lewis 1962). We prefer not to use the epithets sub-
pendula (based on I. latifolia var. subpendula) or rigida
(based on I. rapunculoides var. rigida) for the species
because the former is misleading and the latter is based
on an atypical type specimen.
The immediate relationships of Ixia divaricata are
most likely with I. latifolia. Confusion with that spe-
cies is due in part to some longer-tubed collections with
the tube 1 2—14 mm long (e.g. De Vos 2693 and Leipoldt
4070 ) that also have relatively short, broad leaves, up
to 18 mm wide (a feature of I. latifolia) but fairly slen-
der branches (characteristic of /. divaricata). These are
almost certainly hybrids between the two species, which
grow within sight of one another, although always on
different soils, I. latifolia favouring heavy clay or loam.
Ixia divaricata was associated historically with /.
latifolia , and one of the two specimens cited by Baker
(1892) under var. longifolia of I. scariosa, the name by
which /. latifolia was then known (the other specimen
is I. orientalis and is the lectotype of the name) is I.
divaricata. Although the type of I. latifolia var. angus-
tifolia, from the Witzenberg is I. divaricata, several
other specimens assigned here by Lewis (1962) and De
Vos (1999) are a different species, I. monticola, which
we describe elsewhere (Goldblatt & Manning in prep.).
Although these two taxa have been confused in the lit-
erature they may not be particularly closely allied. The
most important differences are that I. monticola has a
corm with membranous, non-accumulating tunics, few
or no branches, and four or five leaves. I. divaricata, in
contrast, has fibrous tunics of medium texture and a dis-
tinctive branching pattern, and plants from high eleva-
tions typically have two or three leaves in most speci-
mens, although sometimes more in those from middle
elevations.
Another synonym may be Ixia stellata (Andrews)
Klatt (1882), based on I. capitata var. stellata Andrews
(1802), but the quality and detail in the painting on
which the taxon must be based makes a firm identifi-
cation impossible and no preserved material has been
found at K and BM, the herbaria where they would most
likely have been preserved.
We also include the type of Ixia rapunculoides var.
rigida from the Hex River Valley in I. divaricata, which
it closely resembles in the relatively long straight
branches with distally crowded flowers, unlike the
remaining specimens assigned to var. rigida, which
have shorter, flexuose, horizontal branchlets and flow-
ers evenly spaced along their length. This collection is
the only record from the Hex River Valley, where we
assume it no longer occurs as a result of intensive viti-
culture there. Ixia divaricata is still common in the Cold
Bokkeveld, where it is found in sites presently protected
from agriculture, although below Gydo Pass in the Warm
Bokkeveld, it is now threatened by the expansion of
orchards from Ceres and Prince Alfred’s Hamlet to the
foot of the Pass.
Provisionally we also include in Ixia divaricata plants
from upper Moraine Kloof in the Hex River Mountains
( Helme 2864), flowering in January. They have a simi-
lar branching pattern but somewhat smaller, purple flow-
ers with a tube ± 5 mm long, tepals ±10 mm long, and
anthers ±3.2 mm long. The style divides at the top of
the anthers and has branches ±1.3 mm long that extend
above the anther tips, whereas in typical I. divaricata
the style divides opposite the middle of the anthers.
Unfortunately, the specimens lack corms and leaves
without which a fair decision about the status of this
plant, flowering some two months after other collections
of I. divaricata, cannot be made.
Pollination of Ixia divaricata accords with that for
other species of the I. rapunculoides complex. At our
study site for the species at Waboomsrivier in the Cold
Bokkeveld, flowers were visited and presumably polli-
nated by a range of medium-sized and large bees includ-
ing Lasioglossum sp. (Halictidae) and Anthophora diver-
sipes and Amegilla spilostoma (Apidae).
Selected specimens
WESTERN CAPE. — 3219 (Wuppertal): Cedarberg, valley between
Sneeuberg and Tafelberg, wet sandy flats, (-AC), 19 October 1923,
Pocock 408 (NBG); Elandskloof, Ceres, (-CA), 30 September
1944, Barker 3094 (BOL, NBG); Cold Bokkeveld, Waboomsrivier,
sandy vlakte, (-CC), 4 October 1966, Hanekom 776 (K, PRE); Cold
Bokkeveld, 3 miles S of Leeurivier, (-CD), September 1952, Lewis
2512 (PRE, SAM). 3319 (Worcester): Great Winterhoek, Sneeugat,
(-AA), November 1916, Phillips 1874 (NBG); Steindaal, Tulbagh,
October-November 1858, Pappes.n. SAM20937 (NBG); Tulbaghskloof
(Neuwekloof), Tulbaghsthal, foot of the Winterhoeksberg, Witsenberg
and Vogelvalei, (-AA), November 1830, Ecklon & Zeyher lrid. 94
(BOL, K, PRE); Cold Bokkeveld, roadside at Farm Wadrif, in rocky
sandstone ground, (-AB), 30 September 2004, Goldblatt & Porter
12604 (MO, NBG); Schurfdeberg Pass, wet place, (-AB), 30 October
1950, Compton 22277 (NBG); stony flats at the foot of Gydo Pass,
(-AB), 4 October 2004, Goldblatt & Porter 12610 (MO, NBG, PRE);
Witzenberg, Visgat Valley, Farm Wakkerstroom, (-AC), 5 November
2003, Low 9376 (NBG); near Ceres Nature Reserve in wet ground,
(-AD), 8 October 1986, De Vos 2675 (NBG, PRE); top of Michell’s
Pass, (-AD), without date, Marloth 10692 (PRE); Matroosberg, near
Lakenvlei, (-AD), without date, Phillips s.n. SAMI 1876 (NBG);
Ceres, Schoonvlei Industrial Area, (-AD), 28 September 1987, Cloete
& Cillie 103 (NBG, PRE); Hex River Mtns, upper Moraine Kloof
Amphitheatre south of Milner Peak, 1 050 m, (-AD), 4 January 2004,
Helme 2864 (NBG).
Putative hybrids with Ixia latifolia : 3319 (Worcester): Ceres,
Gydouw, (-AB), 3 October 1942, Leipoldt 4070 (PRE); between Ceres
and Hottentotskloof, on Conradie farm, (-AD), 1 8 September 1 987, De
Vos 2691 ( PRE).
Bothalia 38,1 (2008)
21
10. Ixia contorta Goldblatt & J.C. Manning, sp.
nov.
Ixia rapunculoides var. rigida sensu G.J. Lewis: 77 (1962), but
excluding the type.
Plantae (70-) 100-300 mm altae, cormo subgloboso
10-14 mm diam. tunicis fibrosis duris nigrescentibus
persistentibus, foliis 3 inferioribus 2 anguste ensiformi-
bus vel falcatis (2)3-5 mm latis marginibus moderate
incrassatis, caule 1-3-ramoso, ramis contortis, spica
flexuosa inclinata vel subhorizontali, 2-6 flora, floribus
purpureis cupula flava, tubo perianthii 9-13 mm longo,
tepalis subaequalibus 9-12 x (4.5-)6.0-8.0 mm, fila-
mentis ± 3 mm longis in tubo inclusis, antheris 3^4 mm
longis.
TYPE. — Western Cape, 32 1 9 (Wuppertal): Cold Bokke-
veld, low hill east of Farm Waboomsrivier, well-drained
sandy ground, (-CD), 17 September 2006, Goldblatt &
Porter 12854 (NBG, holo.; MO, PRE, iso.).
Plants (70-) 100-300 mm high. Corm subglobose, 10-
14 mm diam., with tunics of medium-textured to fine,
dark, wiry, netted fibres, bearing 1-few sessile cormlets
at base. Leaves 3, lower 2 linear to narrowly sword-
shaped or falcate, (2-)3-5 mm wide, margins and midrib
moderately thickened, straight, about one third as long
as stem, uppermost leaf sheathing lower half of stem.
Stem suberect, usually with 1-3 short, twisted branchlets
mostly held at 60-90° to main axis. Main spike conspicu-
ously flexuose, bent at base and inclined to ± horizontal,
2-6-flowered, lateral spikes 1-6-flowered; bracts translu-
cent, often flushed brown or purple, outer with three dark
veins, mostly 7-9 mm long, bluntly 3-lobed apically,
inner with two dark veins and forked apically. Flowers
upright, purple (or said to be blue) with a yellow cup
sometimes edged with a band of dark purple, unscented;
perianth tube funnel-shaped, 9—1 3(— 14) mm long; tepals
subequal, 9-12 x (4.5-)6.0-8.0 mm. Stamens parallel;
filaments ± 3 mm long, included, inserted 4—5 mm above
base of tube; anthers 3^1 mm long, lower half to one
third included in tube. Style dividing opposite middle
of anthers, branches ± 1 mm long, extending between
anthers. Flowering time : mid-September to October.
Figure 11.
Distribution : Ixia contorta is centred in the Cold
Bokkeveld but extends north to the Cedarberg and east
to Touws River, growing in well-drained, sandy ground
(Figure 12).
Diagnosis and variation : treated by Lewis (1962) as
Ixia rapunculoides var. rigida, this plant from the inte-
rior Western Cape mountains is recognized among those
species with included filaments by the markedly flexuose
spikes and by the main axis strongly flexed below the
first flower, thus inclined to nearly horizontal. The pur-
ple flowers (also sometimes described as blue or lilac)
have a yellow cup sometimes edged in dark purple, and
are held upright on the spikes, unique among the species
with included filaments and anthers held partly within
the perianth tube. The perianth tube is 9- 13 (-14) mm
long and the tepals are 9-12 mm long. The very different
appearance of the flowers suggests that these plants are
not immediately allied to I. rapunculoides and we not
only regard them as representing a separate species, but
FIGURE 11. — Ixia contorta, Goldblatt & Porter 12854 (MO, NBG).
A, whole plant; B, 1/s flower. Scale bar: 10 mm. Artist: J.C.
Manning.
question their relationship to the complex. We describe
this plant here as I. contorta, so named for the twisted
lateral branchlets and flexuose spike, a feature par-
ticularly well-developed in this species. The new name
22
Bothalia 38,1 (2008)
is required because the type of var. rigida is I. divari-
cata (see synonymy of the latter species). Ixia contorta
can be distinguished from I. divaricata by the short,
twisted branchlets, quite different from the long, straight
branches of I. divaricata , the flowers of which are white
to pale pink, rarely purple, and the filaments are usually
shortly excluded from the tube.
Plants are usually fairly short, seldom exceeding 150
mm, but collections from Gydo Pass (e.g. Leipo/dt 3023 )
are taller, some reaching 300 mm. Typical Ixia contorta
has also been collected on Gydo Pass, which indicates to
us that the taller plants with narrow leaves are not sim-
ply a local variant of the species. We suspect they are
hybrids, the result of crossing with I. divaricata, which
also grows there. The longer, narrower leaves, paler
flower colour and slightly larger perianth in these plants
may also reflect the influence of genes of I. divaricata.
Plants from the north of the range near Alberta Farm,
and in the Matjiesrivier Valley in the Cedarberg stand
out in having a longer perianth tube, 12-13 mm in Nanni
81, and 12-14 mm in Gillett 4108. The latter plants,
also exceptional in their slightly wider perianth tube,
were referred to I. rapunculoides var. rigida by Lewis
( 1 962) but evidently puzzled De Vos ( 1 999), who iden-
tified a duplicate of the Gillett collection at PRE as /.
rapunculoides var. namaquana (i.e. /. namaquana). The
latter taxon differs particularly in having flowers facing
to the side, rather than upright, longer floral bracts, and
a straight and erect rather than flexuose, inclined spike
axis. The broad leaves, up to 10 mm in one plant of the
Gillett collection, are exceptional for I. contorta and it
remains possible that the gathering represents a novelty.
Additional collections of I. contorta from the north of its
range are needed to better assess the variation in the spe-
cies.
Selected specimens
WESTERN CAPE.— 3219 (Wuppertal): Clanwilliam, 2 miles
[3.2 km] from top of Uitkyk Pass in Matjies River Valley, (-AC), 9
October 1938, Gillett 4108 (BOL, K, PRE); Cold Bokkeveld, Farm
Onverwaght on road to Alberta, (-BA), 14 October 1994, Nanni 81
(MO); Ceres District, Luiperdskloof 4x4 route near Bloukop, 1 350 m,
(-CB), 13 September 2002, Steyn 174 (PRE); Koue Bokkeveld, Farm
Bokkenfontein, sandy flats near Leeu River, flowers blue, (-CC), 16
September 1976, M. Thompson 2953 (NBG, PRE); Cold Bokkeveld,
Farm Floudenbek, 850 m, (-CD), 13 October 1921, Marloth 10613
(PRE). 3319 (Worcester): Gydouw Pass, (-AB), October 1926, Marloth
12965 (PRE); Gydouw, (-AB), September 1941, Leipoldt 3821 (BOL);
Cold Bokkeveld, between Loch Lynne and Winkelhaak, sandy flats,
(-AB), 11 October 1974, Oliver 51 22A (NBG, PRE). 3320 (Montagu):
Touws River, sandstone koppie, (-AC), September 1924, Levyns 858
(BOL); dry sandy flats W of Touws River, (-AC), 27 September 1924,
Rennie s.n. (BOL).
Putative hybrids with Ixia divaricata : 3319 (Worcester): Ceres,
Gydouw, (-AB), October 1939, Leipoldt 3023 (BOL, K, NBG, PRE,
SAM).
ACKNOWLEDGEMENTS
)
Support for this study by grants 7103-01 and 7799-
05 from the National Geographic Society is gratefully
acknowledged. Collecting permits were provided by the
Nature Conservation authorities of Northern Cape and
Western Cape, South Africa. We thank Ingrid Nanni and
Lendon Porter for their assistance and companionship
in the field and C.D. Michener for identifying bees col-
lected during our study.
REFERENCES
ANDREWS, H. 1802. Ixia capitata var. stellata. The botanists s repo-
sitory 4: t. 232. London.
BAKER, J.G. 1892. Handbook of the Irideae. Bell, London.
BOLUS, H.M.L. 1931. Plants — new and noteworthy. South African
Gardening and Countty Life 2 1 : 368.
DAVIS, P.H. & FIEYWOOD, V.F1. 1973. Principles of angiosperm tax-
onomy. Kriegler, Huntington, New York.
DE VOS, M.P. 1999. Ixia. In M.P de Vos & P. Goldblatt in Flora of
southern Africa 7, part 2, fascicle 1 : 3-87.
DELILE, A.R. 1816. Ixia rapunculoides. In J.P. Redoute, Les Liliacees
8: t. 431. Paris.
GOLDBLATT, P„ BERNHARDT, P. & MANNING, J.C. 2000.
Adaptive radiation of pollination mechanisms in Ixia (Iridaceae:
Crocoideae). Annals of the Missouri Botanical Garden 87: 564 —
577.
GOLDBLATT, P. & MANNING, J.C. 1999. New species of Sparaxis
and Ixia (Iridaceae: Ixioideae) from Western Cape, South Africa,
and taxonomic notes on Ixia and Gladiolus. Bothalia 29: 59-63.
GOLDBLATT, P. & MANNING, J.C. 2007. Pollination of Romulea
syringodeoflora (Iridaceae: Crocoideae) by a long-proboscid fly,
Prosoeca sp. (Diptera: Nemestrinidae). South African Journal of
Botany 73: 56-59.
GOLDBLATT, P. & MANNING, J.C. In prep. Systematics of the south-
ern African Ixia subgenus Morphixia (Iridaceae). 2. Ixia latifolia
and its close allies.
KLATT, F.W. 1882. Erganzungen und Berichtigungen zu Baker’s
Systema Iridacearum. Abhandlungen der Naturforschenden
Gesellschaft zu Halle 15: 44-404.
KLATT, F.W. 1 895. Irideae. In T. Durand & H. Schinz, Conspectus flor-
ae africae, 5. Brussels.
LEWIS, J. 1962. South African Iridaceae. The genus Ixia. Journal of
South African Botany 27: 45-195.
MANNING, J.C., GOLDBLATT, P. & SNIJMAN, D A. 2002. The color
encyclopedia of Cape bulbs. Timber Press, Portland, Oregon.
STUESSY, T.F. 1990. Plant taxonomy. Columbia University, New
York.
Bothalia 38,1: 23-29(2008)
Three new species of Asparagus (Asparagaceae) from South Africa,
with notes on other taxa
S.M. BURROWS* and J.E. BURROWS*
Keywords: Asparagaceae, Asparagus L., South Africa, taxonomy
ABSTRACT
Three new species of Asparagus L. are described from South Africa: Asparagus elephantinus S.M. Burrows, A. hirsutus
S.M. Burrows and A. sylvicola S.M. Burrows; and the past confusion between Asparagus acocksii Jessop and A. lynetteae
(Oberm.) Fellingham & N.L.Mey. is discussed.
INTRODUCTION
Species within the genus Asparagus are notoriously
difficult to identify. The plants are unappealing to col-
lect, and when collected, not enough attention is paid to
the plant’s habit, and especially the rooting system. In
many instances not enough material is collected, making
the identification of the plants difficult. During extensive
field research and herbarium work for a taxonomic revi-
sion of the genus in southern Africa, a number of unde-
scribed taxa have been noted, and the distribution of
several species has been extended. Type localities have
been revisited wherever possible, and material has been
collected from the type locality or as close to the locality
as possible for each taxon. This has been done to ensure
that the correct concept of the taxa is understood.
Since the Buffelskloof Herbarium has yet to attain an
international herbarium acronym, the temporary abbre-
viation BNRH will be used throughout.
Asparagus elephantinus S.M.Burrows, sp. nov.,
A. densiflori similis sed caulis longissimis validis, ramis
ramulisque longis, tuberibus rotundis vel late ellipso-
ideis, spinis validis, pedunculo infra medio articulato,
fructu maiori differt.
TYPE. — Limpopo, 2430 (Pilgrim’s Rest): above J.G.
Strydom Tunnel, Abel Erasmus Pass, R36, Ohrigstad,
(-BC), 8 January 2005, J.E. Burrows & hotter 8781
(PRE, holo.; BNRH, iso.).
Scrambling, multistemmed shrub, evergreen, usu-
ally shallow-rooted, not twining, glabrous throughout.
Rhizome small; roots terete, up to 8 mm diam. with
numerous side rootlets; lateral tubers off thick main
roots, round to broadly ellipsoid, 30-51 x 22^10 mm.
Stems robust, 10-15 mm thick, up to 3 m long, bright
green, completely glabrous, faintly ribbed towards base.
Spines large on stems, up to 20 mm long, smaller else-
where, hard, sharp, straight to curved downwards, pale
orange brown; present below branches, branchlets, clad-
ode fascicles and flowers. Leaf scales on stem triangular,
1.0-5. 5 mm long, very pale with darker median stripe,
* Buffelskloof Nature Reserve Herbarium, RO. Box 710, 1120 Lyden-
burg, South Africa.
MS. received: 2007-06-20.
margins entire, depauperate to absent on branches and
branchlets. Branches spreading, up to 0.6 m long, faintly
papillose-ribbed. Branchlets spreading, up to 150 mm
long, distinctly papillose-ribbed, 5-angled, slightly zig-
zagging. Cladodes 1 — 3(— 6) per fascicle, held at an angle
to branches, flattened, narrowly elliptic to linear, (5-)8-
25(-31) x 1. 5-2.0 mm long, bright green, glossy, base
constricted with a basal foot, margins papillose. Racemes
50-70 mm long, arising from branch or branchlet nodes,
simple or branched, an occasional cladode sometimes
present. Flowers 1-3 per fascicle, sweetly scented. Pedi-
cel 1. 5-3.0 mm long, articulated in lower half. Tepals
6, 3.0—4. 2 x 1.5-1. 9 mm, apex curved inwards, margin
entire. Stamens 6; filaments 2.25-2.65 mm long, fused
length 0.75 mm, free length 1.5-1. 9 mm; anthers orange.
Style 0.4-0. 5 mm long; stigma 3-fid, arms 0. 1-0.2 mm
long. Ovary oblong-ellipsoid, trilocular, 1. 8-3.0 x 1.1-
2.0 mm. Fruit a berry, 2- or 3-lobed, 10-14 mm diam.,
glabrous, red when ripe. All measurements of cladodes,
leaf scales, flowering and fruiting parts taken from pick-
led specimens. Flowering time : January to February.
Figure 1 .
Distribution and habitat : Asparagus elephantinus
is currently known from the Olifants River Valley of
Limpopo Province, from Penge to the Abel Erasmus
Pass (Figure 2), where it occurs mainly on dolomite or
the quartzitic outcrops adjacent to the dolomite forma-
tions. This very robust plant, with 3 m long stems, grows
on rocky ledges, or among rocks in leaf litter. It is prin-
cipally confined to vegetation type SVcb 25, Poung
Dolomite Mountain Bushveld (Mucina & Rutherford
2006). This asparagus is widespread in its area, and is
well protected since it favours rocky ledges and inacces-
sible cliff faces. It is surprising that such a large species
of Asparagus has remained undetected for so long in
such a well-visited area as the Abel Erasmus Pass.
Diagnostic characters and affinities : Asparagus
elephantinus is similar to A. densiflorus (Kunth) Jessop
in that they both have flattened cladodes, with both the
cladodes and branches clearly papillate-ribbed. However,
A. elephantinus differs from A. densiflorus (Table 1).
Etymology. Asparagus elephantinus is named after
the Olifants River Valley, to which it is currently con-
fined; in addition, the name highlights the large size of
the plant.
24
Bothalia 38,1 (2008)
FIGURE 1 . — Asparagus elephantinus, A-G, J-M, J.E. Burrows & Lotter 8781 ; H, I, J.E. & S.M. Burrows 9034. A, stem and first branch; B, rhi-
zome habit; C, raceme; D, cladodc base; E, leaf scale; F, cladode and margin enlargement; G, c/s cladode; H, I, cladodes. J-L, floral details:
J, flower; K, stamen and tepal; L, ovary. M, fruit. Scale bars: A, 12.2 mm; C, F, 3.7 mm; D, 0.3 mm; E, 1.5 mm; G, 0.46 mm; H, 4.9 mm;
I, 1.2 mm; J, K, 0.9 mm; L, 0.6 mm; M, 2.4 mm. Artist: Sandra Burrows.
Specimens examined
LIMPOPO. — 2430 (Pilgrim’s Rest): pass W of Penge on road to
Chuniespoort, 1 200 m, (-AC), 6 March 2005, J.E. & S.M. Burrows
8914 (BNRH); N bank of Olifants R. near new bridge, Penge Mine,
800 m, (-AD), 11 November 2002, J.E. Burrows & M.C. Lotter 7878
(BNRH, PRE); ± 4 km N of Penge, next to Olifants R., 600 m, (-AD),
13 November 2005, J.E. & S.M. Burrows 9304 (BNRH); just above
J.G. Strydom Tunnel, Abel Erasmus Pass, 679 m, (-BC), 28 August
2002, J.E. & S.M. Burrows 7770 (BNRH, PRE).
Bothalia 38,1 (2008)
25
FIGURE 2. — Known distribution of Asparagus elephantinus, •; A.
hirsutus, I; and A. sylvicola , ▲.
Asparagus hirsutus S.M.Burrows, sp. nov., A.
capensi var. capensi similis sed tuberibus ellipsoideis,
ramis non spinescentibus, pilis omnino albis et squamis
foliosis glabris differt; A. minutiflori similis sed pilis
omnino albis, margine squamarum foliosarum integra et
pedicello hirsuto in parte superiore articulato differt.
TYPE. — Limpopo, 2429 (Zebedelia): Atok Platinum
Mine, Zeekoegat Farm, Monametse Village, east of
mineshaft, hill slope, 816 m, (-BD), 16 November 1999,
L.M.D. Riddles 511 (PRE, holo.; BOL, BNRH,J, KEW,
PRU, UNIN, iso.).
Erect, multistemmed, compact, very hairy, evergreen
shrublet of foxtail form, olive to grey-green, 300-500
mm high. Rhizome woody; roots 0. 1-5.5 mm diam.; with
a few lateral ellipsoid tubers present, 36-40 x 14—21
mm; cataphylls present. Stems 1. 2-5.0 mm thick, faintly
ribbed, pale, terete, covered in white hairs, skin peeling
in patches in older specimens to smooth below. Spines
1. 2-6.0 mm long, straight, thin, facing sideward or down-
ward, hairy, only present below branch axils on main
stem. Leaf scales triangular, 1.0-3. 2 x 1.0-1. 5 mm, grey
with dark blotches, becoming smaller distally, and finally
disappearing. Branches 1-5, 117-134 mm long, aris-
ing in clusters from stem, curving upwards, very hairy.
Branchlets absent. Cladodes (6 — )8 — 10 in fascicle, up to
36 in terminal fascicles, clavate to linear, ( 1 .5— )4.0— 6.4
(-10) x 0.5 mm long, olive green, slightly curved, apex
apiculate, 0.5 mm long, 6- or 7-angled, with simple white
hairs (0.2 mm long) along angled ridges, hairs radiating
out to side alternately. Flowers single, borne in cladode
fascicle, no scent detected. Pedicel 0. 5-2.0 mm long,
articulated in top half, hairy, base surrounded by cluster
of small membranous scales. Tepals 6, 3 larger 2. 1-2.5
x 1.0-1. 5 mm, 3 smaller 2-2.1 x 0. 9-1.0 mm, white,
median stripe brownish, with a ridge of hairs, apex acute,
margin sometimes serrated. Stamens 6; filaments 2. 6-3. 5
mm long, fused length 1.5 mm, free length 1.1- 2.0 mm;
anthers orange. Style 1 mm long; stigma 3-fid, arms 0.1-
0.2 mm long. Ovary trilocular, 1.0-1. 5 x 1 mm. Fruit a
berry, 5-7 mm diam., red when ripe, with persistent peri-
anth. All measurements of cladodes, leaf scales, flowering
and fruiting parts taken from pickled material. Flowering
time : September to November. Figure 3.
Specimens examined
LIMPOPO. — 2429 (Zebedelia): Ga Makopane, Zeekoegat Farm
421 KS, near Atok Platinum Mine, hill slope, 850 m, (-BD), 15
October 1997, N.H.G. Jacobsen 5540 (PRE); Zeekoegat Farm 421
KS. near Atok Platinum Mine, 850 m, (-BD), 10 November 1997,
ex cultivation, N.H.G. Jacobsen 5717 (PRE); hills above Atok Mine,
Monametsi R., Sekhukhuneland, (-BD), 29 November 2005, J.E. &
S.M. Burrows 9307 (BNRH, PRE).
Distribution and habitat : this small, extremely hairy
asparagus is currently known only from the hills near the
Atok Platinum Mine at Monametse in Sekhukhuneland
(Figure 2). The plants grow on rocky hillsides, amongst
small rocks, in full sun, or occasionally in the scant
shade of the surrounding vegetation, on medium- to
coarse-grained titanite-bearing tonalite (granite), in veg-
etation type SVcb 28, Sekhukhune Mountain Bushveld
(Mucina & Rutherford 2006). The plant is threatened
and is therefore afforded VUD 1 & 2 conservation status
(D. Raimondo pers. comm. 2006).
Diagnostic characters and affinities', although Aspara-
gus hirsutus is compared with A. capensis L. var. cap-
ensis and A. minutiflorus (Kunth) Baker, the taxon dif-
fers significantly in the following details: A. hirsutus is
an erect foxtail, with straight stems; the tubers are ellip-
soid; the main stem is covered in white hairs; the spines
are densely hairy; the leaf scales are glabrous with
entire margins; the pedicel is hairy and is articulated in
the top half; and the tepals have a ridge of hairs along
the median stripe. A. capensis var. capensis is a much-
branched shrub up to 1.5 m high; tubers absent; the main
stem is glabrous; the spines are sparsely hairy, becom-
ing glabrous; the branches end in a spine; the leaf scales
are hairy with a lacerate margin; the flower is sessile;
and the tepals are puberulous throughout. Although A.
minutiflorus has tubers and a foxtail growth form, the
branches typically arch or sprawl along the ground. It
differs from A. hirsutus in that the cladodes are filiform;
the hairs are glandular and golden brown; the spines
have a few hairs at the base; the leaf scales have lacerate
margins; and the pedicel is glabrous and is articulated in
the lower half.
The known distribution of A. hirsutus is a distinguish-
ing character: whereas A. capensis var. capensis only
occurs in the Cape Province and southern Namibia, A.
minutiflorus has a wider distribution, occuring in dry
areas of Mpumalanga, Limpopo, Swaziland, KwaZulu-
Natal and southern Mozambique.
TABLE 1 . — Differences in morphology of Asparagus elephantinus and
A. densiflorus
26
Bothalia 38,1 (2008)
FIGURE 3. — Asparagus hirsutus, L.M.D. Riddles 511. A-D, floral
details: A, flower; B, stamen and tepal, front view; C, stamen
and tepal, side view; D, ovary. E, fruit; F, cladode; G, c/s cla-
dode; H, I, simple hairs; J, spine and leaf scale; K, leaf scale.
Scale bars: A-C, 1 .6 mm; D, 1 mm; E, 6.6 mm; F, 2.2 mm; I, 0.2
mm; J, 4.4 mm; K, 1.6 mm. Artist: Sandra Burrows.
Etymology: Asparagus hirsutus is not easily confused
with any other Asparagus species, as the extreme hairi-
ness of the plant, (hence the specific epithet), the fox-
tail growth form, and its limited distribution render it
unmistakeable.
Asparagus sylvicola S.M. Burrows, sp. nov., A.
setacei similis sed ramulis late dispersis, cladodiis ram-
ulisque ad angulum 90° caule patentibus, tuberibus fusi-
formibus differt.
TYPE. — Mpumalanga, 2531 (Komatipoort): Bearded
Man, Songimvelo Nature Reserve, Barberton Mtns,
1 400 m, (-CB), 16 April 2005, J.E. Burrows 8931 (PRE,
holo.; BNRH, iso.).
Scandent to twining, evergreen shrublet up to 3 m
high, glabrous throughout. Rhizome creeping; roots
up to 3 mm thick, terete, with rootlets; mature plants
develop spindle-like in-line tubers, 20-100 x 5-17
mm. Stems 2— A mm thick, basally very finely papillate-
ridged, dark olive green to purplish; leaf scales set on
very small swellings. Spine s absent throughout. Leaf
scales triangular, 0. 5-3.0 x 0. 5-1.0 mm long, dark pur-
ple to reddish brown, pale on old stems, triangular, base
with small (0.5 mm) soft protuberance, margin entire.
Branches up to 400 x ± 1 mm, widely placed along
stem, arising at right angles to stem, slightly zigzag-
ging, terete, ridged. Branchlets 0.5 mm thick, up to 120
mm long, terete, widely spaced along branches, arising
at right angles to branch. Secondary branchlets up to 25
mm long, same thickness as cladodes, arising at right
angles to branchlets. Cladodes 1-9 per fascicle, filiform,
terete, falcate, (5-)9-20(-28) mm long, 0.5 mm to hair-
like in width, glossy dark green. Flowers 1 or 2(-4),
borne in terminal cladode fascicle, bud dark purple, no
scent detected. Pedicel 3 mm long, dark purple, articu-
lated in top half, base surrounded by small membranous
bracts. Tepals 6, 3 larger tepals 3. 0-3. 5 x 1. 9-2.1 mm
and 3 smaller tepals 2. 5-3.0 x 1.0-1. 5 mm, apex some-
times notched, underside streaked with purple. Stamens
6; filaments 2.75-3.25 mm long, fused at base; anthers
pale lemon yellow. Style 1.8-2. 5 x 0.5 mm, terete;
stigma 3-fid, arms 0. 7-1.0 mm long. Ovary trilocular,
0. 9-1.1 x 1. 5-2.0 mm. Fruit a berry, 6-10 mm diam.
with persistent perianth and style, black when ripe. All
measurements of cladodes, leaf scales, flowering and
fruiting parts taken from pickled material. Flowering
time: although seldom floriferous, the main flowering
time is from November to January, with the occasional
flower produced throughout the rest of the year. Figure
4.
Distribution and habitat: Asparagus sylvicola was
first noted growing in forested areas in the Barberton
Mountains in Mpumalanga and in the Lubombo
Mountains in Swaziland (Figure 2). A number of spec-
imens previously attributed to A. setaceus (Kunth)
Oberm. are now included in this species, extending
the distribution to include the forested areas of north-
eastern South Africa, from Haenertsberg (Limpopo) to
Siteki (Swaziland). The plant is widespread in these
areas, although seldom common and, since much of the
remaining habitat is protected, so therefore is this spe-
cies.
Diagnostic characters and affinities: Asparagus syl-
vicola may be confused with A. setaceus (Kunth) Jessop
but the most distinctive difference is that the branches
of A. sylvicola radiate out at right angles and the clad-
odes arise in all directions, forming a compact, springy
nonaligned tangle of cladodes, whereas A. setaceus
has its branches and cladodes arranged in a horizontal
plane. The cladodes of A. sylvicola are falcate, and in
some specimens bow-shaped, whereas the cladodes of
A. setaceus are straight to slightly curved. The tubers on
mature plants of A sylvicola resemble small long sweet-
potatoes (spindle-like), and are in-line in the main roots,
whereas the tubers of A. setaceus are ovoid to ellipsoid
and are borne laterally.
Etymology: Asparagus sylvicola (‘forest dweller’) is
so named as it favours forest, either Scarp Forest (FOz
5) or the lower regions of Northern Mistbelt Forest
(FOz 4), (Mucina & Rutherford 2006).
Cultivation: Asparagus sylvicola has been cultivated
as a container plant for several years, and makes a very
attractive horticultural subject.
Specimens examined
LIMPOPO. — 2329 (Pietersburg): Haenertsberg, (-DD), November
1913,/?. Pott 4689 (PRE).
MPUMALANGA. — 2430 (Pilgrim’s Rest): Mariepskop, near
KJaserie Drift, ( DB), 8 April 1969, P. Vorster 616 (PRE, PRU); Mt
Sheba, 1 700 m, (-DC), 2 October 1986, Deal! 2165 (PRE!). 2530
(Lydenburg): Ceylon Forest Reserve, ‘Jantjiesbos’, 1 350 m, (-BA), 7
October 1986, Deal 1 2193 (PRE). 2531 (Komatipoort): Kaalrug road,
Malelane Dist., 350 m, (-CB), 6 February 2003, J.E. & S.M. Burrows
Bothalia 38,1 (2008)
27
FIGURE 4. — Asparagus sylvicola, J.E. Burrows 8931. A-C, K, stem and branch; D, flowering branchlet; E, leaf scale. F-H, flower details: F,
flower; G, front and side view of tepal and stamen; H, ovary. I, fruit; J, cladodes; L, rhizome habit. Scale bars: A-D, 1 1.8 mm; E, 5.3 mm;
F, G, 1 mm; H, 1.8 mm; I, 3.5 mm; J, 2.9 mm; K, 5.9 mm. Artist: Sandra Burrows.
7975 (BNRH); Ida Doya Reserve, Barberton Mtns, (-CB), M.C. Lotter
s.n. (BNRH); Barberton Mtns, Pedlar’s Bush, 1 280 m, (-CC), 29
October 2006, J.E. & S. M. Burrows 9559 (BNRH, PRE); Barberton,
Pedlar’s Bush, 1 100-1 500 m, (-CC), January 1996, T.L. Morgenthal
s.n. (PRE, PUC); Barberton, (-CC), June 1907, Thorncroft s.n. (Herb.
Tvl. Mus. 4349 in PRE).
SWAZILAND. — 2631 (Mbabane): Malagwane Hill, Mbabane,
3570' [1 088 m], (-AA), 22 June 1961, Ben Dlamini s.n. (PRE.
SNDH); ibid., 22 June 1961, B. Dlamini s.n. (PRE, SNHD); ibid., 4
May 1961. B. Dlamini s.n., (PRE, SRGH). 2632 (Bela Vista): Mtibhlati
River, Lubombo Mtns, Siteki area, 340 m, (-CA), 22 November 2002,
J.E. & S.M. Burrows 7895 (BNRH).
28
Bothalia 38,1 (2008)
Distribution of Asparagus acocksii Jessop, and Aspa-
ragus lynetteae (Oberm.) Fellingham & N.L.Mey.
The confusion in the identification between Asparagus
acocksii and A. lynetteae in herbaria, due to the lack of
material or information on the rooting system, has led
both Jessop (1966) and Obermeyer & Immelman (1992)
to accept that A. acocksii occurs disjunctly from the
Eastern Cape to northern KwaZulu-Natal, Mpumalanga
and Sekhukhuneland, Limpopo Province. When Ober-
meyer & Immelman (1992) described the new spe-
cies A. lynetteae from Sekhukhuneland, they failed to
realize that Jessop’s ‘A. acocksii' (Jessop 1966) from
Sekhukhuneland was conspecific with A. lynetteae.
In the type description of Asparagus acocksii, Jessop
(1966) states that the stems are ‘erect to scrambling’,
and that ‘there are single records from both Zululand
and the Transvaal’. Having now visited the type locali-
ties of both A. acocksii and A. lynetteae, the following is
noted. Asparagus acocksii is a very small shrublet, up to
450 mm high. Where the stems are longer than usual (up
to 1 m), they arch downwards but show no inclination
to climb. The rhizome is compact and woody, the roots
terete, up to 5 mm in diameter, with lateral, ovoid tubers,
26-37 x 19-23 mm. A. acocksii was previously consid-
ered to occur from the Eastern Cape to KwaZulu-Natal
and the Transvaal (Mpumalanga and Limpopo).
Asparagus lynetteae is a lax, shrubby scrambler, with
a tendency to climb up to 1.5 m. The rhizome is com-
pact, with numerous thick, ribbed roots, ± 20 mm diame-
ter and more than 1 m long, with terete side roots. Tubers
are absent. Up to now A. lynetteae was considered to be
a narrow Sekhukhuneland endemic, and possibly threat-
ened.
Since most of the collections of Asparagus lynetteae
have been made with either no roots or without reference
to the roots, and since the vegetative parts closely resem-
ble those of A. acocksii, these two taxa have been con-
fused in herbaria and their true geographical distributions
have been clouded. However, re-collections of the plants
and the roots by the authors at the non-Cape ‘A. acocksii'
sites have established that all these plants bear thickened,
terete, tuberless roots and are therefore A. lynetteae,
thus establishing that A. acocksii is restricted to the dry
regions of the Eastern Cape, and that A. lynetteae is no
longer endemic to the Sekhukhuneland area, but is spo-
radic in the bushveld areas of Mpumalanga, Swaziland,
KwaZulu-Natal, and possibly in neighbouring southern
Mozambique. Figure 5.
Asparagus acocksii Jessop in Bothalia 9: 74, t. 7
(1966); Protoasparagus acocksii (Jessop) Oberm.: 243
(1983). Type: Cape, Cradock District, 12 miles NNE of
Fish River Station, (-CB), Acocks 16296 (PRE, holo.!).
Additional specimens examined
EASTERN CAPE.— 3125 (Steynsburg): 21.6 km from Fish
River Station, opp. Grassridge Dam, 1 100 m, (-CB), 16 Feb. 2006,
J.E & S.M. Burrows 9376 (BNRH). 3126 (Queenstown): 5 km from
Queenstown on Cathcart road, 1 150 m, (-DD), 1 February 2006, J.E.
& S.M. Burrows 9452 (BNRH). 3223 (Rietbron): between Beaufort
West and Aberdeen, (-BD), 23 April 1985, MB Bayer 4705 (PRE).
FIGURE 5. — Known distribution of Asparagus acocksii, ■; and A.
lynetteae , •.
3224 (Graaff-Reinet): Graaff-Reinet, (-BC), 7 May 1954, Henrici
4964, (PRE); Graaff-Reinet, Karoo Nature Reserve, Allardice R.,
eastern reserve on western slope along road from pump house,
(-BC), 20 July 1981, C. Reid 1544 (PRE); Lusanne, E of Aberdeen,
(-CA), 11 December 1983, M.B. Bayer 3740A (PRE); Krugerskraal
near Graaff-Reinet, (-CD), November 1984, W.R. Brand s.n. (PRE).
3225 (Somerset East): S of Swaershoek Pass, on road from Cradock
to Somerset East, 1 108 m, (-AD), 24 February 2004, J.E & S.M.
Burrows 8347 (BNRH, PRE); Cradock, 1 km S of Marlow, (-BA), 24
January 1985, M.B. Bayer 4318 (PRE); Mountain Zebra National Park,
eastern region, (-BA), 5 September 1959, P.J. Barnard 513 (PRE).
3325 (Port Elizabeth): ‘Aloes’, (-DC), December 1912, Drege s.n.
(PRE3164).
Asparagus lynetteae Oberm. in Obermeyer &
Immelman, Flora of southern Africa 5,3: 47 (1992).
Type: near Burgersfort, in dense bushveld at Gethlane
Lodge, Mauve, Reid & Smook 5408 (PRE, holo.!).
Additional specimens examined
LIMPOPO. — 2430 (Pilgrim’s Rest): on way to Penge, (-AC), 26
January 1972, G.D. Wright G36 (PRE); Moshira, Motse River, on road
to Penge, 730 m, (-AC), 15 January 2002, J.E. & S.M. Burrows 8783
(BNRH, PRE); pass W of Penge Mine, on road to Chuniespoort, 1 250
m, (-AC), March 2005, J.E. & S.M. Burrows 8913 (BNRH, PRE);
ibid. 6 March 2005, J E. & S M. Burrows 8917 (BNRH, PRE); ibid. 15
January 2005, J.E. & S M. Burrows 8788 (BNRH, PRE); Burgersfort,
5 km NE of Alverton Village, 689 m, (-CA), 5 April 2005, J.E. & S.M.
Burrows 8955 (BNRH); at Gethlane Lodge, Farm Fraaiuitzicht 843
KT, 860 m, (-CD), 20 November 1985, Krynauw 914 (LYD, PRE);
2 miles [3.2 km] N of Ohrigstad, (-DA), 22 November 1933, R.G.N.
Young A616 ( Tvl . Mus. Herb 32648 in PRE).
MPUMALANGA. — 2430 (Pilgrim’s Rest): 13.6 km on Steelport/
Gethlane road near Steelpoort, 840 m, (-CC), 4 August 2002, J.E. &
S.M. Burrows 7751 (BNRH, PRE). 2531 (Malelane): Magnesite Mine,
between Kaapmuiden & Malelane, 630 m, (-CB), 17 April 2005, J.E.
6 S.M. Burrows 8951 (BNRH, PRE); ibid. 6 February 2003, J.E. &
S.M. Burrows 7972 (BNRH, PRE).
SWAZILAND. — 2631 (Mbabane): Mlawula Mountain Resort,
near top entrance gate, 566 m, (-AA), 2 December 2006, J.E. & S.M.
Burrows 9652 (BNRH); Swaziland lowveld near Mliba, (-BC), 14
January 1973, D.N.H. Horter 110 (PRE).
KWAZULU-NATAL. — 2632 (Bela Vista): Ingwavuma District, Ndu-
mu Game Reserve, 200' [61 m], (-CD), 10 February 1954, C.J. Ward
2038 (NPB Herb., PRE); Ingwavuma District, Ndumu Game Reserve,
130' [ 40 m], 15 November 1962, C.J. Ward 4522 (NU, PRE); Ndumu
Game Reserve, Mahemane, 40 m, (-CD), 15 October 1990, Thornhill
283 (PRE). 2731 (Louwsberg): 12 km N of Magudu on tar road to
Pongola, (-BC), 18 February 1982, Reid 491 (PRE); Ngotshe District,
Bothalia 38,1 (2008)
29
Pongola Irrigation Scheme, 800' [244 m], (-CD), 16 October 1963,
C.J. Ward 4766 (PRE); 3 km along Candover road from Mahlangasi,
off Pongola-Nongoma road, 345 m, (-DA), 9 June 2006, J.E. & S.M.
Burrows 9460 (BNRH).
ACKNOWLEDGEMENTS
Our thanks go to the Curators of PRE and K, for
allowing access to the Asparagaceae Section. In particu-
lar we would like to thank Ronell Klopper of PRE, Paul
Wilkens of K, Hugh Glen for the Latin translations,
the Trustees of Buffelskloof Nature Reserve for their
encouragement and financial assistance, and Priscilla
Burgoyne of PRE for bringing Asparagus hirsutus to
our attention.
REFERENCES
JESSOP. J.P. 1966. The genus Asparagus in southern Africa. Bothalia
9 : 31-96.
MUCINA, L. & RUTHERFORD. M.C. 2006. The vegetation of South
Africa, Lesotho and Swaziland. Strelitzia 19. South African
National Biodiversity Institute, Pretoria.
OBERMEYER, A. A. 1983. Protasparagus Oberm. nom. nov.; new
combinations. South African Journal of Botanv 2: 243, 244.
OBERMEYER, A. A. & IMMELMAN, K.L. 1992." Asparagaceae. In
O.A. Leistner, Flora of southern Africa 5,3: 11-82. Botanical
Research Institute, Pretoria.
.
Bothalia 38,1: 31-37 (2008)
A new species of Euclea (Ebenaceae) from ultramaflc soils in Sek-
hukhuneland, South Africa, with notes on its ecology
E. RETIEF*, S.J. SIEBERT** and A.E. VAN WYK***
Keywords: Ebenaceae, Euclea Murray, ecology, new species, Sekhukhuneland, South Africa, summer rainfall region, taxonomy
ABSTRACT
Euclea sekhukhuniensis Retief, Siebert & A.E.van Wyk, a new species with a restricted range in Sekhukhuneland, South
Africa, is described, illustrated and compared with other members of the genus. It is a gregarious geoxylic suffrutex forming
large, much-branched colonies. The species is closely related to the small tree/shrub E. linearis Zeyh. ex Hiem from which it
can be distinguished by its larger fruits, broader leaves and exclusively suffrutex growth form. Geographical range and habitat
preference also differ between the two taxa. E. sekhukhuniensis is endemic to the Sekhukhuneland Centre of Plant Endemism,
where it is confined to the calcareous, heavy-metal soils of the Steelpoort River Valley.
INTRODUCTION
Areas underlain by ultramaflc rocks in Mpumalanga
and Limpopo, and adjacent parts of Zimbabwe, contain a
rich but still poorly studied flora. Examples of taxa newly
described from these substrates include Searsia pyg-
maea (Anacardiaceae) from serpentinite near Barberton
(Moffett 1999), Rhoicissus sekhukhuniensis (Vitaceae)
from norite/pyroxenite near Steelpoort (Retief et al.
2001) and Peristrophe serpenticola from the Great Dyke
(Balkwill & Campbell-Young 2001). Recent vegetation
surveys of the ultramaflc rocks of the Sekhukhuneland
Centre of Plant Endemism (SCPE), a microregional
centre of plant endemism and diversity (Van Wyk &
Smith 2001), have revealed many undescribed plant
taxa endemic to this phytogeographical region (Siebert
et al 2001). Future floristic surveys in this under-col-
lected region are likely to reveal many new distribution
records and possibly further new taxa. Other species
known to be endemic to the SCPE include Raphionacme
villicorona (Apocynaceae), Asparagus sekukuniensis
(Asparagaceae), Acacia ormocarpoides, A. sekhuk-
huniensis, Elephantorrhiza praetermissa (Fabaceae),
Euphorbia barnardii, E. sekukuniensis (Euphorbiaceae),
Gladiolus sekhukhuniensis (Iridaceae), Plectranthus por-
catus, P. venteri (Lamiaceae), Hibiscus coddii subsp.
barnardii (Malvaceae), Searsia batophylla, S. sekhukhu-
niensis (Anacardiaceae), Zantedeschia jucunda and Z.
pentlandii (Araceae).
The SCPE lies to the west of the northeastern sec-
tion of the Great Escarpment of South Africa and is
characterized by a heterogeneous geology, topogra-
phy and climate (Van Wyk & Smith 2001). The core
area of the Centre straddles the border of Mpumalanga
and Limpopo, around the towns of Burgersfort,
Mecklenburg, Roossenekal, Schoonoord and Steelpoort.
The SCPE is best demarcated in geological terms as the
* National Herbarium, South African National Biodiversity Institute,
Private Bag XI 01, 0001 Pretoria.
** A.P. Goossens Herbarium, School of Environmental Sciences
and Development, North-West University, Private Bag X6001, 2520
Potchefstroom, South Africa.
*** H.G.W.J. Schweickerdt Herbarium, Department of Botany,
University of Pretoria, 0002 Pretoria.
MS. received: 2007-04-05.
large, far-eastern outcrop of ultramaflc rocks belonging
to the Rustenburg Layered Suite of the eastern Bushveld
Complex. These rocks are mainly norite, pyroxenite,
anorthosite and ferrogabbro, with localized intrusions of
magnetitite and chromitite (Viljoen & Schiirman 1998).
Topographically the SCPE is a mountainous area bor-
dered by the high ground of the Drakensberg Escarpment
in the north and east, the Highveld Escarpment to the
south and the Springbok Flats to the west. It lies adjacent
to and west of the Wolkberg (Van Wyk & Smith 2001)
and Lydenburg (Schmidt et al. 2002) Centres of Plant
Endemism, both part of the northeastern Drakensberg
Escarpment.
Previously a Euclea taxon with a suffrutex habit and
narrow elliptical leaves from Sekhukhuneland was tenta-
tively considered a hybrid between E. linearis Zeyh. ex
Hiem and E. crispa (Thunb.) Giirke subsp. crispa (De
Winter 1963) — a suspicion based on the overlapping dis-
tribution ranges of these two species in Sekhukhuneland.
However, subsequent detailed field work and compara-
tive morphological studies have shown the putative
hybrid to be a distinct new species, closely related to E.
linearis and endemic to the ultramaflc soils of the SCPE.
The new species is here described as Euclea sekhukhuni-
ensis Retief, Siebert & A.E.van Wyk. This is the second
Euclea species, after E. dewinteri Retief (Retief 1986),
an endemic of the Wolkberg Centre of Plant Endemism,
that is strictly confined to the larger northeastern Drakens-
berg Escarpment.
The genus Euclea comprises ± 20 species, confined
to Africa, Arabia, Socotra and the Comoro Islands, with
its centre of diversity in southern Africa (Dyer 1975;
Bredenkamp 2000). In addition to E. sekhukhuniensis ,
seven species and infraspecific taxa occur in the SCPE,
namely E. crispa subsp. crispa, E. divinorum Hiem,
E. linearis, E. natalensis A. DC. subsp. angustifolia
F. White, E. daphnoides Hiem, E. schimperi A.DC. and
E. undulata Thunb. (Table 1). However, the list is pro-
visional, for the region is still poorly sampled. All these
taxa are evergreen shrubs or trees, except E. sekhuk-
huniensis, which is an evergreen geoxylic suffrutex. A
form of E. crispa (White 1977) along the northeastern
Drakensberg Escarpment (but not entering the SCPE)
exhibits the same growth form as E. sekhukhuniensis.
32
Bothalia 38,1 (2008)
MATERIALS AND METHODS
Euclea specimens housed in the National Herbarium
(PRE), Pretoria, and H.G.W.J. Schweickerdt Herbarium
(PRU), University of Pretoria (acronyms as in Holmgren
et al. 1990) were examined to gather data on morphol-
ogy, phenology and distribution.
Ecological data are based on extensive field obser-
vations of Euclea linearis and E. sekhukhuniensis in
the SCPE (Siebert et al. 2002a, b). Specimens were
sampled from the region during all seasons, environ-
mental factors were noted and plant communities iden-
tified. The specimens Siebert 935, 937 and Van Wyk &
Siebert 13060 were identified as typical E. linearis for
comparing ecological traits with E. sekhukhuniensis.
Plant material and soil samples were taken at ten sites,
five each dominated by either E. linearis or E. sekhuk-
huniensis. Voucher specimens were taken from these
sites; Siebert 937 represents E. linearis and Siebert 938
represents E. sekhukhuniensis (specimens kept at PRU).
Soil analysis was done with X-Ray Fluorescence (XRF)
Spectrometry, Department of Geology, University of
Pretoria and plant analysis with Atomic Absorption
Spectrophotometry (AAS) and Inductively Coupled
Plasma-Mass Spectrometry (ICP-MS) at the Department
of Soil, Climate and Water in Pretoria.
TAXONOMY
Euclea sekhukhuniensis Retief, Siebert & A.E.van
Wyk, sp. nov., E. lineari Zeyh. ex Hiem similis sed
habitu (suffrutex, non frutex vel arbor parva), forma foli-
orum elliptica recta, non anguste elliptica nec lineari nec
falcata, fructu globoso maiore (± 9 mm, non ±5.5 mm
diametro) differt.
TYPE. — Mpumalanga, 2430 (Pilgrim’s Rest): 10 km
NW of Maartenshoop, (-CC), Codd 8796 (PRE, holo.;
K, iso.). Figure I .
Euclea linearis Zeyh. ex Hiem sensu De Winter: 94 (1963) quoad
Codd 8796.
Woody suffrutex, 0.3-1. 5 m high, forming large
colonies of much-branched clones ± 5 m diam. Plants
evergreen, dioecious. Branches ascending, slender and
glabrous, except for a rust-brown granular exudate on
TABLE 1. — Distribution of seven species of Euclea according to quar-
ter-degree grid squares of SCPE
Euclea species/infraspecific
taxa
OQCQQO<mOQPQQ<
mmQQ<;ouuumm<c
O^.OsOxOsOOOOOCEOsO
<N Cl C404rcrcrcrorc<N(Nrc
<N<N<N<N<NCNCN<NCN<NCN<N
E. crispa subsp. crispa ...
E. divinorum • •
E. linearis •
E. natalensis subsp. angus- • • ...
lifolia
E. schimperi var. daphnoi- • •
des
E. schimperi var. schimperi •
E. sekhukhuniensis • .....
E. undulata ..........
Total 163367565522
FIGURE 1. — Euclea sekhukhuniensis , Codd 8796, holo. (PRE).
young growth; bark grey on older stems. Leaves sim-
ple, subopposite, subsessile; blade oblanceolate-elliptic,
straight, 25-75(-90) x 4-8(-10) mm, glabrous, leathery
and smooth, except for a rust-brown granular exudate on
younger leaves, usually yellowish green above and pale
green below; base tapering into a very short petiole (1
mm), apex acute to rounded, margin entire, main vein
and principal lateral veins prominent above and below.
Inflorescences axillary, few-flowered, clusters or short
spikes. Flowers regular. Male flowers: calyx 4-lobed, ± 3
mm long; corolla campanulate, deeply 4-lobed, ± 5 mm
long, pale cream-coloured to pinkish white, appressed
hairy on back; stamens 8, narrowly ovate, ± 3 mm long,
anther thecae pubescent on outer surface. Female flow-
ers: calyx not accrescent, persistent in fruit, 4-lobed, ±
2 mm long; corolla 4-lobed, ± 4 mm long, green, lobes
appressed hairy on back; ovary hairy, borne on a fimbri-
ate, fleshy disc; styles 2. Fruit an indehiscent, globose
berry, ± 9 x 10x8 mm, densely appressed hairy; young
fruits green, ripening through brownish red to purplish
black. Flowering time : October to January. Fruiting
time : November to February. Figure 2.
Diagnostic characters', members of Euclea can be
divided into two groups (De Winter 1963): 1, species
with the corolla shallowly lobed at the apex; and 2, spe-
cies with the corolla cleft at least halfway down or more.
All the Euclea taxa occurring in the SCPE belong to the
latter group. E. sekhukhuniensis and E. linearis are dis-
Bothalia 38,1 (2008)
33
FIGURE 2. — Euclea sekhukhuniensis. Plant Specialist Group 4 (PRE).
A, fruiting branch, x 0.75; B, female flower, x 3.8; C, gynoe-
cium, x 7.5; D, male flower, x 3.8; E, stamen of outer row, x
7.5; F, stamen of inner row, x 7.5. Artist: G. Condy.
tinguished from the others by their narrowly oblanceo-
late-elliptic to linear or linear-falcate leaves and young
leaves and twigs that are covered with a granular, rust-
brown exudate. E. sekhukhuniensis is most closely
related to E. linearis ; however, E. sekhukhuniensis is a
gregarious, evergreen geoxylic suffrutex (White 1977),
whereas E. linearis is a shrub or tree up to 5 m high.
Leaves of E. sekhukhuniensis are straight, broader and
longer than the sickle-shaped leaves of E. linearis and
its fruits are larger than those of E. linearis (Table 2).
Conservation status : Euclea sekhukhuniensis has a
restricted geographical range within which it is locally
fairly common (Figure 3). However, some of its habi-
tat is under immediate threat of rapid urbanization as
a result of increased mining activities in the greater
Steelpoort River Valley and the construction of the De
Hoop Dam on the Steelpoort River. E. sekhukhunien-
sis is not formally protected in any conservation area.
Populations of this species should therefore be closely
monitored and a Red Data List assessment of this spe-
cies prioritized. Its conservation value is considered rela-
tively high, as it could possibly be used in the rehabili-
tation of mine dumps due to its internal mechanism of
excluding heavy metals.
Ecology and speciation: both Euclea sekhukhuniensis
and E. linearis occur on vegetation anomalies — sparsely
vegetated soils that are mineralized (Table 3). This phe-
nomenon is well reported for populations of E. linearis
in wooded grassland on the serpentinites of the Great
Dyke in Zimbabwe (Wild 1965). A distant outlier of
what has been identified as E. linearis is also found on
grassy ridges in fynbos in a limited area in the Calvinia
and Vanrhynsdorp region of the Western Cape, where
it grows on nutrient-poor soils derived from sandstone
of the Table Mountain Group (White 1983). However,
the identity of these plants requires verification. In
the mountainous regions of the northern provinces of
South Africa, E. linearis grows on rocky outcrops and
in dry woodlands on slopes and in valleys of serpen-
tenite (Barberton Supergroup) in the Barberton region,
acidic sandstone (Waterberg Group) of the Waterberg of
Limpopo, and quartzites (Black Reef Formation) along
the northeastern Drakensberg Escarpment (White 1983).
Thus, it appears that E. linearis tends to colonize habi-
tats with harsh soil conditions in mountainous regions
(acidic, nutrient poor and/or rich in heavy metals).
Euclea sekhukhuniensis appears to be an example of
incipient sympatric speciation due to ecological inter-
actions in a new habitat in which restricted gene flow
has evolved through selective reproduction between
individuals of E. linearis that are adapted to a specific
ultramafic substrate. This speculation is supported by
the work of Alados et al. (1999), which demonstrates
that asymmetry and within-plant variance were higher
between specimens of the same species in the contact
zone between ultramafic and normal soils. In the SCPE,
habitat preference has resulted in the two Euclea species
FIGURE 3. — Known distribution of Euclea sekhukhuniensis.
34 Bothalia 38,1 (2008)
TABLE 2. — Leaf and fruit measurements of 1 5 Euclea sekhukhuniensis specimens and only results of 30 randomly selected specimens of E. linearis
from Mpumalanga and Limpopo (all in PRE)
now growing in specific, but different habitats in associ-
ation with specific plant species (Tables 3, 4). According
to Dieckmann & Doebeli (1999), theoretical evidence
suggests a prominent role for ecologically driven specia-
tion in sympatry. Hence, the present study supports the
TABLE 3. — Environmental, structural and floristic characteristics
unique to plant communities dominated by either Euclea
sekhukhuniensis or E. linearis in Steelpoort River Valley
opinion that the ecological species concept is an essen-
tial part of the biological species concept (Grant 1992).
Generally, ecologically driven speciation is the result
of habitat-specific preferences. This has been investi-
gated and confirmed for an endemic species of Impatiens
and its widespread congener (Chung & Kang 1996), as
well as for two endemic species of Dicerandra of the
same region (Menges et al. 1999). In the case of Euclea
sekhukhuniensis, an open niche with an anomalous Ca-
rich substrate (14.68% = 146 800 ppm) in an other-
wise typical environment of brackish soils rich in Mg
(19.97% = 199 700 ppm), probably favoured speciation
(Figure 4A). Similar trends have been perceived between
especially limestone (Ca-rich) and sandstone, once again
for an endemic and its widespread congener (Walck et
al. 1999), as well as two endemics of the same region
(Mustart et al. 1994). Like limestone, the soils inhab-
ited by E. sekhukhuniensis are Ca-rich (1.95Ca:lMg),
more than double that of the soil substrate of E. linearis
(Figure 4A). Furthermore, soils in which E. linearis
grows have higher concentrations of total Cr and Ni
(typical elements of serpentinite) (Figure 4B), with high
Fe, Si and Mg levels (lMg:0.34Ca) (Figure 4A), and it
accumulates relatively high concentrations of Al and Fe
in its roots (Figure 5A). E. sekhukhuniensis accumulates
lower levels of Fe in its roots, but with higher concentra-
tions of Cr and Ni than E. linearis, although these lev-
els are very low and not regarded as hyperaccumulation
(Figure 5A). Overall, it seems that E. sekhukhuniensis is
the better excluder of heavy metals, when considering
the high concentrations of metals in the associated soil.
It is suggested that Euclea sekhukhuniensis was an
ecotype of and has developed from E. linearis as a result
of the genetic properties of the latter to adapt to and col-
onize ultramafic soils such as those derived from serpen-
tinite (Wild 1974). It is hypothesized that E. sekhukhuni-
ensis is a ‘soil-adapted’ neo-endemic which speciated
Bothalia 38,1 (2008)
35
TABLE 4. — Prominent taxa recorded for communities dominated by either Euclea sekhukhuniensis or E. linearis in Steelpoort River Valley
(Siebert et al. 2002b)
recently, after the Pleistocene (Reeves et al. 1983), and
has not yet had the time or routes to migrate out of the
Steelpoort River Valley. However, it is doubtful whether
this will ever happen, as the species probably prefers the
open niches of ultramafic soils where it has a physiologi-
cal mechanism associated with high plant levels of Ca to
tolerate heavy metals (Figure 5B).
Specific epithet and common names: the specific epi-
thet refers to the geographical area where the species is
endemic. Sekhukhuneland is traditionally inhabited by the
Pedi (Monnig 1967) and is currently under the reign of
K.K. Sekhukhune (Paton 1998). Common names for the
taxon include moshigwane (Northem-Sotho), Steelpoort
guarri (English) and Steelpoortghwarrie (Afrikaans).
0 Euclea sekhukhuniensis ED Euclea linearis
FIGURE 4. — Chemical analyses of
five soil samples collected
from root zone (300 mm deep)
for each of Euclea sekhukhu-
niensis and E. linearis.
36
Bothalia 38,1 (2008)
CD
| 3
C
03
Sr 2
o
1
Ca | Mg S
Leaves
P i Ca , Mg S
Stems
Plant organs
P j Ca Mg I S
Roots
0 Euclea sekhukhuniensis , 0 Euclea linearis
FIGURE 5. — Chemical analyses of
plant material from five indivi-
duals each of Euclea sekhukhu-
niensis and E. linearis (young
and old growth).
SPECIMENS EXAMINED
LIMPOPO. — -2429 (Zebediela): Het Fort on Leolo Mountains,
(-BD), Barnard & Mogg 1031 (PRE). 2430 (Pilgrim’s Rest):
Maandagshoek, (-CA), Krilzinger 138 (PRU), Wright G14 (PRE).
MPUMALANGA.— 2430 (Pilgrim’s Rest): Burgersfort, (-CB),
Herman 793 (PRE); Maartenshoop, (-CC), Codd 8796 (PRE);
Frischgewaagd, (-CC), Siebert 411, Van Wyk, Siebert & Retief 13204
(PRU); Thomecliffe Chrome Mine, (-CC), Plant Specialist Group 4
(PRE), Siebert 934, Van Wyk 13035, 13036, Van Wyk & Siebert 13312,
13313 (PRU); Olifantspoortjie, (-CC), Siebert 938 (PRU); Steelpoort,
(-CC), Van Wyk 13299 (PRU).
ACKNOWLEDGEMENTS
The authors are indebted to Maggi Loubser, Geology
Department, University of Pretoria, for assistance with
XRF analysis of the soil samples and to Nina van
Vliet, Department of Soil, Climate and Water, Pretoria,
for assistance with AAS and ICP-MS analysis of plant
material. Francois du Randt of Burgersfort is thanked
for collecting fertile material of the new species. Emsie
du Plessis and Hugh Glen of the South African National
Biodiversity Institute are respectively thanked for proof-
reading the manuscript and for translating the diag-
nosis into Latin. The Andrew W. Mellon Foundation,
University of Pretoria and South African National
Biodiversity Institute provided financial support.
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'
Bothalia 38,1: 39^7 (2008)
The genus Solanum (Solanaceae) in southern Africa: subgenus Lepto-
stemonum , section Giganteiformia
W.G. WELMAN*
Keywords: chemistry, ethnobotany, Giganteiformia (Bitter) Child, Leptostemonum (Dunal) Bitter, morphology, Solanaceae, Solanum L., southern
Africa, taxonomy
ABSTRACT
In the genus Solanum L. (Solanaceae), subgenus Leptostemonum (Dunal) Bitter, section Giganteiformia (Bitter) Child has
four representatives in the Flora of southern Afiica region (South Africa, Namibia, Botswana, Swaziland, Lesotho), namely
S. giganteum Jacq., S. goetzei Dammer, 5. tettense Klotzsch var. renschii (Vatke) A.E.Gor^alves and S. tettense Klotsch var.
tettense. Descriptions, discussions, distribution maps and keys are presented, as well as an illustration of S. goetzei.
INTRODUCTION
In the genus Solanum L., the prickly subgenus Lepto-
stemonum (Dunal) Bitter is represented by eight sec-
tions in southern Africa. Three sections ( Giganteifor-
mia, Melongena, Oliganthes ) contain only indigenous
taxa, and the remaining five sections ( Acanthophora ,
Androceras, Crvptocarpum, Leprophora, Torva) have
only introduced species. Section Giganteiformia (Bitter)
Child is represented by four taxa in southern Africa; all
are indigenous. S. giganteum Jacq. occurs from tropi-
cal Africa (also India and Sri Lanka) to the Western
Cape, S. goetzei Dammer grows from Kenya to northern
KwaZulu-Natal, while S. tettense is found from tropical
Africa to Namibia, Botswana and the northern provinces
of South Africa. Detailed descriptions and discussions
are given for both the section and all four taxa, together
with keys and distribution maps. S. goetzei is illustrated
for the first time. Information on distribution, ecology,
phenology and uses came from specimens in NH and
PRE, unless otherwise stated.
DESCRIPTIONS AND DISCUSSIONS
Section Giganteiformia (Bitter) Child in Feddes
Repertorium 109, 5 & 6: 415, 416 (1998). Lectotype
species: S. giganteum Jacq. (Child 1998).
Series Giganteiformia Bitter: 255 (1921).
Description (based on Child 1998)
Medium-sized shrubs to small trees 2-6 m high,
white-pubescent to tomentose-floccose with sessile to
shortly stipitate stellate hairs (also some simple hairs),
stellate hairs transparent and hyaline, often small, some-
times sparse. Prickles short, 2-6 mm long, ± absent to
many rosoid on stems, smaller on midribs, broad-based,
recurved, laterally compressed, often sparse, sometimes
absent or replaced with bristles. Sympodial units pluri-
foliate with inflorescence remaining erect for some time,
especially on early order shoot generations (then branch-
ing dichasial); leaves and extended inflorescences often
* National Herbarium, South African National Biodiversity Institute,
Private Bag X101, 0001 Pretoria. E-mail: welman@sanbi.org.
MS. received: 2006-08-04.
aggregated terminally. Leaves broadly ovate to obovate,
oblanceolate or ovate-lanceolate, petiolate ( 1 0—40 mm
long), usually unarmed, entire to subrepand, rarely
lobed, tip acute to acuminate or obtuse, basally acute
or rounded, to at least 250 mm long on vigorous veg-
etative shoots, herbaceous, glabrescent above, white or
canescently pubescent to tomentose below. Inflorescence
cymose (in southern Africa) with few to many (20-80)
dense, small, all bisexual flowers; peduncle 2(M10 mm
long, terminal and suberect until finally pushed later-
ally by continued shoot growth, sometimes armed with
small deltoid/rosoid prickles; pedicels 15-18 mm long,
nutant at anthesis, ± erect in fruit. Calyx campanulate,
lobes broadly ovate or deltoid. Corolla stellate, mostly
less than 20 mm diam., mostly blue/purple, usually
deeply divided, 4-, 5- or 6-lobed, lobes narrowly tri-
angular or linear-lanceolate. Stamens : filaments free;
anthers equal, subleptostemonoid, lanceolate, ± attenu-
ate. Ovary crowned with shortly stipitate floor glands
or few stellate hairs; style glabrous. Fruits globose, held
± erect, 6-12 mm diam., juicy, bitter, shiny, red to pur-
plish when ripe. Seeds reticulate, 2. 0-2. 5 mm long, pale
yellow. Chromosome number : n = 12 for S. giganteum
(Bukenya-Ziraba 1996).
Distribution and ecology
Section Giganteiformia consists of ± nine species in
Africa, India and Sri Lanka. All these species are indig-
enous to Africa; only one species, S. giganteum , also
occurs in India and Sri Lanka. Members of this sec-
tion are centred in tropical East Africa from Ethiopia to
Tanzania, but outlying species are found in tropical West
Africa and temperate South Africa. They grow in grass-
land, savanna, forests and forest clearings and edges.
Taxonomy and relationships
In the subgenus Leptostemonum the plants are gener-
ally prickly at least when young; hairs, at least some of
them, are truly stellate; inflorescences are extra-axillary;
anthers are tapering, opening by terminal pores.
Bitter (1921) placed his series Giganteiformia , con-
taining Afro-Asian species, with the neotropical species
in section Torva Nees. Later authors realized that sec-
tion Torva should not include these Afro-Asian plants.
40
Bothalia 38,1 (2008)
Whalen (1984) separated the Giganteum group. He
observed that this group of African species is unusual in
containing some species with plurifoliate and others with
difoliate sympodia. The difoliate species tend to have
smaller inflorescences and sometimes tetramerous flow-
ers. However, many other distinctive characters unify
the group, among them the entire, often markedly dis-
colorous leaves with glabrate upper surfaces and closely
spaced, upwardly arching lateral veins; the small broad-
based prickles (often sparse); the inflorescence branches
with closely spaced small, pendent flowers; the tendency
to floral tetramery; and the small, juicy, erect red berries.
Jaeger & Hepper (1986) kept them together in section
Torva for convenience, but emphasized the differences
between the neotropical, mainly Central American, spe-
cies of section Torva (lobate leaves, bifoliate geminate
sympodial units, white flowers and firmer, larger, green
to yellow fruits) and the African species of that section.
Thus, they concluded. Solarium torvum Sw. and S. gigan-
teum Jacq. should not be in the same section.
Child (1998) noted that the appearance of his sec-
tion Giganteiformia closely resembles that of species of
section Brevantherum (of the subgenus Brevantherum)
in branching pattern, in the attitude of the flowering
and fruiting inflorescence and in leaf form. However,
Child (1979) also noted that most species in section
Giganteiformia are prickly, unlike section Brevantherum ,
but on the other hand, some species have a thick stel-
late indumentum like species of section Brevantherum.
(Section Brevantherum has pleiochasial, multilateral
corymbose to subumbellate cymes with a long common
axis or peduncle but with reduced rhachides. The inflor-
escence remains erect and shoot continuation is delayed
until well after anthesis.) Bitter (1921) noticed that the
pseudostipular leaves in the axils of cauline leaves in
Solanum giganteum are similar to those of S. mauritia-
num and some other species of section Brevantherum.
Species of section Giganteiformia may therefore be
regarded as parallel to the section Brevantherum, at least
in respect of branching pattern and habit. According to
Child (1998), plurifoliate sympodial units with di- to
trichasial branching, lanceolate or ovate unlobed leaves
and pleiochasial inflorescences may be regarded as ple-
siomorphic characters within the genus Solanum.
Levin et al. (2006) concluded that most of the Old
World species of subgenus Leptostemonum belong
to a single species-rich clade. Their investigations of
Solanum kwebense (from tropical and southern Africa)
and S. schimperianum (from tropical Africa) suggest
that within their large Old World clade, the S. giganteum
group appears monophyletic.
Key to species of section Giganteiformia in southern
Africa (based on Gonqalves 2005)
la Leaves ± glabrous below at maturity; prickles absent; inflor-
escence a branched cyme; seeds 3-5 x 2. 5-4.0 mm . . . S. goetzei
lb Leaves markedly hairy below at maturity; prickles mostly
present; inflorescence an unbranched or paniculiform
cyme; seeds 2. 5-3. 8 x 2-3 mm:
2a Leaves white floccose-tomentose below; anthers 2-4 mm
long S. giganteum
2b Leaves grey- or yellow-tomentose below; anthers 4-7 mm
long S. tettense
1 . Solanum giganteum Jacq., Collectanea austri-
aca ad botanicum 4: 125 (1791); Jacq.: 11, t. 328 (1793);
Dunal: 258 (1852); Wright: 94 (1904); Wright: 229
(1906); Bitter: 256 (1921); Compton: 516 (1966); Palmer
& Pitman: 1984 (1973); Symon: 117, t. 37 (1981);
Whalen: 215, fig. 10 (1984); Beentje: 580 (1994); Van
Wyk & Van Wyk: 118 (1997); Gonqalves: 88 (2005).
Type: cultivated in Hortus Vindobonensis, Vienna, origi-
nally from the Cape, South Africa (Gonqalves 2005).
S. niveum Vahl ex Thunb.: 36 (1794). Type: from Cape of Good
Hope, South Africa (Gonfalves 2005).
S. farinosum Wall, ex Roxb.: 255 (1824). Type: from India (Wright
1904).
Description (based on Gonqalves 2005)
Short-lived, soft-wooded, much-branched undershrub,
shrub or sometimes a small tree, up to 6 m high; sympo-
dia plurifoliate. Hairs stellate, white, very fine, floccose,
± sessile, regular, with many short rays. Prickles stout,
straight or slightly curved, ± flat and triangular, 1-5 mm
long, often white hairy in lower half. Branches white-
tomentose, prickles scattered, sometimes quite unarmed,
sometimes ± glabrescent. Leaves usually closely set at
ends of branches, evergreen, rarely drought deciduous;
petiole white-tomentose, 10-85 mm long; stem leaves
sometimes bearing 1 or 2 leaf-like, elliptic to obovate
pseudostipules at base, 1 0-40 x 5-20 mm; lamina mem-
branous, elliptic to broadly ovate, obovate, lanceolate or
oblanceolate, 50-250 x 20-100 mm, apex usually ± acu-
minate, base cuneate to sub-rounded, narrowing to peti-
ole and ± unequal-sided, softly textured, markedly dis-
colorous, at first velvety whitish silver-tomentose on both
surfaces, soon glabrescent and dark green shiny above,
persistently tomentose and rarely with 1 or 2 short prick-
les beneath, with 9-12 pairs of closely pinnate lateral
nerves; margin entire or ± repand-sinuate. Cymes termi-
nal or subterminal, becoming lateral, 45-100 mm long,
corymbiform to ± paniculiform, dense, 20 to > 60-flow-
ered, densely white-tomentose, sometimes ± glabrescent
in fruit; peduncle 15-45 mm long; flowers and fruit often
found on same plant, even in one inflorescence. Flowers
faintly scented, (4)5(6)-merous, ± nodding; pedicels
5-20 mm long, slender, reflexed in flower, in fruit elon-
gated up to 25 mm, ± thickened, erect. Calyx 4-6 mm
long, campanulate or cyathiform, in fruit saucer-shaped,
densely white-tomentose outside, unarmed, ± accrescent;
lobes lanceolate-triangular to deflate or ovate-triangular,
1-4 x 1-2 mm, obtuse or acute, sometimes ± acuminate.
Corolla mauve to blue or purple, rarely white, midvein
of each lobe green, rotate; limb 10-16 mm across; lobes
lanceolate to oblong, 5-7 mm long, acute to acuminate,
tomentose outside, few stellate hairs on midvein and
near apex inside, widely spreading to reflexed. Stamens
yellow; filaments ± 0.5 mm long; anthers 2-A mm long,
linear or lanceolate-elliptic in outline, with small termi-
nal pores, ± incurved. Ovary ± globose, ± 1 mm diam.,
mostly glabrous; style 5-8 mm long, exceeding stamens,
straight or ± curved at apex, mostly glabrous. Fruits often
numerous, ± globose, 5-10 mm diam., smooth, glossy,
green ripening through orange to bright red, finally
purplish red. Seeds numerous, compressed, obliquely
reniform to suborbicular in outline, 2. 5-3. 8 x 2-3
mm, shallowly reticulate, straw-coloured to ± whitish.
Chromosome number. 2n = 24 (Bukenya-Ziraba 1996).
Bothalia 38,1 (2008)
41
Taxonomy and diagnostic characters
Solanum giganteum shows some resemblance to the
invasive shrub, S. mauritianum Scop., which is a native
of South America and a Declared Weed in South Africa
(Henderson 2001). However, that species has no prick-
les, has yellow fruit and is very densely velvety or felty
hairy in almost all parts. Gon9alves (2005) stated that in
5. giganteum the very fine snow-white tomentum on the
underside of the leaves (tending to become greyish or
yellowish in old herbarium specimens), contrasting with
the almost glabrous upper surface, is highly distinctive.
Distribution
Solanum giganteum is a widespread, mainly Afro-
montane species that has a disjunct distribution in Africa
south of the Sahara from Nigeria and Cameroon in the
west to Ethiopia in the northeast and down to the Cape
Peninsula (Gbile 1979). In southern Africa it has been
recorded in Swaziland and South Africa where it occurs
in all provinces except the Free State and Northern Cape
(Figure 1). It also grows in southern India and Sri Lanka
(Deb 1979). Gon9alves (2005) reported that S. giganteum
is widespread throughout tropical and southern Africa,
usually as a highland species, recorded from Ethiopia
southwards throughout East Africa to South Africa
(Western Cape) and westwards to Nigeria, Cameroon and
Gabon, extending to the Canary Islands (Tenerife).
Africa can be divided into 18 major phytochoria accord-
ing to White’s system (1976); Solanum giganteum is wide-
spread mainly in the Afromontane Archipelago-like centre
of endemism in western, eastern and southern Africa.
Ecology
In southern Africa Solanum giganteum usually grows
in dense to partial shade in forests, forest margins, under-
growth and clearings, among trees and often on river
banks, in ravines and other moist places. It is common
in high rainfall areas, up to 2 000 mm annual rainfall
and is a component of woodland and grassland at a wide
range of altitudes from 5-2 000 m. It can grow on steep
or gentle slopes of all aspects and prefers humus-rich,
well-drained brown or red sandy or loamy soils, also
stony soils. The geology has been described as granite,
Swaziland rocks, middle Ecca sandstone. The flower-
ing time is in summer from October to April. Fruiting
specimens have been collected throughout the year, but
mainly from December to July; fruit remain on the plant
for at least six months. Larger birds such as bulbuls,
doves and loeries feed on the fruit, particularly in late
winter when food is in short supply. It can also occur as
a weed in disturbed areas. Wells et al. (1986) listed 5.
giganteum as an occasional ruderal or silvicultural weed
that grows in dry to moist soil, in temperate to subtropi-
cal areas with summer, winter or all year rainfall. In the
Flora zambesiaca area, it grows in forest edges, river-
ine forest and among rocks on granite outcrops; from sea
level to 1 650 m (Gon9alves 2005).
Medicinal and horticultural uses
Pappe (1850) reported that the application of the
woolly lower surface of the leaves of Solanum gigan-
teum ( S . niveum ) to festering ulcers ‘cleanses them, and a
FIGURE 1. — Distribution of Solanum giganteum, •; and 5. goetzei,
O, in the FSA region.
cure is afterwards effected by applying the upper surface.
Hence the Dutch name Geneesblaren. The fresh juice
of the berries and leaves, when formed into an ointment
with lard or fat, is also in use amongst the farmers for the
same purpose’. Watt & Breyer-Brandwijk (1962) noted
that the Xhosa and Mfengu in the Eastern Cape use the
berry to curdle milk. The berry has also been used as a
remedy for throat abscesses. Doses of the fresh immature
and mature fruit have produced no ill effect in the rab-
bit. Various parts of the plant have given negative tests
for saponin. Fox & Norwood- Young (1982) had a report
from Modjadje’s Reserve in Duiwelskloof, Limpopo
Province, that the berry of this plant is used to curdle
milk, and also that the ‘red bitter apple’ is considered edi-
ble. The fruit is used for throat ulcers by the Zulu, Xhosa
and Mfengu; the fruit also curdles milk. The leaves are
used for festering sores, either directly applied as a dress-
ing or used in ointments (Hutchings et al. 1996).
The fruits and leaves of Solanum giganteum are used
in traditional medicine in Uganda. The leaves are used
for the treatment of insomnia and the leaves and fruits
for ulcers (Bukenya-Ziraba 1996).
Solanum giganteum is often seen in botanical gardens,
especially in the northern hemisphere. It is cultivated
in gardens and parks in South Africa and elsewhere as
an attractive ornamental shrub or small tree with shiny
dark green leaves and showy bright red fruit. It forms a
spectacular subtropical bedding plant. It can be used as a
background plant in herbaceous borders and also as part
of a hedge (Nichols 2002). Seeds germinate easily when
cleaned out of the fruit. Seedlings grow rapidly and the
plant should be fruiting in the second year. Bailey &
Bailey (1977) listed it as a garden subject for southern
California. Symon (1981) reported that it is occasion-
ally grown in gardens in Australia, but is not known to
be naturalized. In English-speaking countries it is known
as African holly.
Chemotaxonomy and chemistry
Maiti et al. (1979) found that Solanum giganteum
contained sufficient quantities of total alkaloids to war-
42
Bothalia 38,1 (2008)
rant further studies in order to develop it as a source of
raw material for the steroid industry. The fruit contained
1.9% solasodine as dry weight. Dan & Dan (1984)
reported that solanogantine and solanogantamine (both
3-aminosolanidane derivatives) were isolated as major
leaf constituents of S. giganteum. Hutchings et al. (1996)
reported that solanogigine and three 3-aminosolanidanes
namely solanogantine, solanogantamine and isosolano-
gantamine, had been isolated from this plant. Solasodine
had been found in the fruit and leaves.
Notes
Gbile (1986) did epidermal studies on Solarium gigan-
teum and stated that straight upper epidermal walls had
previously been observed in mountain species of other
sections of Solanum : 'For some unknown reason, most
mountain species have straight upper epidermal walls.
Some workers, however, observed that straight-walled
epidermal cells are commoner in xeromorphic plants
than in mesomorphic ones, which typically have undu-
late cell walls’.
Gbile & Sowunmi (1979) described the pollen of
Solanum giganteum from Nigeria as subprolate and tri-
angular, polar axis ± 30 pm, equatorial diameter ± 25
pm, with the exine pattern faintly distinct.
Solanum giganteum appears on the official Tree
Lists of South Africa (no. 669.4) and Zimbabwe (no.
1014). Various common names for this species have
been recorded in southern Africa e.g. healing-leaf tree,
geneesblaarboom (Afrikaans), icuba lasendle (Xhosa).
Gonqalves (2005) listed red bitter apple or red bitter
berry for the Flora zambesiaca area.
2. Solanum goetzei Dammer , Botanische Jahrbiicher
28: 473 (1901); Wright: 218 (1906); Bitter: 269 (1921);
Jaeger: 352 (1985); Beentje: 580 (1994); Gon9alves: 91
(2005). Types: syntypes from Tanzania (Gonqalves 2005).
S. muha Dammer: 186 (1906). Type: syntypes from Tanzania
(Gotifalves 2005).
Description (based on Gon9alves 2005)
Erect, much-branched perennial herb or shrublet,
rarely scandent, up to 2 m high; prickles absent; sym-
podia difoliate; hairs stellate, whitish, sometimes viola-
ceous or reddish tinged, minute, ± sessile, regular, with
many short rays. Branches terete, floccose-tomentose
at first, gradually glabrescent. Leaves solitary or partly
subgeminate; petiole 5-45 mm long, base purple; lamina
thin, soft, membranous, lanceolate to obovate, 30-225
x 10-90 mm, apex acute or acuminate, rarely ± obtuse,
base cuneate, gradually narrowing into the petiole and ±
unequal-sided, somewhat subrepand to scarcely undu-
late, rarely ± entire, tending to dry blackish, initially
with ± abundant whitish hairs, soon becoming sparsely
hairy to quite glabrous with age, dark green above,
underside paler, with 5-9 pairs of lateral nerves. Cymes
soon leaf-opposed or leaf-remote, forked 1 or 2 times or
rarely unbranched, racemiform, 15-30 mm long, 3-24-
flowered, ± pulverulent-tomentose; peduncle 1-18 mm
long; densely stellate-tomentose, rhachis 1-8 mm long.
Flowers (4)5-merous, ± nodding, unscented; pedicels 5-
1 1 mm long, slender, purple, often at first densely hairy,
glabrescent, except at base, in fruit elongated to 16 mm,
somewhat thickened distally, ascending or erect. Calyx
2-5 mm long, somewhat accrescent, campanulate or
cupular, ± hairy; 5-lobed, lobes ± unequal, ovate-trian-
gular to triangular-elongate or broadly obovate, 0. 5-3.0
x 0.5-1 .0 mm, acute or mucronate to narrowly long-acu-
minate, in fruit enlarged to 5 x 2 mm, finally ± reflexed.
Corolla bluish to pale violet or lilac, sometimes white,
campanulate-stelliform; limb 8-15 mm across; deeply
5-lobed, lobes ± lanceolate, 3-8 x 1.0-3. 5 mm, acute, ±
densely hairy outside mainly on median part, glabrous
except for a few stellate hairs scattered along midvein
or only near apex inside, erect to reflexed. Stamens 5,
subequal; filaments glabrous, 0. 5-1.0 mm long; anthers
lanceolate-elliptic in outline, 3-5 x 0.8- 1.2 mm, slightly
obtuse, ± emarginate at apex, yellow. Ovaiy ± globose,
0. 7-1.0 mm diam., glabrous or with few minute glands
near apex; style 4. 5-8. 5 mm long, glabrous, exceeding
stamens, often arcuate at apex, glabrous or with few min-
ute glands near base, white; stigma subglobose, green.
Fruit globose, 6-10 mm diam., green, shining bright to
deep red when ripe, soft. Seeds few, compressed, some-
what obliquely reniform, 3-5 x 2. 5—4.0 mm, surface
reticulate-tuberculate, pale yellowish. Chromosome num-
ber. unknown. Figure 2.
Taxonomy and diagnostic characters
Solanum goetzei is closely related to S. schuman-
nianum Dammer from the upland forests of Kenya and
Tanzania and also to S. anomalum from West Africa.
This is the species which Ross (1972) referred to as
Solanum sp. no. 30 ( Ward 3840) from Tongaland. The
shrubby, unarmed habit and thin, soft, dark green, gla-
brescent lanceolate to obovate leaves as well as the very
large seeds of S. goetzei are unique among the species of
Solanum found in southern Africa.
Distribution
This species has been recorded from Kenya, Tanzania,
Malawi and Mozambique down to the Ingwavuma,
Ubombo and Hlabisa (Tongaland) Districts of KwaZulu-
Natal (Figure 1). According to White’s system (1976),
Solanum goetzei is endemic to the Zanzibar-Inhambane
and Tongaland-Pondoland regional mosaics; these form an
intermittent strip of forest along the east coast of Africa.
Ecology’
Whalen (1984) observed that his Giganteum group
is divided between montane forest and savanna habi-
tats in Africa. The species with difoliate sympodia (e.g.
Solanum goetzei) are probably derived within the group
and occupy open savannas, grasslands and forest mar-
gins. According to Jaeger (1985), this species grows in
disturbed or open places in the forests of East Africa
between a few metres above sea level and 1 200 m alti-
tude. Beentje (1994) reported that S. goetzei grows in
forest, riverine forest and coastal bushland in Kenya. In
the Flora zambesiaca area, Gonqalves (2005) reported
that this species grows in dry forest margins and the
understorey, Combretum-Terminalia and mopane wood-
land, savanna woodland and coastal bushland, sometimes
on termite mounds or around granite outcrops, ruderal
places and other areas of disturbance, particularly along
Bothalia 38,1 (2008)
43
FIGURE 2. — Solarium goetzei, A-D, M.C. Ward 1328 (PRE); C.J. Ward 3840 (PRE). A, habit, x 1; B, flower, x 3; C, fruit, x 1.5; D, seed, x 1.5.
Artist: G. Condy.
roads. In South Africa it grows on well-drained sandy
soil at forest margins, in clearings and along forest paths,
also in woodland, in deep as well as light shade, from
about sea level to 100 m altitude. In KwaZulu-Natal,
flowering and fruiting material have been collected from
November to May.
Medicinal and other uses
Dammer (1901) cited Goetze who wrote that the
roots are used as medicine against toothache. Magogo &
Glover 1030 (PRE) from Tanzania noted that ‘a hot poul-
tice made from the leaves is used to reduce swellings and
to draw out abscesses; also used to draw out whitlows on
the fingers’. Jaeger (1985) stated that Solanum goetzei is
used as a leaf vegetable in Kenya, while Beentje (1994)
noted that, also in Kenya, a poultice of the leaves is used
to draw out abscesses.
Toxicity and chemistry
The chemistry and toxicity, if any, of this plant have
not yet been investigated.
3. Solanum tettense Klotzsch in Peters, Natur-
wissenschaftliche Reise nach Mossambique, Botanik 6,1 ;
237 (1861); Dammer: 355 (1895); Wright: 212 (1906);
Bitter: 276 (1921); Gonqalves: 86 (1997); Gonqalves: 92
(2005). Type: Mozambique, Tete Prov., Tete (‘Tette’), W.
Peters s.n. (B, holo.; BM, K, iso.) [Gonqalves 1997]).
44
Bothalia38,l (2008)
Description (based on Gon9alves 1997)
Erect, semi-woody herb or shrub, laxly to much
branched from base, sometimes scrambling, up to 3 m
high; covered with grey to yellowish/brownish dense
tomentum of ± sessile, short-radiate, stellate hairs, or
bearing a long central ray, sometimes apically glandu-
lar, minute to large hairs all over (at least when young),
and also of simple, apically glandular, spreading hairs
sparsely intermixed, then pruinose, finally glabrous on
some parts; prickles few to many, laterally compressed,
broad-based, straight or curved, ± stout, pale yellow to
brown, 1-5 mm long, stellate-tomentose in lower half
or glabrous, sometimes absent; sympodia difoliate.
Stems and branches ± terete, glabrescent, bark smooth
to ± rough, pale yellow/brown-grey to dark grey, some-
times with conspicuous lenticels. Leaves solitary or
appearing geminate, sometimes closely set at terminals;
petiole 4-40 mm long, densely to ± hairy, prickles usu-
ally absent; lamina membranous, papery, leathery or
± fleshy, ± discolorous, obovate to lanceolate, 15-140
x 5-80 mm, apex obtuse-rounded to ± acuminate,
base rounded to cuneate, narrowing into the petiole, ±
unequal, with 4-7 pairs of curved-ascending lateral
nerves, greyish/yellowish below, tomentose to subgla-
brescent; green above, stellate-hairy to glabrescent;
rarely 1-few prickles on midrib; margins repand-sinu-
ate to entire. Inflorescences terminal, cymes becoming
lateral, unbranched to several times forked, 30-50 mm
across, few to many-flowered, often dense; peduncle
0- 30 mm long; rachis 7 — 45 mm long. Flowers 4 or
5(-7)-merous, ± nodding; pedicels 3-13 mm long, elon-
gated in fruit, up to 18 mm long, ± thickened distally,
erect. Calyx greyish/yellowish, densely to ± hairy out-
side, 2. 5-6.0 x 3-6 mm, ± accrescent, campanulate or
cupular; lobes lanceolate to broadly obovate, 1.0-3. 5 x
1- 2 mm, apically rounded, long-acuminate, 5x2 mm
in fruit, ± reflexed, glabrous inside. Corolla white to
mauve, blue, violet or purple, 6-13 mm long, ± rotate;
limb 6-22 mm across, lobes oblong-ovate to linear, 4-
11 x 1.5-4. 5 mm, apex acute/obtuse, greyish or whitish
hairy outside, mainly on median part, mostly glabrous
inside, erect to reflexed. Stamens ± equal, glabrous,
exserted; filaments 0-1.5 mm long, whitish; anthers yel-
low to orange-yellow, 4-7 mm long, ± lanceolate in out-
line, opening by 2 small, oblique pores. Ovaty globose/
ellipsoid, 1.0-1. 5 mm diam. /length, glabrous; style 5-11
mm long, slender, longer than stamens, straight or ± api-
cally curved, glabrous; stigma small, capitate, obtuse or
slightly 2-lobed. Fruits globose, 5-10 mm diam., green
turning yellow-brown to deep red when ripe, fleshy,
glossy, glabrous, in axillary clusters or pseudo-terminal
cymes. Seeds numerous, compressed, obliquely reni-
form, reticulate-tuberculate, 3. 0-3. 5 x 2-3 mm, ± pale
yellow, drying blackish. Chromosome number. 2n = 24
(Bukenya-Ziraba 1996).
Key to varieties (from Gompalves 1 997)
Stellate hairs often bearing a long central ray intermixed with
simple, long, spreading, ± abundant hairs, both apically
glandular var. tettense
Stellate hairs eglandular, sometimes intermixed with simple,
short, abundant, apically glandular hairs var. renschii
3a. var. tettense
See description above.
3b. var. renschii (Vatke) A. E. Gong, in Kirkia 16,1:
89, fig. 1 (1997); Gon9alves: 93, t. 18 (2005). Type:
Kenya, Central Prov., Machakos or Kitui Dist., Ukamba
(Ukambani area), J.M. Hildebrandt 2735 (B, holo.; W,
iso.) [Gon9alves 1997].
S. renschii Vatke: 328 (1882). Type: as above.
S. kwebense N.E.Br. ex C.H. Wright: 225 (1906). Types: Botswana,
Ngamiland, Kwebe Hills, Lugard 50 (K, lectosyn.), Mrs Lugard 62 (K,
syn.) [Gonsalves 1997],
S. luederitzii Schinz: 264 (1912). Type: Namibia, Hereroland, Liide-
ritz la [Podlech & Roessler 1969].
5. upingtoniae Schinz: 266 (1912). Type: Namibia, Amboland, Os-
hando, Schinz 868 [Podlech & Roessler 1969].
S. tenuiramosum Dammer: 244 (1912). Type: Botswana, Masalan-
yane Pan (Massaringani Vlei), Seiner 11.271 (B, holo.) [Gonsalves
1997],
S. chondropetalum Dammer: 335 (1915). Type: Namibia, Damara-
land, Naugubais, Dinter 1448 [Podlech & Roessler 1969].
Taxonomy and diagnostic characters
Wright (1906) and Bitter (1921) recognized Solanum
kwebense, S. renschii and S. tettense as three distinct
species. Lebrun & Stork (1997) recognized both S. kwe-
bense and S. renschii (S. tettense), whereas Podlech &
Roessler (1969) recognized only S. kwebense. Studies
by Gon9alves (1997) revealed that both the vegetative
and inflorescence characters of these three species show
a strong intergradation. Therefore these taxa constitute
a single polymorphic species with great morphological
and ecological diversity. S. tettense is the correct name
for this species. The two varieties cannot be separated on
geographical or ecological terms.
The diagnostic characters of Solanum tettense in rela-
tion to the other two members of section Giganteiformia
in southern Africa, are outlined in the key above. S.
tettense can be distinguished from other Solanum species
in the same area by its pedunculate cymes and entire,
discolorous leaves without prickles.
Gonsalves (2005) observed that Solanum tettense is
somewhat intermediate between section Giganteiformia
and section Oliganthes', it is relatively common in the
Flora zambesiaca area, but is easily overlooked or mis-
identified.
Distribution
Solanum tettense is widespread throughout tropical
and southern Africa; it is recorded from Ethiopia south-
ward through East Africa to South Africa, and westward
to Zaire, Angola, Botswana and Namibia. In southern
Africa, var. renschii is found in Namibia, Botswana
and the Limpopo and Mpumalanga Provinces in South
Africa (Figure 3). In southern Africa var. tettense occurs
only in Botswana (Figure 3). According to White’s sys-
tem (1976), 5. tettense is widespread in the Somalia-
Masai regional centre of endemism, but is also found in
the Zambezian regional centre of endemism and in the
Kalahari-Highland regional transition zone.
Bothalia 38,1 (2008)
45
Ecology’
In southern Africa, Solarium tettense grows on well-
drained red or brown, shallow to deep, dry to damp,
sandy or loamy soils that can be stony or quartzitic. It
is often found on weathered granite, gneiss or dolomite
outcrops and inselbergs, also on calcified dolomite or cal-
crete soils. This species grows on flat areas such as the
edges of pans or on flood plains, but also on moderate
or steep slopes of all aspects, on sandy dunes and rocky
hillsides. It has been collected on termitaria, in disturbed
and overgrazed areas, also on roadsides. S. tettense grows
in full sun but more often in the semi-shade or shade of
taller shrubs or trees, also in dense thickets. Larger leaves
develop in shady habitats. It becomes invasive in over-
grazed vegetation with a reduced grass cover.
Solatium tettense grows in vegetation types ranging
from grassland and savanna to various kinds of decidu-
ous or evergreen woodland and bushland. Trees and
shrubs such as Acacia , Boscia , Combretum and Grewia
are commonly associated with S. tettense.
The rainfall in the distribution area of Solatium
tettense var. renschii in southern Africa, is 400-1 000
mm per year and it grows at an altitude of 335-1 370 m.
Flowering time'. October to March, mainly from January
to March. Fruiting time : November to May, mainly from
January to April.
Gonqalves (2005) stated that in the Flora zambesi-
aca area, including Botswana, Solanum tettense grows
in mixed woodland, mopane and wooded grassland or
thickets, extending into miombo on termite mounds, also
streamsides, rocky places and areas of disturbance in
moist and semi-arid situations at 100-1 600 m altitude.
Medicinal and other uses
Maguire noted on the label of his specimen B.
Maguire 2272 in PRE (collected at Karakuwise in
Namibia in 1953), that the fruits provide ingredients for
the arrow poison of the Khoi-San (Bushmen). Barnard
noted on the label of his specimen Barnard 153 in
FIGURE 3. — Distribution of Solanum tettense var. tettense, O; and var.
renschii, ■, in the FSA region.
PRE (collected in Sekhukhuneland near Lydenburg
in Mpumalanga Province in November 1934), that
'the roots are cooked and placed in a calabash (fruit of
Lagenaria siceraria ), in which a small opening has been
made. The calabash is then fastened over a suppurat-
ing wound to draw out the pus’. Collectors have noted
that Solanum tettense is eaten by the large antelope, the
eland. The Tswana common name mwarasupe is listed
by Miller on the label of his specimen B/469 in PRE,
collected in Botswana. The Kamba in Kenya use the
roots against typhoid (Beentje 1994).
Toxicity
Pienaar et al. (1976) proved that Solanum tettense can
be poisonous to cattle. A neurological disease of cattle
(named maldronksiekte by farmers), occurring in a local-
ized, badly overgrazed area of the Limpopo Province
of South Africa, was experimentally reproduced at
Onderstepoort Veterinary Institute by feeding S. tettense
var. renschii plants (known as rooibessie among fann-
ers) to cattle. According to Vahrmeijer (1981), the dis-
ease is characterized by temporary loss of balance and
transient epilepsy-like seizures precipitated by a vari-
ety of stimuli, such as exercise, handling (dipping and
loading) and fright. The animal staggers about with an
extended and slightly twisted neck, and in serious cases
falls to the ground. After a while it rises again as if noth-
ing were wrong. Losses are suffered when animals are
injured during falls. When not disturbed, most affected
animals appear to be completely normal. The poison
affects the central nervous system and causes permanent
damage to, and interferes with the function of the cere-
bellum. Maldronksiekte is a chronic intoxication with
a latent period of at least 50 days between ingestion of
the plant and the appearance of typical clinical signs.
Donkeys, goats and sheep are apparently not affected by
the poison. The chemistry of S. tettense var. renschii was
not investigated in the above study.
SPECIMENS EXAMINED
(southern Africa only)
Specimens held at PRE, unless otherwise indicated.
The numbers in brackets indicate the identity of the
specimens: (1) Solanum giganteum ; (2) S. goetzei ; (3a)
S. tettense var. tettense; (3b) S. tettense var. renschii.
Abner 71 (3b). Acocks 11519 (1). Acocks & Hafstrom 1376 (3b). Allen
346 (3b). Ankiewicz 8 (2).
Barnard 153, 529 (3b). Barnard & Mogg 857, 1013 (1); 1082 (3b).
Bayliss 1132, 7052 (1). Brink 294 (1). Bruce 46 (3b). Buerger 1020
(3b). Buitendag 1162 (1) Lowveld Botanical Garden, PRE. Burgoyne
3076, 3423, 3241 (3b).
Codd 5921 (1); 4147, 5984. 8884 (3b). Coetzee 1154 (1). Compton
25550. 30071 (1). Crampton 113 (1). Curson 465, 481, 526 (3b).
Davidse 6814 (1). Devenish 1580 (1). De Winter 2817 (3b); 8273
(1). De Winter & Leistner 5095, 5516 (3b). Dinter 5312, 7448 ( 3b).
Dlamini s.n. PRE31390 (1).
Edwards 1318 (1).
Flanagan 472 (1).
Galpin 2854, 7815, 9039 (1). Germishuizen 7540, 7765, 9552, 9770
(3b). Gerstner 5747 (1); 6047 (3b). Giess 9277, 12592, 15016 (3b).
Giess & Muller 11808, 13977 (3b). Giess, Watt & Snyman 11129 (3b).
Giffen 1086, 1437 (1). Goldblatt & Manning 8399 (1).
46
Hansen 3355 (3b). Harbor TRV14089 (3b). Hardy, Retief & Herman
5363 (1). Haynes 967a (1). Hemm 597 (1 ). Henning 5 (3b). Hilner 475
(1). Hines 638 (3b). Holt 135 (1). Hobohm PRE41601 (3b). Hoffmann
LH204 (3b). Hulley, Olckers & Hill 379 (3b). Huntley 1049 (3b); 1306
(1).
Jacobsen 729, 2906 ( 1 ); 2200 (3b). Junod 4291 (1).
Kerfoot & Falconer 41, 156 (3b). Killick 401, 1707 (1). Kluge 718 (1)
Lowveld Botanical Garden, PRE. Kotze 84 (3b).
Lawson 324, 340 (2) NH. Leendertz 554, 689 (1). Leistner 3160 (3b).
Leistner, Oliver, Steenkamp & Vorster 41 (3b). Lent 46 (3a). Le Roux
819 (3b). Liebenberg 2950 (1); 4821, 4918 (3b). Louw 15 ( 1 ).
MacDevette 883 (2) NH. Maguire 2272 (3b). Mannheimer &
Mannheimer CM530 (3b). Marloth 4311, 5677 (1). Marriott PRE22643
(1). Meeuse 10457 (3b). Merxmiiller & Giess 30455 (3b). Meyer 1031
(1). Miller B/469, B/1197 (3b). Mogg 11511, 12455, 15004, 36077 ( 1).
Moll & Morris 693 (1). Morris 711 (1). Moss 2085 (3b). Morze 2085
(1). Muller & Biegel 2302 (3b). Muller & Scheepers 225 (1). Munro
TRV23163 (1).
Naude 1/94 (3b). Nienaber EN325 (1).
Obermeyer TRV28143, TRV29223 (1). Obermeyer, Schweickerdt &
Verdoorn 57 (3b). Onderstall 916 (1) Lowveld Botanical Garden, PRE.
Paterson TRV25847 (1 ). Pegler 709, 2890 (1). Pentz 299 (1). Pienaar
367 (3b). Pole Evans 238 (1); 1933, 4536 (3b). Pott PRE59768 (1).
Prior PRE41 307 ( 1).
Raal 809, 1388 (3b). Repton 621, 1005 (1). Rodin 3959, 4519 (1).
Rogers PRE3791, 14455, 23467(1).
Scheepers 676 (1). Schoenfelder 1004 (3b). Sim 1272, 20000, 20006
(1). Smith 357A, 6868 (1); 3101 (3b). Stalmans 555 (1). Stephens 7 (1).
Story 6481 (3b). Straub 158 (3b). Strey 3477 (3b); 3843 (1); 4786 (2)
NH, PRE.
Theron 1511 (1); 2879 (3b). Thode A1292 (1). Thorncroft TRV3930
(1). Tolken & Hardy 894 (3b).
Van der Schijff 3503 (3b); 4600, 4874, 7357 (1). Van der Spuy 10 (3b).
Van Rooyen 3401 (3b). Van Son TRV29022 (3b). Van Vuuren 89 (1).
Van Warmelo 275 (3b). Van Wyk 3681 (1). Van Wvk & Van Wvk 1599
(1). Venter 3795 ( 1); 12820 (3b). Volk 966. 2911 (3b).
Wahl TRV15368 (1). Ward 87, 399, 1328 (2) NH; 608, 2961 (1); 3226,
3840, 7053 (2). Watt & Breyer-Brandwijk 1027 (1). Welman 432 (1).
Westfall 1009 (1). Wild & Drummond 6857 (3a).
Zwanziger 758 ( 3b).
ACKNOWLEDGEMENTS
The late Dr R.N. Lester, Birmingham, United King-
dom, gave much appreciated assistance with this project.
Ms G. Condy of SANBI is thanked for the line drawing
of Solatium goetzei. Ms H. Steyn of SANBI is thanked
for the distribution maps. The Curators of NH and PRE
are thanked for loans of their specimens. The referees
are thanked for valuable advice.
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WRIGHT, C.H. 1904. Solanaceae. In W.T. Thiselton-Dyer, Flora cap-
ensis 4,2: 87-121.
WRIGHT, C.H. 1906. Solanaceae. In W.T. Thiselton-Dyer, Flora of
tropical Africa 4,2: 207-261.
Bothalia 38,1: 49-55 (2008)
Two new species of Babiana (Iridaceae: Crocoideae) from western
South Africa, new names for B. longiflora and B. thunbergii , and com-
ments on the original publication of the genus
P. GOLDBLATT*, J.C. MANNING** and R. GEREAU***
Keywords: Babiana Ker Gawl., new names, new species, range extension, replacement names, section Teretifolieae , South Africa, taxonomy
ABSTRACT
Babiana symmetrantha and B. virescens are two new species of section Teretifolieae of this southern African genus,
now comprising 90 species. Babiana symmetrantha blooms in August and early September and is restricted to the summit
of the Langberg, which lies along the border of Namaqualand and the western Karoo near Loeriesfontein. One of only three
species of the section with radially symmetric flowers, it is acaulescent and has linear, almost plane leaves covered with long
hairs, a perianth tube 45— 60(— 75) mm long, and subequal, spreading tepals. Babiana virescens blooms in early winter, in
May and June, and occurs in seasonally moist open ground on gentle slopes and valley bottoms in southern Namaqualand
between Nuwerus and Bitterfontein. It is distinguished by the slightly twisted, oblong leaves with thickened, pale margins,
and the glabrous cataphylls and leaf sheaths. The stem, which may be produced up to 1 00 mm above the ground, is simple
or has a single short branch and the spikes bear 2-6 greenish flowers, the tepals of which are unusually narrow and longer
than the perianth tube. We also propose a new name, B. tubaeformis, for the homonym B. longiflora , and provide a new
combination B. hirsuta based on the 1783 name Antholyza hirsuta Lam., which replaces B. thunbergii Ker Gawl. (1804).
Lastly, we present an argument that Babiana is correctly attributed to Ker Gawler alone and that its protologue dates from
1802, contrary to some current authorities.
INTRODUCTION
Botanical exploration in western southern Africa
since the completion of our monograph of Babiana in
February 2005 (Goldblatt & Manning 2007) has resulted
in the discovery of two additional species of this south-
ern African and largely western South African genus.
Including the new species described below, Babiana
now comprises 90 species, all but two of which are
endemic to the southern African winter rainfall zone.
The new species are B. symmetrantha, from the isolated
Langberg massif between Namaqualand and the western
Karoo, which is unusual in its linear leaves and radi-
ally symmetric flower with an elongate perianth tube
45-60(-75) mm long, and B. virescens from the granite
hills of southern Namaqualand, which has narrow tepals,
usually a well-developed aerial stem, and unusual, twis-
ted leaves with thickened, hyaline margins. Both fall
in section Teretifolieae G.J. Lewis emend. Goldblatt &
J.C. Manning. Formal descriptions, illustrations, and a
discussion of the biology and relationships of the two
species are presented below. Recent collections have also
extended the known range of B. ambigua, with a first
record for the Northern Cape. Then, two names in cur-
rent use in the genus, need to be replaced. Babiana lon-
giflora is a homonym, and we propose the replacement
name, B. tubaeformis, for the species, and Antholyza
hirsuta Lam. (1783) is a legitimate earlier epithet for B.
thunbergii Ker Gawl. (1804), hence we provide the new
combination B. hirsuta for this common plant of the
* B.A. Krukoff Curator of African Botany, Missouri Botanical Garden,
RO. Box 299, St. Louis, Missouri 63166, USA.
** South African National Biodiversity Institute, Private Bag X7,
Claremont 7735, Cape Town.
*** Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri
63166, USA.
MS. received: 2007-08-03.
west coast of South Africa. Finally, the question of the
author of the genus is re-examined in the light of some
confusion as to the correct citation.
TAXONOMY
Babiana symmetrantha Goldblatt & J.C. Man-
ning, sp. nov.
Plantae 100-200(-300) mm altae foliis inclusis, caule
subterraneo vel supra terrain producto simplici vel unira-
moso, foliis anguste linearibus usque ad 200 mm longis
2— 5(— 8) mm latis, spica 2- vel 3-flora, bracteis viridibus,
30-35 mm longis bractea interiore ad apicem furcata,
floribus hypocrateriformibus actinomorphis violaceis
basem versus colore atroviolaceo notatis tubo exteme
albo, die odorem Violae exhalentibus, tubo perianthii
45— 60(— 75) mm longo, tepalis aequalibus patentibus lan-
ceolatis (22-)28-35 x 5-7 mm, filamentis ± 10 mm lon-
gis, antheris ± 5 mm longis ad apicem connatis, ovario
glabro, stylo diviso infra basem antherarum, ramis styli
4-5 mm longis.
TYPE. — Western Cape, 3019 (Loeriesfontein): sum-
mit plateau of the Langberg among dolerite boulders,
(-DB), 5 September 2006, Goldblatt & Porter 12772
(NBG, holo.; K, MO, PRE, S, iso.).
Plants 100-200(-300) mm high, including leaves,
forming clumps. Corms globose, 8.5-11.0 mm diam.,
with fibrous tunics. Stem subterranean or reaching above
ground when growing through vegetation, producing
slender cormlets in lower axils, simple or 1 -branched,
enclosed by neck of brittle, brown fibres. Leaves basal,
linear, exceeding flowers, up to 200 x 2— 5(— 8) mm, flat,
sparsely pubescent. Spike 2- or 3-flowered; bracts green,
50
Bothalia 38,1 (2008)
becoming dry and light brown at tips, 30-35 mm long,
finely pubescent, long-attenuate, inner slightly shorter
than outer, forked apically and 2-keeled. Flowers acti-
nomorphic, salver-shaped, tepals outspread when fully
open, violet with darker violet marks near base of tepals,
sometimes lower lateral tepals with white marks out-
lined with violet, sweetly scented of violets during day;
perianth tube cylindric, 45-60(-75) mm long, straight,
slightly expanded in upper 10 mm, hollow above but
with thick walls tightly enclosing style in lower half,
containing nectar; tepals subequal, lanceolate, (22-)28-
35 x 5-7 mm, inner tepals narrower than outer. Stamens
symmetrically arranged, erect; filaments ± 10 mm long,
exserted ± 7 mm, purple; anthers arched inward, ± 6 mm
long, contiguous at tips, white; pollen white. Ovary gla-
brous; style erect, purple, dividing below base of anthers,
style branches 4-5 mm long, expanded in upper third,
arching between anthers. Capsules and seeds unknown.
Flowering time : mid-August to mid-September. Figure
1.
Distribution and ecology, restricted to the summit
plateau of the Langberg west of Loeriesfontein in the
Northern Cape (Figure 2), Babiana symmetrantha
appears to be a narrow geographic and edaphic endemic.
The Langberg, an isolated, flat-topped mountain ± 1 000
m in elevation, is capped by a dolerite sill and B. sym-
metrantha occurs in heavy red clay among dolerite boul-
ders in succulent Karoo vegetation.
Discussion: the species has inner floral bracts that are
forked at the apex, placing it in section Teretifolieae , one
of the three sections of the genus currently recognized
(Goldblatt & Manning 2007). It is one of just three spe-
cies of the section with a radially symmetric perianth,
the others being Babiana pygmaea (Burm.f.) Baker and
B. radiata Goldblatt & J.C. Manning, but B. symmetran-
tha is distantly allied to these species and is more likely
related to the B. framesii L. Bolus-5, sambucina Ker
Gawl. group, which also has an underground stem and
elongate perianth tube, but bilabiate and often scented
flowers. Apart from the radially symmetric flowers,
B. symmetrantha is unusual in its linear, almost plane
leaves that show little hint of the pleating present in
most species. We assume that the long-tubed flowers are
adapted for pollination by long-proboscid flies as they
have the long, hollow perianth tube associated with the
Prosoeca peringueyi pollination system (Goldblatt et al.
1995; Manning & Goldblatt 1996; Goldblatt & Manning
2000). Flowers in this guild are typically unscented but
floral fragrance is present, and possibly atavistic, in a
few species, including B. framesii and some popula-
tions of B. sambucina subsp. longibracteata (G.J. Lewis)
Goldblatt & J.C. Manning.
Babiana symmetrantha was only discovered in 2006
when we mounted an expedition to explore the botany of
the Langberg. The mountain stands some 1 000 m high,
well above the surrounding low granite domes and sandy
flats of eastern Namaqualand and the western Karoo.
Geologically the mountain consists of rocks of early
Karoo System age, mostly bedded Ecca shales, intruded
by dolerite sills. The flat, slightly tilted summit plateau
consists of weathered dolerite boulders and heavy, red
FIGURE 1 . — Babiana symmetrantha. Goldblatt & Porter 12772 (NBG):
A, whole plant. B, C, flower: B, front view, C, side view. D,
outer (left) and inner (right) floral bracts. Scale bar: 10 mm.
Artist: John Manning.
clay soil derived from weathered dolerite. The vegeta-
tion above the granite belt at the base of the mountain
resembles that of the western Karoo rather than that of
Namaqualand. Ixia rapunculoides , Moraea bifida and M.
tripetala (Iridaceae) are three of the more common geo-
phytes on the plateau, and occur also on the Bokkeveld
Plateau and on doleritic clays east of Loeriesfontein,
both sites ± 70 km to the southeast. Another disjunct is
Boophone haemanthoides (Amaryllidaceae), which also
occurs on dolerites of the Bokkeveld Plateau as well as
on the western coastal plain of Western Cape Province.
The fairly common Namaqualand species, Babiana
JJabellifolia , occurs at lower elevations, together with
Moraea serpentina and M. schlechteri , both of which
Bothalia 38,1 (2008)
51
FIGURE 2. — Known distribution of Babiana symmetrantha, •; and B.
vires cens, O.
are common in Namaqualand but rare or absent from
the western Karoo to the east. The Langberg is linked
to the western Karoo highlands by a second isolated
range, the Kubiskou Mountains, a short distance west of
Loeriesfontein. The vegetation of this range has not been
well explored but its lower and middle slopes have char-
acteristic western Karoo species such as Babiana crispa,
B. spathacea and Ixia marginifolia, whereas I. sobolifera
and Geissorhiza heterostyla grow on the dolerite-topped
summit. Of these species, only the two last named occur
on the Langberg.
Additional specimen examined
NORTHERN CAPE. — 3019 (Loeriesfontein): summit plateau of
the Langberg among dolerite boulders, (-DB), 16 September 2006,
Goldblatt & Porter 12839 (MO, NBG).
Babiana virescens Goldblatt & J.C. Manning, sp.
nov.
Plantae 70-200 mm altae, caule usitate eramoso, foliis
lanceolatis minute velutinis leviter plicatis marginibus
incrassatis anguste hyalinis, spica 3- ad 7-flora arcuata,
bracteis viridibus 20-30 mm longis subaequalibus, brac-
tea exteriore velutina bractea interiore ad apicem furcata,
floribus zygomorphis pallide viridi-griseis tepalis inferi-
oribus flavis purpureo-notatis odoratis, tubo perianthii
18-23 mm longo oblique infundibuliformi, tepalis inae-
qualibus tepalo dorsali 32-38 x 6. 5-8.0 mm, inferioribus
25-30 x ± 4—5 mm, filamentis suberectis 12-16 mm lon-
gis, antheris 7-8 mm longis, ovario glabro, ramis styli
4—6 mm longis.
TYPE. — Western Cape, 3018 (Kamiesberg): Farm Kama-
gap, 1 km along Bitterfontein-Pofadder road, hillside on S
side, (-CD), 26 May 2007, Manning 3093 (NBG, holo.;
MO, iso.).
Plants 70-200 mm high. Conn subglobose, 15-20
mm diam., outer tunics of fine-textured, reticulate
fibres, persisting as loose neck. Stem reaching to ground
level or up to 100 mm above ground, erect but strongly
indexed above uppermost leaf sheath, unbranched or
with one short branch, glabrous below, becoming vel-
vety near spike; cataphylls and leaf sheaths glabrous.
Leaves 3 or 4, oblong to lanceolate, slightly twisted dis-
tally, blade suberect at ± 30°, 40—1 00( — 1 50) x (5—) 1 0— 1 8
mm, thinly velvety, lightly pleated, midrib and margins
slightly thickened, margins hyaline, minutely hairy;
seedling leaves linear-lanceolate and coiled, hairy. Spike
3- 7-flowered, arching horizontally, secund; bracts green
with brown tips, outer ovate, 20-30 mm long, margins
narrowly hyaline, thinly and minutely velvety, especially
distally, apical margins markedly ciliate, truncate or
emarginate-apiculate, inner bracts about as long as outer,
forked for 5-8 mm, with broad hyaline margins in lower
half. Flowers zygomorphic, pale grey-green or flushed
with greyish lilac, lower lateral tepals with yellowish
green median blotches and purple streaks, with sweet,
spicy scent; perianth tube obliquely funnel-shaped, nar-
row part hollow to base, containing nectar, 18-23 mm
long; tepals unequal, narrowly lanceolate, margins crisp-
ulate, dorsal suberect, recurving distally, 32-38 x 6.5-
8.0 mm, upper laterals narrower, joined to lower tepals
for 5-7 mm forming lip, lower tepals 25-30 x 4—5 mm.
Stamens unilateral; filaments suberect, 12-16 mm long;
anthers 7-8 mm long, whitish. Ovary glabrous; style
dividing opposite middle third of anthers, style branches
4- 6 mm long. Capsules subglobose, 8-9 mm long. Seeds
unknown. Flowering time: late May to early June, fruit-
ing in August to September (Figure 3).
Distribution and ecology, a local endemic of south-
ern Namaqualand, between Nuwerus and Bitterfontein,
where it occurs on granite hills on gentle slopes and val-
ley bottoms in gravelly ground in open, succulent kar-
roid shrubland, sometimes between granite outcrops but
never wedged in rock crevices (Figure 2).
Discussion: Babiana virescens accords with section
Teretifolieae in its apically forked inner floral bracts but
is unusual in its well-developed aerial stem, since most
members of the section have the stem reaching only
shortly above the ground or completely subterranean.
The bilabiate flowers stand out in having particularly
narrow, ± linear tepals, the dorsal 32-38 x 6. 5-8.0 mm
and the remaining tepals 4-5 mm wide. Although quite
large, the greenish flowers are not conspicuous but have
a characteristic spicy-acrid, chemical odour that is com-
mon in the genus. The flowers close at night and cease
to produce fragrance, lasting four or five days when not
pollinated.
This new species may be most closely allied to
Babiana torta G.J.Lewis and B. namaquensis Baker on
account of the slightly coiled leaves and early flowering
but it is easily distinguished from them by its greenish
flowers with particularly narrow tepals and by the sub-
glabrous or very short-hairy leaves and the aerial stem
up to 100 mm high. The slightly thickened leaf margins
are most evident in herbarium specimens, drying whit-
ish or pale. Both blue-flowered B. torta and pale mauve-
to white-flowered B. namaquensis have distinctly hairy
leaves without conspicuous margins and typically occur
in rock outcrops, granite in the former but often lime-
stone in the latter, with their corms wedged in crevices.
Babiana torta typically occurs in granite rock outcrops,
whereas B. virescens always occurs in open ground and
52
Bothalia 38,1 (2008)
FIGURE 3. — Babiana virescens.
Manning 3093 (NBG): A,
whole plant. B, C, flower: B,
front view, C, side view. D,
outer (left) and inner (right)
floral bracts. Scale bar: 10
mm. Artist: John Manning.
relatively deep, loamy gravel. Both species have been
collected within a few kilometres of each other east of
Bitterfontein, confirming their status as distinct species.
Babiana virescens shares grey-green flowers having
unusually narrow tepals with B. gariepensis Goldblatt
& J.C. Manning, a poorly known species from northern
Namaqualand. This species is acaulescent, has leaves
without evidently thickened margins, and flowers with
a perianth tube 20-24 mm long, slender below, curv-
ing outward and wider in upper 7 mm, and a dorsal
tepal 28-32 x ± 5 mm. Additional material in flower
is needed to better understand this apparently rare
Richtersveld endemic that is currently known from three
sites, Koeskop in the Richtersveld National Park in the
north, Cornellsberg, and from Grasvlakte in the southern
Richtersveld. The larger dorsal tepal, 32-38 x 7-8 mm,
and aerial stem readily distinguish B. virescens from B.
gariepensis.
Babiana virescens appears to have first been collected
in May 1 993 by Cape Town botanist Dee Snijman. The
collection was referred to B. torta , at the time poorly
documented (Lewis 1959) but now known to be a rela-
tively common, early flowering species of southern and
central Namaqualand (Goldblatt & Manning 2007),
which typically has pale blue flowers with broad tepals.
Our first collection of B. virescens ( Goldblatt & Porter
12145) was also referred to B. torta in our monograph
of the genus, leading us to describe the leaves of that
species as variously hairy to almost hairless. With these
specimens removed from B. torta , the leaves of that spe-
cies are now known to be uniformly softly hairy.
Bothalia 38,1 (2008)
53
Additional specimens examined
WESTERN CAPE. — 3018 (Kamiesberg): Kliprand road close
to turnoff from Bitterfontein. (-CD), gentle slope, 19 May 1993,
Snijman 1304 (NBG). 3118 (Vanrhynsdorp): 3 km south ofNuwerus,
road to Vredendal, granitic soil in valley bottom, (-AB), 2 September
2002 (fruiting), Goldblatt & Porter 12145 (MO, NBG), [flowered
Portland. USA. December 2005. Goldblatt & Porter 12145A (MO), 16
September 2006 (fruiting), Goldblatt & Porter 12853 (MO)].
The two new species may be accommodated in the
key to section Teretifolia of Babiana in Goldblatt &
Manning (2007: 12, 13) as follows:
2a Stem aerial; flowers cream-coloured to pale yellow with
brown to dull purple centre; leaves lanceolate to ovate;
perianth tube 15-25 mm long; ovary hairy B. pygmaea
2b Stem subterranean; flowers blue or violet; leaves linear; peri-
anth tube > 25 mm long; ovary hairless:
2a’ Flowers with red centre and throat; perianth tube 30-75
mm long; tepals 10-12 mm wide; anthers violet B. radiata
2b’ Flowers without red centre; perianth tube 45-75 mm long;
tepals 5-7 mm wide; anthers white B. symmetrantha
32a Flowers pale greyish green; leaf margins thickened . . . B. virescens
32b Flowers pale blue, mauve or whitish; leaf margins usually
unthickened:
RANGE EXTENSION
Babiana ambigua (Roem. & Schult.) G.J. Lewis
The range of Babiana ambigua is largely coastal,
extending from near Lambert’s Bay in the north to
Riversdale in the southeast (Goldblatt & Manning
2007). Populations also occur inland on the Gifberg,
the Cold Bokkeveld and the Olifants River Mountains.
Wherever it has been collected, the species occurs on
coarse sand or sandy granitic gravel. A new collec-
tion from sandy ground at the top of Botterkloof Pass
extends the range of the species to some 150 km north of
the Cold Bokkeveld and 50 km inland from the Gifberg.
The record is surprising as B. ambigua does not, accord-
ing to available records, occur in the Cedarberg, which
lies between Botterkloof and the next nearest popula-
tions of the species to the south. This is the first record
of B. ambigua from Northern Cape. The vegetation
in the localized area of coarse sandy soil at the top of
Botterkloof Pass is dominated by clumps of Willdenowia
(Restionaceae) and includes several annual species of
Scrophulariaceae. The otherwise largely coastal and low-
land sandveld species, Gladiolus speciosus and B. rin-
gens have also been found at this site, the latter a sight
record supported by a photograph (Manning & Goldblatt
1997:91).
Additional specimen
NORTHERN CAPE. — 3119 (Calvinia): top of Botterkloof Pass,
sandy ground, (-AD), 31 August 2006 (fruiting), Goldblatt & Porter
12731 (MO, NBG).
NEW NAMES FOR BABIANA LONGIFLORA AND
B. THUNBERGII -
After examining the draft checklist of Iridaceae at the
Royal Botanic Gardens, Kew website, World Checklist
of selected plants families (Govaerts pers. comm.), we
have learned that the southwestern Cape species Babiana
longiflora Goldblatt & J.C. Manning (2004) is a homo-
nym for B. longiflora (L.f.) Steud. We propose the new
name B. tubaeformis for this rare plant. Also, there is a
valid and legitimate earlier synonym, Antholyza hirsuta
Lam. (1783), for B. thunbergii Ker Gawl. (1804), a com-
mon species of coastal sands of western South Africa.
We provide a new combination B. hirsuta and present
a more complete synonymy for this species than that
given in our revision of the genus (Goldblatt & Manning
2007).
Babiana tubaeformis Goldblatt & J.C. Manning,
nom. nov., pro B. longiflora Goldblatt & J.C. Manning
in Bothalia 34: 94 (2004), nom. illeg. non B. longiflora
(L.f.) Steud.: 686 (1840) (= Tritonia cooperi subsp. coo-
peri).
Babiana hirsuta (Lam.) Goldblatt & J.C. Manning,
comb. nov.
Antholyza hirsuta Lam., Encyclopedie methodique, Botanique,
vol. 1: 201 (1783). Gladiolus sulcatus Lam.: 119 (1791), nom. illeg.
superfl. pro A. hirsuta Lam. Gladiolus mollis Vahl: 119 (1805), as a
new name for Antholyza hirsuta Lam. Type: without locality or collec-
tor, ‘antholyza hirsuta. encycl.’ (Herb. Lamarck, P, holo.!).
Babiana thunbergii Ker Gawl.: 233 (1804), as a new name for
Antholyza plicata L.f.: 96 (1782). Anaclanthe plicata (L.f.) N.E.Br.:
269 (1932).
Anaclanthe namaquensis N.E.Br.: 269 (1932). Antholyza nam-
aquensis (N.E.Br.) Goldblatt: 444 (1971).
NOMENCLATURE: WHO IS THE AUTHOR OF BABIANA1
The genus Babiana was formally described in August
1802 in Curtis’s Botanical Magazine by John Gawler,
now known to the botanical community as John Ker
Gawler (Gawler subsequently changed his name to
Bellenden Ker). The article that accompanied a painting
of Babiana plicata Ker Gawl. (now B. fragrans (Jacq.)
Steud.) included formal Latin and English descriptions
of Babiana and the single species, Babiana plicata
(itself a replacement name for Gladiolus plicatus Thunb.
non L.). Babiana plicata is thus the type species of the
genus. Ker Gawler (1802a) believed he was the author
of this southern African genus, which now includes
some 90 species (Goldblatt & Manning 2007), and so
did contemporary and later authorities, notably Steudel
(1840), Klatt (1882) and Baker (1877, 1892, 1896). Both
Klatt and Baker were specialists in the systematics of
Iridaceae. Lewis (1959), who revised Babiana almost
50 years ago, also treated the genus as having been
described by Ker Gawler in August 1 802.
Dandy (1969) proposed the conservation of Babiana
against Beverna Adans. (1763) and the Committee for
Spermatophyta (McVaugh 1971) recommended its con-
servation. Babiana thus appeared in the list of conserved
generic names in the subsequent International Code of
Botanical Nomenclature (Stafleu et al. 1972: 290) as
Babiana Ker Gawl. (1802a). Then, in the list of con-
served generic names in the Sydney Code (Voss et al.
1983: 330), the genus suddenly appears as Babiana Ker
Gawl. ex Sims (1801).
The reason appears to be as follows. In an article in
Curtiss Botanical Magazine, plate 539 (November
1801), Ker Gawler provided a formal account of the pre-
54
Bothalia 38,1 (2008)
viously and validly published Ixia conica Salisb. This is
followed by a short narrative about the Cape (i.e. west-
ern South African) species of Iridaceae written by Sims,
the editor of the magazine and author of the article. After
first discussing the history of I. conica and its possible
synonyms, Sims proceeded to comment on the current
state of the taxonomy of Jussieu’s natural order [family]
Irides (sic). Sims then related the difficulty of circum-
scribing genera in the family because of the conflicting
patterns of variation in perianth tube length and shape
and the symmetry of the flower (our paraphrasing). Sims
continued: ‘so that, notwithstanding the pains he has
taken, our friend Mr. Gawler, has not been as yet able to
reduce the genus [Ixia] to its proper standard, and is best
able for the present only to divide it into several sec-
tions, distinguished from each other by characteristical
marks, with notices of such as appear to him most likely
to become the foundation of future separate genera. With
these observations we hope soon to have an opportunity
of treating our botanical readers’. [We note here that
Ker Gawler subsequently described Tritonia (1802b),
Anomatheca (1804) (now a synonym of the conserved
Freesia Klatt), Geissorhiza (1804), and Hesperantha
(1804), the last three genera in Konig & Sims’s Annals
of Botany, all based on species until then included in
Gladiolus and Ixia.]
Continuing, Sims (1801) wrote: ‘In the mean time,
there is one division, the species composing which
are at the present dispersed in the three genera of Ixia ,
Gladiolus, and Antholyzai, so very distinct at first sight
from the rest, by their smooth sheathing petioles, ter-
minated in general by a plicate and villose leaf, with
their bulbs situated unusually deep in the earth, that Mr.
Gawler has with the greatest propriety united these into a
separate genus, with the name of Babiana ’. Sims goes on
to list the species that the genus ‘will contain’ [when it
has been described and combinations made is evidently
intended here] in the above three genera that belong in
Babiana. Sims here clearly ascribed the genus name
Babiana to Ker Gawler and provided a diagnosis that is
clearly credited to the investigations of Ker Gawler (‘so
very distinct .... that Mr. Gawler has . . .’ ) and may be
either a direct quote or a paraphrase. Thus, if this article
is accepted as the valid place of publication, Babiana
must be attributed to Ker Gawler alone under article 46.2
of the International Code of Botanical Nomenclature
(McNeill et al. 2006). Foster (1979: 167) evidently
accepted this interpretation by the listing of Babiana Ker
Gawler (1801) in Index Nominum Genericorum.
In August 1802, just nine months after Sims’s com-
ments about the taxonomy of Ixia, Ker Gawler (1802a)
formally described Babiana in Curtis's Botanical
Magazine. There is no mention of the November 1801
article in which Sims informed the readers of the same
magazine about the imminent publication of a new genus
Babiana. Clearly neither Ker Gawler nor Sims consid-
ered the name to have been published, as implied by the
phrase ‘This genus will contain . . We thus argue that
Sims (1801) merely mentioned Babiana in anticipation
of its future acceptance and therefore did not validly
publish it under Article 34.1(b) of the Code (McNeill et
al. 2006).
Because “The Code does not provide for conserva-
tion of a name against ... an ‘isonym’, the same name
with the same type but with a different place and date
of valid publication and perhaps with a different author-
ship . . . than is given in the relevant entry in App. II,
III, or IV” (Art. 14.4 Note 1), we can and should change
the citation of Babiana Ker Gawl. ex Sims (1801) in
the Sydney Code and subsequent Codes to Babiana Ker
Gawl. (1802).
ACKNOWLEDGEMENTS
Support for this study by grant 7799-05 from the
National Geographic Society is gratefully acknowl-
edged. Collecting permits were provided by the Nature
Conservation authorities of Northern and Western Cape,
South Africa. We thank Mrs A. Visser for permission to
collect on her land and Lendon Porter for his assistance
and companionship in the field. We acknowledge Bertil
Nordenstam, Dan Nicolson, John McNeill, and Kanchi
Gandhi for comments on the nomenclature of Babiana,
the two last-named, not, however, in agreement with our
conclusion.
REFERENCES
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BAKER, J.G. 1 877. Systema iridacearum. Journal of the Linnean Society
of London, Botany 16: 61-180.
BAKER, J.G. 1 892. Handbook of the Irideae. Bell, London.
BAKER, J.G. 1 896. Irideae. In W.T. Thiselton-Dyer, Flora capensis 6:
7-1 7 1 . Reeve, London.
BROWN, N.E. 1932. Contributions to a knowledge of the Transvaal
Iridaceae. Part II. Transactions of the Royal Society of South
Africa 20: 261-280.
DANDY, J.E. 1969. Nomina conservanda proposita. Proposal 265.
Taxon 18: 464, 465.
FOSTER, R.C. 1979. Iridaceae. In E.R. Farr, J.A. Leussink & F.A.
Stafleu, Index Nominum Genericorum (plantarum), vol. 1. Regnum
Vegetabile 100: vii-xxvi, 1-630.
GOLDBLATT, P. 1971. Cytological and morphological studies in the
southern African Iridaceae. Journal of South African Botany 37:
317-460.
GOLDBLATT, P. & MANNING, J.C. 2000. The long-proboscid fly polli-
nation system in southern Africa. Annals of the Missouri Botanical
Garden 87: 146-170.
GOLDBLATT, P. & MANNING, J.C. 2004. Taxonomic notes and
new species of the southern African genus Babiana (Iridaceae:
Crocoideae). Bothalia 34: 87-96
GOLDBLATT, P. & MANNING, J.C. 2007. A revision of the southern
African genus Babiana, Iridaceae: Crocoideae. Strelitzia 18. South
African National Biodiversity Institute, Pretoria and Missouri
Botanical Garden, Missouri.
GOLDBLATT, P„ MANNING, J.C. & BERNHARDT, P. 1995.
Pollination biology of Lapeirousia subgenus Lapeirousia
(Iridaceae) in southern Africa: floral divergence and adaptation
for long-tongued fly pollination. Annals of the Missouri Botanical
Garden 82: 517-534.
KER GAWLER, J. 1802a. Babiana plicata. Sweet-scented babiana.
Curtiss Botanical Magazine 16: t. 576.
KER GAWLER, J. 1 802b. Tritonia squalida. Sweet-scented tritonia.
Curtiss Botanical Magazine 16: t. 581.
KER GAWLER, J. 1804. Ensatorum ordo. In J. Konig & J. Sims .Annals
of Botany 1:21 9-247.
KLATT, F. W. 1 882. Erganzungen und Berichtigungen zu Baker’s Systema
Iridacearum. Abhandhingen der Naturforschenden Gesellschaft
zu Halle 15: 44 404.
LAMARCK, J.B.A.P. de. 1783. Encyclopedie methodique, Botanique,
vol. 1. Panckoucke, Paris.
LAMARCK, J.B. A.P. de. 1791. Tableau encyclopedique et methodique,
vol. 1 . Panckoucke, Paris.
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LEWIS, G.J. 1959. The genus Babiana. Journal of South African Bota-
ny, Suppl. 3.
LINNAEUS, C. (fil.). 1782 [as 1781], Supplementum plantarum. Orpha-
notropheus, Brunswick.
MANNING, J.C. & GOLDBLATT, P. 1996. The Prosoeca peringueyi
(Diptera: Nemestrinidae) pollination guild in southern Africa:
long-tongued flies and their tubular flowers. Annals of the Missou-
ri Botanical Garden 83: 67-86.
MANNING, J.C. & GOLDBLATT, P. 1997. Nieuwoudtville, Bokkeveld
Plateau and Hantam. South African Wild Flower Guide 9. Bota-
nical Society of South Africa, Cape Town.
MCNEILL, J„ BARRIE, F.R., BURDET, H.M., DEMOULIN, V., HAWKS-
WORTH, D.L., MARHOLD, K., N1COLSON, D.H., PRADO, I,
SILVA, P.C., SKOG, J.E., WIERSEMA, J.H. & TURLAND, N.J.
2006. International Code of Botanical Nomenclature (Vienna Code)
adopted by the 17th International Botanical Congress, Vienna,
Austria, July 2005. Regnum Vegetabile 146: i-xviii, 1-568. Koeltz
Scientific Books, Konigstein.
MC VAUGH, R. (sec. ). 1 97 1 . Report of the Committee for Spermatophy-
ta. Conservation of generic names, XIV. Taxon 20: 384-389.
SIMS, J. 1801. Ixia conica. Orange-coloured ixia. Curtiss Botanical
Magazine 15: t. 539.
STAFLEU, F.A., BONNER, C.E.B., MCVAUGH, R., MEIKLE, R.D.,
ROLLINS, R.C., ROSS, R., SCHOPF, J.M., SCHULZE, G.M.,
DE VILMORIN, R. & VOSS, E.G. 1972. International Code of
Botanical Nomenclature. Regnum Vegetabile 82: 1-426.
STEUDEL, E.G. 1840. Nomenclator botanicus, vol. 1. Cottae, Stuttgart.
VAHL, M. 1805. Enumeratio plantarum, vol. 2. Moller, Copenhagen.
VOSS, E.G., BURDET, H.M., CHALONER, W.G., DEMOULIN, V,
HIEPKO, P., MCNEILL, J., MEIKLE, R.D., NICOLSON, D.H.,
ROLLINS, R.C., SILVA, P.C. & GREUTER, W. 1983. International
Code of Botanical Nomenclature. Regnum Vegetabile 111: i-xv,
\-A12.
_
.
'
Bothalia 38,1: 57-63 (2008)
The genus Wellstedia (Boraginaceae: Wellstedioideae) in southern
Africa
E. RETIEF^ and A.E. VAN WYK* **
Keywords: Boraginaceae, palynology, phytogeography, southern Africa, taxonomy, Wellstedia Balf.f., Wellstediaceae, Wellstedioideae
ABSTRACT
This regional taxonomic revision of the genus Wellstedia Balf.f., a member of the family Boraginaceae s.l. (including
Hydrophyllaceae s.str.), is part of a series of publications on the Boraginaceae in southern Africa. Wellstedia comprises six
species, five in Socotra, Somalia and Ethiopia with the remaining one, W. dinteri Pilg., occurring in southern Africa. W. dinteri
Pilg. subsp. dinteri occurs in Namibia and the Northern Cape, whereas the newly instated subspecies W. dinteri subsp. gracilior
(D.R.Hunt) Retief & A.E.van Wyk, based on W. dinteri Pilg. var. gracilior D.R.Hunt, is confined to Namibia only. The disjunct
distribution of Wellstedia and numerous other plant and animal taxa between the arid regions of northeastern Africa and southern
Africa is usually explained by the postulated periodic existence of an arid corridor between the two regions during the arid
phases of the Pleistocene and even earlier. Wellstedia is treated here in Wellstedioideae, a subfamily of Boraginaceae s.l. but is
sometimes placed in a family of its own, Wellstediaceae Pilger. Morphologically Wellstedia displays strong similarity to genera
of the Ehretioideae and also to certain members of the Hydrophyllaceae. The genus is characterized by a perennial, dwarf shrub
habit, densely hairy leaves, 4-merous flowers, a terminal, bifid style and a 1- or 2-seeded capsule. A key to the two subspecies,
diagnostic characters, a distribution map and illustrations of various macro- and micromorphological features are provided.
INTRODUCTION
Wellstedia was described in 1884 by Balfour after
a visit to the island of Socotra. He decided on the
name Wellstedia in honour of the British Lieutenant
J.R. Wellsted, who explored Socotra for the Indian
Government in 1834 (Thulin & Johansson 1996).
Balfour believed that W. socotrana Balf.f., the only
species in the genus at the time, had its closest affinity
with members of Boraginaceae s.str. Besides this spe-
cies, five more are now recognized on the African conti-
nent— four in Somalia and Ethiopia, and one in Namibia
and South Africa. Such floristic (and faunistic) disjunc-
tions between the arid regions of northeastern Africa and
southern Africa are usually ascribed to the periodic exis-
tence of a linking arid corridor between the two regions
in the past (Van Wyk & Smith 2001). In 1912 Pilger
placed Wellstedia in its own subfamily, Wellstedioideae
(Boraginaceae). Novak (1943), however, decided on a
separate family, Wellstediaceae. Since then, the family
classification of Wellstedia has been a matter of contro-
versy. Merxmuller (1960), unaware of Novak, also estab-
lished a new family, Wellstediaceae, thereby creating a
later homonym. In 1 967 the family was also recognized
by Friedrich-Holzhammer in a Prodromus on the flora of
South West Africa [Namibia], an approach more recently
followed by Lebrun & Stork (1997).
Different modem views regarding the delimitation of
Boraginaceae exist: splitting Boraginaceae s.l. into two
separate families, Boraginaceae s. str. and Heliotropiaceae
with Hydrophyllaceae not included (Diane et al. 2002);
or recognizing several segregate families, Boraginaceae
* National Herbarium, South African National Biodiversity Institute,
Private Bag X101, 0001 Pretoria.
' Affiliation: Department of Botany, University of Pretoria, 0002
Pretoria.
** H.G.W.J. Schweickerdt Herbarium, Department of Botany,
University of Pretoria, 0002 Pretoria.
MS. received: 2004-06-02.
5. str., Cordiaceae, Ehretiaceae, Heliotropiaceae, Hydro-
phyllaceae, Lennoaceae and Wellstediaceae (Lebrun &
Stork 1997; Gottschling et al. 2001; Gottschling 2003).
In the present contribution, Wellstedia is regarded as a
monotypic genus of the subfamily Wellstedioideae with-
in Boraginaceae s.l. (including Hydrophyllaceae s.str.,
Codonoideae, Ehretioideae, Heliotropioideae and Boragi-
noideae) (Retief 2003). Ferguson (1999) and the Angio-
sperm Phylogeny Group II (2003) are followed here in
regarding the tribes Phacelieae and Hydrophylleae as
part of Boraginaceae s.l. The capsular fruit of Wellstedia
is unknown elsewhere in Boraginaceae s.str. and has
been used to motivate the recognition of a monotypic
family Wellstediaceae. However, with members of
Hydrophyllaceae, all with capsular fruits, included in a
broadly defined Boraginaceae, this argument is no longer
of importance. Pollen and other characters such as a ter-
minal style, 4-merous flowers, trichomes with multicellu-
lar bases and cymose inflorescences similar to other tra-
ditional members of Boraginaceae, support the placement
of Wellstedia in a subfamily within Boraginaceae s.l.
The aim of this paper is to present a taxonomic revi-
sion of the genus Wellstedia in southern Africa, includ-
ing Namibia, Botswana, South Africa, Lesotho and
Swaziland. Diagnostic characters, an identification key,
illustrations and a distribution map are provided. This
paper forms part of a revision of the Boraginaceae in
southern Africa. The genus description is based on mate-
rial from southern Africa only.
MATERIALS AND METHODS
Herbarium specimens in BM, BOL, E, GRA, K,
NBG, NH, NU, PRE, PRU, SAM and WIND (acronyms
as in Holmgren et al. 1981) were studied to gather data
on morphological characters, phenology and geographi-
cal distribution. Pollen and various plant parts were stud-
ied with an ISI-SX-25 scanning electron microscope.
Measurements of pollen grains were done from aceto-
58
Bothalia 38,1 (2008)
lysed grains mounted in glycerine jelly. Acetolysis fol-
lowed the standard method of Erdtman (1960). Tapetal
orbicules were obtained from herbarium specimens and
prepared for viewing with a JEOL 6000 F in-lens field
emission scanning electron microscope (for procedure
followed see Retief et al. 2001, 2002).
HISTORICAL OUTLINE
In 1912, Pilger described W. dinteri from specimens
collected by botanist and botanical explorer Moritz
Kurt Dinter (1868-1945) in Namibia. Although Pilger
knew the genus only from Balfour’s (1884) description
and illustration of W. socotrana, he was convinced that
Dinter had found a Wellstedia in the southwestern part
of Africa. He was also of the opinion that Wellstedia
belongs to Boraginaceae with affinity to Coldenia L.
of the Ehretioideae, and not Verbenaceae, a family also
metioned by Balfour ( 1 884) when he described the first
species. The capsular fruit with two seeds, however,
distinguishes Wellstedia from all other members of
Boraginaceae and a new subfamily Wellstedioideae was
described by Pilger (1912).
Hunt (1969) divided W dinteri into two varieties.
Variety dinteri is characterized by flowers and cap-
sules congested on short lateral branches, whereas var.
gracilior has flowers and capsules on lax branchlets,
appearing as if solitary, axillary, rarely close together.
When Friedrich-Holzhammer (1967) revised the genus
for Prodromus einer Flora von Siidwestafrika , she did
not recognize the two infraspecific taxa proposed by
Hunt (1969) and regarded W. dinteri as belonging to
Wellstediaceae. Thulin & Johansson (1996), in a revi-
sion of the genus, also did not maintain the two varieties
in W. dinteri. They classify the genus in Wellstedioideae
and proposed the New World genus Tiquilia Pers.
(Boraginaceae: Ehretioideae) as a possible sister to it.
In the present revision, the status of W. dinteri Pilg. var.
gracilior D.R.Hunt is raised to that of subspecies, a deci-
sion based mainly on differences in macromorphology
and habitat.
PHYTOGEOGRAPHY
The two subspecies of Wellstedia dinteri in southern
Africa (Figure 1) are associated with the Nama-Karoo,
Succulent Karoo and Savanna Biomes, according to the
biome map in Van Wyk & Smith (2001: 8). W. dinteri
subsp. dinteri is recorded as growing in shale-derived
soil mixed with fine gravel (Davidse & Loxton 6339). W.
dinteri subsp. gracilior, however, occurs on dolomite or
limestone ( Acocks 15641) or on black soil derived from
dolomite.
Arid regions of the African continent are phytogeo-
graphically linked by a large number of taxa with dis-
junct distributions. This pattern is particularly well
developed between arid regions in southern Africa
and northeastern Africa (Horn of Africa and adjacent
Arabian Peninsula). Wellstedia is a typical example of
a taxon with a disjunct distribution between the arid
areas of northeastern Africa and southern Africa: W. din-
FIGURE 1. — Known distribution of Wellstedia dinteri subsp. dinteri,
• ; and W. dinteri subsp. gracilior , O.
teri occurring in Namibia and Northern Cape (Figure
1), whereas W. filtuensis Hunt & Lebrun is endemic to
Ethiopia, with W. sotnalensis Thulin & A.Johanss, W.
robusta Thulin and W. laciniata Thulin & A.Johanss.
confined to Somalia. The disjunct pattern is usually
explained by the postulation of an arid corridor (or cor-
ridors) linking these regions during arid phases of the
Pleistocene and even earlier (Verdcourt 1969; Goldblatt
1978; Van Wyk & Smith 2001). Various other authors,
for example, De Winter (1971), Thulin ( 1 994) and Thulin
& Johansson (1996) also commented on the recurring
pattern of disjunct distributions. Gaillonia A. Rich, ex
DC. (Rubiaceae), as circumscribed by Thulin (1998), is
another example of a genus with a disjunct distribution
comparable to that of Wellstedia (Thulin 1994).
Disjunct distributions in Africa and the New World are
rather unusual, but for southern Africa, as many as seven
families and many more genera are involved (Goldblatt
1978). Codon L., an exclusively southern African genus,
has its nearest relatives, members of Hydrophyllaceae
s.str., in North America. However, Wellstedia also shows
similarities in pollen and habit characters with some of
the genera of Hydrophyllaceae s.str. A possible explana-
tion for this disjunct distribution pattern is the existence
of a Tertiary North Atlantic land bridge (Tiffney 1985),
whereas over-water dispersal seems unlikely, except dur-
ing the early stages of continental separation.
CHARACTERS OF TAXONOMIC SIGNIFICANCE
Habit
Members of the Boraginaceae are mainly herbaceous,
but shrubs and trees do occur. Wellstedia is characterized
by a woody, dwarf shrub habit, a growth form also dis-
played by Tiquilia. of the subfamily Ehretioideae — sug-
gested as a potential sister group of Wellstedia (Thulin &
Johansson 1996). The woody habit and other morpholog-
ical similarities with the Ehretioideae support the classi-
fication of Wellstedia as a member of the Boraginaceae
s.l. instead of placing it in a family of its own.
Bothalia 38,1 (2008)
59
Leaves
The narrowly obovate to ovate leaves of the southern
African species with its two subspecies are densely pubes-
cent (Figure 2A, B). The trichome complement consists of
rigid setae with multicellular bases, usually 3-layered, scat-
tered on the blade (Figure 2C) and fine setae with undevel-
oped bases. Leaves of Wellstedia dinteri subsp. gracilior
are smaller in size and different in colour, greyish white
compared to yellowish green in W. dinteri subsp. dinteri.
FIGURE 2. — Morphology of Wellstedia. A-C, leaf: A, W. dinteri subsp. dinteri , Acocks 15615 , upper leaf surface; B, W. dinteri subsp. gracilior ,
Muller 1329 , upper leaf surface; C, W. dinteri subsp. dinteri , Liebenberg 5179, multicellular base of a seta. D-F, flower: D, W. dinteri subsp.
gracilior, Muller 1329, calyx and corolla; E, W. dinteri subsp. dinteri, Leistner 2589, anthers; F, W. dinteri subsp. dinteri, Oliver & Muller
6397, stigma and style. G-K, fruit: W. dinteri subsp. dinteri, Oliver & Muller 6397: G, capsule splitting open; I, seed; J, indumentum of
capsule. W. dinteri subsp. gracilior, Muller 1329: H, capsule; K, indumentum of capsule. L, W. dinteri subsp. dinteri, Oliver & Muller 6397,
part of capsule and calyx lobe. Scale bars: A, 75 pm; B, 259 pm; C, 89 pm; D, 365 pm; E, 24 pm; F, 16 pm; G, 460 pm, H, 431 pm; I, 314
pm; J, 259 pm; K, 254 pm; L, 509 pm.
60
Bothalia 38,1 (2008)
FIGURE 3. — Pollen morphology of Wellstedia dinteri subsp. dinteri. A-E, Leistner 2589: A, pollen grain in equatorial view, tapetum clothed with
orbicules; B, grain showing mesocolpial concavity; C, grains in different views; D, compound aperture; E, mesocolpial concavity, tectum
reticulate. F, Oliver & Muller 6397, grain ± in polar view. Scale bars: A, 4 pm; B, 2.9 pm; C, 23 pm; D, 2.1 pm; E, 1.2 pm; F, 2.5 pm.
Flowers
Flowers of Wellstedia are 4-merous (Figure 2D), a
state which is rare in Boraginaceae s.l. It is often used
to support the recognition of a separate family, Well-
stediaceae. However, the flowers of Coldenia , a mono-
typic genus and also a member of the Ehretioideae,
is similarly 4-merous. This is another link between
Wellstedia and a member of Boraginaceae sd. The calyx
is deeply lobed and accrescent in fruit (Figure 2L). The
outer surface of the corolla is densely pubescent (Figure
2D), but the inner surface is glabrous (Figure 2E). A ter-
minal style is present (Figure 2F), persistent (Figure 2L)
and slightly bifid with capitate stigmas, the latter covered
with exudate when receptive (Figure 2F).
Fruit and seed
All members of Wellstedia are characterized by hairy,
mussel-like capsules (Figure 2G, H, J, K), but differ in
dissepiment morphology (Thulin & Johansson 1996).
The seeds (Figure 21) are truncate, and pitted above with
a circle of long, rigid trichomes in the upper part.
Pollen
Pollen grains (Figure 3A-F) of Wellstedia are tricol-
porate, isopolar and with mesocolpial concavities (some-
times regarded as ‘pseudocolpi’) and a reticulate tectum,
showing strong similarity with genera of the subfam-
ily Ehretioideae (Retief & Van Wyk 2001). Wellstedia
and members of Ehretioideae furthermore show simi-
larity in their pollen morphology with members of
the tribes Hydrophylleae and Phacelieae of the family
Hydrophyllaceae, here regarded as part of Boraginaceae
sd. Doughnut-shaped tapetal orbicules (Retief et al.
2001: fig. 2), occur in both Wellstedia and Codon, con-
firming the close relationship of these genera.
TAXONOMIC TREATMENT
Wellstedia Balf.f in Proceedings of the Royal
Society of Edinburgh' 12: 402 (1884); Gurke: 131 (1897);
Pilg.: 558 (1912); E.Phillips: 633 (1951); Friedr.-Holzh.:
121 (1967) as Wellstediaceae; D.R.Hunt: t. 3665, 3666
(1969); R. A. Dyer: 514 (1975); Thulin & Johansson: 80
(1996); Retief: 183 (2000). Type: W. socotrana Balf.f.
Dwarf shrubs, densely pubescent with trichomes
appressed; trichome complement consists of rigid setae
with prominent multicellular bases or fine setae with
undeveloped bases. Branches decumbent or ascending.
Leaves spirally arranged; blade narrowly obovate to
spathulate, decurrent, often upper part (‘blade’) forms an
Bothalia 38,1 (2008)
61
FIGURE 4. — Wellstedia dinteri
subsp. dinteri (= W. dinteri
var. dinteri), Acocks 18101.
A, habit, x 1;B, part of lower
leaf surface, x 6; C, branch-
lets with flowers and fruits, x
3; D, flower, x 8; E, flower, x
12; F, corolla, x 12; G, corolla
opened out, x 12; H, placen-
tation and developing seeds
(semi-diagrammatic), x 24.
Artist: E.M. Stones, with kind
permission; taken from Hunt
(1969).
abscission, with lower part (‘petiole) persistent, becom-
ing spine-like. Flowers small, regular, in dense, scorpioid
cymes or solitary, well separated on lax branches. Calyx
deeply 4-lobed. Corolla pink or white, with a short, cam-
panulate tube, slightly constricted at throat, membranous
with 4 ovate lobes ± as long as tube or shorter. Stamens
4, borne on corolla throat, shorter than corolla lobes;
filaments linear, exserted; anthers subglobose, ± as long
as filaments, 2-locular, dehiscing introrsely. Disc absent.
Ovary bilocular, compressed, densely hairy, with a sin-
gle ovule in each loculus; style terminal, bifid at apex;
stigmas small, capitate. Fruit a capsule, variously pubes-
cent, loculicidally dehiscent. Seeds ± triangular, truncate
and pitted above with a circle of long, rigid trichomes in
upper part below pitted area, acute and shortly pubescent
below.
Key to subspecies
Flowers and capsules congested on short, lateral branchlets; leaves
yellowish green, 10-50 x 3. 5-6.0 mm; leaf blade surface with
multicellular-based setae prominent and an under layer of fine
setae with undeveloped bases 1 . W. dinteri subsp. dinteri
Flowers and capsules well spaced on lax branchlets; leaves greyish
white, 7-15 x 2.0-4.5 mm; leaf blade surface densely pubes-
cent, with fine setae with undeveloped bases and scattered
setae with multicellular bases 2.W. dinteri subsp. gracilior
1 . Wellstedia dinteri Pilg. subsp. dinteri in
Botanische Jahrbiicher 46: 559 (1912); Friedr.-Holzh.: 1
(1967) pro parte; D.R.Hunt: 4 (1969). Syntypes: Namibia,
Karasberge, Sandverhaar, Dinter 1193 (Bf); Natnib, Dinter
1379 (B|); Auchab, Range 691 (B|; SAM!); Karibis,
Dinter 1250 (B|; SAM!, lectotype, designated here).
62
Bothalia 38,1 (2008)
FIGURE 5. — Wellstedia dinteri
subsp. gracilior (= W. dinteri
var. gracilior), Strey 2140. A,
habit, x 1; B, branchlet with
flowers and fruit, x 3; C; older
branchlet with fruit, x 3; D,
flower, x 1 0; E, corolla, x 1 0;
F, corolla opened out, x 10.
Artist: E.M. Stones, with kind
permission; taken from Hunt
(1969).
Illustration: Hunt: t. 3666 (1969). Figure 4.
Dwarf shrub, up to 300 mm high, with pseudodis-
tichous branching. Stems procumbent, decumbent or
ascending. Leaves yellowish green, 10-50 x 3. 5-6.0
mm; leaf blade with multicellular-based setae, distinctly
spaced, and an under layer of fine trichomes prominent.
Corolla pink, white or pale rose in centre, white along
margins; tube ± 1 mm long. Flowering time'. April to
May, August to October.
Distinguishing characters', dwarf shrub; leaves 10-50
x 3. 5-6.0 mm; flowers and capsules mostly congested
on short, lateral branchlets; leaf blade with multicellular-
based setae, setae distinctly spaced, and an under layer
of fine trichomes prominent.
Distribution and habitat'. Wellstedia dinteri subsp.
dinteri is found in Namibia and Northern Cape (Figure
1). It occurs on arid, gravelly hillocks, in shallow soil
between rocks and stones.
2. Wellstedia dinteri subsp. gracilior ( D.R.Hunt )
Retief & A.E.van Wyk, comb, et stat. nov.
W. dinteri Pilg. var. gracilior D.R.Hunt in Hooker’s leones
Plantarum 37: t. 3667 (1969). Type: Namibia, Buellspoort, Strey 2140
(K!, holo.; BOL!, PRE!, WIND!).
Illustration: Hunt: t. 3667 (1969).
Procumbent, dwarf shrub with branches radiating
from centre, ± 100-150 mm high, 300-400 mm diam.
Leaves greyish white, 7-15 x 2^4.5 mm; leaf blade
surface densely pubescent with fine setae, bases unde-
Bothalia 38,1 (2008)
63
veloped, and scattered setae with multicellular bases.
Corolla white; tube ± 1.5 mm long. Flowering time :
August to February. Figure 5.
Distinguishing characters : procumbent, dwarf shrub;
leaves 7-15 x 2. 0-4. 5 mm; flowers and capsules mostly
solitary on lax branchlets; leaf blade surface densely
pubescent, with fine setae, bases undeveloped, and scat-
tered setae with multicellular bases.
Distribution and habitat : Wellstedia dinteri subsp.
gracilior is found in Namibia (Figure 1). It occurs on
rocky hillsides, black rocks and broken veld on dolo-
mite.
SPECIMENS EXAMINED
The numbers in brackets signify the identity of the
specimens: (1) W. dinteri subsp. dinteri ; (2) W. dinteri
subsp. gracilior.
Acocks 15615 (1) PRE; 15641 (2) PRE; 18101 (1) K. PRE. WIND.
Bean. Vlok & Viviers 1817 (1) BOL.
Craven 2532, 2597 (1) WIND.
Davidse 6339 (1) PRE. De Winter 3270 (1) K. PRE. WIND. Dinter
1250 (1) SAM; 4845 (1) BOL, K, PRE. SAM.
Galpin 14163 (1) E, PRE. Giess 14521 (1) WIND. Giess & Miiller
12165 (1) PRE. WIND; 14324 (1) WIND. Giess. Volk c£ Bleissner
5324 (2) K, PRE. WIND; 6829, 6978 (1) PRE. WIND; 8763 (1) PRE.
Goldblatt & Manning 8856 (1) PRE, WIND; 8763 (1) PRE. Giinster
9366 (2) WIND.
Kubirske & Coetzee 98 (1) WIND.
Jurgens 22412 (1) PRU; 22710 (1) PRE.
Leistner 2589 (1) PRE, WIND. Liebenberg 5179 (1) K, PRE. WIND;
5179 A (1) PRE.
Merxmuller & Giess 3444 (2) K, PRE. WIND; 3642 (1) K, PRE,
WIND. Miiller 265 (1) PRE, PRU, WIND; 1329 (2) PRE, WIND;
WIND26709 (1) WIND.
Oliver & Muller 6397 (2) K, PRE. Oliver. Midler & Steenkamp 6322
(1) K, PRE.
Pearson 4669 (1) BOL, GRA, SAM; 8576 ( 1) K, BOL, PRE.
Range 691 (1) SAM. Rodin 2843 (1) BOL, K, PRE.
Schlieben 9084 (1 ) BM, K, NBG, PRE. Strey 2140 (2) K, PRE. WIND.
Strohbach , Kubirske & Sheuyange 3011 (1) WIND. Smook 7562 (2)
PRE. Strohbach & Chivell 3474 (1 ) WIND.
Thompson & Le Roux 206 ( 1 ) NBG.
Van Jaarsveld. Forester & Jacobs 8428 (2) NBG. Van Wyk 8709 (1)
PRE. PRU, WIND. Volk 12277 (1) WIND.
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Angiosperm Phylogeny Group classification for the orders and
families of flowering plants: APG II. Botanical Journal of the
Linnean Society 141: 399 — 436.
BALFOUR, I.B. 1 884. Diagnoses plantarum novarum phanerogamarum
socotrensium. Proceedings of the Royal Society of Edinburgh 12:
402 — 4 1 1 .
DE WINTER, B. 1971. Floristic relationships between the northern
and southern arid areas in Africa. Mitteilungen der Botanischen
Staatssammlung, Miinchen 10: 424-437.
DIANE, N., FORTHER, H. & HILGER, H.H. 2002. A systematic
analysis of Heliotropium, Toumefortia , and allied taxa of the
Heliotropiaceae (Boraginales) based on ITS1 sequences and
morphological data. American Journal of Botany 89: 287-295.
DYER, R.A. 1975. The genera of southern African flowering plants,
vol.l, Dicotyledons. Botanical Research Institute, Pretoria.
ERDTMAN, G. 1960. The acetolysis method, a revised description.
Svensk Botanisk Tidskrift 54: 561-564.
FERGUSON, D M. 1999. Phylogenetic analysis and relationships
in Hydrophyllaceae based on ndh¥ sequence data. Systematic
Botany 23: 253-268.
FRIEDRICH-HOLZHAMMER, M. 1967. Wellstediaceae. Prodromus
einer Flora von Sudwestafrika 121: 1 .
GOLDBLATT, P. 1978. An analysis of the flora of southern Africa: its
characteristics, relationships and origins. Annals of the Missouri
Botanical Garden 65: 369 — 436.
GOTTSCHLING, M. 2003. Phylogenetic analysis of selected Bora-
ginales. Ph.D. thesis, Freie Universitat, Berlin. Unpublished.
GOTTSCHLING, M„ HILGER, H.H., WOLF, M. & DIANE, N. 2001.
Secondary structure of the ITS1 transcript and its application in
a reconstruction of the phylogeny of Boraginales. Plant Biology
3: 629-636.
GURKE, M. 1897. Borraginaceae. Die natiirlichen Pflanzenfamilien 4,
3a: 71-131.
HOLMGREN, P.K., KEUKEN, W. & SCHONFIELD, E.K. 1990. Index
herbariorum, edn 8. Regnum Vegetabile 120.
HUNT, D.R. 1969. Wellstedia socotrana, W. dinteri var. dinteri, W.
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LEBRUN, J. & STORK, A.L. 1 997. Gamopetales: Clethraceae a Lamia-
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MERXMULLER, H. 1960. Wellstediaceae. Mitteilungen der Botani-
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NOVAK, F.A. 1943. Wellstediaceae. Prat, Rostlinopsis 9: 530.
PHILLIPS, E.P. 1951. Boraginaceae. The genera of South African flow-
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.
Bothalia 38,1 : 65-88 (2008)
Notes on African plants
VARIOUS AUTHORS
ASTERACEAE-GNAPHALIEAE
METALASIA HELMEI, A NEW MEMBER OF A SMALL CLADE FROM THE WESTERN CAPE?
The genus Metalasia R.Br. of the Asteraceae, tribe
Gnaphalieae, was revised by Karis (1989). At that
stage, 52 species and three subspecific taxa were rec-
ognized. An unknown Metalasia was recently collected
by the second author on the Kwadousberg to the east of
Worcester, in the Western Cape. After thorough investi-
gation, it turned out that this newly collected plant does
not fit into any currently known species of Metalasia ,
and that it constitutes a distinct, undescribed species.
Metalasia helmei P.O. Karis, sp. nov.
Fruticulus sine brachyblastis axillaribus. Folia basi
plerumque ± applanata versus apicem canaliculata, non
tortilia, 3. 2-6. 5 mm longa. Synflorescentiae (4— )8— 1 8
mm latae, densae, capitulis (2-) 10-25. Capitula floribus
5. Bracteae involucri ± 15, 6- vel 7-seriatae, intrinsecus
gradatim longiores, erectae autem intimae erecto-pat-
entes, extimae foliaceae vel subfoliaceae, intrinsecus
scariosae vel petaloideae, brunneae vel atrobrunneae,
intimae albae. Cypselae anguste oblongae vel anguste
ovoideae, 2.2-3. 0 mm longae, glabrae, costis 5-10 inae-
qualiter distributis. Pappi setae serratae, apicaliter clava-
tae.
TYPE. — Western Cape, 3319 (Worcester): Worcester
Div., Kwadousberg, Farm Witvlakte 175, on plateau at
head of Keurboskloof, 1 400 m, S 33°31' E 19°43', (-DA),
10-10-2005, Helme 3647 (NBG, holo.!; BOL!, K!, MO!,
PRE!, S!).
Sparsely to well-branched shrublet up to 0.4 m high.
Branches mostly erect, sometimes ascending, without
brachyblasts, young whitish tomentose, older becom-
ing less tomentose and with leaf scars; foliage on young
shoots dense to very dense, but becoming less so with
age. Leaves involute-ericoid, not twisted, very nar-
rowly triangular to narrowly triangular or ovate-acumi-
nate, upper part 3. 2-6. 5 x 0.75-1.0, base 1.0-1. 9 mm,
acute or rounded apically, mucronate, tips straight or
sometimes declinate, glabrous beneath, erect-spreading
or rarely spreading, straight or occasionally subsquar-
rose, or slightly curved, often slightly involute and ± flat
at base and more involute and canaliculate towards tip.
Synflorescences (4 — )8— 1 8 mm wide, campanulate, dense,
with (2-) 10-25 capitula arranged in obscure clusters or
sometimes solitary. Capitula 5-flowered, sessile, free
from one another. Involucre hairy for 3/5-2/3, cyathiform
but conspicuously widened at petaloid part, apically
2. 3-3. 6 mm wide, slightly wider at anthesis; bracts ± 15
in 6 or 7 series, gradually longer inwards, hyaline mar-
gins often absent or sometimes narrow or obsolete, out-
ermost-inner erect, innermost erect-spreading; outermost
± half as long as innermost and foliaceous to subfolia-
ceous, narrowly ovate or elliptic, acute, mucronate; outer
scarious, brown to dark brown (sometimes reddish),
elliptic to narrowly elliptic to narrowly oblanceolate,
acute to obtuse, mucronate or not; inner scarious to sub-
petaloid, brownish to whitish, oblanceolate or narrowly
oblong, obtuse; innermost petaloid, white, narrowly
oblanceolate, with folded and/or plicate apical margins.
Corolla narrowly cylindrical, 3^1 mm long, purple or
reddish above. Cypselae narrowly oblong to narrowly
ovoid, 2. 2-3.0 mm long, with ± 5-10 unevenly distrib-
uted ribs, brown, glabrous. Pappus bristles slightly ser-
rate, apically clavate. Flowering time : August-October.
Figure 1.
Diagnostic characters'. Metalasia helmei is a small
shrublet immediately recognized by its mostly erect-
spreading, untwisted and rather short leaves, by the 5-
flowered capitula, and the dark brown, scarious, erect,
outer involucral bracts which contrast with the innermost
white ones which are erect-spreading. Furthermore, the
fruits have 5-10 unevenly distributed ribs, and the pap-
pus bristles are apically clavate. M. helmei is similar to
many other small, shrubby Metalasia species in habit,
but in its foliage it is most similar to M. agathosmoides
Pillans which also has 5-flowered capitula, but the latter
differs by its axillary brachyblasts, and the apically con-
cave, pink, petaloid, involucral bracts.
DISCUSSION
Karis (1989) made a morphologically-based cladistic
analysis of Metalasia , and M. agathosmoides formed a
clade with M. fastigiata (Thunb.) D.Don on account of,
among other things, the nonhomoplasious feature ‘cypse-
las with 5-7 endocarpous ribs’. Regrettably, this descrip-
tion is erroneous since the reinforced cells of the fruit,
constituting the ribs, are formed by the seed coat and
not by the innermost layer of the carpels. However, it is
obvious that M helmei shares these fruit features with M.
agathosmoides and M. fastigiata , but it shares the unusual
foliage (Figure 1) with the former alone. Consequently,
it is highly probable that Metalasia helmei and M. aga-
thosmoides are sister species. There is a slight disjunc-
tion between M. helmei and the more widely distributed
M. agathosmoides which has its closest locality at the
Bonteberg, some 20-25 km to the northeast (Figure 2).
As can be seen from the map, the disjunctions between
the (known) populations of M. agathosmoides are more
pronounced than between the species. Karis (1989, data
from the herbarium sheets) reported a range in altitude
66
Bothalia 38,1 (2008)
FIGURE 1. — Metalasia helmei, Helme 3138 (S). A, portion of plant; B, branch with leaves; C, leaf from above (left) and from side (right); D,
capitula; E, F: all involucral bracts from one capitulum, with E, outermost from upper left, to innermost F, on lower right; G, three cypselae,
each displayed from the ventral (left) and lateral (right) sides (note ribs). H, I, pappus: H, bristle; 1, tip magnified. Scale bars: A( 10 mm; B,
2 mm; C-G, 1 mm; FI, 0.5 mm; I, 0.1 mm. Artist: Emma Hulten.
Bothalia 38,1 (2008)
67
FIGURE 2. — Known distribution of Metalasia helmei, • ; and M. aga-
thosmoides , O, 0 (data from Karis 1989).
for M. agathosmoides of 1 250-1 700 m, but Karis sub-
sequently collected material of this species at 900-950 m
in the Cedarberg. M. fastigiata is also 5-flowered, as in
M. helmei and M. agathosmoides , but differs from both
those species by its larger size, by the twisted, rather
sharp-pointed leaves, as well as the synflorescences with
more numerous capitula. M. fastigiata also displays a
much wider ecological range. A study based on molecu-
lar and morphological data (Stangberg & Karis in prep.)
might indicate whether the M. agathosmoides-fastigiata
clade now should be extended to contain also M. helmei,
and, if the latter is sister to M. agathosmoides. If so, the
high altitudinal range might be due to common ances-
try. Metalasia compacta Zeyh. ex Sch.Bip. has leaves
that approach those of M. helmei, i.e. with a ± flat and
slightly involute base, but more conspicuously involute
towards the tip. However, M. compacta differs by its
robust habit, its pure white-tomentose younger branches,
its twisted leaves, as well as its 6-flowered capitula that
are arranged in larger synflorescences.
Distribution and habitat: Metalasia helmei is known
only from the Kwadousberg, where it is fairly common
in Sandstone Fynbos on well-drained, deep, coarse white
sands from 1 400-1 800 m (Figure 2). The upper parts
of this 12 km2 range receive a light dusting of snow on
average twice every winter. The area is relatively arid,
and rainfall probably ranges for 400-700 mm per annum.
The habitat is not under any direct threat, although there
are old agricultural lands near the type locality, and there
are estimated to be more than 10 000 plants. The entire
known range of this nonresprouting species constitutes a
single locality. Due to having less than five localities and
occupying an area of less than 5000 km2, it qualifies for
Rare in terms of the Orange List criteria (Victor & Keith
2004).
Additional material examined
Western Cape. — 3319 (Worcester): Worcester Div.,
Kwadousberg, Blaaskloof 182, summit ridge near jeep
track, 1 700 m, (-DA), 24 Oct. 2004, Helme 3138
(NBG!, S).
ACKNOWLEDGEMENT
Financial support from the Swedish International
Development Cooperation Agency — Swedish Research
Links (348-2004-5631 to POK) is acknowledged.
REFERENCES
KARIS, RO. 1989. Systematics of the genus Metalasia (Asteraceae-
Gnapahalieae). Opera Botanica 99: 1-150.
STANGBERG, F.K & KARIS, P.O. in prep. Generic delimitation of the
Metalasia clade: a total evidence-based approach.
VICTOR, J.E. & KEITH, M. 2004. The Orange List: a safety net for
biodiversity in South Africa. South African Journal of Science
100: 139-141.
P.O. KARIS* and N. HELME**
* Department of Botany, Stockholm University, SE-106 91 Stockholm,
Sweden.
** P.O. Box 22652, 7975 Scarborough, South Africa.
MS. received: 2007-08-31.
ASPHODELACEAE: ALOOIDEAE
BULB IN E TRIEBNERI. AN EARLIER NAME FOR BULBINE ALBA, AS WELL AS ADDITIONAL AND NEW LOCALITIES IN EASTERN
AND NORTHERN CAPE, SOUTH AFRICA
The genus Bui bine Wolf comprises ± 73 species
occurring in Africa and Australia. While only six species
are found in Australia (Watson 1987; Keighery 2004),
67 occur in southern Africa, with only five of these also
extending into tropical Africa (Williamson 2003; Klopper
et al. 2006). The genus is therefore essentially a south-
ern African entity. Bulbine is characterized by miniature
to low succulent plants with lax or dense racemes (some-
times somewhat corymbose) of mostly yellow (rarely
white, orange or pink) flowers with bearded filaments
(Smith & Meyer 2000). Bulbine triebneri Dinter is the
only known member of the genus with white flowers.
Following an earlier treatment of the genus Bulbine
(Baijnath 1977), three white-flowered species with other
distinguishing features were segregated into a new genus,
Jodrellia Baijnath (Baijnath 1978). Although B. triebneri
shares the character of white flowers with Jodrellia, it
is a true Bulbine, sharing a close affinity with the other
frutescent species. It is distinguished from its nearest
68
Bothalia 38,1 (2008)
FIGURE 3. — Distribution of Bulbine triebneri, based on specimens at
PRE, NBG and BOL, •; specimens listed in Baijnath (1977),
▲ ; new localities in the Eastern and Northern Cape, O.
relative B. frutescens (L.) Willd. by its soft-textured and
glaucous, subterete leaves (Van Jaarsveld 2001 ).
Recently, Van Jaarsveld (2001) described a new spe-
cies, Bulbine alba Van Jaarsv. from the Western Cape.
Examination of the type material of B. triebneri and B.
alba showed them to be conspecific, therefore B. alba
becomes a synonym of B. triebneri. B. triebneri was
based on Dinter 7899 (B, holo! -K, photo.!), material
that was collected by Von Triebner at Eendoom, east of
Warmbad in Namibia and cultivated in Windhoek (Von
Poellnitz 1943).
Bulbine triebneri is said to be widely distributed in the
Little Karoo and southern Great Karoo (Van Jaarsveld
2001). The current known distribution, based on that of
the synonym B. alba, is restricted to the Western Cape,
from Laingsburg in the west to Oudtshoom and Beaufort
West in the east (Figure 3). It is locally abundant and
grows on shale ridges and scree in the Succulent Karoo
and along the southern border of the Nama-Karoo (Van
Jaarsveld 2001). Baijnath (1977) has shown this taxon
to have a much wider distribution in the Western and
Eastern Cape and that it also extends into the Northern
Cape Province.
A healthy population of Bulbine triebneri was recently
found in natural vegetation in the Urquhart Caravan
Park, Graaff-Reinet, Eastern Cape. At this locality, the
plants grow in full sun on a south-facing scree slope next
to the road leading to the bottom of the wall of the Van
Ryneveld’s Pass Dam. According to the management of
the park, this area has never been under cultivation and
it is therefore a natural occurrence of the species. The
area falls in the Southern Karoo Riviere vegetation type
(Mucina & Rutherford 2006). The plants, with their pale
bluish green leaves, were well camouflaged among the
grass and other vegetation of the slope while their flowers
were closed early in the day, and only a few plants right
next to the road were initially noticed. Upon returning to
the site in the late afternoon, it was found that the flowers
had opened and that the entire slope was in fact covered
with plants of this species.
The species was also observed in the Willowmore area
of the Eastern Cape (S.P. Bester pers. comm.) in North
Kammanassie Sandstone Fynbos (Mucina & Rutherford
2006). Unfortunately, the observer was not able to col-
lect material for herbarium specimens at this locality at
the time and was unable to relocate the population on a
subsequent visit, probably because the plants were not
flowering. Another flowering population was observed
in October 2001, ± 5 km north of Pofadder on the
Onseepkans road, Northern Cape (G. Williamson pers.
comm.). The plants were growing on the western aspect
of a low quartzitic outcrop. Regrettably, no specimens
could be made, but an illustration was made from field
notes by G. Williamson (Figure 4). Barker collected the
species in the same area in August 1954 and a specimen
is housed at NBG (Baijnath 1977).
Bulbine triebneri was also recently collected on the
Ouberg Pass, Northern Cape. Plants were found in full
sun on a lower west-facing rocky scree slope, in Tanqua
Escarpment Shmbland (Mucina & Rutherford 2006).
A few seeds were collected from the Graaff-Reinet
population. They germinated rapidly in Pretoria and the
seedlings flowered within their first season. The plants,
although growing somewhat more robustly in cultivation
than in the field, retained their distinguishing characters.
The most interesting feature of this species is that the
flowers seemingly open only in the very late afternoon
for two to three hours at most, a pattern similar to some
grassland species of Trachyandra Kunth. The only other
Bulbine species where past midday flowering has been
reported thus far is Bulbine torsiva G.Will. (Williamson
1996). By the time the flowers of B. triebneri open,
those of the other yellow-flowered Bulbine species have
already closed or are in the process of closing. Flowers of
the genus are devoid of nectar and are visited mainly by
bees for pollen. The specific time of day when the flow-
ers are open and their white colour, suggest that B. trieb-
neri might have undergone an evolutionary adaptation to
a pollinator that differs from those of most other species
of Bulbine. It would be useful to investigate the plant-
pollinator relationships of the genus as a whole.
Additional material examined
EASTERN CAPE. — 3224 (Graaff-Reinet): Graaff-Reinet, Urquhart
Park, on slope next to road leading to dam wall (Van Ryneveld’s Pass
Dam), (-BA), R.R. Klopper & A.W. Klopper 214 (PRE).
NORTHERN CAPE. — 3220 (Sutherland): Ouberg Pass, near lower
end of pass (-AD), H.M. Steyn 810 (PRE).
ACKNOWLEDGEMENTS
Ms Hester Steyn, Data Management Unit, SANBI,
Pretoria, is thanked for producing the distribution map.
Ms Emsie du Plessis, of the Scientific Publications
Section, SANBI, Pretoria, is thanked for comments on
an earlier draft of this paper. We would also like to thank
an anonymous referee for suggesting improvements to
the manuscript.
REFERENCES
BAIJNATH, H. 1977. Taxonomic studies in the genus Bulbine Wolf,
sensu lat. Ph.D. thesis, University of Reading. Unpublished.
Bothalia 38,1 (2008)
69
FIGURE 4. — Bulbine triebneri, G.
& F. Williamson 5984. A,
whole plant, longest leaf 248
x 3 mm; B, sectioned leaf, 3
x 2.5 mm; C, leaf surface with
dark green veins; D, pedicel
bract flattened, 1.8 mm wide
at base, 2.5 mm long; E, flow-
er, side view, pedicel ± 4.3
x 0.5 mm; F, outer tepal 6 x
1 .8 mm; G, inner tepal 6x2.8
mm; H, outer stamen 3.7 mm
long with anther 1 mm long;
I, inner stamen 3.7 mm long
with anther 0.6 mm long; J,
glandular trichomes, longest
1.2 mm; K, ovary, style and
stigma, overall 3.6 mm long.
Artist: G. Williamson.
BAUNATH, H. 1978. Jodrellia, a new genus of Liliaceae from tropical
Africa. Kew Bulletin 32: 571-578.
KEIGHERY, G. 2004. A new species of Bulbine (Asphodelaceae) from
Western Australia. Nuytsia 15: 241-244.
KLOPPER, R.R., CHATELAIN, C„ BANNINGER, V., HABASHI,
C., STEYN, H.M., DE WET, C„ ARNOLD, T.H., GAUTIER,
L., SMITH, G.F. & SPICHIGER, R. 2006. Checklist of the flow-
ering plants of sub-Saharan Africa. An index of accepted names
and synonyms. Southern African Botanical Diversity Network
Report No. 42. SABONET, Pretoria.
MUCINA, L. & RUTHERFORD, M.C. (eds). 2006. The vegetation
of South Africa, Lesotho and Swaziland. Strelitzia 19. South
African National Biodiversity Institute, Pretoria.
SMITH, G.F. & MEYER, N.L. 2000. Asphodelaceae. In O.A. Leistner,
Seed plants of southern Africa: families and genera. Strelitzia 10:
582-586. National Botanical Institute, Pretoria.
VAN JAARSVELD, E. 2001. South African succulent plants: two new
species and two new combinations. Haseltonia 8: 37-41.
VON POELLNITZ, K. 1 943. Die Bulbine-arten Deutsch-Sudwestafrikas.
Feddes Repertorium 52: 111-114.
WATSON, E.M. 1987. Bulbine. Flora of Australia 45: 236-241.
WILLIAMSON, G. 1996. New Bulbine species (Asphodelaceae) from
the northwestern Cape. Haseltonia 4: 13-23.
WILLIAMSON, G. 2003. Bulbine. In G. Germishuizen & N.L. Meyer,
Plants of southern Africa: an annotated checklist. Strelitzia 14:
989-992. National Botanical Institute, Pretoria.
R.R. KLOPPER*, A.W. KLOPPER**, H. BAIJNATH***
and G.F. SMITH+
* Research and Scientific Services Directorate, South African National
Biodiversity Institute, Private Bag X101, 0001 Pretoria.
** Molecular Ecology and Evolution Programme, Department of
Genetics, University of Pretoria, 0002 Pretoria.
*** School of Biological and Conservation Sciences, University
of KwaZulu-Natal, Westville Campus, Private Bag X54001, 4000
Durban.
+ Biosystematics Research and Biodiversity Collections, South African
National Biodiversity Institute, Private Bag X101, 0001 Pretoria/
Acocks Chair, H.G.W.J. Schweickerdt Herbarium, Department of
Botany, University of Pretoria, 0002 Pretoria.
MS. received: 2007-05-10.
70
Bothalia 38,1 (2008)
ASPHODELACEAE
NOTES ON THE NOMENCLATURE AND TYPIFICATION OF ALOE NATALENSIS (ALOOIDEAE)
INTRODUCTION
The morphologically highly variable A. arborescens
Mill, is widely distributed in southern and south tropical
Africa (Smith et al. in review) and is important medici-
nally (Liao et al. 2006) and horticulturally (Van Jaarsveld
2002) far beyond its African range. Select variants
[most importantly, A. arborescens Mill. var. natalensis
(J.M.Wood & M.S. Evans) A. Berger] remain important
in the context of biotechnological (Kawai et al. 1993)
and pharmacological (Obata et al. 1993; Teradaira et al.
1993; Tsuda et al. 1993) research, particularly in Japan.
In view of the recent taxonomic dismantling of A. arbo-
rescens (Van Jaarsveld & Van Wyk 2005), and consider-
ation of infraspecific diversity for a CITES assessment
(Smith et al. in review), we revisited its synonymy. This
revealed both typification and protologue citation errors
in respect of A. natalensis J.M.Wood & M.S. Evans.
NOMENCLATURE OF ALOE NATALENSIS
The name Aloe natalensis first appeared in the litera-
ture in a parochial report on the activities of the Colonial
Herbarium in Durban, South Africa (today the KwaZulu-
Natal Herbarium, NH). At that stage, the authors of the
report (Wood & Evans 1901a) were uncertain, due to dis-
ruptions related to the South African [Anglo-Boer] War,
whether their new taxa descriptions would first appear
in print in the 39th volume of the Journal of Botany,
British and Foreign to which they had earlier submitted
it as part of an ongoing series called ‘New Natal Plants’.
As it turned out, the herbarium report was published first
(before May 1901), as confirmed by the editor of the
Journal of Botany , James Britten (see editorial note on
page 169, vol. 39). Furthermore, the Journal of Botany
publication (Wood & Evans 1901b) was only a verba-
tim repeat of Wood & Evans’s earlier account. Although
neither publication cites a voucher specimen, the name
Aloe natalensis is considered to have been validly pub-
lished in the Colonial Herbarium report (Wood & Evans
1901a) in terms of the International Code of Botanical
Nomenclature ( ICBN) (McNeill et al. 2006).
In volume 3 of his Natal plants , Wood (1902) profiles
Aloe natalensis more comprehensively and provides
a detailed illustration. Under the heading ‘Habitat’ he
cites the following: ‘Natal: Midlands, 800 to 3,000 feet
alt, usually in rocky situations; Inanda and Noodsberg,
Wood , cultivated in Natal Botanic Gardens, Wood No.
4342' . This latter specimen was prepared from a plant
grown in the Natal Botanic Gardens (today the Durban
Botanic Gardens) and labelled ‘ Aloe arborescens Mill.’
in the case of the NH specimens, and ‘A. purpurascens '
in the case of a duplicate sent to Kew. Notably, Wood,
who as Curator of NH from 1882 till 1915 (Schrire
1983), never changed the name of the NH specimen
label to A. natalensis , despite citing it as a voucher
for the name in several publications, the first of which
was Wood (1902). He continued to uphold his concept
of natalensis at species level in various publications
(Wood 1908; 1912; 1915) until the time of his death.
Considering the manner in which Wood cited specimens
for other species treated in the Natal plants series, it
would appear that the unnumbered reference to 'Wood'
may imply a Wood specimen submitted to the Herbarium
without a collector’s number. Such unnumbered Wood
specimens [of A. arborescens ] are to be found in the NH
collection, but none antedate the Colonial Herbarium
report. Therefore it is more likely that the citation of
'Wood’ in Natal plants (Wood 1902) refers to sight
records of this taxon at both ‘Inanda and Noodsberg’,
localities referred to in the A. natalensis tableau. Berger
(1908) referred to ‘Wood n. 5019! und 5020!’ in his
treatment of 'A. arborescens Mill. var. natalensis (Wood
et Evans) Berger’ — material that had earlier been depos-
ited at NH under Wood 4342. Berger evidently cited
the separate herbarium accession numbers rather than
Wood’s single collection number in view of the anoma-
lous referencing situation with which he was confronted.
Both specimens of Wood 4342 have survived at NH and
each provides representative vegetative, and reproduc-
tive, material. One of these (in NH5019 ) is more com-
plete in that it has a seed vial labelled ‘ Aloe natalensis'
attached to it.
Kew holds a duplicate of Wood 4342 but as the mate-
rial was received by that institution on 4 September
1890, and as Wood never subsequently travelled to Kew,
it is reasonable to assume that this particular specimen
was not used in the course of describing A. natalensis a
decade later: duplicates were after all available to Wood
and Evans at NH.
Seeing that no voucher was cited in the protologue,
a lectotype is here designated. From the specimens that
were at the disposal of the authors, we chose the most
complete duplicate of Wood 4342, namely the one with
the NH accession no. 5019. The duplicate in NH and the
one at K are considered to be isolectotypes.
Despite the unambiguous statement of Britten, sub-
sequent regional treatments of Aloe L. have incorrectly
cited the protologue, in respect of both place and date
of publication (Reynolds 1950; Glen & Hardy 2000).
In their treatment of the genus Aloe for the Flora of
southern Africa, Glen & Hardy (2000) cite the follow-
ing protologue publication: A. natalensis J.M.Wood
& M.S. Evans: 9 (1900) [cited in their bibliography
as: ‘Durban Botanic Society Report on Natal Botanic
Gardens for the year 1900’. This was published in 1901].
They give the following type citation: KwaZulu-Natal,
Inanda, Wood 4342 (NH, holo.!; K!; PRE, photo.!). The
cited PRE photograph was not found nor could the ori-
gin of any of the material seen be traced to Inanda.
We conclude that the authorship and place of valid
publication of Aloe natalensis should be cited as fol-
lows:
Bothalia 38,1 (2008)
71
Aloe natalensis J.M.Wood & M.S. Evans in J.M.
Wood in Colonial Herbarium. Report for the year 1900:
9 (1901a).
The type of Aloe natalensis should be cited as fol-
lows:
TYPE. — Natal, 2931 (Stanger): ex hort., Botanic
Gardens, Durban, (-CC), 10-6-1890, Wood 4342 in
NH5019 (NH, lecto.), Wood 4342 in NH5020 (NH,
isolecto.), Wood 4342 (K, isolecto.).
ACKNOWLEDGEMENTS
Dr O.A. Leistner, previously of the Publications
Section, SANBI, Pretoria, is thanked for discussing this
problem with us and commenting on an earlier draft of
this paper. The Board of Trustees of the Royal Botanic
Gardens, Kew kindly made available, images of both
their isolectotype, and associated correspondence.
REFERENCES
BERGER, A. 1908. Liliaceae-Asphodeloideae-Aloineae. 8. Aloe L. In
A. Engler, Das Pflanzenreich. Regni Vegetabilis Conspectus , IV.
38. III. II: 159-326. Engelman, Leipzig.
GLEN, H.F. & HARDY, D.S. 2000. Aloaceae (First part): Aloe. In G.
Germishuizen, Flora of southern Africa, vol. 5, part 1, fasc. 1:
1-167. National Botanical Institute, Pretoria.
KAWAI, K„ BEPPU, H„ KOIKE, T„ FUJITA, K. & MARUNOUCHI,
T. 1993. Tissue culture of Aloe arborescens Miller var. natalen-
sis Berger. Phytotherapy Research 7: S5-S10.
LIAO, H.M., SHENG, X.Y. & HU, Z.H. 2006. Ultrastructural studies
on the process of aloin production and accumulation in Aloe
arborescens (Asphodelaceae) leaves. Botanical Journal of the
Linnean Society 150: 241-247.
McNEILL, I, BARRIE. F.R., BURDET, H.M., DEMOULIN, V.,
HAWKSWORTH, D.L., MARHOLD. K.. NICOLSON, D.H.,
PRADO, J„ SILVA, P.C., SKOG, J.E., WIERSEMA, J.H. &
TURLAND, N.J. (eds). 2006. International Code of Botanical
Nomenclature (Vienna Code) adopted by the seventeenth
International Botanical Congress Vienna, Austria, July 2005.
A.R.G. Gantner Verlag, Ruggell, Liechtenstein. [Regnum
Vegetabile 146].
OBATA, M„ ITO, S„ BEPPU, H„ FUJITA, K. & NAGATSU, T. 1993.
Mechanism of anti-inflammatory and antithermal bum action of
CPase from Aloe arborescens Miller var. natalensis Berger in
rats and mice. Phytotherapy Research 7: S30-S33.
REYNOLDS, G.W. 1950. The aloes of South Africa. The Trustees, The
Aloes of South Africa Book Fund, Johannesburg.
SCHRIRE, B.D. 1983. Centenary of the Natal Herbarium, Durban,
1882-1982. Bothalia 14: 223-236.
SMITH, G.F., KLOPPER, R.R. & CROUCH, N.R. (in review). Aloe
arborescens Mill. (Asphodelaceae: Alooideae) and CITES.
Haseltonia.
TERADAIRA, R„ SHINZATO, M„ BEPPU, H. & FUJITA, K. 1993.
Antigastric ulcer effects in rats of Aloe arborescens Miller var.
natalensis Berger extract. Phytotherapy Research 7: S34 S36.
TSUDA, H„ MATSUMOTO, K„ ITO, M„ HIRONO, I., KAWAI, K„
BEPPU, H., FUJITA, K. & NAGAO, M. 1993. Inhibitory effect
of Aloe arborescens Miller var. natalensis Berger (Kidachi
aloe) on induction of preneoplastic focal lesions in the rat liver.
Phytotherapy Research 7: S43-S47.
VAN JAARSVELD, E.J. 2002. Aloe arborescens and its nine cultivars.
Veld & Flora 88: 63-65.
VAN JAARSVELD, E.J. & VAN WYK, A.E. 2005. A new subspecies
of Aloe arborescens from the Mzimnyati River, KwaZulu-Natal.
Aloe 42: 40^12.
WOOD, J.M. 1902. Natal plants, vol. 3: t. 258. Bennett & Davis,
Durban.
WOOD, J.M. 1908. Revised list of the flora of Natal. Transactions of
the South African Philosophical Society 18: 121-280.
WOOD, J.M. 1912. Notes and corrections for ‘Natal plants’, vols 1 to 6.
Natal plants, vol. 6. Bennett & Davis, Durban.
WOOD, J.M. 1915. List of trees, shrubs, and a selection of herbaceous
plants growing in the Durban Municipal Botanic Gardens with a
few remarks on each. Bennett & Davis, Durban.
WOOD, J.M. & EVANS, M.S. 1901a. Aloe natalensis Wood & Evans.
In J.M. Wood, Colonial Herbarium. Report for the year 1900:
9/10. Bennett & Davis, Durban.
WOOD, J.M. & EVANS, M.S. 1901b. New Natal plants. Journal of
Botany, British and Foreign 39: 169-172.
N.R. CROUCH*, G.F. SMITH** and R.R. KLOPPER***
* Ethnobotany Unit, South African National Biodiversity Institute,
P.O. Box 52099, 4007 Berea Road/School of Chemistry, University of
KwaZulu-Natal, 4041 Durban, South Africa. E-mail: Crouch@sanbi.org
** Biosystematics Research & Biodiversity Collections, South African
National Biodiversity Institute, Private Bag XI 01, 0001 Pretoria/
Acocks Chair, H.G.W.J. Schweickerdt Herbarium, Department of
Botany, University of Pretoria, 0002 Pretoria. E-mail: SmithG@sanbi.org
*** Research and Scientific Services Directorate, South African
National Biodiversity Institute, Private Bag X101, 0001 Pretoria. E-
mail: Klopper@sanbi.org
MS. received: 2007-07-18.
ERRATA IN BOTHALIA 37,2 (2007)
MOFFETT, R.O. 2007. Name changes in the Old
World Rhus and recognition of Searsia (Anacardiaceae).
Bothalia 37 ,2: 165-175.
Page 169: Searsia krebsiana: replace ( Pres l ex Engl.)
with ( C.Presl ex Engl.).
Rhus krebsiana'. replace Presl ex Engl, with
C.Presl ex Engl.
Page 170: Searsia natalensis: replace (Bernh. ex Krauss)
with {Bernh. ex C. Krauss).
Rhus natalensis-. replace Bernh. ex Krauss
with Bernh. ex C. Krauss.
Page 171: Searsia pterota: replace (Presl) with (C.Presl).
Rhus pterota Presl in Botanische Bemer-
kungen: 44 (1884) should be:
Rhus pterota C.Presl in Botanische Bemer-
kungen: 41 (1844).
Page 173: Searsia volkii: replace (Siisseng.) with (Suess.).
Rhus volkii. replace Siisseng. with Suess.
Page 174: PRESL, K.B. 1884. Botanische Bemerkun-
gen: 43-45. Haase, Prague should be:
PRESL, K.B. 1844. Botanische Bemerkun-
gen: 40^12. Haase, Prague.
72
Bothalia 38,1 (2008)
HYACINTHACEAE
DRIMIOPSIS LINIOSETA, A NEW SPECIES FROM THE SEKUKHUNELAND CENTRE OF ENDEMISM, SOUTH AFRICA
The most recent taxonomic treatise on Drimiopsis
Lindl. & Paxton is by Muller-Doblies & Muller-Doblies
(1997). They recognized nine infrageneric taxa in south-
ern Africa and provided keys to their identification.
Lebatha (2005) subsequently revised both the subtropical
and tropical African material. In her study it became evi-
dent that a plant collected by the first author and Sharon
Turner in Sekukhuneland, South Africa was new to sci-
ence as it could not be keyed out using Muller-Doblies
& Muller-Doblies (1997). It also displayed a unique set
of morphological character states when compared to the
other known species.
Different opinions exist regarding the generic status
of Drimiopsis in relation to Resnova Van der Merwe and
Ledebouria Roth. In this regard we subscribe to Lebatha
et al. (2006) who view any formal generic reclassifica-
tion in the Ledebouriinae as premature, for example as
recently proposed by Manning et al. (2004). Hence this
new species is placed in Drimiopsis.
Drimiopsis linioseta A.J.Hankey & P.D. Lebatha,
sp. nov., D. reilleyanae U.Miill.-Doblies & D.Mull.-
Doblies similis, sed praesentia pseudopetioli, et setis
solum seriebus rectis differt.
TYPE. — Mpumalanga, 2529 (Witbank): between Roos-
senekal and Lydenburg, west-facing slopes opposite La
Ronelle guest house, (-BB), 25 November 1999, Hankey
& Turner 900 (PRE, holo.; K, iso.).
D. liniopapilla Lebatha, nom. inval., non rite publication (2005).
Plants 100-140 mm high, leaves appearing before or
with the flowers, bulbaceous. Bulbs hypogeal, ovoid to
oblong, 3CM-0 x 3CM-5 mm, gregarious, nonstolonifer-
ous, whitish; apices truncate; bulb scales loosely packed,
without threads when tom; outer scales white and fleshy;
dead bulb scales absent. Leaves 1 or 2, conduplicqte,
cordiform to broadly lanceolate, 30-80 x 100-140 mm,
without threads when tom, mostly unspotted adaxi-
ally, immaculate or variously streaked and mottled pur-
ple/brown abaxially; apex acuminate, base attenuate,
amplexicaul on scape; indumentum present, in form of
hairs arranged in rows on adaxial and abaxial lamina
surface, pseudopetiole and scape; lamina margin entire
becoming crenulate towards base, soft, pseudopeti-
olate; pseudopetiole off-white and horizontally banded
with purple/brown, especially toward base, ± as long
as lamina. Inflorescence 1(2) per bulb, a dense erect
raceme and flaccid post anthesis, ± as long as leaves;
rachis ovoid-terete, 20-40 mm long; peduncle dark
green-purplish; bracts in mature inflorescence vestigial.
Flowers 20-30, densely distributed becoming more lax
post anthesis, small, 2-3 mm long, actinomorphic, sex-
tepalous, hyacinth-scented, inner and outer lobes becom-
ing green to purple-brown, post anthesis; hypanthium
inconspicuous with base rounded, pedicellate; pedicel
glabrous, 1 mm or less. Tepals dimorphic, outer tepal
lobes purple-brown, apices and margins whitish in bud
becoming whitish green at anthesis, partially spreading,
longitudinally cucullate, apically conduplicate; inner
tepal lobes purple-brown with whitish green margins and
yellowish green conduplicate apices, hardly spreading at
anthesis, connivent, shorter than outer whorl, longitu-
dinally cucullate. Stamens 6, greenish to whitish, erect,
epitepalous, as long as pistil; filaments free, deltoid to
acuminate; anthers dorsifixed. Gynoecium tricarpellate;
ovary sessile, globose, whitish to greenish; ovules 2 per
locule; style as long as ovary, terete; stigma trilobed.
Flowering time : late spring and early summer. Figure 5.
Etymology, the specific epithet linioseta is named for
the conspicuous rows of setae that adorn all parts of the
plant except the flowers.
Diagnostic features and discussion', the phylogenetic
relationships within Drimiopsis are unclear at pres-
ent. A cladistic analysis by Lebatha (2005) grouped D.
linioseta with undescribed entities as well as D. atro-
purpurea N.E.Br. and D. pusilla. U.Mull.-Doblies &
D.Mull.-Doblies. However, these groupings are tenta-
tive due to the absence of synapomorphies and bootstrap
support. In terms of appearance, D. linioseta may also
be related to D. reilleyana. Drimiopsis linioseta is char-
acterized by leaves possessing hairs exclusively in rows
that trace the veins in the leaf (Figure 6). This charac-
ter state distinguishes D. linioseta from D. reileyana as
well as the prominent pseudopetiole, longer inflores-
cence and larger stature and smaller flowers (Table 1).
D. comptonii U.Mull.-Doblies & D.Mull.-Doblies also
possess epidermal structures in rows, as well as scattered
in between. D. linioseta is 100-140 mm high at anthesis
with larger conduplicate laminae, whereas D. comptonii
is a dwarf species ± 30-60 mm high at anthesis, with flat
to somewhat recurved laminae. The hairs in D. compto-
nii also possess a swollen base which does not occur in
D. linioseta. Furthermore, D. linioseta also differs from
D. comptonii in the flowers that are densely distributed
in the inflorescence, are minutely pedicellate ± 1 mm
long, and yellowish green as opposed to sparsely distrib-
uted purple-blue flowers with pedicels ± 0.4 mm long.
Other species that possess hairs on the leaves are D.
atropurpurea , D. pusilla and D. reilleyana, but here the
distribution of the hairs differs (Table 1).
In the past, Drimiopsis linioseta may have been mis-
identified as D. burkei subsp. burkei. However, as men-
tioned, D. linioseta differs not only by the possession
of rows of hairs, but also by the possession of a pseu-
dopetiole, a state not found in D. burkei subsp. burkei.
In this regard, D. linioseta resembles D. atropurpurea.
However, the possession of well-defined rows of hairs
again as compared to longer more scattered hairs, as well
as flower colour — purple in D. atropurpurea — sets them
apart (Table 1 ).
Bothalia 38,1 (2008)
73
FIGURE 5. — Drimiopsis linioseta, Hankey & Turner 900. A, bulb and plant, x 1; B, inflorescence and leaf; C, indumentum; D, flower; E, inner
tepals and stamen; F, outer tepal and stamen; G, opened perianth. H, gynoecium: left, from above; right, from side. I, capsule: top, from
above; bottom, from side. Scale bars: B, C, 20 mm; D, G, H, 1 mm; E, F, 2 mm; I, 7 mm. Artist: S. Burrows.
74
Bothalia 38,1 (2008)
TABLE 1 . — Principal differences between Drimiopsis linioseta and other species of Drimiopsis possessing an indumentum
The following couplet can be inserted into the key
provided in Mtiller-Doblies & Mtiller-Doblies (1997) to
assist in the identification of this species:
3c Leaves 1 or 2, setose, with setae arranged in longitudinal
rows, pseudopetiole ± length of lamina D. linioseta
FIGURE 6. — Drimiopsis linioseta, Hankey & Turner 900. SEM photo
of linear setae on adaxial leaf surface.
Distribution and habitat'. Drimiopsis linioseta has
been recorded from the Sekukhuneland area of Mpuma-
langa, where it is confined to the vicinity of Roossenekal
and Tonteldoos (Figure 7). It grows among rocks and
boulders, and in humus-rich pockets in the shade of
trees and shrubs often at the base of large boulders. The
known area of distribution places it in the Sekukhuneland
Centre of Endemism (Van Wyk & Smith 2001) and the
vegetation type Sekukhune Montane Grassland (Mucina
& Rutherford 2006).
Conservation status
Due to the limited distribution range and the ever-
present threat of habitat loss from mining in this area,
we consider the species at present to be best described as
vulnerable and give it a VU D2 ranking according to the
IUCN (2000) criteria.
thanked for the SEM contribution, Ronell Klopper for
her assistance in the National Herbarium and Sandie
Burrows for the line drawing.
REFERENCES
IUCN 2000. IUCN Red List Categories. Prepared by the IUCN Species
Survival Commission, IUCN Council, Switzerland, 9 February
2000.
LEBATHA, P.D. 2005. A systematic revision of Drimiopsis (Hyacin-
thaceae). Ph.D. thesis, North-West University, Potchefstroom.
Unpublished.
LEBATHA, P.D., BUYS, M.H. & STEDJE, B. 2006. Drimiopsis,
Resnova and Ledebouria : a tale of three genera. Taxon 55: 643-
652.
MANNING, J.C.. GOLDBLATT, P. & FAY, M.F. 2004. A revised
generic synopsis of Hyacinthaceae in sub-Saharan Africa, based
on molecular evidence, including new combinations and the
new tribe Pseudoprospereae. Edinburgh Journal of Botany 60:
533-568.
MUCINA, L. & RUTHERFORD, M.C. 2006. The vegetation of South
Africa, Lesotho and Swaziland. Strelitzia 19. South African
National Biodiversity Institute, Pretoria.
MULLER-DOBLIES, U. & MULLER-DOBLIES, D. 1997. A par-
tial revision of the tribe Massonieae (Hyacinthaceae). Feddes
Repertorium 108: 49-96.
VAN WYK, A.E. & SMITH, G.F. 2001 . Regions offloristic endemism in
southern Africa. A review with emphasis on succulents. Umdaus
Press, Hatfield, Pretoria.
A.J. HANKEY*, M.H. BUYS** and P.D. LEBATHA***
ACKNOWLEDGEMENTS * Walter Sisulu National Botanical Garden, South African National
Biodiversity Institute, P.O. Box 2194, 1731 Wilro Park, Gauteng, South
We thank Hugh Glen for assistance in translating Africa,
the diagnosis into Latin and the Botswana College of ** Compton Herbarium, South African National Biodiversity Institute,
Agriculture for financial support. We are grateful to Prlvate Bag X7’ 7735 Claremont and Department ot Botany and
r ■ .i Zoology, University of Stellenbosch, South Africa.
Mervyn Lotter for assisting With determining the con- *** Bca Herbarium, Department of Basic Sciences, Botswana College
servation status of the species, and Lize Agenbag for 0f Agriculture, Botswana,
comments on the conservation status. Lourens Tiedt is MS. received: 2007-06-12.
Bothalia 38,1 (2008)
75
ROSACEAE
VALIDATION OF THREE CLIFFORTIA TAXA IN BOTHALIA 37,1 (2007)
In the recent article on Cliffortia (Whitehouse & Fel-
lingham 2007), the following new names were invalidly
published, as explicit indications of the herbaria hold-
ing the holotypes were accidentally omitted: Cliffortia
sparsa, C. cuneata var. cylindrica and C. filifolia var.
arenaria. As this is contrary to the International Code of
Botanical Nomenclature: Art. 37.7 (McNeill et al. 2006),
the correct citations, including location of the holotype,
are given here:
Cliffortia sparsa C. Whitehouse, sp. nov.. in Bo-
thalia 37: 16 (2007).
TYPE.— Western Cape, 3419 (Caledon): Caledon
District, Hottentots Holland Nature Reserve, around
Nuweberg Forest Station, 560 m, (-AA), 25 Oct. 2001,
Whitehouse 283 (BOL, holo.!; NBG!).
Cliffortia cuneata var. cylindrica C. Whitehouse,
var. nov., in Bothalia 37: 18 (2007).
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, holo.!; NBG!).
Cliffortia filifolia var. arenaria C. Whitehouse, var.
nov., in Bothalia 37: 20 (2007).
TYPE. — Western Cape, 3422 (Mossel Bay): George
District, sand dunes E of Wilderness, (-BA), 3 Dec. 1951,
Esterhuysen 19336 (BOL, holo.!; NBG!, PRE!).
REFERENCES
McNEILL, J, BARRIE, F.R., BURDET, H.M., DEMOULIN, V.,
HAWKSWORTH, D.L., MARHOLD, K., NICOLSON, D.H.,
PRADO, J„ SILVA, P.C., SKOG, J.E., WIERSEMA, J.H. &
TURLAND, N.J. (eds). 2006. International Code of Botanical
Nomenclature (Vienna Code). Regnum Vegetabile 146: 1-568.
WHITEHOUSE, C.M. & FELLINGHAM, A.C. 2007. New species and
notes on the genus Cliffortia (Rosaceae). Bothalia 37: 9-22.
C.M. WHITEHOUSE*
* The Royal Horticultural Society Garden, Wisley, Woking, Surrey
GU23 6QB, England, UK.
OXALIDACEAE
A NEW SPECIES OF OXALIS FROM THE HANTAM-ROGGEVELD PLATEAU, NORTHERN CAPE, SOUTH AFRICA
Oxalis L. is a large genus of more than 800 species,
and although world-wide in distribution, most of the
species, some 90%, are concentrated in South-Central
America (± 500 spp.) and South Africa (±210 spp.).
Among the South African species, all but a half-dozen
are concentrated in the winter rainfall region, in the
Fynbos and Succulent Karoo Biomes (Oberlander et al.
2002).
Although now over 60 years old, the revision of the
South African species of Oxalis by Salter (1944) remains
the only complete study of the genus in the subregion.
Salter travelled extensively through the southwestern
Cape and Namaqualand but the adjacent Hantam and
Roggeveld escarpments remained mostly inaccessible to
him, which is why a species as common and attractive as
the one described here eluded him.
Oxalis odorata J.C. Manning & Goldblatt, sp. nov.
Planta acaulescens stolonifera, foliis basalibus, folio-
lis (2)3 vel 4(5) ellipticis vel oblanceolatis ad anguste
oblanceolatis, conduplicato-falcatis, emarginatis, (7-)
10—1 5(— 25) x 2-5(-8) mm, pallide griseo-viridibus adax-
iale minute papillato-puberulis, glabris, subglabris vel
sparse pubescentibus, atroviridibus abaxiale callosis dis-
persis punctiformibus nigris, floribus lilacinis vel albis
distale pallide lilacinis tubo viridi fauce viridi vel pur-
pureo, grateodoratis, sepalis lanceolatis, acutis vel obtu-
sis 4—5 x 1.0-1.25 mm, marginibus sparse ciliatis distale
et callis aurantiacis angustis ad apicem et in medio, pet-
alis oblique oblanceolatis 15-25 x 5-7 mm, filamentis
edentatis, antheris ellipsoideis.
TYPE. — Northern Cape, 3220 (Sutherland): 7 km
south of Sutherland, top of Verlatekloof Pass, Farm
Jakkalsvlei, seasonally wet clay flats, (-BC), 2 June
2007, Manning 3095 (NBG, holo.; MO, iso.).
Deciduous, acaulescent bulbous herb, not tufted. Bulb
narrowly ovoid, 15-20 x 7-10 mm; outer tunics papery,
brown, minutely striatulate. Rhizome 25-100 mm long,
bearing brown, papery scales and developing a slender
stolon in each scale axil; aerial stem absent. Leaves 4-
18, basal; petioles 10-40(-100) mm, glabrous or thinly
patent-pubescent, reddish, outer dilated or scale-like
below basal articulation; leaflets (2)3 or 4(5), elliptic or
oblanceolate to narrowly oblanceolate, conduplicate-fal-
cate, emarginate, (7—) 1 0— 1 5(— 25) x 2-5(-8) mm, pale
matte greyish green and minutely papillate-puberulous
adaxially, glabrous, subglabrous or thinly pubescent and
darker green abaxially with scattered, punctiform, black
calli. Peduncles 1 -flowered, 12-30 mm long, mostly
a little shorter than leaves, glabrous or thinly patent-
pubescent, with 2 subopposite, subulate, glabrous bracts
inserted in distal half 1— 5(— 7) mm below flower. Flowers
tristylous, lilac or white flushed pale lilac, with pale
76
Bothalia 38,1 (2008)
green, narrowly funnel-shaped tube, veins darker green
or purple in throat, sweetly scented; sepals lanceolate,
acute or obtuse, 4—5 x 1.0-1.25 mm, adpressed-pubes-
cent or subglabrous with few short hairs towards base,
margins sparsely ciliate distally, with narrow orange calli
at apex and towards midline; petals adhering for ± 6 mm,
obliquely oblanceolate, 15-25 x 5-7 mm; lamina obtuse,
± twice as long as claw, glabrous or ciliate on outer mar-
gin, with or without several elongated calli near apex.
Stamens : filaments erect, shorter 4—5 mm long, longer
6-7 mm long, longest 8-9 mm long and exserted ± 3
mm from tube, minutely glandular-pubescent, edentate;
anthers ellipsoid, ± 0.75 mm long, yellow. Ovary densely
hairy in distal half with a mixture of short, glandular
hairs and longer, unicellular, eglandular hairs; locules 2-
ovulate; styles minutely glandular-pubescent. Capsules
and seeds unknown. Flowering time : mid-May to mid-
June, rarely to late June. Figure 8.
Distribution and ecology, known from the summit of
the Hantamsberg at Calvinia and from the Roggeveld
Escarpment, from near Middelpos and Sutherland
(Figure 9). The species is locally common, forming large
populations on seasonally wet flats or around the mar-
gins of ephemeral pools in shallow soil on dolerite sills,
multiplying and spreading through the production of
axillary stolons from the scales along the rhizome.
Diagnostic characters and affinities', the pale lilac
flower of Oxalis odorata with a slight greenish or pink
eye, is sweetly scented during the day. Floral fragrance in
Oxalis is extremely rare, and its occurrence is not men-
tioned by Salter (1944). The leaves of O. odorata are
highly distinctive, with a pale, greyish green, matte upper
surface densely covered with minute papillae, contrast-
ing with the smooth or sparsely pubescent, darker green
underside. Electron microscopic examination of the leaf
surface shows the adaxial epidermal cells to be relatively
small, highly domed, and covered with a thick coating of
epicuticular wax platelets (Figure 10A, B), which cause
the characteristic greyish bloom on the leaf upper surface.
The abaxial epidermal cells, in contrast, are larger, with
a thick, laevigate cuticle lacking distinctive epicuticular
wax deposits, and those over the midline are longitudi-
FIGURE 8. — Oxalis odorata , Man-
ning 3095. A, whole plant; B,
single leaf flattened out show-
ing adaxial surface. C-F, flor-
al details: C, detached sepal;
D, detached petal; E, stamens
and ovary; F, ovary (rear two
styles not shown). Scale bar:
A-C, 10 mm; D-F, 2 mm.
Artist: John Manning.
Bothalia 38,1 (2008)
77
FIGURE 9. — Known distribution of Oxalis odorata.
nally elongated (Figure 10C). Stomata are restricted to
the adaxial leaf surface, where they are protected by the
conduplicate folding of the blade.
Salter (1944) divides the South African species of
Oxalis with 1 -flowered peduncles into eight sections.
Among these, O. odorata keys out to section Angustatae
Salter subsection Lineares Salter on the basis of its rela-
tively narrow leaflets more than twice as long as broad.
It is somewhat anomalous here in several features, nota-
bly its acaulescent habit and the elliptical leaflets that
lack paired apical calli. In section Latifoliatae Salter,
however, three other species of Oxalis are known with
similar discolorous, elliptical to ovate leaflets and mauve
flowers with some sort of darker eye in the mouth of the
tube ( O . callosa R.Knuth, O. petiolulata L. Bolus and O.
stokoei Weintroub) and the relationships of O. odorata
are more likely to lie with them. Although not men-
tioned by Salter (1944), the pale greyish upper leaf sur-
face is characteristic of this small group of species. In
this group, O. petiolulata from the Cedarberg has ebrac-
teate peduncles and leaflets with broadly ovate leaflets
bearing a single line of black calli along the margin.
This contrasts with the bracteate peduncles and leaflets
with scattered, punctiform calli found in the remaining
two species as well as in O. odorata. Among these, O.
stokoei from the Hex River Mountains is distinguished
by its unusual, subfibrous outer bulb tunics that split into
matted segments.
Thus, O. odorata is most likely to be confused with
O. callosa, another early-blooming species endemic
to the Bokkeveld Plateau and Hantam, and which also
produces stolons from the rhizome. Several vegetative
and floral characters separate the two. The bulbs in O.
callosa are ± ovoid and the species invariably produces
trifoliolate leaves with obovate leaflets 6-8 x 4-6 mm,
thus 1. 5-2.0 times as long as wide, with the upper sur-
face ± smooth and the lower surface densely adpressed-
pubescent and marked with reddish or orange calli; the
calli on the sepals are marginal; and the flowers have a
conspicuous dark red eye in the mouth. In contrast, O.
odorata has narrowly ovoid bulbs and variably foliolate
leaves, usually with 3 or 4 leaflets but not uncommonly
up to 5, which are narrowly elliptical-oblanceolate and
mostly 10-15 x 2-5 mm, thus (3— )4— 8 times as long
as wide, with the upper surface papillate-puberulous
and the lower subglabrous or sparsely pubescent and
marked with black calli; sepals with the calli concen-
trated towards the midline; and flowers with the cen-
tral eye only weakly developed. In addition, O. callosa
is not uncommonly caulescent and favours damp clay
flats over shale, although it will also grow in dolerite
outcrops, whereas O. odorata is strictly acaulescent and
restricted to seasonally wet or waterlogged habitats over
dolerite sills.
Additional specimens examined
NORTHERN CAPE. — 3119 (Calvinia): summit of Hantamsberg,
(-BD), 31 May 1999, Manning 2223 (NBG). 3120 (Williston): 66
km from Calvinia on Blomfontein road to Middelpos, (-CC), 4 June
FIGURE 10. — Scanning electron
micrographs of leaflet surface
micromorphology. A, adax-
ial surface (stoma arrowed);
B, leaflet margin; C, abaxial
surface with longitudinally
elongated epidermal cells with
narrower epidermal cells along
midline. Scale bars: 30 /u m.
78
Bothalia 38,1 (2008)
1980, Goldblatt 5533 (MO, NBG, PRE). 3220 (Sutherland): top of
Verlatekloof Pass, (-DA), 26 June 1977, M.B. Bayer 906 (NBG); 25
May 2003, Manning 2808 (NBG).
ACKNOWLEDGEMENTS
Our thanks to Elizabeth Parker for facilitating the trip
to collect type material of Oxalis odorata. The micro-
graphs were taken at the Electron Microscopy Unit of
the University of Cape Town and the plate was prepared
by Michelle Smith. Material was collected under a per-
mit from Northern Cape Nature Conservation.
REFERENCES
OBERLANDER, K.C., DREYER, L.L. & ESLER, K.J. 2002. Bio-
geography of Oxalis (Oxalidaceae): a preliminary study. Bothalia
32:97-100.
SALTER, T.M. 1 944. The genus Oxalis in South Africa. The Journal of
South African Botany, Suppl. vol. 1 .
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: 2007-07-09.
IRIDACEAE
ROMULEA LUTEA AND R. TUBULOSA (CROCOIDEAE), TWO NEW SPECIES FROM NAMAQUALAND, SOUTH AFRICA
Romulea Maratti, with ± 95 spp., is one of the larger
genera in subfamily Crocoideae, the largest of the seven
subfamilies now recognized in Iridaceae v Goldblatt
& Manning in press). The genus is distributed through
eastern sub-Saharan Africa, the Mediterranean and
the Near East, with its centre of diversity in the winter
rainfall region of southern Africa, where 76 species are
now known to occur (De Vos 1972, 1983; Manning &
Goldblatt 2001, 2004, 2006), including the two described
here. The current taxonomy of the genus recognizes two
subgenera and five sections, based largely on characters
of the corm (Manning & Goldblatt 2001, 2004), which
provide essential information for accurate identification
of many species. Flower structure, with few exceptions,
is conservative (Goldblatt et al. 2002).
Since the last review of the genus (Manning &
Goldblatt 200 1 ), three new species have been discovered
and named (Manning & Goldblatt 2004, 2006) and field
work in Namaqualand in 2007 has resulted in the discov-
ery of a further two novelties, which we describe here.
Romulea lutea J.C. Manning & Goldblatt, sp. nov.
Plantae 50-100 mm altae, caule subterraneo vel bre-
viter supra terrain producto, cormo oblique complanato
crista lunata ciliata, tunicis duris atrobrunneis, foliis 3,
1 basali, laminis anguste 4-sulcatis, inflorescentia flori-
bus usque ad 4, bracteis 8-10 mm longis bractea externa
subobtusa viridi vel purpurea suffusa marginibus trans-
lucentibus, interna acuta viridi marginibus latis trans-
lucentibus pallide brunneis striatis, floribus profunde
cupuliformibus aureis exteme viridibus cupulo 8-9 mm
profundo, tubo perianthii infundibuliformi ± 3 mm longo
parte inferiore ± 1 mm longo, tepalis oblanceolatis 11-
13 x 4—5 mm, staminibus flavis filamentis ± 4 mm lon-
gis minute pilosis in dimidio inferiore, antheris ± 4 mm
longis, stylo in parte superiore in ramos ± 1 mm longos
diviso, capsulis oblongo-ellipsoideis ± 8 mm longis,
pedunculis recurvatis.
TYPE. — Western Cape, 3118 (Vanrhynsdorp): coastal
sandveld west of Koekenaap at Farm Skaapvlei, local
in drainage line, (-AC), 24 August 2007, Goldblatt &
Manning 12868 (NBG, holo.; MO, PRE, iso.).
Plants 50-100 mm high; stem subterranean or shortly
aerial. Corm subglobose, asymmetric, obliquely flat-
tened towards base with crescent-shaped basal ridge;
tunics hard, smooth, dark brown, split into numerous
fine parallel fibrils on basal ridge and short fibrils up to
2 mm long above. Cataphylls 3, membranous, flushed
greenish above ground. Leaves 3, lowermost one basal
in plants with aerial stem, three to four times as long as
flowering stems, blades narrowly 4-grooved, 60-150 x
± 1 mm. Inflorescence of up to 4 solitary flowers; outer
bract obtuse, 8-10 mm long, green or flushed purple,
with narrow, translucent membranous margins, inner
bract acute, thin-textured, green with broad translucent
margins flecked pale brown, about as long as outer.
Flowers deeply cup-shaped, cup 8-9 mm deep, golden
yellow but pale green on reverse, ± 20 mm diam.,
faintly scented; perianth tube funnel-shaped, 3 mm long
with lower narrow portion ± 1 mm long; tepals oblan-
ceolate, 11-13 x 4^5 mm. Stamens yellow; filaments
inserted at base of cup, free, minutely pilose in lower
half, 4 mm long; anthers parallel, 4 mm long. Style
dividing opposite upper third of anthers, branches ± 1
mm long, divided for about half their length. Capsules
oblong-ellipsoid, ± 8 mm long, peduncles recurved.
Seeds unknown. Flowering time: August to early
September. Figure 11.
Distribution and ecology, thus far known only from
the Western Cape coast west of Koekenaap and north
of the Olifants River Mouth, where we encountered
several populations in seasonally moist drainage lines
and depressions (Figure 12) on the adjacent farms
Skaapvlei, Graafwater, and Brakvlei, although only
material from the type locality was collected. The flow-
ers open at about midday and close in the mid-afternoon
and have a faint, somewhat sour fragrance.
Diagnosis and relationships'. Romulea lutea is a
relatively unspecialized member of section Ciliatae
of subgenus Romulea and is distinguished from other
members of the section primarily by its relatively small,
completely yellow flowers. Plants are typically stemless
or with a short aerial stem, and in the latter instance it
is evident that they have a solitary basal leaf plus two
Bothalia 38,1 (2008)
79
FIGURE 11. — Romulea lutea, P Goldblatt & J.C. Manning 12868. A, whole plant; B, outer bract; C, inner bract; D, stamens and style; E, t/s leaf.
Scale bar: A-C, 10 mm; D, 2 mm; E, 0.25 mm. Artist: John Manning.
cauline leaves. In stemless individuals, however, all
three leaves appear to be basal and careful dissection
is necessary, when it becomes evident that the inflores-
cences emerge from the axils of the upper two leaves. R.
lutea resembles R. tabularis Beg. in general appearance,
including the solitary basal leaf with unspecialized blade
anatomy (Figure HE), but this species has white or pale
blue flowers with a yellow cup, typically a well-exserted
aerial stem, and more membranous inner bracts. It also
tends to occupy much wetter situations, typically water
meadows, seepages, and shallow temporary pools. R.
lutea also resembles R. schlechteri Beg., another white
or cream- (rarely yellowish) flowered species that occurs
along the west coast as far north as Piketberg. Apart from
flower colour, R. schlechteri differs from R. lutea in hav-
ing two basal leaves and leaf blades in which the lateral
primary veins lack complete sclerenchyma sheaths (De
Vos 1972, fig. 13: 7). This difference in the number of
basal leaves separates R. lutea from the other yellow-
flowered members of section Ciliatae with relatively
firm-textured inner bracts and complete bundle sheaths
(R. citrina Baker, R. elliptica M.P.de Vos and R. montana
Beg.), all of which have two rather than a solitary basal
leaf. The hue of the flowers in R. lutea is also somewhat
different, being golden yellow rather than canary yellow,
and lacking any trace of dark markings in the throat.
80
Bothalia 38,1 (2008)
Romulea tubulosa J.C. Manning & Goldblatt, sp.
nov.
Plantae 30-40 mm altae, caule subterraneo, cormo
subgloboso oblique complanato infra, tunicis duris atro-
brunneis, foliis 3-5 arcuatis vel sinuatis (20-)30^45 x ±
0.5 mm anguste 4-sulcatis, sulcis ciliatis, inflorescentia
floribus usque ad 4, bracteis 10-12 mm longis, bractea
externa marginibus apicibusque membranaceis brun-
neis striatis, bractea interna ad apicem furcata, floribus ±
hypocrateriformibus pallide flavis tepalis in parte dimidio
atrobrunneis striatis rubro-brunneis exteme, ± 15 mm
diam., tubo perianthii 13-14 mm longo ± cylindrico,
tepalis patentibus ellipticis 9-10 x 2. 0-2. 5 mm, stamini-
bus flavis filamentis ±1.5 mm longis glabris inclusis vel
± 0.5 mm exsertis, antheris ± 3 mm longis, stylo in parte
superiore in ramos ± 1 mm longos diviso, capsulis semi-
nibusque ignotis.
TYPE. — Northern Cape, 3018 (Kamiesberg): Kamies-
berg, ± 2 km SE of Leliefontein near Naras Farm,
local in renosterveld, (-AC), 28 August 2007, R. & R.
Saunders sub Manning 3117 (NBG, holo.).
Plants 30—40 mm high; stem subterranean. Corm sub-
globose, asymmetric, obliquely flattened towards base
with crescent-shaped basal ridge; tunics hard, smooth,
dark brown, split into numerous fine parallel fibrils on
basal ridge and fibres up to 4 mm long above. Cataphylls
3, membranous, flushed greenish above ground. Leaves
3-5, arching or sinuous, up to three times as long as
flowering stems, blade narrowly 4-grooved, (20-)30-45
x ± 0.5 mm, ciliate along grooves. Inflorescence of up to
4 solitary flowers; outer bract obtuse, 10-12 mm long,
green, with broad, translucent membranous margins wid-
ening above into prominent, brown-flecked, membra-
nous apex, inner bract notched apically, central portion
firm-textured and green, with broad translucent margins
and apex flecked dark brown, slightly longer than outer,
central portion with ± 12 closely spaced veins. Flowers ±
salver-shaped, 15-18 mm diam., pale canary yellow with
median brown streak in throat, flushed reddish brown on
reverse; perianth tube subcylindrical, 13-14 mm long,
widening slightly from base to apex; tepals spreading,
elliptical, 9-10 x 2. 0-2. 5 mm, outer slightly wider than
inner. Stamens yellow; filaments inserted ± 1 mm below
mouth of tube, shortly decurrent, glabrous, ±1.5 mm
long thus exserted up to ± 0.5 mm; anthers just exserted,
parallel, ± 3 mm long. Style dividing opposite middle of
anthers, branches ± 1 mm long, divided for ± half their
length. Capsules and seeds unknown. Flowering time:
August to early September. Figure 13.
Distribution and ecology : currently known from a sin-
gle locality on the Kamiesberg in central Namaqualand,
just southeast of Leliefontein near the farm Naras
(Figure 12). The species is inconspicuous and appears
to be relatively rare, occurring in small communities in
open ground in renosterveld. No estimate of the number
of individuals is available.
Diagnosis and relationships', a member of section
Ciliatae of subgenus Romulea, the pale yellow, salver-
shaped flowers of R. tubulosa are unique in the genus.
The distinctive perianth, with ± cylindrical tube longer
than the tepals, places it among just six known species
FIGURE 12. — Known distribution of Romulea lutea, •; and R. tubu-
losa, O.
in the genus with similar ± salver-shaped flowers, all
of which have a white or, more usually, pink to purple
perianth. Differences in vegetative features, including
corm tunics, bract structure and leaf anatomy, suggest
that this distinctive flower form has evolved repeatedly
in the genus from the ancestral, funnel-shaped type, and
is therefore no certain indicator of relationship. Romulea
tubulosa appears to be no exception to this pattern.
Three of the known species with such flowers, R. albi-
flora J.C. Manning & Goldblatt, R. hantamensis (Diels)
Goldblatt and R. syringodeoflora M.P.de Vos, are mem-
bers of subgenus Spatalanthus, whereas the remaining
three species, R. kamisensis M.P.de Vos, R. singularis
J.C. Manning & Goldblatt and R. stellata M.P.de Vos
belong in subgenus Romulea (Manning & Goldblatt
2001). The flowers of R. tubulosa , with tepals just 9-10
x 2. 0-2. 5 mm, are among the smallest in the genus and
comparable in size but not colouring to those of R. stel-
lata. The oblique corms with finely fimbriate lower tunic
margins of R. tubulosa place it among the latter group of
species.
Within subgenus Romulea section Ciliatae, the dis-
tinctive bracts, with broad, translucent margins and
apices, offer a better indication of the relationships
of R. tubulosa than flower shape, and suggest that it is
the fifth member of a group of species endemic to the
higher lying parts of Namaqualand and the Richtersveld,
centred on the Kamiesberg. This alliance comprises R.
maculata J.C. Manning & Goldblatt (white flowers and
purple-speckled cataphylls), R. neglecta (Schult.) M.P.de
Vos (magenta flowers), R. pearsonii M.P.de Vos (yel-
low flowers) and R. rupestris J.C. Manning & Goldblatt
(white flowers). The leaf anatomy of R. tubulosa is con-
sistent with this putative relationship, namely the pres-
ence of vascular girders joining the primary bundles to
the epidermis, and of sclerenchyma strands along the rib
margins (Figure 13E). Both of these anatomical features
are rare in subgenus Romulea but are characteristic of
this alliance (Manning & Goldblatt 2001). The funnel-
shaped flowers and relatively long floral bracts, 15-30
mm long, of these four members of the group are typical
Bothalia 38,1 (2008)
81
FIGURE 13. — Romulea tubulosa, R.
& R. Saunders sub Manning
3117. A, whole plant; B. outer
bract; C, inner bract; D, sta-
mens and style; E, t/s leaf.
Scale bar: A-C, 10 mm; D, 2
mm; E, 0.25 mm. Artist: John
Manning.
of the genus, and differ markedly from the salver-shaped
flowers of R. tubulosa, with a cylindrical perianth tube
13-14 mm long, and short bracts 10-12 mm long.
Within the alliance, the brown-flecked bract margins
and especially the yellow flowers, a colour that appears
to be derived in the genus, suggest that R. tubulosa may
be most closely allied to R. pearsonii. Apart from R.
tubulosa, this species also has the shortest bracts in the
group, 15-25 mm long.
Species of Romulea with salver-shaped flowers have
in most instances been shown to be adapted for pollina-
tion by long-proboscid flies in the family Nemestrinidae,
and conform to the Prosoeca peringueyi pollination
guild in their magenta to purple flowers (Goldblatt et
al. 2002; Goldblatt & Manning 2007). Although pol-
lination by long-proboscid insects is likely for R. tubu-
losa, the yellow perianth suggests that different insect
species are involved, most likely bee flies in the family
Bombyliidae. The general resemblance in form, colour
and in perianth tube length between the flowers of R.
tubulosa and those of co-occurring Zaluzianskya ben-
thamiana (Scrophulariaceae) suggests that they belong
to the same pollination guild.
ACKNOWLEDGEMENTS
We are indebted to Rod and Rachel Saunders for their
discovery of Romulea tubulosa. This study was sup-
ported by grant 8428-07 from the National Geographic
Society (US). Collecting permits were provided by the
Nature Conservation authorities of Western and Northern
Cape, South Africa.
REFERENCES
DE VOS, M.R 1972. The genus Romulea in South Africa. Journal of
South African Botany, Suppl. 9.
DE VOS, M.R 1983. Romulea. In O.A. Leistner, Flora of southern
Africa 7, part 2, fasc. 2: 10-73. Botanical Research Institute,
Pretoria.
82
Bothalia 38,1 (2008)
GOLDBLATT, P, BERNHARDT, P. & MANNING, J.C. 2002, Floral
biology of Rormilea (Iridaceae: Crocoideae): a progression from
a generalist to a specialist pollination system. Adansonia 24:
243-262.
GOLDBLATT, P. & MANNING, J.C. 2007. Pollination of Romulea
syringodeoflora (Iridaceae: Crocoideae) by a long-proboscid fly,
Prosoeca sp. (Diptera: Nemestrinidae). South African Journal of
Botany 73: 56-59.
GOLDBLATT, P. & MANNING, J.C. In press. The iris family: natural
history and classification. Timber Press, Oregon.
MANNING, J.C. & GOLDBLATT, P. 2001. A synoptic review of
Romulea (Iridaceae: Crocoideae) in sub-Saharan Africa, the
Arabian Peninsula and Socotra including new species, biologi-
cal notes, and a new infrageneric classification. Adansonia 23:
59-108.
MANNING, J.C. & GOLDBLATT, P. 2004. Two new species of Romu-
lea (Iridaceae: Crocoideae) from the western Karoo, Northern
Cape and notes on inffageneric classification and range exten-
sions. Bothalia 34: 17-22.
MANNING, J.C. & GOLBLATT, P. 2006. New species of Iridaceae
from the Hantam-Roggeveld Centre of Endemism, and the Bok-
keveld. Northern Cape, South Africa. Bothalia 36: 139-145.
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, PO. Box 299, St. Louis, Missouri 63166, USA.
MS. received: 2007-11-05.
ASPHODELACEAE: ALOOIDEAE
NEW EVIDENCE IN SUPPORT OF A DISJUNCT DISTRIBUTION OF ALOE KARASBERGENSIS
Aloe karasbergensis Pillans was described by Pillans
in 1928. It is a member of Aloe Section Paniculatae
Salm-Dyck ex Kunth, which consists of six species
namely A. buhrii Lavranos, A. karasbergensis , A. kom-
aggasensis Kritzinger & Van Jaarsv., A. kouebokke-
veldensis Van Jaarsv. & A.B.Low, A. reynoldsii Letty
and A. striata Haw. A. karasbergensis and A. komaggas-
ensis were reduced to subspecific status under A. striata
by Glen & Hardy (1987, 2000). Lavranos (2004) argued
that these two taxa should be recognized as separate spe-
cies, since all six taxa in A. Section Paniculatae are eas-
ily distinguishable on the basis of vegetative and floral
characters.
Aloe karasbergensis grows in semi-desert sand and
stony mountain slopes in Namibia and the Northern
Cape Province (Figure 14), especially in areas with very
low and erratic rainfall (Glen & Hardy 2000). The distri-
bution details provided by Reynolds (1950) indicate that
this species occurs in southern Namibia from the Kubub
Mountains south to the Richtersveld and Springbok area
in South Africa, and then again near Kenhardt and south
of Prieska. The distribution map provided by Glen &
FIGURE 1 4. — Distribution of Aloe karasbergensis , based on specimens
at PRE,®; confirmed record in central Northern Cape, O.
Hardy (2000) clearly echoes this disjunct distribution.
A. karasbergensis has also recently been found further
West, on the Aurus Mountains, inside Diamond Area No.
1 in Namibia (E. van Jaarsveld pers. comm.)
Even though the only specimen in the National
Herbarium, Pretoria (PRE) of Aloe karasbergensis from
the central Northern Cape was collected by the revered
J.P.H. Acocks, author of Veld types of South Africa , there
has been some doubt on the validity of the disjunct dis-
tribution range of A. karasbergensis , as reflected in
Reynolds (1950) and Glen & Hardy (2000).
A recent collection by the second author confirms that
the species indeed occurs in the central Northern Cape
Province.
NORTHERN CAPE. — 3021 (Vanwyksvlei): Carnarvon area, Farm
Konkakloof, (-DD), PJ. du Preez 3670 (BLFU).
REFERENCES
GLEN, H.F. & HARDY, D.S. 1987. Nomenclatural notes on three
southern African representatives of the genus Aloe. South African
Journal of Botany 53: 489-492.
GLEN, H.F. & HARDY, D.S. 2000. Aloaceae (First part): Aloe. In G.
Germishuizen, Flora of southern Africa, vol. 5, part 1, fascicle 1 :
1-1 59. National Botanical Institute, Pretoria.
LAVRANOS, J.J. 2004. Aloe striata Haworth and its associates: reflec-
tions on consistency in taxonomic treatment. Aloe 41 : 50, 51.
REYNOLDS, G.W. 1 950. The aloes of South Africa. The Aloes of South
Africa Book Fund, Johannesburg.
R.R. KLOPPER*, PJ. DU PREEZ** and G.F. SMITH***
* Research and Scientific Services Directorate, South African National
Biodiversity Institute, Private Bag XI 01, 0001 Pretoria. E-mail:
Klopper@sanbi.org.
** Department of Plant Sciences, Faculty of Natural and Agricultural
Sciences, University of the Free State, PO. Box 339, 9300 Bloem-
fontein. E-mail: Dpreezpj.sci@ufs.ac.za.
*** Biosystematics Research & Biodiversity Collections, South
African National Biodiversity Institute, Private Bag X101, 0001
Pretoria/Acocks Chair, H.G.W.J. Schweickerdt Herbarium, Department
of Botany, University of Pretoria, 0002 Pretoria. E-mail: SmithG@
sanbi.org.
MS. received: 2007-08-03.
Bothalia 38,1 (2008)
83
COLCHICACEAE
FURTHER NEW COMBINATIONS IN COLCHICUM
The recent sinking of the largely African genus An-
drocymbium Willd. into Colchicum L. (Manning et al.
2007) required the publication of more than 60 new
names and combinations in the latter genus. Two taxon
names were inadvertently omitted, and a further name in
Androcymbium resulting from the transfer of Merendera
schimperiana Hochst. (Persson & Del Hoyo 2007)
requires a new combination in Colchicum. Despite dif-
fering opinions regarding generic circumscriptions
(Persson 2007), the new combinations are effected here
for curatorial purposes.
Colchicum leistneri (U.MiiU.-Doblies & D.Miill.-
Doblies) C. Archer, comb. nov.
Androcymbium leistneri U.Miill.-Doblies & D.Mull.-Doblies in
Feddes Repertorium 109: 557 (1998). Type: Northern Cape, 2824
(Kimberley): Nooitgedacht, ± 10 mi. [16 km] SE of Barkly West, ±
3800 ft. [1 160 m], (-DA), 17.07.1959, Leistner 1969 (K, holo.; KMG,
PRE!, iso.).
Colchicum palaestinum (Baker) C. Archer, comb.
nov.
Androcymbium palaestinum Baker in Journal of the Linnean
Society, Botany 17: 445 (1879). Lectotype (designated by Greuter
1967: 253): ‘Palaestina prope Hieosolymam’, Roth s.n. (G-Bs, iso.).
Colchicum schimperianum (Hochst.) C. Archer,
non C. schimperi Janka:117 (1882), comb. nov.
Androcymbium schimperianum (Hochst.) K.Perss. & Del Hoyo
in Botanische Jahrbiicher 127: 228 (2007). Merendera schimperiana
Hochst.: 1126 (1842). Lectotype (designated by Persson & Del Hoyo
2007: 228): Ethiopia, Begemdir, ‘In montanis inter Enschedcap et
Schoata, 2000 ped. infra Enschedcap’, 4.12.1838, Schimper 1126 (P.
holo.; BR, G, K, M, P, iso.) [images of all types (excepting G) observed
on website: http://www.aluka.org, accessed 21.02.2008].
REFERENCES
BAKER, J.G. 1 879. A synopsis of Colchicaceae and the aberrant tribes of
Liliaceae. Journal of the Linnean Society, Botany 17: 405-510.
GREUTER, W. 1967. Contributiones floristicae austro-aegaea 10-12.
Candollea 22: 233-253.
HOCHSTETTER, C.F. 1842. In schaed. Schimperi iter Abyssinicum,
sect, secunda. [Printed notes, distributed to herbaria with
Schimper’s specimens, comprising descriptions and type cita-
tions of new taxa.]
JANKA, V. 1882. Colchicum schimperi. Megjegyzesek Boissier
Flora Orientalisanak otodik kotetenek elso fuzetehez. Magyar
Novenytani Lapok 6: 113—120.
MANNING, J., FOREST, F. & VINNERSTEN, A. 2007. The genus
Colchicum L. redefined to include Androcymbium Willd. based
on molecular evidence. Taxon 56: 872-882.
MULLER-DOBLIES, U. & MULLER-DOBLIES, D. 1998. De
Liliifloris Notulae. 6. De decuria prima specierum novarum
generis Androcymbium sect. Androcymbium (Colchicaceae) in
Africa Australi s.l. Feddes Repertorium 109: 551-572.
PERSSON, K. 2007. Nomenclatural synopsis of the genus Colchicum
(Colchicaceae) with some new species and combinations.
Botanischer Jahrbiicher 127: 165-243.
PERSSON, K. & DEL HOYO, A. 2007. Androcymbium schimpe-
rianum. In K. Persson, Nomenclatural synopsis of the genus
Colchicum (Colchicaceae) with some new species and combina-
tions. Botanischer Jahrbiicher 127: 165-243.
C. ARCHER*
* National Herbarium, South African National Biodiversity Institute,
Private Bag X101, 0001 Pretoria. E-mail: archerc@sanbi.org.
MS. received: 2008-01-10.
APOCYNACEAE
ANEW SPECIES OF HUERNIA (ASCLEPIADOIDEAE-CEROPEGIEAE) FROM ANGOLA
The genus Huemia R.Br. consists of 50 species
(Bruyns 2007), so that the present new species brings
this number to 51. Huemia is widely distributed in sub-
Saharan Africa and six species also occur in the Arabian
Peninsula, as far east as the former Peoples’ Democratic
Republic of Yemen (South Yemen). In Angola, Huemia
is represented by H. calosticta Bruyns, H. lopanthera
Bruyns, H. oculata N.E.Br., H. similis N.E.Br., H. urceo-
lata L.C. Leach, H. verekeri Stent and H. volkartii Peitsch.
ex Werderm. & Peitsch. Of these, H. calosticta, H. lop-
anthera and H. similis are 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 distributed in southern Africa, from Angola to
Mozambique (Bruyns 2005). The new species described
here is from the central coastal parts between Lucira and
Benguela, beyond the limits of the Namib Desert.
With seven species, Huemia is the largest genus
among the stapeliads in Angola. So far, apart from the
seven species of Huemia, Australluma peschii, Duvalia
polita, Hoodia currorii, H. mossamedensis , H. parviflora ,
H. pedicellata, Orbea huillensis, O. lutea, O. maculata,
Stapelia kwebensis, S. parvula, S. schinzii, Tavaresia
angolensis, T. barklyi and T. thompsoniorum are the other
1 5 stapeliads that are known in Angola (Bruyns 2005 and
more recent new records).
Huernia calosticta Bruyns, sp. nov., H. oculata simil-
issima sed cum corolla latiora, maculata marronina et
papillis longioribus, corona interiore latiore et praesentia
tuberculi nectarei differt.
TYPE. — Angola, north of Lucira, 600 m, Bruyns
10733 (BOL, holo.; E, iso.).
Dwarf succulent, forming clumps up to 200 mm diam.
Stems 20-200 x 8-20 mm (excluding teeth), erect to
spreading, grey-green; tubercles 6-10 mm long, spread-
ing, deltoid, slightly laterally flattened and joined into 5
angles along stem, tapering into slender caducous tooth.
84
Bothalia 38,1 (2008)
FIGURE 15. — Huernia calosticta, Bruyns 10733. A, side view of dissected flower; B, papillae inside corolla from near mouth of tube; C, side view
of gynostegium; D, face view of gynostegium; E, pollinarium. Scale bars: A, 5 mm; B, 0.5 mm (at A); C, D, 1 mm (at C); E, 0.25 mm (at
C). Artist: RV. Bruyns.
Inflorescence with several flowers developing in grad-
ual succession from short, stout peduncle, with nar-
rowly attenuate bracts 4—8 mm long at base; pedicel
8—10 xl.5 mm, spreading and holding flower facing
outwards and slightly upwards; sepals 11-12 mm
long, 1 mm broad at base, narrowly ovate-attenuate,
slightly papillate on exterior. Corolla shallowly bowl-
shaped, 8-12 x 20-25 mm; outside cream-coloured
becoming faintly greenish towards tips of lobes,
with small irregular maroon spots becoming smaller
towards tips of lobes, covered with low conical and
rough papillae, with 4 or 5 raised longitudinal veins
running down from lobes; inside cream-coloured with
round dark maroon spots radially elongated around
corona and nearly circular further out, spots very fine
on lobes and on steep part of tube, larger in flatter
part for 5-6 mm around corona, covered with min-
ute, slender, bristly papillae usually tipped with small
bristle, papillae usually same colour as background;
tube bowl-shaped, ± 7 mm deep, widening from base
to mouth; lobes 6 mm long, 12 mm broad at base,
broadly deflate, somewhat acuminate, spreading.
Corona 4x5 mm, pale yellow speckled finely with
maroon, sessile; outer lobes ± 0.5 mm long, descend-
ing to surface of corolla so that apex adpressed to it,
± semicircular-emarginate in outline; inner lobes ±
1 .5 mm long, adpressed to backs of anthers, dorsiven-
trally flattened, with broad transverse dorsal gibbos-
ity ± 2 mm wide at base, beyond anthers descending
slightly towards yellow broadly obtuse tips with white
bristles. Figure 15.
Huernia calosticta is known from near the coast
north of Lucira and from near Catengue, southeast of
Benguela (Figure 16). Both of these localities are to
the north of and outside the arid Namib region.
Plants occur in bushland dominated by Acacia ,
Commiphora and Adansonia digitata, often with a few
other succulents such as Ceraria carissoana, Pachypo-
dium lealii , Stapelia kwebensis, various species of
Kalanchoe, Euphorbia subsalsa and Sansevieria
cylindrica. In the vicinity, Huernia urceolata, Stapelia
kwebensis and Tavaresia angolensis were observed.
Huernia calosticta is closely allied to H. ocu-
lata and is vegetatively indistinguishable from it. In
H. calosticta the corolla faces outwards but slightly
upwards, rather than horizontally as in H. ocu/ata.
Inside, the corolla has dark maroon spots on a cream-
coloured background, unlike the alternating pale and
dark bands of H. ocu/ata. These spots are very fine on
the lobes and on the steep part of the tube, becoming
abruptly broader on the flat-tened base of the tube for
a radius of 5-6 mm around the corona. The inside of
Bothalia 38,1 (2008)
85
the corolla is covered with fine papillae which are ±
twice the length of those in H. oculata.
The corona is broader in Huernia calosticta than in H.
oculata and lacks the very slight stipe found at the base
in H. oculata. While the outer corona lobes are similar in
both, the two differ in the inner corona lobes. In H. calo-
sticta the inner lobes are broader, rising slightly over the
anthers and then descending towards their apices, rather
than rising up well above the style apex as in H. oculata.
The inner corona lobes have especially broad dorsal gib-
bosities, that are ± twice as broad as those in H. oculata.
Another difference is the presence here of a fairly con-
spicuous, erect tubercle beneath each guide-rail, a fea-
ture absent in H. oculata.
Other material examined
ANGOLA. — Near Catengue, SE of Benguela, 780 m, Bruyns
10756 ( BOL).
REFERENCES
BRUYNS, RV. 2005. Stapeliads of southern Africa. 2 vols. Umdaus
Press, Hatfield, Pretoria.
BRUYNS, P.V. 2007. A new species of Huernia (Asclepiadoideae-
Ceropegieae) from Angola. Bothalia 37: 15-17.
P.V. BRUYNS*
*Bolus Herbarium, University of Cape Town, 7701 Rondebosch.
MS. received: 2007-07-02.
ASTERACEAE
TR1PTER1S CALCICOLA. ANEW CALCIPHILOUS SPECIES FROM WESTERN CAPE, SOUTH AFRICA
The small tribe Calenduleae of the family Asteraceae
is centred in southern Africa, where it is concentrated
in the winter rainfall parts of the Western and Northern
Cape Provinces of South Africa. About 110-120 spe-
cies are known, currently distributed among 12 gen-
era (Nordenstam 2006, 2007), but the generic lim-
its and relationships within the tribe are still far from
completely resolved. The last revision of the southern
African species recognized the six genera Castalis
Cass., Chrysanthemoides Fabr., Dimorphotheca Vaill.
ex Moench., Garuleum Cass., Gibbaria Cass, and
Osteospermum L. (Norlindh 1943). The largest of these,
Osteospermum, was divided on the basis of cypsela
morphology into subgenus Osteospermum , with 12 sec-
tions, and subgenus Tripteris (Less.) T.Norl., with three
sections (Norlindh 1943). Following a morphological
cladistic analysis of the tribe, the genus Castalis was
included in Dimorphotheca along with Osteospermum
sect. Blaxium (Cass.) T.Norl.; and Osteospermum subge-
nus Tripteris and O. section Oligocarpus (Less.) T.Norl.
were each recognized at generic level as Tripteris
Less, and Oligocarpus Less, respectively (Nordenstam
1994a, b). As a result of these adjustments, the tribe
Calenduleae then comprised the eight genera Calendula
L., Chrysanthemoides, Dimorphotheca, Garuleum ,
Gibbaria, Oligocarpus, Osteospermum and Tripteris.
This treatment remained current until recently, when
analysis of DNA sequence data suggested that several
of the genera were paraphyletic as circumscribed. As a
result, the circumscriptions of some of the genera were
substantially narrowed in an attempt to define them as
monophyletic lineages (Nordenstam 2006; Nordenstam
et al. 2006). The species Gibbaria ilicifolia was seg-
regated as the new monospecific genus Nephrotheca
B.Nord. & Kallersjo, and the following elements were
removed from Osteospermum and Tripteris into new
or existing genera: the two species of Tripteris sect.
Unifenestrata T.Norl. were transferred to the new genus
Nephrotheca ; the new genus Norlindhia B.Nord. was
erected to accommodate the two species of Tripteris sect.
Efenestrata T.Norl. subsect. Confusa T.Norl. plus a third,
newly described species, N. aptera B.Nord. (Nordenstam
2006); the new genus Monoculus B.Nord. was described
for the two species previously placed in Tripteris sect.
Unifenestrata ; the monotypic genus Inuloides B.Nord.
was described for the single species of Tripteris sect.
Efenestrata subsect. Tomentosa T.Norl.; and a single
species of Osteospermum was transferred to each of the
genera Dimorphotheca and Oligocarpus.
Currently, therefore, the tribe Calenduleae comprises
the genera Calendula (± 15 spp.), Chrysanthemoides
(2 spp.), Dimorphotheca (20 spp.), Garuleum (8 spp.),
Gibbaria (1 sp.), Inuloides (1 sp.), Monoculus (2 spp.),
Nephrotheca (1 sp.), Norlindhia (3 spp.), Oligocarpus
(2 spp.), Osteospermum (± 45 spp.) and Tripteris (20
spp.) (Nordenstam 2007). Despite these taxonomic
adjustments, the genus Osteospermum is still evidently
paraphyletic as currently circumscribed (Nordenstam et
al. 2006). The distinction between Osteospermum and
Chrysanthemoides in particular has been blurred by the
discovery of a species that is intermediate between them
in cypsela morphology (Wood & Nordenstam 2003).
Few synapomorphies are available for the recogni-
tion of most of these new segregates, several of which
are mono- or oligotypic. It is clear that additional genera
will need to be recognized as further taxa are included
in a molecular analysis if this treatment is to be carried
to its logical conclusion (Nordenstam 2007). An alter-
native, and in our opinion a more useful treatment, is to
adopt a synthetic view of the genus Osteospermum as
constituting the monophyletic clade sister to the genus
Dimorphotheca as resolved in the phylogenetic analysis
reproduced by Nordenstam et al. (2006), thus includ-
ing the genera Calendula, Chrysanthemoides , Gibbaria,
Inuloides, Monoculus, Nephrotheca, Norlindhia, Oligo-
carpus and Tripteris. In this circumscription the two
genera Dimorphotheca and Osteospermum are separa-
ted from Garuleum by the synapomorphy of a shortly
bilobed style with an annular collar of hairs in the disc
florets, and are in turn distinguished from one another by
86
Bothalia 38,1 (2008)
several morphological, micromorphological and chemi-
cal characters: Dimorphotheca by large flower heads
with ray florets more than twice as long as the involucre,
± uniseriate involucre, ray florets that are white, orange
or purple (rarely pure yellow) and flushed dark beneath,
fertile or functionally male disc florets, uniseriate corolla
hairs, and the presence of dimorphecolic acid; and
Osteospermum by the smaller flower heads with ray flor-
ets usually up to twice as long as the involucre, mostly
bi- or triseriate involucre, yellow or orange-yellow ray
florets, functionally male disc florets, biseriate corolla
hairs, and the presence of significant quantities of calen-
dic acid (Nordenstam 1994b).
Until a complete classification of the tribe is avail-
able, however, we follow current practice and describe
the following new species in the genus Tripteris s. str.
(Nordenstam 2006), with which it accords in its 3-
winged cypselae with the apical air chamber enclosed
by three translucent windows. In this narrow sense,
Tripteris comprises ± 20 species of perennial (rarely
annual) herbs, subshrubs or shrubs that are widely dis-
tributed through Africa, with about half the species
occurring in the winter rainfall parts of South Africa
and southern Namibia. An attractive new species from
coastal limestone outcrops at Jacobsbaai on the Saldanha
Peninsula in the Western Cape represents the first lime-
stone endemic in the genus, which we describe here as
T. calcicola.
Tripteris calcicola J.C. Manning & Goldblatt, sp.
nov.
Herba multicaulis perennis, caulibus florentibus 200-
300 mm altis Tripteris aghillanae affinis sed ab ea habitu
rhizomatoso (non suffrutex pulviniformis), rhizomate
multiramoso lignoso ± 5 mm diam., foliis latioribus 30-
40 mm (non 5-20 mm) latis, bracteis involucralibus ± 20
(non 12 ad 15) atque flosculis radiati involucro ± triplo
(non duplo) longioribus distinguitur.
TYPE. — Western Cape: 3217 (Vredenburg): Jacobs-
baai, inland of Mauritzbaai, limestone outcrops, (-DD),
15 August 2007, Manning 3100 (NBG, holo.; MO, iso.).
Multi-stemmed, rhizomatous perennial, 200-300 mm
high; rhizome much-branched, ± 5 mm diam., produc-
ing clusters of tufted shoots that elongate the follow-
ing year and flower before dying down; stems suberect,
sparsely branched, nodding in fruit, glabrous on first
season’s growth but densely glandular-pubescent on sec-
ond season’s growth, pale green with glandular heads of
trichomes purple. Leaves alternate throughout, congested
basally, erect, decreasing in size acropetally, lower
leaves broadly obovate to oblanceolate, tapering below
into long, petiole-like base, midrib prominent abaxially,
petiole 35-45 mm long; blade 3CM10 x 20-30 mm, leath-
ery, puberulous with glandular and eglandular trichomes,
these longer along margins, margins sparsely and weakly
± 10-dentate, subobtuse-mucronulate, upper leaves
oblong-lanceolate, 10-20 x (3-)5-7 mm. Capitula het-
erogamous, radiate, ± 60 mm diam., solitary on shortly
leafy, peduncle-like stems. Involucre shallowly cam-
panulate, 10-12 mm diam.; involucral bracts ± 20, uni-
seriate, lanceolate, 6-7 x 2-3 mm, acute, densely glan-
dular-pubescent, glandular heads of trichomes purple,
with scarious margins 0.5-1 .0 mm wide. Receptacle flat,
glabrous. Ray florets female-fertile, 20-22; tube sparsely
glandular-pubescent, ± 0.8 mm long, up to 9 setting
fruit; lamina spreading but recoiling in afternoon, nar-
rowly elliptic, 4-veined, ± 3x as long as involucre, 20-
25 x 4-5 mm, straw-yellow marked with dark maroon
adaxially in basal 5-6 mm, flushed reddish abaxially,
veins red. Anthers vestigial, free, reduced to 4, subulate
staminodes ±0.8 mm long, yellow. Style terete, on short
stylopodium, branching ± 1 mm above mouth of tube,
branches narrowly elliptic-lanceolate, obtuse, ± 1.5 mm
long, yellow, lateral margins stigmatic. Ovary obovoid,
3-angled, ± 2 mm long, shortly glandular-pubescent.
Cypselae homomorphic, 3-winged, 10-12 x 7 mm, body
narrowly turbinate, ± 6 mm long, glandular-pubescent,
with apical, trifenestrate airchamber, windows ovate, 3.5
x 2.0 mm, wings translucent, ± 2.5 mm wide. Disc flo-
rets functionally male, numerous; corolla narrowly fun-
nel-shaped, 4—5 mm long, yellow; tube glandular-hairy,
± 3 mm long; lobes suberect, triangular, ± 2 mm long,
with marginal veins joining at sinuses and continuing
down tube. Anthers 2 mm long, dark purple with yellow
filaments; anther base tailed, tails equalling filament col-
lar; anther appendage ovate. Ovary compressed-ovoid
with lateral ribs, ± 1 mm long, glandular-pubescent.
Style terete, on short stylopodium, shortly bilobed api-
cally, lobes deltoid, acutely papillate with basal fringe of
longer trichomes. Flowering time : August. Figure 17.
Distribution and ecology, so far known only from the
vicinity of Jacobsbaai just north of Saldanha Bay on the
west coast of the Western Cape (Figure 18), the species is
a component of Saldanha Limestone Strandveld (Mucina
& Rutherford 2006), a rare coastal vegetation type that is
almost entirely limited to the Saldanha Peninsula and is
rich in Red Data species. Plants of Tripteris calcicola are
restricted to exposed outcrops of limestone, where they
grow in shallow, loamy soils overlying limestone pave-
ment. Co-occurring endemic shrublets include Diosma
cf. guthriei (Rutaceae), Felicia elongata (Asteraceae)
and Nenax hirta subsp. calciphila (Rubiaceae).
The woody rhizomes of Tripteris calcicola creep
along the surface of the rock through the shallow soil
layer, sending up numerous aerial shoots with tufts of
leaves during the growing season. These shoots do not
die back during the ensuing summer but elongate over
the second growing season during the winter, and flower
in the early spring before dying back completely after
fruiting. This distinctive growth form results in a sin-
gle ramet spreading over an area of one or two square
metres, producing numerous, tufted shoots each year
plus several suberect flowering stems. The flower heads
expand fully during the mid-morning but the ray florets
curl back by mid-afternoon. The flower heads have a
remarkable similarity in size, colour and marking to the
local form of Ursinia anthemoides (for illustration see
Manning 2003: 256).
Diagnosis and relationships', the large, 3-winged
cypselae with 3-windowed apical air chambers place
this species among those treated as Osteospermum sub-
genus Tripteris section Trifenestrata by Norlindh (1943)
and as the genus Tripteris s. str. by Nordenstam (1994a).
Among these species it appears to be most closely allied
to T. aghil/ana DC., a cushion-forming subshrub with
Bothalia 38,1 (2008)
87
FIGURE 1 7. — Tripteris calcicola, Manning 3100 (NBG). A, vegetative shoots; B, flowering shoot; C, involucral bracts. D-F, details of ray floret; D.
floret; E. staminode; F, style branches. G-I, disc floret: G, floret; H, single anther; I, style branches. J, fruiting head; K, cypsela. Scale bars:
A, B, J, 10 mm; C, 1.5 mm; D, G, K, 2 mm; E, F, H, I, 0.5 mm. Artist: J. Manning.
large, homomorphic cypselae, basally congested leaves,
and large flower heads in which the ray florets are
strongly marked at the base with dark purple bands and
the anther appendages at least are also dark purple. Ray
florets with dark bases are known elsewhere in the sub-
genus only in the somewhat distantly related Monoculus
monstrosus (Burm.f.) B.Nord. (= Tripteris clandes-
tinei Less.), an annual species placed in Osteospermum
subgenus Tripteris section Unifenestrata by Norlindh
(1943). T. calcicola differs from T. aghillana in distri-
bution, ecology, habit and morphology. T. aghillana is
widespread through the interior of the Western Cape,
reaching the coast only at Cape Agulhas, and although
also a subshrub with annual flowering stems, it is not
rhizomatous, and the persistent woody bases of the stems
are carried above ground so that the plant develops a
rounded, cushion-like habit. This is in contrast to the
subterranean woody rhizomes of T. calcicola , in which
only the leafy shoots project above the soil surface and
the plant develops a creeping, mat-like habit. T. calcicola
Bothalia 38,1 (2008)
FIGURE 18. — Known distribution of Tripteris calcicola.
is further distinguished by its broader leaves, 30—40 mm
wide vs 5-20 mm; more numerous involucral bracts, ±
20 vs 12-15; and longer ray florets, ± three times as long
as the involucre vs ± twice as long.
ACKNOWLEDGEMENTS
This species was discovered and brought to our atten-
tion by Jacobsbaai resident and conservationist, Koos
Claassens, and we are most grateful to him and his wife,
Elize. Rupert Koopman assisted in collecting the type,
which was made under a permit from Cape Nature, and
Roy Gereau kindly provided the Latin diagnosis.
REFERENCES
MANNING, J.C. 2003. Jewels of the veld. Struik, Cape Town.
MUCINA, L. & RUTHERFORD, M.C. 2006. The vegetation of South
Africa, Lesotho and Swaziland. Strelitzia 19. South African
National Biodiversity Institute, Pretoria.
NORDENSTAM, B. 1994a. New combinations in the Calenduleae.
Compos itae Newsletter 25: 46—49.
NORDENSTAM, B. 1994b. Tribe Calenduleae. In K. Bremer,
Asteraceae: cladistics and classification : 365-376. Timber
Press, Portland, Oregon.
NORDENSTAM, B. 2006. Generic revisions in the tribe Calenduleae
(Compositae). Compositae Newsletter 44: 38-49.
NORDENSTAM, B. 2007. Tribe Calenduleae Cass. In K. Kubitzki, The
families and genera of vascular plants. VIII Asterales. Springer,
Berlin.
NORDENSTAM, B„ KALLERSJO, M. & ELDENAS, P. 2006.
Nephrotheca, a new monotypic genus of Compositae-
Calenduleae from the southwestern Cape Province. Compositae
Newsletter 44: 32-37.
NORLINDH, T. 1943. Studies in the Calenduleae. I. Monograph of the
genera Dimorphotheca, Castalis, Osteospermum, Gibbaria and
Chrysanthemoides. Bloms, Lund.
WOOD, A. & NORDENSTAM, B. 2003. An interesting new species
of Osteospermum (Asteraceae-Calenduleae) from the Western
Cape Province, South Africa, providing a link to the genus
Chrysanthemoides. South African Journal of Botany 69: 572—
578’.
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: 2007-10-10.
Bothalia 38,1: 89-102 (2008)
Vegetation and vegetation-environment relationships at Grootbos
Nature Reserve, Western Cape, South Africa
M. MERGILI* and S. PRIVETT**
Keywords: Afromontane Forest, Agulhas Plain, biodiversity, CCA, fynbos, GIS, Milkwood Scrub Forest, numerical vegetation analysis,
TWINSPAN, Western Cape
ABSTRACT
The private Grootbos Nature Reserve is located at the Western edge of the Agulhas Plain in the Cape Floristic Region
of South Africa, an area characterized by high habitat and floristic diversity. The Reserve is covered in near-natural fynbos
shrublands with a few patches of forest and wetland. The main objective of this study was to classify the vegetation into
discrete units and relate them to the prevailing environmental conditions. The vegetation was analysed by numerical means
(TWINSPAN, DCA, CCA) and mapped on GIS. At the vegetation type level. Forest & Thicket and Fynbos formed distinctive
clusters, whereas the wetland releves were intermixed, but without relationships to one of these units. Fire incidence served
as the major determinant of the forest-fynbos boundary. The Forest & Thicket grouping was separated into Thicket (as
transitional to fynbos), Afromontane Forest and Milkwood Scrub Forest. Two broad complexes were distinguished within the
Fynbos grouping, the Alkaline Sand Fynbos Complex corresponding to Coastal Fynbos, and the Acid Sand Fynbos Complex
corresponding to Mountain Fynbos. They discriminated along gradients of pH, soil depth and rock cover. The complexes were
further subdivided into formations by using one or a few subjectively chosen dominant species as indicators. The transitions
between these formations were rather continuous than discrete. The vegetation type and complex levels correspond well to
existing fynbos-wide classifications. Comparing the formations to the results of other vegetation studies is problematic even
on the scale of the Agulhas Plain, due to the high regional plant diversity in the Fynbos Biome.
INTRODUCTION
An important, and well-established aspect of the spe-
cies-rich Cape flora is the difficulty in resolving regional
vegetation associations into meaningful, easily identifi-
able floristic groupings (Campbell 1986). Various authors
have tried to classify the habitat types and the plant com-
munities of the Fynbos Biome in the previous decades,
some looking at broad descriptions of major vegetation
types (Taylor 1978; Moll et al. 1984; Campbell 1985;
Cowling & Heijnis 2001) and others at detailed descrip-
tions of communities (McKenzie et al. 1977; Boucher
1978; Taylor 1983; Richards et al. 1995; Cowling et al.
1996; McDonald et al. 1996; Taylor 1996). The Agulhas
Plain, covering an area of 270 000 hectares of semi-arid
lowland fynbos and renosterveld, has been prioritized
as an area of high irreplaceability and high vulnerabil-
ity, with exceptionally rich coastal lowland ecosystems
(Cowling et al. 1999). Cowling et al. (1988) applied
the mountain vegetation classification approach of
Campbell (1985) to the Agulhas Plain. This classifica-
tion was produced using the Braun-Blanquet method
of table sorting and was based on structural and higher
taxon characteristics of the vegetation. Nine zonal com-
munities, at various hierarchical levels, were recognized
and mapped. This phytosociological approach provided
a qualitative description of community environmental
characteristics on the Agulhas Plain. The only study on
the Agulhas Plain to provide a quantitative assessment
of the importance of environmental factors was Richards
et al. (1995) who explored vegetation-environment rela-
* Corresponding author: Institute of Geography, University of Innsbruck,
Innrain 52, 6020 Innsbruck, Austria and Mountain Research: Man and
Environment, Austrian Academy of Sciences, Technikerstrasse 21a,
6020 Innsbruck, Austria. E-mail: martin.mergili@uibk.ac.at.
** Grootbos Private Nature Reserve, P.O. Box 148, 7220 Gansbaai,
South Africa. E-mail: sean@greenfiitures.co.za.
MS. received: 2005-12-15.
tionships in a 30 ha study area, the Soetanysberg, ± 15
km west of Cape Agulhas (34 45' S; 19°50' E). However,
no comparable studies have been undertaken on the west
side of the Agulhas Plain.
The major objective of this study was to fill this gap
through a detailed mapping and numerical analysis of
the vegetation of Grootbos Nature Reserve (GNR),
which is located at the western edge of the Agulhas
Plain, between the villages of Stanford and Gansbaai
(Figure 1 A). The geographic coordinates are 34°32'30" S
and 19°24'50" E. The privately owned reserve consists of
seven formerly separate farms covering an area of 1 700
ha. It is run as an upmarket ecotourism lodge. The land-
scape is sloping, with a maximum elevation of ± 475 m
along the slopes of the Swartkransberg (Figure IB). The
underlying rock type of the more elevated hills is quartz-
itic Table Mountain Sandstone. On the lower slopes,
deposits of sandy aeolian material overlay the bedrock.
Outcrops of the Bredasdorp Limestone are exposed in
some places.
The climate, which can be characterized as maritime
mediterranean, shows a strong seasonality in precipita-
tion. May to August are the wettest months (most pre-
cipitation is carried by northwesterly winds), whereas
the southeasterly winds prevailing in the summer months
are dry. There has been no long-term precipitation record
at GNR. The mean annual rainfall from 1996-1998 was
730 mm. The mean maximum daily temperatures fluc-
tuate between 25°C in February and 14°C in July. The
GNR is frost-free.
Since 1995 the property has been managed for con-
servation and ecotourism. Over the last decade a team
of botanists has been actively sampling and collect-
ing plant specimens over the entire reserve. At the lat-
est count, 660 species of indigenous plants have been
recorded on the property, with still more being found
90
Bothalia 38,1 (2008)
FIGURE 1. — A, study area located between Cape Town and Cape
Agulhas, 5 km east of Walker Bay seashore; B, western part
of Grootbos Nature Reserve gently sloping towards Walker
Bay, eastern part hilly with maximum elevation slightly below
500 m.
each year (Privett & Lutzeyer in press). Fifty-one spe-
cies are included in the Red Data List of threatened
species (Hilton-Taylor 1996), and three species ( Erica
magnisylvae, Cliffortia cmthospermo ides , Lachenalia
lutzeyerii) were recently discovered and are considered
endemic to the Reserve. The vegetation of GNR, largely
fynbos shrubland with some patches of forest and wet-
land, is amongst the best surveyed in the Cape Floristic
Region and lends itself to a detailed quantitative analysis
of vegetation-environment relationships.
METHODS
Data collection
Vegetation sampling at GNR was carried out in winter
1997 and in spring 2004. Seventy-one 50 m2 (10 x 5 m)
rectangular releves were analysed for their floristic com-
position and their major environmental features. They
were subjectively chosen in order to represent homoge-
neous patches of vegetation and divided into 10 cells,
2.0 x 2.5 tn each. The floristic composition was investi-
gated for each cell, including all identifiable species. All
taxonomy followed Goldblatt & Manning (2000). The
cover was recorded as a percentage. Soil samples were
taken at random localities within each releve, in depths
of 5-30 cm depending on soil depth. Environmental data
were recorded for each releve: topographic parameters
(slope, elevation, aspect), vegetation age, soil depth,
rock cover, pH, resistance and nutrient levels of the soil.
Flow accumulation (wetness index), solar radiation and
the exposure to the fire-bearing southeasterly winds were
computed from a digital elevation model (DEM). All
releves were used for analysis, including those located
in transitional vegetation. The GPS coordinates of each
releve were stored and the southwestern comer was
marked with a concrete lintil or an iron peg to enable
further monitoring.
In addition to the regular releves (core dataset), 127
sites were analysed to a lesser extent (these and the core
dataset together are referred to as the extended data-
set). This was done without setting up a formal plot and
only the dominant and easily identifiable species were
recorded. The coordinates were located with a GPS and
stored for mapping. No environmental data were taken
directly for these additional releves, but the parameters
extracted from the DEM were stored.
Nwnerical vegetation analysis
A classification and certain ordinations were applied
to the sampled dataset. The classification was performed
using TWINSPAN (Hill 1994). For the classification of
the core dataset, the pseudospecies cut levels were set to
0, 5, 10, 20 and 50, for the extended dataset to 0, 2, 5,
10, 20, 50. The first pseudospecies of the extended data-
set was excluded from the classification. Splitting was
allowed down to two species (default: five) for both data-
sets. An indirect ordination (Detrended Correspondence
Analysis, DCA) was performed for the extended dataset.
A direct ordination (Canonical Correspondence Analysis,
CCA) was applied to the core dataset (due to the avail-
ability of environmental data). Both ordinations were run
with the whole dataset and then repeated only including
releves classified as Fynbos. The default parameters sug-
gested by the programs (DECORANA for the DCA and
MVSP for the CCA) were applied to the analyses.
Vegetation mapping
A set of colour orthophotos (spatial resolution: 0.75 m)
as well as a GIS dataset representing preliminary vegeta-
tion units mapped in 1997 were available. Attempts to
extract vegetation units from the imagery by numerical
means failed as the different fynbos types showed very
similar reflection properties, while the reflection proper-
ties varied very much within each type. Consequently,
GNR was explored during several excursions and
assessed according to the subjective impression of the
authors, supported by the releve information. In addition,
a large number of localities were recorded by GPS in
order to locate the transitions between vegetation units.
All the point data were transfered to a GIS and mapped
onto the composite orthophoto. The information provided
by the orthophotos was combined with the point data,
the results of the numerical vegetation analysis and the
existing GIS dataset to generate a comprehensive map of
meaningful vegetation units.
Bothalia 38,1 (2008)
91
RESULTS
Numerical vegetation analysis
The TWINSPAN classification clearly supported the
presence of two major vegetation types on GNR: one
fynbos (Cape Fynbos Shrublands) and the other non-
fynbos (Forest & Thicket). The wetland releves were not
clearly assigned to any of these groups, and as a result
were considered as an independent vegetation type. The
Fynbos grouping was further split into an Acid Sand
Fynbos Complex and Alkaline Sand Fynbos Complex,
the Forest & Thicket grouping in Milkwood Scrub
Forest, Affomontane Forest and Thicket. No further sig-
nificant splitting was possible in the wetland grouping.
The level of vegetation complexes was the maximum
of detail supported by the classification. However, for
the fynbos, some smaller units were clearly recognizable
in the field, mostly dominated by a few or even only one
species and with the bulk of the species shared among all
the units of the corresponding complex. These detailed
units were conceptualized as formations, which are sum-
marized in Table 1 .
The ordination results corresponded well to the clas-
sification results. In the DCA for the extended data-
set, Afromontane Forest, Milkwood Scrub Forest and
Fynbos were clearly separated along the first axis
(eigenvalue = 0.91), with the Thicket and the Wetland
releves intermediate between Milkwood Scrub Forest
and Fynbos (Figure 2A). The two fynbos complexes
were discriminated along the second axis (eigenvalue =
0.65). The DCA only including the releves classified as
fynbos (Figure 2B; eigenvalues: 0.68 for the first axis
and 0.62 for the second axis) showed a clear cluster-
ing of some formations established in the classification
( Thamnochortus fraternus Restioid Fynbos, Erica ses-
siliflora Ericaceous Fynbos) and the post-fire releves of
Protea repens Proteoid Fynbos. The transitions between
the remaining formations appeared to be highly continu-
ous.
The CCA (Figure 3A) illustrated the clear and discrete
discrimination of Fynbos and Forest depending on veg-
etation age, which was almost coincident with the first
axis (for eigenvalues and canonical coefficients com-
pare Table 2). The forest complexes were discriminated
by elevation, slope, aspect and certain soil characteris-
tics. The fynbos releves were aligned continuously along
the second axis, discriminated by gradients of elevation,
slope, pH, soil depth and rock cover. The Alkaline Sand
Fynbos Complex and the Acid Sand Fynbos Complex
were clearly recognizable, but closely together. The
exclusion of all non-fynbos releves from the CCA
(Figure 3B) did not lead to a clearer distinction of these
clusters. No clear structure was recognizable within the
two subclusters either. Only the Thamnochortus frater-
nus Restioid Fynbos and the Protea repens Proteoid
Fynbos formed ± proper clusters. A number of environ-
mental variables showed similar explanatory value. In
addition to the site scores, the scores of selected species
were plotted (Figure 4). The species were grouped in the
same manner as the releves they dominate. The red data
species appear to be associated with rather extreme con-
ditions as they occupy the edges of the plot.
Vegetation units
Affomontane Forest
Five patches of Afromontane Forest are present at the
GNR, all of them situated in the forest valley (Figure
5A). The CCA suggested an association with shal-
low, neutral to slightly acid soils rich in K and Mg. The
relief is characterized by steep slopes protected from
the southeasterly winds. Affomontane Forests form tall
canopies with heights of more than 10 m. The canopy is
frequently dominated by Rapanea melanophloeos. The
well-developed subcanopy is 3-10 m high, compris-
ing Celtis africana , Chionanthus foveolatus, Diospyros
whvteana, Kiggelaria africana and Sideroxylon inerme.
Most of these species have the potential to grow to can-
opy height. Due to the dark interior of the forest, the
ground layer is usually sparse with Asplenium adian-
tum-nigrum, Droguetia iners and Ehrharta erecta as the
most common species. Climbers are present ( Asparagus
aethiopicus, A. scandens and Cynanchum obtusifolium).
The species diversity is low at the 50 m2 level, averaging
13.2 identifiable species per releve.
The range of temperate forest ecosystems referred
to as Affomontane Forest includes the mountains of the
southern Cape but stretches far into tropical Affica where
they occur at higher altitudes (Midgley et al. 1997). The
Afromontane Forests of the Agulhas Plain are ecologi-
cally similar but floristically distinct from those consid-
ered by Campbell (1985). Instead they show affinities to
the dune forests of the Tongaland-Pondoland Forest (Moll
& White 1978). These forests, and also the Afromontane
Forest along the south coast (Knysna, Tsitsikamma), have
significantly higher species diversities than the forests
on the GNR. They host taxa not present on GNR, such
as Podocarpaceae and Cyatheaceae. One reason for the
declining diversity towards the southwest may be that
more and more species disappear as the climatic condi-
tions become harsher and the forest patches smaller.
Milkwood Scrub Forest
Four patches of Milkwood Scrub Forest are present
at low elevations, associated with deep, slightly acid to
alkaline, sandy, colluvial soils with high contents of Ca
and P. In contrast to the Afromontane Forest, the CCA
did not indicate a negative spatial coincidence with the
southeasterly winds. The high levels of mineral com-
ponents appear to be a distinctive feature of this type of
ecosystem. Ca in particular is considerably richer in the
forest than anywhere else on the GNR. In addition, the
soils are very fertile due to plant-induced organic enrich-
ment (Thwaites & Cowling 1988), which makes them
suitable for agriculture and thus susceptible to anthropo-
genic disturbance.
A single tree layer, usually dominated by Sideroxylon
inerme, attains a height of 6 m and a very dense cover.
Euclea racemosa does form part of the canopy in some
places, whereas Chionanthus foveolatus, Gymnosporia
buxifolia and the winter-deciduous Celtis africana occur
as single trees or small groups. The soil is covered by a
50-300 mm high, sparse to dense (where sufficient light
is available) herb layer dominated by Droguetia iners
and Ehrharta erecta. The only shrub species in the full
92
Bothalia 38,1 (2008)
TABLE 1. — Summary of vegetation units recognized in this study. Average species numbers of complexes on 50 m 2 level in brackets
Vegetation unit
Diagnostic spp.
Common species
Average value
a b c
cospermum patersonii, Protea obtu- Erica irregularis, Ischyrolepis eleocharis, Thamnochortus
sifolia, P. repens , Thamnochortus erectus, Otholobium bracteolatum, Anthospermum aethiopi-
cum
fratemus', < 3: Erica coccinea (yel-
low-flowered variant)
Neutral Sand Proteoid Fynbos > 1 : Leucadendron coniferum or
Leucospermum patersonii ; < 3:
Erica coccinea (yellow-flowered
variant); < 2: Protea obtusifolia, P.
repens , Thamnochortus fratemus
> 1 : Protea repens; P. obtusifolia
< P. repens
> 1 : Protea obtusifolia ; < 2: Tham-
nochortus fratemus ; P. obtusifolia
= > Erica coccinea (yellow-flow-
ered variant)
> 2: Erica coccinea (yellow-flow-
ered variant); < 2: Thamnochortus
fratemus ; E. coccinea (yellow-
flowered variant) < Protea obtusi-
folia, Leucadendron conifemm
and Leucospermum patersonii
Thamnochortus fratemus > 1 : Thamnochortus fratemus
Restioid Fynbos
Protea repens Proteoid Fyn-
bos
Protea obtusifolia Proteoid
Fynbos
Erica coccinea Ericaceous
Fynbos
Leucadendron coniferum, Leucospermum patersonii
disticha, Diosma subulata var. subulata, Passerina vulgaris
Protea obtusifolia. Erica coccinea (yellow-flowered variant),
Leucadendron coniferum, Leucospermum patersonii
Erica coccinea (yellow-flowered variant), Erica irregularis,
Cullumia squamosa, Oedera capensis, Indigofera brachys-
tachya
Thamnochortus fratemus, Erica coccinea (yellow-flow-
ered variant), Protea obtusifolia, Leucadendron conifemm,
Indigofera brachystachya
Acid Sand Fynbos Complex (35.0)
TWINSPAN indicators: Mimetes cucullatus, Leucadendron salignum. Erica glabella
Acid Sand Proteoid Fynbos < 2: Leucadendron coniferum ,
Transitional Proteoid Fynbos
Erica sessiliflora Ericaceous
Fynbos
Elegia thyrsifera Restioid
Fynbos
Mimetes cucullatus, Leucadendron salignum, Aulax umbel-
Leucospermum patersonii , Protea lata. Protea cynaroides, L. xanthoconus, Protea longifolia,
obtusifolia ; < 3: Erica sessiliflora , Penaea mucronata, Morelia quercifolia, Indigofera brachy-
Elegia thyrsifera stachya, Elegia juncea, Pseudopentameris macrantha,
Bobartia indica, Erica glabella, Trichocephalus stipularis
Leucadendron coniferum, Leucospermum patersonii, Protea
obtusifolia. Mimetes cucullatus, Leucadendron tinctum, L.
salignum, Erica glabella
> 1 : Leucadendron coniferum or
Leucospermum patersonii'. Erica
sessiliflora, Elegia thyrsifera < L.
coniferum , L. patersonii
> 2: Erica sessiliflora ; E. thyr-
sifera, Leucadendron coniferum,
Leucospermum patersonii < = E.
sessiliflora
> 2: Elegia thyrsifera', E. thyrsif-
era = > Leucadendron coniferum,
Leucospermum patersonii; E.
thyrsifera > Erica sessiliflora
Erica sessiliflora. E. glabella, Leucadendron coniferum,
Mimetes cucullatus, Leucospermum patersonii, Drosera
capensis, Cliffortia ferruginea, Psoralea arborea, Berzelia
lanuginosa, Staavia radiata
Elegia thyrsifera, Helichiysum patulum, Thamnochortus
erectus, Morelia quercifolia, Leucospermum patersonii,
Leucadendron coniferum. Erica sessiliflora
335 59 6.5
376 40 5.6
297 55 5.6
ND ND ND
a, elevation (m); b, soil depth (cm); c, pH; ND, no data.
Numbers close to diagnostic species: pseudospecies levels of TWINSPAN-classification for extended dataset.
Bothalia 38,1 (2008)
93
O Afromontane Forest
^ Milkwood Scrub Forest
(■) Thicket
A Wetland
# Alkaline Sand Fynbos Complex
O Acid Sand Fynbos Complex
A i w+ <&o o
I { t
# Dune Asteraceous Fynbos
© Dune Asteraceous Fynbos (disturbed)
# Neutral Sand Proteoid Fynbos
# Protea repens Proteoid Fynbos
(?) Protea repens Proteoid Fynbos (post-fire)
A Protea obtusifolia Proteoid Fynbos
A Erica coccinea Ericaceous Fynbos
A Thamnochortus fraternus Restioid Fynbos
A; j j
A ; ! |
<•> Acid Sand Fynbos (post-fire)
O Acid Sand Proteoid Fynbos
A Erica sessiliflora Ericaceous Fynbos
O Elegia thyrsifera Restioid Fynbos
3 Transitional Proteoid Fynbos
FIGURE 2. — A, Detrended Corre-
spondence Analysis (DCA)
for complete extended dataset
resulted in clusters of Afro-
montane Forest, Milkwood
Scrub Forest, Alkaline Sand
Fynbos Complex and Acid
Sand Fynbos Complex; B,
DCA for fynbos releves of
extended dataset resulted in
discrimination of two fynbos
complexes.
shade of the forest is Myrsine afiricana, which may grow
higher than 1 m. The abundance of lianas ( Asparagus
aethiopicus, Cynanchum obtusifolium) and epiphytes
(mainly cryptogams) is considerable. Dead vertical
branches of the spiny A. aethiopicus are particularly
responsible for parts of the forest having a thicket-like
character (Figure 5B). The species diversity is low at the
50 m2 level, averaging only 1 1 identifiable species.
Thicket
The fynbos-forest boundary is rarely sharply defined,
but is often made up of a thicket of varying height and
density. This vegetation unit does not necessarily corre-
spond to the Subtropical Thicket described by Midgley
et al. (1997), although it may share certain characteris-
tics with it. Thickets are characterized by a mixture of
forest and fynbos elements. They occur at sites where
94
Bothalia 38,1 (2008)
%\
• I*
• Dune Asteraceous Fynbos
© Dune Asteraceous Fynbos
(disturbed)
■ Neutral Sand Proteoid Fynbos
# Protea repens Proteoid Fynbos
<•? Protea repens Proteoid Fynbos
(post-fire)
A Protea obtusifolia Proteoid Fynbos
A Erica coccinea Ericaceous Fynbos
A Thamnochortus fraternus
Restioid Fynbos
© Acid Sand Fynbos (post-fire)
O Acid Sand Proteoid Fynbos
A Erica sessiliflora Ericaceous Fynbos
9 Transitional Proteoid Fynbos
, q»i
<# i n jf|
*AAi
i A [
FIGURE 3. — A, results of Canonical Correspondence Analysis (CCA)
for entire core dataset indicate clear separation between Fynbos
complexes, Afromontane Forest and Milkwood Scrub Forest,
whereas Wetland releves are intermixed; B, results of CCA
for fynbos releves of core dataset show continuous transition
between formations.
fire frequency or intensity have been reduced over a cer-
tain period, but not sufficiently to support forest: in the
buffer zones between fynbos and the Milkwood Scrub
Forest, in protected ravines as a successional stage from
fynbos to Afromontane Forest and on some south-fac-
ing slopes of the forest valley. The latter (adjacent to
the Afromontane Forest) is dominated by the fern P ter is
dentata, which attains a very high cover in these places.
These ecosystems are therefore considered as separate
formations ( Pteris dentata Shrubland, compare Figure
8). Forest edge thickets and valley thickets, in contrast,
are floristically similar so that a separation into two for-
mations is not supported. The broad-leaved subtropical
shrub species (Cowling et al. 1997), mainly Euclea race-
mosa, Olea capensis, O. exasperata and various species
of Searsia (Moffett 2007) frequently dominate thickets
on GNR together with Salvia africana-lutea. Among the
fynbos elements, large individuals of Leucadendron con-
iferum and Thamnochortus erectus are most common.
Certain forest elements, most commonly Sideroxylon
inerme, may join the thicket but some thicket species
may also occur as trees in the Milkwood Scrub Forest.
Wetlands
The catchment areas supplying the GNR are not large
enough to support permanent streams under the prevail-
ing precipitation regime. In winter, some springs can
develop and small rivers may persist until the end of
October. As a result, the distribution of true wetlands is
extremely limited at the GNR and is confined to a few
suitable habitats. Nevertheless the diversity of differ-
ent wetland habitats is considerable and it is difficult to
point out one type of wetland characteristic of the GNR.
As the different types are located adjacently along sev-
eral environmental gradients, they will be treated as one
entity. Wetlands are related to soils with a high content
of organic matter, indicated by the dark colour. The
ordinations placed the wetland releves in between for-
TABLE 2. — Eigenvalues and canonical coefficients of Canonical
Correspondence Analyses (CCA) for complete core dataset and
fynbos releves of core dataset
Complete core dataset Core dataset — fynbos
axis 1 axis 2 axis 1 axis 2
Bothalia 38,1 (2008)
95
FIGURE 4. — Canonical Correspondence Analysis (CCA) biplot score
for some selected species. Aspa glo, Aspalathus globulosa, Eric
eye. Erica coccinea (yellow-flowered variant); Eric gla, Erica
glabella ; Eric irr, Erica irregularis ; Eric mag. Erica magnisyl-
vae; Eucl rac, Euclea racemosa', Glad deb. Gladiolus debilis',
Leuc con, Leucadendron coniferum', Leuc pat, Leucospermum
patersonii ; Leuc tin, Leucadendron tinctum; Eric ses. Erica
sessiliflora; Meta mur, Metalasia muricata; Mura sat, Muraltia
satureioides; Oxal sp., Oxalis sp.; Pass vul, Passerina vulgaris',
Pela sub, Pelargonium suburbanum; Pent sp., Pentaschistis
sp.; Prot cyn, Protea cynaroides; Prot obt. Protea obtusifolia',
Prot rep. Protea repens', Saty car, Satyrium cameum; Tham ere,
Thamnochortus erectus', Tham fra, Thamnochortus fratemus.
est and fynbos without clear relationships to any of the
measured variables. The pH ranges from 6. 1-7.9 but the
releve with the highest value is situated below a slope
that may have an alkaline character. The largest patch
is located at the bottom of the forest valley. South-fac-
ing slopes are covered in a dense, 1.5 m high canopy
of Pteris dentata, (recognized as formation Pteris den-
tata Femland), whereas slightly north-facing slopes
support mats of Cliffortia ferruginea and dispersed
stands of Artemisia afra, Leonotis leonurus and some
thicket species. Gunner a perpensa and Senecio halimi-
folius are dominant directly in the temporary stream.
Species such as Mariscus thunbergii, Hippia frutescens
and Zantedeschia aethiopica are present in all types.
Transitions to Thicket are common. One characteristic
feature of the wetlands is their low alpha-diversity with
an average of only 9.3 species per releve.
Alkaline Sand Fynbos Complex
The Alkaline Sand Fynbos Complex covers most of
the lower parts of the GNR. Two hundred species were
recorded in 42 releves, averaging at 24.9 species per
releve. The substrate is mainly wind-blown, colluvial
sand of varying depth, pH and nutrient levels. Dune
Asteraceous Fynbos, Neutral Sand Proteoid Fynbos
and Protea repens Proteoid Fynbos are associated with
these habitats. Some limestone outcrops show a dif-
ferent vegetation but they are too small to house many
of the rare limestone endemics found a few kilometres
to the east on larger patches. Three different limestone
formations, which are frequently intermixed, have been
recorded: Protea obtusifolia Proteoid Fynbos, Erica coc-
cinea Ericaceous Fynbos and Thamnochortus fratemus
Restioid Fynbos.
Dune Asteraceous Fynbos covers the entire western
part of the GNR. It forms an extensive matrix of low eri-
coid/restioid fynbos, higher ericoid fynbos and thicket
dominated by broad-leafed shrubs (Figure 6 A). The for-
mation is characterized by low elevation, alkaline, deep
soils (> 100 cm) and low rock cover. It corresponds well
with the Dune Asteraceous Fynbos described by Cowling
et al. (1988) for the Agulhas Plain and with the Eastern
Type of Coastal Fynbos (Kruger 1979). Large ericoids
( Metalasia muricata and Passerina vulgaris ) and non-
proteoid broad-leafed shrubs ( Chrysanthemoides monili-
fera) are the structural dominants, in some places joined
or replaced by fabaceous shrubs, Otholobium bracteola-
tum, Aspalathus forbesii. Erica irregularis and certain
non-ericaceous ericoids ( Anthospermum aethiopicum,
Phylica ericoides) are widespread. Thamnochortus
erectus is the most conspicuous restioid but the lower
Ischyrolepis e/eocharis has a very high cover in some
sites. Lower individuals of the subtropical shrub species
are very common all over the Dune Asteraceous Fynbos
( Euclea racemosa, Olea capensis subsp. capensis, O.
exasperata and Searsia laevigata). One characteristic
feature of the Dune Asteraceous Fynbos is the lack of
proteoids. If they occur, especially Leucadendron coni-
ferum and Protea obtusifolia, they indicate a transition
to other formations.
Neutral Sand Proteoid Fynbos is associated with
intermediate, but varying environmental variables. The
pH ranges from 4. 8-7. 8. The soils are usually shallower
than those supporting Dune Asteraceous Fynbos, but the
depths range from 0 to > 100 cm. Wind-blown alkaline
sands lying over the acidic Table Mountain Sandstone
provide intermediate habitats between the Dune
Asteraceous Fynbos and the Acid Sand Fynbos Complex.
The formation shows some association with the Protea
susannae-Leucadendron coniferum Proteoid Fynbos
described by Cowling et al. (1988) and Richards et al.
(1995), and the Neutral Sand Proteoid Fynbos described
by Mustart et al. (2003). These units prefer rather deep,
colluvial neutral sands at least partly derived from
Bredasdorp Limestone. Protea susannae, however, does
not occur on the GNR. Instead, Leucadendron coniferum
and Leucospermum patersonii are dominant with shift-
ing composition. Relatively old, almost monospecific
stands of L. coniferum resemble low forests with canopy
heights of up to 5 m (Figure 6B), but with an extremely
high density of thin stems, making them almost impen-
etrable. In young stands (up to 10 years after fire), the
two proteoids are approximately equally abundant. An
ericoid layer with variable density is present. As in the
96
Bothalia 38,1 (2008)
FIGURE 5. — A, patches of Afro-
montane Forest concentra-
ted in south-facing ravines
(left and right); Pteris den-
tata Shrubland in centre; B,
interior of Milkwood Scrub
Forest. Dense undergrowth of
Droguetia iners can only deve-
lop in light places, e.g. after
anthropogenic disturbance.
Photographs: M. Mergili.
Dune Asteraceous Fynbos, Thamnochortus erectus is the
most conspicious restioid element, whereas low restioids
occur in the ground layer, together with various species
of the sedge Ficinia. North- and east-exposed slopes of
the forest valley are covered in a different vegetation
unit. Owing to the high frequency of Leucadendron coni-
ferutn and Leucospermum patersonii and the scarcity of
species indicating a different unit, it has been assigned to
the Neutral Sand Proteoid Fynbos, but as a different unit
in Figure 7.
Protea repens Proteoid Fynbos occupies the lower
slopes of the Swartkransberg heading northwards towards
the broad Steynsbos Valley, as well as large parts of the
Steynsbos Valley itself. It forms 2.5 m high, medium-
dense to dense stands of Proteoid Fynbos, associated
with extraordinarily high values for electrical resistance
of the deep (> 1 m), slightly acid soils. The pH ranges
from 6. 4-6. 7 and at the transition to the Acid Sand
Fynbos Complex it is 5.6. It is inappropriate to com-
pare this formation to the Protea repens Proteoid Fynbos
described by Cowling et al. (1988) for the Agulhas Plain
as it differs structurally and floristically. The formation
is clearly dominated by Protea repens , but it shares most
of its species with the Dune Asteraceous Fynbos and the
Neutral Sand Proteoid Fynbos. Leucadendron coniferum
is frequently intermixed. The ericoids Cliffortia ilicifo-
lia and Passerina vulgaris are very frequent, growing up
to more than 1 .5 m, as does the restioid Thamnochortus
erectus. A sparse to medium-dense undergrowth attains
heights of some tens of centimetres. Parts of the forma-
tion were in an early post-fire stage at the time of the
survey, showing a high cover of Aspalathus microphylla
and lacking visible individuals of Protea repens.
Bothalia 38,1 (2008)
97
FIGURE 6. — A, Dune Asteraceous
Fynbos: in foreground Olho-
lobium bracteolatum, the large
shrub at left, Metalasia muri-
cata. Note thicket element
Olea capensis subsp. capensis
in centre; B, fourteen-year-old
stand of Leucadendron coni-
femm (Neutral Sand Proteoid
Fynbos). Photographs: M.
Mergili.
Protea obtusifolia Proteoid Fynbos grows on lime-
stone outcrops and is associated with shallow to mod-
erately deep, alkaline soils and with low rock cover.
The pH ranges from 6.9-8. 1. The formation is charac-
terized by 2 m high, sparse to medium-dense stands of
P. obtusifolia often intermixed with Leucadendron coni-
ferum and sometimes with Leucospermum patersonii.
The bulk of the species occurs throughout the Alkaline
Sand Fynbos. The formation is frequently intermixed
with other formations of the Alkaline and even the Acid
Sand Fynbos, sometimes on a very fine scale. Despite
floristic differences, it shows affinities to the Protea
obtusifolia-Leucadendron meridianum Proteoid Fynbos
of the Agulhas Plain (Cowling et al. 1988) and to the
Leucadendron meridianum-Protea obtusifolia Proteoid
Fynbos of the Soetanysberg (Richards et al. 1995).
Erica coccinea Ericaceous Fynbos is associated with
alkaline soils rich in Na and Ca. It is not related to the
Ericaceous Fynbos of Campbell (1985) and Cowling
(1988) and has its centre of distribution on rather steep
limestone outcrops. It is characterized by a high cover
of E. coccinea (yellow-flowered variant), which is
frequently joined by Indigofera brachystachva and
Cullumia squarrosa. Transitions to other formations of
the Alkaline Sand Fynbos are common.
Thamnochortus fraternus Restioid Fynbos (Figure
8A) grows on steep limestone slopes supporting shal-
low, rocky, Mg-rich soils with low resistance and a pH
between 7.6 and 8.0. It is structurally dominated by
< 1 m high stands of the limestone endemic T. frater-
nus. The usually sparse ground cover is made up of
shrubs such as Indigofera brachystachya or Cullumia
squarrosa, and also Erica coccinea (yellow-flowered
variant), the latter indicating a transition to the E. coc-
cinea Ericaceous Fynbos. Protea obtusifolia and small
individuals of Leucadendron coniferum may appear
98
Bothalia 38,1 (2008)
■H Afromontane Forest
SHI Milkwood Scrub Forest
iSH Thicket
S8?5 Wetland
Acid Sand Fynbos Complex
3 post-fire regenerating
Acid Sand Proteoid Fynbos
[ I undifferentiated
IlSil with Leucadendron xanthoconus
with Aulax umbellata
££§ Ebca sessiliflora Ericaceous Fynbos
Transitional Proteoid Fynbos
SS undifferentiated
OS) vrith Leucospenmum patersonii
with Olea capensis
Elegla thyrsfera Restloid Fynbos
ITU with Thamnochortus erectus
with Erica sessiliflora
Alkaline Sand Fynbos Complex
f~~ ■ post-fire regenerating
Dune Asteraceous Fynbos
Neutral Sand Proteoid Fynbos
undifferentiated
filfi dry
13 Erica coccwea Ericaceous Fynbos
Ml Thamnochortus fratemus Restioid Fynbos
Qn'j Protea obtusifotia Proteosd Fynbos
ilSHl Protea repens Proteoid Fynbos
0 500 1 000 2.000 m
1 I l I I l I I I
as well. The formation is associated to Kruger’s (1979)
limestone fynbos.
Acid Sand Fynbos Complex
The Acid Sand Fynbos Complex is indicated by the
presence of one or more Proteaceae species associated
with shallow, acidic soils derived from Table Mountain
Sandstone ( Mimetes cuccullatus and Leucadendron
salignum are the most frequent). The higher hills of the
GNR are entirely covered in this complex (Figure 8B).
Acid Sand Fynbos has the highest species diversity
among the vegetation complexes of Grootbos, averaging
at 35 species per releve with a maximum of 48 species.
One hundred and forty-nine species were recorded in the
12 releves of the core dataset altogether. Four formations
of Acid Sand Fynbos were separated in this study. Part
of the complex was in an early post-fire stage at the time
of the survey and was dominated by Aspalathus ciliaris,
Pseudopentameris macrantha, Thesium strictum and
Othonna quinquedentata .
Acid Sand Proteoid Fynbos grows predominantly
on hilltops and steeper slopes, with its largest patch on
the Swartkransberg. It is associated with high eleva-
tion, shallow, rocky and acidic (pH 5. 2-5. 6) soils with
low levels of all nutrients except K, and north-facing
slopes. The formation shows environmental and struc-
tural affinities to both the Leucadendron xanthoconus-
Leucospermum cordifolium Ericaceous Fynbos and the
FIGURE 7. — Vegetation map of
Grootbos Nature Reserve,
based on orthophotos and
field studies.
Aulax umbellata-Protea compacta Proteoid Fynbos of
the Soetanysberg (Richards et al. 1995). It is floristically
and structurally characterized by the absence of species
that have their centre of distribution in the Alkaline Sand
Fynbos Complex and by a low to medium-dense proteoid
layer usually less than 1.5 m tall, including several aci-
dophilous Proteaceae with a changing composition and
without clear dominance of one species. Mimetes cucul-
latus and Leucadendron salignum are the most common,
joined by Aulax umbellata , Leucadendron tinctum , L.
spissifolium , L. xanthoconus , Protea acaulos, P. cynar-
oides, P. longifolia and P. speciosa. Several smaller spe-
cies of Erica and further ericoid shrubs form a sparse to
medium, low-height ground layer. The restioid compo-
nent is variable but Elegia juncea in particular is abun-
dant in some places, as is the large ‘graminoid’ Iridaceae
Bobartia indica. Some of the non-sprouting Proteaceae,
in particular Aulax umbellata and Leucadendron xan-
thoconus', may gain dominance in some places as well.
According to Campbell (1986) and Cowling el al. (1988),
part of the Acid Sand Proteoid Fynbos distinguished in
this study should instead be recognized as Asteraceous
Fynbos as those authors only recognize a community as
Proteoid Fynbos if it contains more than 10% cover of
reseeding proteoids. The resprouting Mimetes cucullatus
and Leucadendron salignum , in contrast, do not fulfil this
requirement.
Erica sessiliflora Ericaceous Fynbos grows on damp,
south-facing slopes and in valleys. The Acid Sand
Bothalia 38.1 (2008)
99
FIGURE 8. — A, Thamnochortus
fraternus Restioid Fynbos on
limestone ridge; B, Acid Sand
Fynbos on southwestern slope
of Swartkransberg; Acid Sand
Proteoid Fynbos with Mimetes
cucullatus and /In/ax umbella-
ta in foreground; Transitional
Proteoid Fynbos with high
cover of Leucadendron coni-
ferum in background. Photo-
graphs: M. Mergili.
Fynbos elements are joined by several moisture indica-
tors, such as Berzelia lanuginosa, Cliffortia ferruginea ,
Drosera capensis and Psoralea arborea. Structurally
dominant is Erica sessiliflora, which can be up to 1.5
m high and can attain cover values of more than 75%.
Mimetes cucullatus, Leucadendron coniferum and E.
glabella are common. The largest patch occurs on the
southern slope of a mountain on the Steynsbos prop-
erty, but extended canopies of E. sessiliflora also occur
on the northern slope of the Swartkransberg, indicating
that the species can also cope with less moist conditions.
Transitions to other fynbos formations are manifold.
The formation corresponds well to the Wet Ericaceous
Fynbos described by Campbell (1986) and Cowling et
al. (1988).
Elegia thyrsifera Restioid Fynbos occupies three
patches of fynbos on GNR. It is structurally domi-
nated by Elegia thyrsifera and Thamnochortus erec-
tus which can both exceed a height of 2 m. Proteoids
( Leucadendron coniferum, Leucospermum patersonii
and partly Mimetes cucullatus) are abundant but not
dominant. Helichrysum patulum and Morelia quercifolia
show high cover values in some places. The formation
is associated with north-facing slopes and corresponds to
Campbell’s (1986) Restioid Fynbos although it is diffi-
cult to be assigned to one of the subseries. The sites on
the GNR may represent a gradient from Mesic Restioid
Fynbos with a higher share of ericaceous Ericaceae
(mainly E. sessiliflora ) to Dry Restioid Fynbos. Floris-
tically it does not correspond to the Dry Restioid Fynbos
of the Agulhas Plain (Cowling et al. 1988).
Transitional Proteoid Fynbos constitutes a mixture of
species centred in the Alkaline Sand Fynbos, and spe-
cies centred in the Acid Sand Fynbos. The transitional
100
Bothalia 38,1 (2008)
character is well indicated by the CCA. The pH, how-
ever, does not exceed 6. Various subtypes of this forma-
tion are present on GNR. Some patches are structurally
similar to the Neutral Sand Fynbos but acidophilous
proteoids, usually Mimetes cucullatus, indicate the more
acidic conditions. Damp ravines on the southern slope
of the Swartkransberg support a thicket-like vegetation
with a high cover of Olea capensis, but still with fynbos
character. Another transition zone exists between Protea
obtusifolia Proteoid Fynbos and Acid Sand Fynbos,
leading to very complex situations with P. obtusifolia
and A u lax umbel lata growing almost together, but rep-
resenting completely different soil nutrient regimes. The
major difference to the transition described above is that
it is not based on a gradual decrease of soil depth and
pH-value, but on a fine-grained mosaic of young wind-
blown, shallow, calcareous soils, older limestone ridges
and underlying Table Mountain Sandstone. Extremely
complex situations prevail at the western slope of a
mountain at Steynsbos, where Leucadendron coniferum,
Leucospermum patersonii and P. obtusifolia coexist with
some of the acid sand proteoids.
Vegetation mapping
The vegetation units illustrated in the map (Figure 7)
largely correspond to the units established in the analy-
sis. Only a few changes were made (e.g. patches of fyn-
bos in early post-fire stages). An additional level of detail
was introduced in some cases. Most of GNR is cov-
ered in Fynbos (1 620 ha or 95.3%). Forest & Thicket
account for 78 ha or 4.6% (Afromontane Forest 4.1 ha
or 0.2%, Milkwood Scrub Forest for 43 ha or 2.5% and
Thicket for 30.9 ha or 1 .8%), whereas only 1 .4 ha (0.1%)
are covered in Wetland. Dune Asteraceous Fynbos cov-
ers more than half of GNR (914 ha or 53.8%), and the
whole Alkaline Sand Fynbos Complex occupies 1 385
ha or 81.5%. Even the second largest formation of this
complex, the Neutral Sand Proteoid Fynbos (191 ha or
11.3%), approaches the same amount of cover as the
whole Acid Sand Fynbos Complex (235 ha or 13.8%).
Figure 7 also gives an idea of the fragmentation of the
vegetation units. The interpretation of this information
has to be approached with caution because the fragmen-
tation may be caused by different factors, including the
shape of GNR. Dune Asteraceous Fynbos and Neutral
Sand Proteoid Fynbos cover continuous, hardly frag-
mented areas with average patch sizes of 305 and 96 ha,
respectively. Protea repens Proteoid Fynbos and Erica
sessiliflora Ericaceous Fynbos also exceed average patch
sizes of 25 ha. In contrast, the distribution of Limestone
Fynbos is rather patchy, with 27 patches ranging from
tens of hectares of Protea obtusifolia Proteoid Fynbos
to tiny limestone outcrops covered with Thanmochortus
fraternus Restioid Fynbos. The average patch size (6
ha) is not representative in this case. In the Forest the
patch size of the highly fragmented Afromontane Forests
remains below 1 ha (0.81), whereas the patches of
Milkwood Scrub Forest occupy slightly more than 10 ha
on average.
DISCUSSION
Three levels of vegetation units were established on
the GNR: the vegetation type level and the vegetation
complex level were based on environmental conditions
and species groupings (TWINSPAN), and the formation
level on dominant species.
The units of the vegetation type level largely cor-
respond to the biome level of the Broad Habitat Units
established by Cowling & Heijnis (2001), based on
environmental variables. The Fynbos and Forest Biomes
occur on GNR, the Thicket Complex identified in this
study does not correspond to the Thicket Biome of
Cowling & Heijnis. The complexes of this study corre-
spond partly to the primary units. Campbell (1985) con-
sidered Forest and Thicket as one group, as in this study.
Wetlands are neither considered by Campbell (1985)
nor by Cowling & Heijnis (2001). Cowling et al. (1988)
classified them as azonal vegetation.
The separation of the fynbos on GNR into an Alkaline
and an Acid Sand Fynbos Complex seems sufficiently
supported by the classification and the ordinations. The
two complexes correspond to the Coastal Fynbos (or
Lowland Fynbos) and Mountain Fynbos, respectively
(Acocks 1953; Taylor 1978; Kruger 1979; Moll et al.
1984). According to Cowling et al. (1988), a separation
of fynbos in this way has to be rejected because none of
the stuctural units (Campbell 1985) are restricted to one
of those groups, and because the floristic changes within
the Mountain Fynbos are as significant as between
Mountain Fynbos and Lowland Fynbos. However, this
study shows that it appears to be highly relevant on a
local scale, where the gamma-diversity does not play the
role it does on a broader scale while the beta-diversity is
still high.
The division of the complexes into proper commu-
nities presents problems. Most subcomplex vegetation
units are based on one or a maximum of two dominant
species which give the landscape a very characteristic
appearance. Nevertheless, the classic concept of plant
communites as an association of several characteris-
tic species which differentiates it from other commu-
nites should not be applied to these entities. The struc-
tural component in such a classification is evident, and
the groups (here called formations) show a fairly strong
correlation to the classification systems of Campbell
(1985) and Cowling et al. (1988) where structural fea-
tures of the vegetation were included more systemati-
cally. The difficulty of purely floristic classification sys-
tems in the Fynbos Biome, due to high gamma-diversity,
is illustrated by comparing the findings of this study to
the vegetation study of Richards et al. (1995) for the
Soetanysberg. Despite environmentally comparable con-
ditions and a distance of < 50 km, substantial floristic dif-
ferences, also among the dominant species, are evident.
The expected dependence of fynbos vegetation on
certain environmental factors was confirmed by the
study. The major explanatory variables in the CCA of
Richards et al. (1995) in their vegetation study of the
Soetanysberg were pH, rock cover, soil depth and soil
texture. Apart from soil texture, which was investigated
in more detail than in this study, the explanatory vari-
ables are the same as for GNR. Elevation, which plays a
major role here but not in the Soetanysberg, is probably
mainly a surrogate for the aeolian sediment accumula-
Bothalia 38,1 (2008)
101
tion budget, wind speed and the distribution of different
substrate types, as the correlation values (e.g. 0.78 with
pH) indicate.
The clustering of the releves in the DCA and the CCA
was rather poor (compare Figures 2B; 3B) and the tran-
sitions between the fynbos units many. Only the Acid
Sand Proteoid Fynbos formed a clear cluster, together
with some releves of Transitional Proteoid Fynbos. This
finding is partly in line with the study of Richards et al.
(1995): the releves connected to low pH (sandstone)
were poorly clustered in the CCA biplot, but they were
clearly separated from the releves on limestone. On
GNR, the transition between the limestone formations
and the remaining formations was continuous with the
Neutral Sand Proteoid Fynbos as intermediate forma-
tion, but rather with affinities to the limestone forma-
tions. In contrast, the Protea susannae-Leucadendron
coniferum Proteoid Fynbos of Richards et al. (1995), a
formation that is associated with neutral sands according
to Cowling et al. (1988), did not differ substantially in
pH from the sandstone formations, but was clearly dis-
criminated from the limestone formation ( Protea obtusi-
folia-Leucadendron meridianum Proteoid Fynbos).
The poor clustering in the fynbos of the GNR may
be explained by a variety of factors. The location of the
releves (some of them were placed in transitional zones)
and the high level of detail in the study (leading to a
considerable amount of noise) may serve as one expla-
nation, the topographic and geological patterns on GNR
as another; alkaline sands cover the entire western part of
the GNR, and limestone ridges of varying shape and size
are widely dispersed. This leads to wind-blown alkaline
sands of variable depth over large parts of acid substrate
and to an extremely fine-scaled pattern of different physi-
cal and chemical substrate properties. This is especially
true for the foothills and the lower slopes of the higher
hills of GNR, leading to very complex vegetation pat-
terns; in some places Aulax umbellata, a strong indica-
tor of acidic conditions, and Protea obtusifolia, a strong
indicator of alkaline conditions, grow immediately adja-
cent to each other. Only the highest parts of the GNR, the
Acid Sand Fynbos Complex, remain untouched and do
not contain calcareophilous species. In such a fine mosaic
of different environmental conditions, the vegetation is
sensitive to influences other than topographic and sub-
strate variables. The problems with grouping fynbos eco-
systems based on reseeding proteoids (Richardson & Van
Wilgen 1992; Privett et al. 2001) are well established,
as these organisms are susceptible to local extinction
(‘drifting clouds of species abundance’). Therefore the
structural dominants of the vegetation of a certain place
may shift from fire interval to fire interval, as Privett et
al. (2001) have shown for the Cape Peninsula. However,
this phenomenon only occurs at a subcomplex level. The
boundaries of the fynbos complexes are not affected as
steep environmental gradients prevent mixing of the spe-
cies pools (high beta-diversity).
Figure 9 represents the floristic diversity of the veg-
etation of GNR at the complex level. Fynbos and Forest
constitute completely different species pools. Less than
ten taxa occur in both vegetation types, whereas Acid
Sand Fynbos and Afromontane Forest share no taxa at
all. In contrast the complexes within each vegetation
FIGURE 9. — Species diversity of vegetation complexes of Grootbos
Nature Reserve. Size of each box and number inside indicate
total species number of complex, whereas width of lines and
adjacent numbers indicate number of species shared by two
complexes. AcSF, Acid Sand Fynbos Complex; A1SF, Alkaline
Sand Fynbos Complex; MSF, Milkwood Scrub Forest; AF,
Afromontane Forest; WL, Wetland.
type share a considerable number of their taxa. Wetland
shares only a few taxa with the other groupings, but
the low diversity of the wetland may contribute to this
phenomenon. Although the Alkaline Sand Fynbos hosts
more taxa (200) than the Acid Sand Fynbos (149), the
diversity of the latter is more than four times higher if
normalized to the area (0.63 and 0.14 taxa per hectare
respectively). Even though this ratio may be of limited
value, it confirms the general patterns obtained from the
releve data.
As a conclusion, it can be stated that the vegetation
patterns of GNR were investigated in great detail, lead-
ing to a differentiated picture of the spatial distribution
of the vegetation units and the vegetation-environment
relationships. The major implications for further man-
agement is the potential impact of fire on vegetation
structure, in particular the vegetation units dominated by
reseeding proteoids. Careful planning to mimic natural
fire frequencies and conditions will be necessary in order
to prevent local extinction. In order to protect Red Data
species, particular care has to be taken when undertaking
controlled bums of extreme habitats, such as steep lime-
stone ridges and sandstone slopes.
SHORT NOTE
In February 2006, after the submission of this paper,
the fynbos ecosystems of the entire GNR were burnt by
a fire. Since then, the number of species recorded has
increased from 660 to 732 (Privett & Lutzeyer in press)
and two further new species ( Capnophyllum sp. nov. and
Pterygodium sp. nov.) were recorded.
ACKNOWLEDGEMENTS
Thanks to the Lutzeyer family and the staff of
Grootbos Nature Reserve for their support during the
102
field work. Special mention of G. Siebrits who under-
took the original vegetation survey on Grootbos in
1997. Grateful thanks also to the Dean’s Office of
the Faculty for Natural Sciences of the University of
Innsbruck and to the Office for International Relations,
University of Innsbruck, for their grants enabling the
field work in spring 2004, as well as to B. Erschbamer
of the Department of Botany, University of Innsbruck
for her valuable comments and to A. Erhard1 (Institute
of Geography, University of Innsbruck), R De Necker
(Department of Geography, University of Stellenbosch)
and S. Smuts (Department of Botany, University of Cape
Town) for their support.
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Bothalia 38,1: 103-110 (2008)
Resource demand estimates for sustainable forest management: Mnga-
zana Mangrove Forest, South Africa
C.H. TRAYNOR* * and T.R. HILL*t
Keywords: Bruguiera gymnorrhiza (L.) Lam., construction timber. Eastern Cape, forest product utilization, mangroves, Rhizophora mucronata
Lam., rural building styles, sustainable forest management
ABSTRACT
Since democratization in 1 994, South African forest policies have promoted sustainable forest management. However,
implementation has been problematic due to limited information concerning forest product utilization. This paper
investigates and quantifies timber use from the Mngazana Mangrove Forest, Eastern Cape Province, South Africa. Three
local communities utilize stems of the mangrove species Rhizophora mucronata Lam. and Bruguiera gymnorrhiza (L.) Lam.
for building construction. There were two distinct building shapes, circular and rectangular. On average, 155 stems were used
for circular buildings and 378 stems for rectangular buildings. Most buildings were constructed using mangroves as well
as indigenous timber from coastal scarp forests. The proportion of mangrove stems in buildings varied from 0 to 95%. The
annual demand for mangroves was estimated to be 1 8 400 stems. Due to the high annual demand, projected human population
growth rates have a minor influence upon future demand values. For effective sustainable forest management, the standing
stock at Mngazana should be restricted to the two mangrove species utilized for building construction, and a forest inventory
performed so that demand for building can be compared to supply.
INTRODUCTION
Over the last decade, global advancements in for-
est management have progressed towards Sustainable
Forest Management (SFM), a term first incorporated
into Agenda 21 and the Forest Principles which were
both outputs of the United Nations Conference on Envi-
ronment and Development (UNCED) 1992. Agenda 21
called for enhanced sustainable management of all for-
ests, whereas the Forest Principles stated that ‘forest
resources and forest lands should be sustainably man-
aged to meet the social, economic, ecological, cultural
and spiritual needs of present and future generations’.
It also specified that these needs include forest prod-
ucts such as wood and wood products (United Nations
General Assembly 1992: 291, 292). Since its inception,
the SFM concept has evolved through international for-
est policy dialogue, as well as country and eco-regional
initiatives. These activities have resulted in global altera-
tions to forest policies and forest management strategies
(FAO 1994).
Although South Africa did not participate in UNCED,
global developments have influenced its policies and
since democratization in 1994, forest policy has striven
to broaden access and sustainable use of indigenous
forests (Lawes et al. 2004a). The new South African
Constitution promotes conservation, ecologically sus-
tainable development and the use of natural resources
within justifiable social and economic develop-
ment (RSA 1996a). Furthermore, the White Paper on
Sustainable Forest Development in South Africa (RSA
1996b), and the National Forest Act (RSA 1998), both
* Discipline of Geography, School of Environmental Sciences,
University of KwaZulu-Natal, Private Bag X01, 3209 Scottsville,
Pietermaritzburg, South Africa.
*t Corresponding author, Tel.: +27 (0) 33 260 5341; Fax.: + 27 (0) 260
5344. E-mail address: hillt@ukzn.ac.za.
MS. received: 2006-03-09.
acknowledge the vital role of forests in the livelihoods of
rural communities. These policies recognize the obliga-
tion which the country has, to use its natural resources to
further the development of the poor (Willis 2004). Thus,
South African policy promotes the sustainable use of
forests. Forest biomes and their management have been
well researched (Muir 1990; Oribi et al. 2002; Lawes
et al. 2004a), but implementation of management plans
sometimes poses a problem. There is also a paucity of
information on the usefulness of the forests, especially
to local people and on actual removal of wood and other
products (Lawes et al. 2004).
SFM approaches have been applied to mangrove
forests (International Tropical Timber Organization
2002), but no examples exist for South Africa’s man-
groves, even though their forest products are utilized
by local communities (Bruton 1980; Ward et al. 1986;
Steinke 1999; Rajkaran et al. 2004). Mangroves in South
Africa are located along the Indian Ocean coastline
in the provinces of KwaZulu-Natal and Eastern Cape.
Over 1 7 years, a reduction of 7% in mangrove area has
been reported for the Eastern Cape, and easily accessi-
ble forests that lie outside of protected areas are under
great pressure from resource users (Adams et al. 2005).
Mangrove ecosystems have a positive influence on
coastal protection, nutrient cycling and export, sediment
trapping, and serve as breeding and nursery grounds for
fish (Lugo & Snaedakar 1974; Hogarth 1999; Mumby et
al. 2004). South African mangrove habitats are threatened
and as a result, many authors have called for greater con-
servation and management efforts (Branch & Grindley
1979; Day 1981; Ward et al. 1986; Beijak et al. 1997;
Steinke et al. 1995).
The Mngazana Mangrove Forest (MMF) in the
Eastern Cape was selected for study, as mangrove trees
have been harvested throughout the forest (Rajkaran et
al. 2004) and local communities use them with other
104
Bothalia 38,1 (2008)
FIGURE 1 . — Location of the Mnga-
zana Mangrove Forest and
study villages. Eastern Cape,
South Africa.
indigenous wood species as building construction mate-
rials. In the villages surrounding the mangroves there are
limited employment and income opportunities. People
have restricted means to purchase building construction
materials and thus over 90% of dwellings are traditional
huts or structures made from natural materials (Statistics
South Africa 2001). Every South African citizen has the
right to access adequate housing (RSA 1996a) and the
communities at Mngazana are dependent upon the local
natural resources, including the mangrove forest, to meet
their housing needs. At Mngazana the buildings are typi-
cal of rural styles constructed from indigenous wood;
a wooden framework is made to which stone and mud
walls and roofing materials are added (Liengmel983;
Cunningham 1985; Shackleton et al. 2002).
This paper investigates the SFM concept focusing
on the estimation of empirical values for forest product
utilization. Through the case of the MMF, the intrica-
cies of precisely calculating demand for housing con-
struction timber is presented and the usefulness of these
estimates to SFM considered. Firstly, the total numbers
of mangrove stems within standing structures is quanti-
fied. Secondly, the current annual demand for mangrove
stems for building construction is evaluated. Thirdly,
future demand for mangrove stems is predicted under
different scenarios. Finally, the contribution the demand
estimates can make to a mangrove SFM strategy and the
wider relevance of the case is discussed.
The context of the Mngazana Mangrove Forest
The topography of the Mngazana area is a tidal flat,
up to 1 500 m wide in places, attributed to the geological
formation, the Ecca and Beaufort Groups of the Karoo
Sequence (Harrison et al. 1999). These groups are com-
posed of conglomerates, sandstones and mudstones
which produce weakly structured, shallow soils when
weathered (Nicolson 1993). The average annual rainfall
is 1 200 mm, and 70% occurs in summer. The vegetation
includes a coastal strip of dune forest, mangroves, salt-
marsh and inland grassland and forest patches.
The MMF is a riverine mangrove forest that is situ-
ated within the Mngazana Estuary (31°42'S 29°25'E),
Eastern Cape. It is the third largest mangrove forest in
South Africa (Ward & Steinke 1982) covering an area of
± 145 ha (Adams et al. 2005) and is composed of the
mangrove species Avicennia marinia (Forssk.) Vierh.,
Bruguiera gymnorrhiza (L.) Lam. and Rhizophora
mucronata Lam. The estuary is biologically important
with the highest recorded invertebrate diversity for east
coast estuaries (Branch & Grindley 1979) and rated first
in the region using a botanical importance rating system
(Colloty 2000) and 22nd in South Africa for its biodiver-
sity importance (Turpie et al. 2002). The MMF has been
heavily disturbed, and Rajkaran et al. (2004), using GIS
analysis, estimated that over 40% of the forest showed a
ratio of one adult tree to two harvested stumps, indicat-
ing that the harvesting intensity was extremely high.
The area is inhabited by Xhosa-speaking Mpondos.
Prior to democracy in 1 994, the area was classified as a
homeland by the apartheid government and consequently
is poorly developed (Ashley & Ntshona 2003). There
are three villages surrounding the Mngazana Estuary;
Mqualeni, Cwebeni and Tekweni (Figure 1). In these vil-
lages, ± 30% of the population have received no school-
ing, less than 1 0% of the labour force are employed and
over half of the homesteads are female-headed (Statistics
South Africa 2001). The land is in trust to the Tribal
Authorities but owned by the state, and community
members may utilize the land subject to legislation and
local rules. The communities at Mngazana and a non-
profit organization hope to devise a SFM plan for the
mangrove forest that will permit mangrove harvesting.
Bothalia 38,1 (2008)
105
METHODS
Investigations focused upon Rhizophora mucronata
and Bruguiera gymnorrhiza as these species are utilized
exclusively for building construction. They are har-
vested by cutting the main stem at its base; this entire
stem minus branches is used in building construction.
It has been suggested that R. mucronata can regenerate
after cutting ‘if cut high enough on the stem that live
branches (with leaves) are spared’ (Walters 2005a: 344),
but coppicing of cut stems has never been observed at
Mngazana for R. mucronata and B. gymnorrhiza (Adams
et al. 2005) and thus, under the current cutting regime,
one stem equates with one tree destructively harvested.
Therefore, demand (and supply) calculations can use
numbers of individual trees rather than weight and vol-
ume measurements of wood.
Mangrove utilization for building construction pur-
poses has been determined through empirical observa-
tions supplemented by information from semi-structured
interviews with homestead occupants and group discus-
sions. As house construction activities only occur every
few years, a respondent’s recollections concerning quan-
tities of materials used may not be very precise, so empiri-
cal observations are generally more reliable (Shackleton
& Shackleton 2004). The villages of Mqualeni, Cwebeni
and Tekweni were selected for study. There are 574
homesteads in these villages (Statistics South Africa
2001) and of these, 108 (19%) were randomly selected
and surveyed, 32 in Cwebeni, 48 in Tekweni and 28 in
Mqualeni. The semi-structured interviews with home-
stead occupants were in Xhosa (the vernacular language
spoken in the area) with English translation. The inter-
views were designed to elicit information regarding
mangrove characteristics, utilization and building lon-
gevity. Within each homestead the number and shape
of individual buildings were recorded. Buildings con-
structed with mangroves were identified and the number
of mangrove poles utilized in each structural unit (wall
poles, roof poles and roof laths) was counted. The diam-
eter of mangrove stems used in each structural unit was
determined by sampling a random ten exposed stems
and measuring the diameter using metal callipers to the
nearest millimetre. The length of stems used for different
units was obtained through group discussions. The pres-
ence of alternative building materials was also recorded.
Three group discussions with community members were
held from October 2003 to October 2004 and relevant
information from these has been included for compara-
tive purposes. Groups were composed of members of the
Mngazana Mangrove Management Forum and interested
villagers from all three villages, between 1 1 to 20 people
attended each meeting.
Human population growth rates were estimated from
the 2001 South African census (Statistics South Africa
2001) and the population estimates of the United Nations
(United Nations 2005). The following terminology will
be used throughout this paper: homestead: an area allo-
cated for a family’s dwellings, usually consisting of sev-
eral different individual buildings; mangrove used as a
prefix to structural unit, building or homestead denotes
use of mangroves in construction. Utilization was cal-
culated at different scales of structural unit, building,
homestead and village using the procedure summarized
in Figure 2.
RESULTS
Mangrove utilization
Group discussions and direct observations indicated
that the main use of mangroves is for building construc-
tion: Rhizophora mucronata and Bruguiera gymnor-
rhiza are the only species utilized for this purpose. An
extremely small minority of homesteads used these spe-
cies for non-building construction purposes such as fenc-
ing. Avicennia marinia was collected by some home-
steads for fuel wood; dead wood was preferred to green
wood. There was a low frequency of collection because
non-mangrove tree species were also utilized for fuel
wood. This study focuses upon construction use because
it is the predominant use of green mangrove wood.
Mangrove stems in standing structures
Buildings constructed from mangroves are circular or
rectangular. Both of these shapes are constructed from
structural units of wall poles, wall laths and roof poles.
Wall poles are positioned vertically and then laths laid at
ninety degrees to these to produce a framework.
The average number of mangrove stems used in
building units constructed entirely from mangroves was
calculated for all three villages (Table 1). A circular
building if constructed entirely from mangroves would
utilize 155 stems, ± 1 10 of these would be wall laths and
the remainder wall or roof poles. Rectangular buildings
composed of only mangrove stems utilize more than
double the number of stems (± 378 stems), because the
wall laths require 335±151 stems. Requirements for
wall and roof poles are approximately similar. The aver-
age diameter of poles was 6.1±2.0 cm (n = 787) and the
average diameter of laths was 3.3±1.0 cm (n = 550).
Group discussions revealed that poles and laths over 2 m
long were utilized.
However, mangroves are utilized in conjunction
with other timbers, the percentage of mangroves within
these structural units shows variation (Figures 3, 4).
Mangroves compose a high percentage of building mate-
FIGURE 2. — Summary of method used to calculate demand for man-
grove stems within each village.
106
Bothalia 38,1 (2008)
TABLE 1 . — Average no. mangrove stems ( ± std dev.) per building unit constructed entirely
from mangroves in three villages studied
rials in the roof poles of circular buildings (up to 100%)
and the wall laths of rectangular buildings (75% or
more). In other structures, mangroves account for 50% or
less of materials. The percentage of mangroves utilized
showed variation between the villages studied. Mqualeni
Village showed the most consistent use of mangroves
and they were utilized in all structural units composing
± 30% to 80% of all stems. In Cwebeni and Tekweni, the
percentage of mangroves used shows wider variations.
All the roof poles in circular buildings in these villages
are mangroves, but mangroves make up 10% or less of
wall poles and laths. Rectangular buildings in these two
villages display different patterns; mangroves make up
75% or more of wall laths but only up to 25% of wall
and roof poles.
A one-way between-groups analysis of variance was
conducted to explore the impact of village location upon
the number of mangrove stems used in different-shaped
buildings. There was a statistically significant differ-
ence at the p = < 0.05 level for the number of mangroves
stems per circular building between villages (F(2, 59) =
10.7, p = 0.00). Post-hoc comparisons using the Turkey
HSD test indicated the mean score for Maqualeni (M
= 72.80, SD = 64.76) was significantly different from
Cwebeni (M = 33.00, SD = 28.70) and Tekweni (M =
FIGURE 3. — Percentage of mangrove stems used within structural units
in circular mangrove buildings.
20.17, SD = 7.50). There was no statistically significant
difference at the p = < 0.05 level in the number of man-
grove stems used in rectangular buildings between vil-
lages (F(2, 36) = 2.32, p = 0.11).
Typical homesteads in the villages studied consist
of circular and rectangular buildings, constructed from
a variety of materials including indigenous timber and
blocks.
Thus a conversion factor was utilized to estimate the
number of mangrove stems utilized per homestead (Table
2). This conversion factor was the average number of
mangrove stems in mangrove buildings multiplied by
the average number of buildings containing mangroves
in homesteads (that utilized mangroves). Accounting for
these variations, mangrove homesteads used an average
of 89 stems in Tekweni, 346 stems in Mqualeni and 397
stems in Cwebeni. The percentage of homesteads utiliz-
ing mangroves was 54% in Tekweni, 79% in Mqualeni
and 88% in Cwebeni.
At the village scale, the total number of homesteads
using mangroves was 67 in Mqualeni, 96 in Tekweni
and 272 in Cwebeni and the estimated total number of
mangrove stems utilized was 8 581 stems in Tekweni,
23 108 stems in Mqualeni, 108 034 stems in Cwebeni.
Thus in total, 139 723 mangrove stems were utilized in
standing structures within buildings at the time of the
study.
Current annual demand for mangrove stems (2004)
The percentage of new buildings compared to the
total number of buildings was 3% in Tekweni, 16% in
Cwebeni and 18% in Maqualeni. As the different-shaped
buildings utilize different amounts of mangrove stems.
TABLE 2. — Conversion of mangrove utilization figures from buildings
to homestead level
C, circular building; R, rectangular building.
Bothalia 38,1 (2008)
107
FIGURE 4. — Percentage of mangrove stems used within structural
units in rectangular mangrove buildings.
FIGURE 5. — Demand for mangroves in 2004 under different building-
shape scenarios.
the shape of buildings constructed has an important
influence upon demand for mangrove stems. The ratio of
circular to rectangular mangrove buildings in the villages
studied was 1:1, thus the current demand for stems can
be calculated using this building shape ratio. Circular
buildings utilize fewer mangrove stems than rectan-
gular buildings, thus demand from circular buildings
represents the minimum number of stems necessary to
fulfil housing requirements and demand from rectangu-
lar buildings represents maximum demand. Using these
demand scenarios, the minimum demand would be for
3 700 stems, the 1:1 demand would be for 18 400 stems
and the maximum demand for 33 500 stems (Figure 5).
Group discussions suggested that the longevity of
mangrove buildings was 10 years and homestead sur-
veys suggested an average of 20 years, thus, the average
is 15 years.
Predicting future demand scenarios
Future demand for mangrove stems can be estimated
using human population growth predictions derived
from past growth rates or demographic models. Past
rates (1996-2001) of annual human population growth
in all three villages was 3% (Statistics South Africa
2001). However, this census data may underestimate
the impact of HIV/AIDS upon the population and as the
most direct demographic consequence of HIV/AIDS is
an increase in mortality (Whiteside 2001) demograph-
ics will change. The United Nations demographic mod-
els take account of the effect of HIV/AIDS and in South
Africa annual human population growth rates were esti-
mated to be 0.78% for the period 2000 to 2005 (United
Nations 2005).
The influence of a decade of human population
growth on mangrove stem demand can be estimated
using the different human population growth forecasts
and various building-shape scenarios (Figure 6). These
estimates suggest that under the 0.78% human popula-
tion growth rate, demand in all building scenarios over
the next ten years will increase gradually but it will
not exceed more than 0.5% of the current demand fig-
ure. Under the 3% human population growth forecast,
demand for mangrove stems increases by 6% of current
estimated values, and in the maximum demand scenario
of rectangular buildings, an additional 2 628 stems per
year would be required in 2015 compared to current val-
ues.
DISCUSSION
The investigations have demonstrated that differ-
ent-shaped buildings require a different total number of
resource stems. In buildings constructed entirely from
FIGURE 6. — The predicted demand for mangrove stems under different
population growth forecasts and building-shape scenarios. -«-3%
growth (rectangular); -^0.78% growth (rectangular);-*-3% growth
(circular and rectangular); — 0.78% growth (circular and rectangu-
lar); -*-3% growth (circular); ^^0.78% growth (circular).
108
Bothalia 38,1 (2008)
mangroves, rectangular buildings require ± 100% or over
200 more stems than circular buildings. Liengme (1983)
found that rectangular buildings used ± 50% more poles
than circular buildings, the main difference attributable
to rectangular buildings having two rather than one row
of roof poles. In the current study, the major variation
between building shapes was in the number of wall laths
required.
Buildings in the current study are rarely constructed
entirely from mangroves and species combinations have
also been documented in other parts of South Africa
(Liengme 1983). The proportion of mangroves used var-
ies with structural unit and building shape. Mangroves
are the preferred materials for circular building roof
poles and rectangular building wall laths. Shackleton et
al. (2004) reported that proportions of species could alter
within walls and roofs. In the current study, mangroves
composed a higher percentage of rectangular than circu-
lar building stems.
The average distance from surveyed homesteads to
the mangrove boundary was 393 m in Mqualeni, 1 733 m
in Tekweni and 2 040 m in Cwebeni. Mangrove use var-
ied spatially between the villages; mangroves are used
for all structural units in both circular and rectangular
buildings in Mqualeni Village, however in Tekweni they
are rarely used as roof poles and never used as circular
building wall laths. Circular buildings have curved walls
and group discussions revealed that indigenous species
from the coastal scarp forests were preferred for this pur-
pose because they were more flexible than mangroves.
In Mqualeni Village, mangroves were often used for cir-
cular wall laths as the mangrove resource was nearby, ±
400 m from homesteads. Similar examples of resource
selection for less preferred species due to higher avail-
ability and ease of collection have been documented by
Nomtshongwana (1999) and Walters (2005b). In the cur-
rent study, the relationship between mangrove use and
distance to the mangroves was confounded by the fact
that mangrove use was also affected by utilization of
alternative building materials such as indigenous timbers
and sand blocks.
Utilization evaluation -at the homestead scale shows a
variety of mangrove and n©Ti-mangrove materials used
for both circular and rectangular buildings. Homesteads
in Tekweni use approximately a quarter the amount of
mangroves compared to homesteads in the other villages
studied. Tekweni is the furthest distance from the man-
grove forest and also has proximity to the coastal scarp
forest that can provide building construction materials
(Obiri 1997). Analysis of use at the village scale indi-
cates that observed differences at the homestead scale are
generally enlarged at the village scale. Tekweni Village
which has homesteads with the lowest utilization values,
the lowest percentage and actual number of mangrove
homesteads, is estimated to have used less than 9 000
mangrove stems. Cwebeni Village at the other extreme,
has high homestead utilization values and a high per-
centage and actual number of mangrove homesteads.
These factors combine to produce utilization figures of
approximately 100 000 stems.
Demand for a resource may constantly change and
the dynamic needs to be understood (Ellery et al. 2004).
Both the annual and predicted demand for mangrove
stems is influenced greatly by building shape. The annual
demand for mangrove stems is based upon the percent-
age of sampled homesteads that were observed con-
structing new mangrove buildings, the value in Mqualeni
was over 50%. This high value may indicate that this
category of homestead was over-sampled. However, as
Mqualeni is a small village, its influence on the overall
demand for mangroves from Mngazana Forest is rela-
tively limited. The high annual demand indicates that
many homesteads are constructing new buildings on
their land. These tend to be required due to a change in
individual circumstance often with young adults requir-
ing the privacy of a separate building.
The influence of human population growth rates
of 0.78% or 3% upon demand is limited as the current
demand rates are high when compared to the demand
changes due to population growth which are relatively
small. The differences in values between current demand
in 2004 and predicted demand in 2015 due to human
population growth suggest that there are factors other
than human population growth which are influencing
the construction of new buildings. These factors may
include maturing children who desire their own private
room or marriage of family members who remain on the
homestead and family members returning to live at the
homestead such as migrant workers. Although demand
is predicted to show minor changes due to human popu-
lation growth rates, it could change drastically due to the
shape of the new buildings being constructed.
In the past, fuel wood models were devised that pre-
dicted future demand scenarios and an impending ‘fuel
wood crisis’, however this never materialized; the mod-
els failed partly because they did not appreciate the com-
plexity of rural energy and focused upon supply without
accounting for demand changes (Shackleton et al. 2004).
Humans demonstrate adaptability to resource decline
and increasing scarcity and Shackleton (1993) reported
that where demand for the preferred fuel wood — dead
wood — outstripped supply, harvesters responded by
selecting the non-preferred live wood resources and even
developed strategies to circumvent legislation. Factors
such as these may well influence the demand scenarios
presented for Mngazana, particularly as although man-
groves are generally preferred for building construction,
the inhabitants are not strictly reliant upon them and can
obtain alternative timber resources from the indigenous
terrestrial forests or use mud or sand blocks for construc-
tion.
A previous study concerning wood utilization for
building construction focused upon quantities used per
building (Liengme 1983). This study demonstrates that
the number of stems required per building, the propor-
tion of a species used in different structural units, the
building shape and the proportion of buildings con-
structed from different materials must all be evaluated so
that a realistic estimate of wood use can be determined.
Sustainable Forest Management at Mngazana must
recognize that the local communities utilize the man-
groves for building construction and that the people
have a right to access mangrove wood products. Given
the socio-economic status of the local communities at
Bothalia 38,1 (2008)
109
Mngazana, the demand for low-cost building materials
will persist. Although there are alternatives to mangrove
wood, it is highly valued and will probably continue
to be used preferentially for building in the immedi-
ate future. Mangrove forests in other developing coun-
tries are under similar pressures to those of Mngazana
(Semesi 1992; Dahdouh-Guebas et al. 2000) and the
most immediate value of mangroves is placed on their
wood products (Alongi 2002). Additionally, natural for-
ests as opposed to plantation forests are often viewed as
open access resources and as a result may be intensively
harvested (Walters 2005b). Given these conditions, the
extent to which these demands can be met needs to be
analysed based upon a forest inventory.
SFM implies that there should be no decrease in
wood products and that the capacity to regenerate be
maintained. However, studies have demonstrated that
‘small-scale, local woodcutting can be a significant form
of ecological disturbance in mangroves’ (Walters 2005a:
345), and it has been shown that forest structure was dra-
matically altered.
A desktop extrapolation, to determine the sustain-
ability of current extraction rates, was made using pub-
lished data from Adams et al. (2005). They reported
that the density of trees greater than one metre high in
the Mngazana Estuary was 230 Bruguiera gymnorrhiza
trees and 489 Rhizophora mucronata trees per hectare.
As the Mngazana Estuary covers an area of 145 ha this
suggests that there are ± 104 255 trees > 1 m height of
B. gymnorrhiza and R. mucronata. With an estimated
annual demand of 1 8 400 stems, the annual off-take
would be approximately 17.6%. The resource would
last for five and a half years at the current demand,
without any recruitment. This is only sustainable in
the very short term. Such a desktop extrapolation can
be used as a guideline within an adaptive management
context. Further investigations concerning recruitment
and growth rates are required to accurately assess sus-
tainability. As Mngazana has already been subjected to
past harvesting, the forest must be assessed to deter-
mine whether the present community structure is func-
tioning in a desirable state or if current stands need to
be improved. Harvesting needs to be planned so that the
ecosystem functioning and biodiversity are maintained.
Studies suggest that mangroves can be managed sus-
tainably for their wood, the Matang Mangrove Forest in
Malaysia has been managed since 1906 for commercial
purposes and continues to be productive (Hogarth 1999).
SFM should also aim to broaden the range of ben-
efits derived from the mangroves, particularly of non-
consumptive uses. Activities such as mangrove honey
production, could demonstrate that non-consumptive
mangrove uses have the potential to generate incomes.
External factors that may affect demand also need to be
considered: at Mngazana these include the link between
mangrove use and use of other indigenous timbers and
alternatives to mangroves such as sand blocks. In the
village of Cwebeni, a group was formed to produce and
locally sell sand blocks for building construction. These
blocks are durable, highly regarded as building materi-
als and, if competitively priced, could reduce demand
for mangrove stems. Other factors may indirectly affect
demand for mangroves, for example clearing of the
coastal scarp forest for agriculture will reduce the sup-
ply of construction timber from these forests, builders
may turn to the mangrove forests to meet their needs
and thereby increase demand for mangrove stems. In the
past, forest clearing for agriculture has reduced the sup-
ply of indigenous hardwood poles from coastal scarp for-
est in northern KwaZulu-Natal (Cunningham & Gwala
1986).
ACKNOWLEDGEMENTS
This project was funded by USAID, the National
Department of Environmental Affairs and Tourism
(Marine and Coastal Management) and the Southern
African Consortium of Universities for Development and
Environment — Sustainable Land Use Project: SLUSE-SA.
Thanks to the Institute of Natural Resources, particularly F.
Lewis and A. Msimang who are facilitating the Mangrove
Management Plan. The Discipline of Geography, Uni-
versity of KwaZulu-Natal (Pietermaritzburg Campus)
provided logistical support and B. Gijsbertsen of the
Carto-graphic Unit produced the location map. Thanks
to Eamoch Mtambeki for translation and guidance in the
field. Special thanks to members of the Mqualeni, Tekweni
and Cwebeni villages who participated in the study.
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OBITUARY
EILY EDITH AGNES GLEDHILL (NEE ARCHIBALD) (1914-2007)
Eily Gledhill (Figure 1) was bom on 7 September
1914 in Walmer, Port Elizabeth, as Eily Edith Agnes
Archibald. Her father was a merchant in Port Elizabeth
and her mother, Lilian Irving, a keen collector of succu-
lent plants, particularly of the genus Haworthia.
Upon finishing her schooling at St Dominic’s Priory,
Walmer, Eily enrolled at Rhodes University where
she completed her B.Sc.Cl.L. degree with Botany and
Mathematics as majors. Later she obtained an M.Sc.
degree in Botany with a thesis on jointed cactus; and
went on to London University to do her Ph.D. thesis on
plant populations. When Eily returned to South Africa
she joined the Division of Chemistry of the Department
of Agriculture. In 1941 she was appointed as lecturer at
the Botany Department of Rhodes University where she
continued the research in which she had been engaged
at Chemical Services — relating soil survey results to
plant distribution. In the 1950s, she was occupied with
trace element research on citrus and pineapple cultiva-
tion in the Eastern Cape. It was at Rhodes University
where Eily met and married Jack Gledhill, head of the
Department of Physics and Electronics.
Eily Gledhill’s botanical work included a survey
of the Alexandria District as well as ecological sur-
veys of the Fish River Catchment Area in 1946 and
of the Greater Addo Elephant National Park in 1954.
During this time she was engaged in assembling the ±
9 000 specimens that are today lodged in the Selmar
Schonland Herbarium in the Albany Museum, in the
National Herbarium in Pretoria and in the herbarium of
the Royal Botanic Gardens at Kew. Many of these are
signed cryptically ‘E2A2’ (Lubke & Brink 2004) which
refers to the initials of her maiden name. With Dr M.A.
Pocock and others she started the Rhodes University
Herbarium to house the collections made by students.
This has now been amalgamated with the Selmar
Schonland Herbarium.
Eily described a number of new species and worked
on a revision of the genus Albuca. She also published
‘The genus Dioscorea in the Cape Province west of
East London’ in 1967. Many of her numerous publica-
tions were illustrated by herself with accurate black-
and-white drawings. Furthermore, her book published
in 1969 and with a second edition in 1981, The Eastern
Cape veld flowers, in which she describes 554 spe-
cies from 125 families, each illustrated by her fine line
drawings, is still of great value to students of botany
and the public at large (Figure 2). In the preface she
writes: ‘If this guide assists owners of land and visitors
to know more about the plants of the eastern Cape, it
will have achieved its object, for any one who becomes
interested in this unique flora cannot fail to realise why
it needs to be protected’. She coined the term bonte-
veld which has become recognized as a veld type and is
widely used.
Eily Gledhill’s interests were not only confined to
plants. In 1939, she joined St John Ambulance and
taught first aid and child care courses throughout her
active life. Through this Eily was honoured in 1992 by
Queen Elizabeth II, the sovereign head of the order, by
her promotion to Dame of the Most Venerable Order of
the Hospital of St John of Jerusalem.
Eily was a member of the British Ecological Society,
the Royal Society of South Africa, the Botanical Society
of South Africa, the Van Riebeeck Society and the South
African Ornithological Society.
She also became more and more interested in local
history and contributed to The 1820 Settlers Illustrated,
edited by Guy Butler (1974) and to Rex and Barbara
Reynolds’s (1974) book, Grahamstown: from cottage
to villa. She was instrumental in the restoration of many
buildings including the Observatory Museum and in
1980 she researched and published with her husband a
book, In the steps of Piet Retief about the famous trek-
ker’s activities in Grahamstown as trader and farmer.
FIGURE 1. — Eily Edith Agnes Gledhill (1914—2007). By courtesy of St
John Ambulance, Grahamstown. Copied by FotoFirst.
112
Bothalia 38,1 (2008)
She realized the tourist potential of Grahamstown and
published many pamphlets and booklets on frontier forts
and founded the Grahamstown Guild of Tour Guides,
leading many tours herself. Makana handicrafts, which
created jobs for township women and Abalizi, a low-cost
housing project, were also founded by her.
Eily Gledhill died on 30 May 2007 in Grahamstown
where she had lived and worked for most of her life.
She was a woman of great intellect and high principles
but my (E. B.) most prized recollection of her was her
delightful, quirky sense of humour which always light-
ened the working day at the herbarium whenever she
visited her special friend, Grace Britten.
She is commemorated in Haworthia ei/vae Poelln.
(Gunn & Codd 1981).
REFERENCES
GLEDHILL, E. 1969. The Eastern Cape veld flowers. Department of
Nature Conservation of the Provincial Administration of the
Cape of Good Hope, Cape Town.
GLEDHILL, I. 2007. Eily Gledhill: a tribute. Grocott’s Mail 08.06.2007: 3.
GUNN, M. & CODD, L.E. 1981. Archibald, Eily Edith Agnes. Botanical
exploration of southern Africa. Balkema, Cape Town.
LUBKE, R. & BRINK, E. 2004. One hundred years of botany at Rhodes
University. South African Journal of Science 1 00: 609-6 1 4.
PUBLICATIONS BY E.E. A. GLEDHILL [NEE ARCHIBALD]
ARCHIBALD, E.E. A. 1939. The development of the ovule and seed
of jointed cactus (Opuntia aurantiacus Lindley). South A frican
Journal of Science 36: 1 95-2 11.
1 940. Haworthia lockwoodii. The Flowering Plants of South A frica
20: t. 792.
Bothalia 38,1 (2008)
113
- 1946. Haworthia tauteae. The Flowering Plants of Africa 25: t. 1103.
- 1948. Plant populations. 1. Anew application ofNeyman’s Contagious
Distribution. Annals of Botany 12, n.s.: 221-235.
- 1950. Plant populations. 2. The estimation of the number of individu-
als per unit area of species in heterogeneous plant populations.
Annals of Botany 14, n.s.: 7-21.
- 1951a. Sterculia alexandri. The Flowering Plants of Africa 28: t.
1103.
- 1951b. Pro tea acaulis var. cockscombensis. The Flowering Plants of
Africali: t. 1115.
- 1951c. Notes on Sterculia alexandri Harv. Journal of South African
Botany 17: 73-75.
- 1952. A possible method for estimating the area covered by the basal
parts of plants. South African Journal of Science 48: 286-292.
- 1954 [1955], An ecological survey of the Addo Elephant National
Park. Journal of South African Botany 20: 137-1 54.
- 1956a. Neopatersonia uitenhagensis. The Flowering Plants of Africa,
vol. 31: t. 1204.
- 1956b. Albuca crudenii. In E.E.A. Archibald, L.E. Codd, R.A. Dyer,
A.D.J. Meeuse & D. van Druten, New and interesting records of
South African flowering plants. Bothalia 6: 542-544.
- 1960. Some species of Eriospermum in the Eastern Cape Province.
Journal of South African Botany 26: 93-1 37.
- 1967. The genus Dioscorea in the Cape Province west of East London.
Journal of South African Botany 33: 1 — 46.
GLEDHILL, E. 1969. The Eastern Cape veld flowers. Department of
Nature Conservation of the Provincial Administration of the
Cape of Good Hope, Cape Town, (also in Afrikaans)
- 1974. Contributions in G. Butler, The 1820 Settlers Illustrated. Human
& Rousseau, Cape Town.
- 1974. Text in Grahamstown: from cottage to villa by R. Reynolds &
B. Reynolds. Phillip, Claremont, Cape Town.
- 1981. The Eastern Cape veld flowers, edn 2. Department of Nature &
Environmental Conservation, Cape Town, (also in Afrikaans)
E. BRINK* and G. GERMISHUIZEN**
* Selmar Schonland Herbarium, Department of Botany, Rhodes
University, P.O. Box 94, 6140 Grahamstown, South Africa.
** South African National Biodiversity Institute, Private Bag X101,
0001 Pretoria.
MS. received: 2008-01-24.
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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
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The history of the
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BOTHALIA
Volume 38,1
May 2008
CONTENTS
1 . Systematics of the southern African genus Ixia (Iridaceae). 1 . The I. rapunculoides complex. P. GOLD-
BLATT and J.C. MANNING 1
2. Three new species of Asparagus (Asparagaceae) from South Africa, with notes on other taxa. S.M.
BURROWS and J.E. BURROWS 23
3. A new species of Euclea (Ebenaceae) from ultramafic soils in Sekhukhuneland, South Africa, with notes
on its ecology. E. RETIEF, S.J. SIEBERT and A.E. VAN WYK 31
4. The genus Solarium (Solanaceae) in southern Africa: subgenus Leptostemonum , section Giganteiformia.
W.G. WELMAN 39
5. Two new species of Babiana (Iridaceae: Crocoideae) from western South Africa, new names for B.
longiflora and B. thunbergii, and comments on the original publication of the genus. P. GOLDBLATT,
J.C. MANNING and R. GEREAU 49
6. The genus Wellstedia (Boraginaceae: Wellstedioideae) in southern Africa. E. RETIEF and A.E. VAN
WYK 57
7. Notes on African plants:
Apocynaceae. A new species of Huemia (Asclepiadoideae-Ceropegieae) from Angola. P.V.
BRUYNS 83
Asphodelaceae. Notes on the nomenclature and typification of Aloe natalensis (Alooideae). N.R.
CROUCH, G.F. SMITH and R.R. KLOPPER ' 70
Asphodelaceae: Alooideae. Bulbine triebneri, an earlier name for Bulbine alba , as well as additional
and new localities in Eastern and Northern Cape, South Africa. R.R. KLOPPER, A.W.
KLOPPER, H. BAIJNATH and G.F. SMITH 67
Asphodelaceae: Alooideae. New evidence in support of a disjunct distribution of Aloe karasbergensis.
R.R. KLOPPER, P.J. DU PREEZ and G.F. SMITH 82
Asteraceae-Gnaphalieae. Metalasia helmei, a new member of a small clade from the Western Cape?
P.O. KARIS and N. HELME 65
Asteraceae. Tripteris calcicola, a new calciphilous species from Western Cape, South Africa. J.C.
MANNING and P. GOLDBLATT 85
Colchicaceae. Further new combinations in Colclnicum. C. ARCHER 83
Errata in Bothalia 37,2 (2007): MOFFETT, R.O. 2007. Name changes in the Old World Rhus and
recognition of Searsia (Anacardiaceae) 71
Hyacinthaceae. Drimiopsis linioseta, a new species from the Sekukhuneland Centre of Endemism,
South Africa. A.J. HANKEY, M.H. BUYS and P.D. LEBATHA 72
Iridaceae. Romulea lutea and R. tubulosa (Crocoideae), two new species from Namaqualand, South
Africa. J.C. MANNING and P. GOLDBLATT 78
Oxalidaceae. A new species of Oxalis from the Hantam-Roggeveld Plateau, Northern Cape, South
Africa. J.C. MANNING and P. GOLDBLATT 75
Rosaceae. Validation of three Cliffortia taxa of Cliffortia species in Bothalia 37,1 (2007). C.M.
WHITEHOUSE 75
8. Vegetation and vegetation-environment relationships at Grootbos Nature Reserve, Western Cape, South
Africa. M. MERGILI and S. PRIVETT 89
9. Resource demand estimates for sustainable forest management: Mngazana Mangrove Forest, South
Africa. C.H. TRAYNOR and T.R. HILL 103
10. Obituary: Eily Edith Agnes Gledhill (nee Archibald) (1914-2007). E. BRINK and G. GERMISHUI-
ZEN Ill
Abstracted, indexed or listed in • AETFAT Index • AGRICOLA • AGRIS • BIOSIS: Biological Abstracts/RRM • CABS • CABACCESS • CAB
ABSTRACTS • ISI: Current Contents, Scisearch, Research Alert • Kew Record of Taxonomic Literature • Taxon : reviews and notices.
ISSN 006 8241
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