- n
ISSN 0006 8241 = Bothalia
Bothalia
A JOURNAL OF BOTANICAL RESEARCH
Vol. 26,2 October 1 996
TECHNICAL PUBLICATIONS OF THE NATIONAL BOTANICAL INSTITUTE,
PRETORIA
Obtainable from the National Botanical Institute, Private BagXIOl, Pretoria 0001, Republic of South
Africa. A catalogue of all available publications will be issued on request.
BOTHALIA
Bothalia is named in honour of General Louis Botha, first Premier and Minister of Agriculture of the
Union of-South Africa. This house journal of the National Botanical Institute, Pretoria, is devoted to
the furtherance of botanical science. The main fields covered are taxonomy, ecology, anatomy and
cytology. Two parts of the journal and an index to contents, authors and subjects are published
annually.
A booklet of the contents to Vols 1-20 is available.
STRELITZIA
A series of occasional publications on southern African flora and vegetation, replacing Memoirs of
the Botanical Survey of South Africa and Annals of Kirstenbosch Botanic Gardens.
MEMOIRS OF THE BOTANICAL SURVEY OF SOUTH AFRICA
The memoirs are individual treatises usually of an ecological nature, but sometimes dealing with
taxonomy or economic botany. Published: Nos 1-63 (many out of print). Discontinued after No. 63.
ANNALS OF KIRSTENBOSCH BOTANIC GARDENS
A series devoted to the publication of monographs and major works on southern African flora.
Published: Vols 14-19 (earlier volumes published as Supplementary volumes to the Journal of
South African Botany). Discontinued after Vol. 19.
FLOWERING PLANTS OF AFRICA (FPA)
This serial presents colour plates of African plants with accompanying text. The plates are prepared
mainly by the artists at the National Botanical Institute. Many well known botanical artists have
contributed to the series, such as Cythna Letty (over 700 plates), Kathleen Lansdell, Stella Gower,
Betty Connell, Peter Bally, Fay Anderson, Ellaphie Ward-Hilhorst and Gillian Condy. The Editor is
pleased to receive living plants of general interest or of economic value for illustration.
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, synonymy, literature and limited specimen citations, as well as taxonomic and
ecological notes.
Contributions to the FSA also appear in Bothalia.
PALAEOFLORA OF SOUTHERN AFRICA
A palaeoflora on a pattern comparable to that of the Flora of southern Africa. Much of the
information is presented in the form of tables and photographic plates depicting fossil populations.
Now available:
Molteno Formation (Triassic) Vol. 1. Introduction. Dicroidium, by J.M. & H.M. Anderson.
Molteno Formation (Triassic) Vol. 2. Gymnosperms (excluding Dicroidium ), by J.M. &
H.M. Anderson.
Prodromus of South African Megafloras. Devonian to Lower Cretaceous, by J.M. & H.M.
Anderson. Obtainable from: A. A. Balkema Marketing, Box 317, Claremont 7735, RSA.
BOTH ALIA
A JOURNAL OF BOTANICAL RESEARCH
Volume 26,2
Scientific Editor: O.A. Leistner
Technical Editor: B.A. Momberg
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INSTITUTE
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Oct. 1996
Editorial Board
D.F. Cutler
B.J. Huntley
P.H. Raven
J.P. Rourke
M.J. Werger
Royal Botanic Gardens, Kew, UK
National Botanical Institute, Cape Town, RSA
Missouri Botanical Garden, St Louis, USA
Compton Herbarium, NBI, Cape Town, RSA
University of Utrecht, Utrecht, Netherlands
CONTENTS
Volume 26,2
1. A synopsis of Peristrophe (Acanthaceae) in southern Africa. K. BALKWILL 83
2. Studies in the Ricciaceae of sub-Saharan Africa. S.M. PEROLD 95
3. FSA contributions 6: Orchidaceae: Holothrix. K.L. IMMELMAN 125
4. FSA contributions 7: Verbenaceaet : Vitex. C.L. BREDENKAMP and D.J. BOTHA 141
5. Notes on African plants:
Amaryllidaceae. Should Crinum forbesii (Lindl.) Schult. & Shult.f. be reinstated? R.H.
ARCHER and C. ARCHER 153
Asphodelaceae. Generic numbers in the subfamily Alooideae. G.F. SMITH and A.E. VAN
WYK 158
Campanulaceae. Transfer of eleven varieties of Lightfootia nom. illeg. to Wahlenbergia. W.G.
WELMAN 157
Proteaceae. A new species of Serruria from the northern part of the Western Cape. J.P. ROURKE 154
6. Exormotheca bulbigena sp. nov. (Hepaticae, Marchantiales) and its relationship to E. holstii in
southern Africa. T. BORNEFELD, O.H. VOLK and R. WOLF 159
7. Morphological and biochemical genetic evidence for hybridization in the genus Centella (Apiaceae),
with notes on phylogenetic and taxonomic implications. M.T.R. SCHUBERT, B-E. VAN WYK,
H.F. VAN DER BANK and M. VAN DER BANK 167
8. Red Data List of southern African plants. 1. Corrections and additions. C. HILTON- TAYLOR . 177
9. Book reviews 183
10. National Botanical Institute: list of staff and publications, 31st May 1996. Compiler: B.A. MOM-
BERG 185
11. Guide for authors to Bothalia 201
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Bothalia 26,2: 83-93 (1996)
A synopsis of Peristrophe (Acanthaceae) in southern Africa
K. BALKWILL*
Keywords: Acanthaceae, Peristrophe, southern Africa, synopsis, taxonomy
ABSTRACT
A synopsis of Peristrophe Nees (Acanthaceae) in southern Africa is provided. Peristrophe comprises nine species, one with
two subspecies in southern Africa. A key for identification and descriptions of species not included in recent literature are
provided. Morphology of tertiary bracts, indumentum on stems and distribution is illustrated and diagnostic characters,
distribution, habitat, flowering time and conservation status are discussed for each species.
INTRODUCTION
I undertook a revision of Peristrophe Nees in southern
Africa (Balkwill 1985) and while doing so, published an
account of the Peristrophe grandibracteata complex
(Balkwill et al. 1988) and some new species (Balkwill et
al. 1985; Balkwill & Getliffe Norris 1989). I thought that
the discussions under these species and that of Brummitt
(Wood et al. 1983: 451) clarified the fate of previously
recognised species, but from Welman (1993), it seems that
this is not the case. At present, no satisfactory key to the
southern African species is available in the literature and
there is likely to be a long delay before further accounts
of Acanthaceae are published in the Flora of southern
Africa series. I have therefore decided that it would be
constructive to publish a synopsis to aid in the identifica-
tion of species of this genus.
Peristrophe Nees in Wall., Plantae Asiaticae Rari-
ores 3: 112 (1832); Endl.: 707 (1839); Meisn.: 297 (1840);
Nees: 374 (1841); Nees: 492 (1847); Harv.: 286 (1868);
Benth. & Hook.f.: 1071 (1876); C.B.Clarke: 554 (1885);
Lindau: 331 (1895a); C.B.Clarke in Burkill & Clarke: 242
(1899); C.B.Clarke: 84 (1901); Hutch. & Dalziel: 264
(1936); Heine: 424 (1963); Agnew: 609 (1974); R.A.Dyer:
593 (1975). Type species: Peristrophe baphica (Spreng.)
Bremek. in Nova Guinea new ser. 8: 149 (1957) [ = P.
tinctoria (Roxb.) Nees comb, illeg.].
Suffruticose annuals or evergreen perennials up to
2 m high. Leaves petiolate, simple, opposite, widely
ovate to lanceolate, acuminate to acute, entire, attenuate
at base, herbaceous, usually with eglandular trichomes,
especially on midrib. Inflorescence of monochasial
cymes (inflorescence units) enclosed by a pair of terti-
ary bracts, with (1)2 or 3(4) inflorescence units umbel-
lately arranged, often compounded. Bracts : secondary
bracts 2, free, narrowly triangular to lanceolate, sessile or
leaflike; tertiary bracts lanceolate to broadly ovate, acu-
minate. Flowers perfect, zygomorphic, occasionally cleis-
togamous. Calyx with short tube and lanceolate lobes,
* C.E. Moss Herbarium, Department of Botany, University of the Wit-
watersrand. Private Bag 3, Wits 2050, South Africa.
MS. received: 1996-03-29.
margins usually membranous, inner surface pubescent,
trichomes appressed and eglandular. Corolla bilabiate,
resupinate; tube narrowly cylindric below, subcampanu-
late above, with 2 pairs of longitudinal hairy ridges
within, with eglandular and sometimes glandular
trichomes without; lip in lower position elliptic or
ovate, minutely emarginate; lip in upper position nar-
rowly elliptic, 3-fid; flowers lilac to purple, occasion-
ally white, with dark purple honey guides on white
background on lip in upper position. Stamens 2, ex-
serted; filaments epipetalous, linear, usually white or
yellow; anthers bithecous, thecae superposed and sepa-
rated, approximated or slightly overlapping, introrse,
purple. Disc shallowly cupular, with two small awns,
nectariferous. Gynoecium bicarpellate; stigma bilobed;
style exserted, filiform, glabrous to sparsely strigose;
ovary superior, with septum in median longitudinal
axis, ovoid or ellipsoid, bilocular, with two ovules in
each locule. Fruit a stipitate capsule, clavate or obtrul-
late, placentae inelastic at the bases and bearing reti-
nacula, dehiscing loculicidally. Seeds discoid, usually 2
in each locule, rough and tuberculate.
The genus Peristrophe occurs in Africa and the East
Indies and comprises about 25 species, of which nine oc-
cur in southern Africa. Peristrophe is very closely related
to Dicliptera Juss. and differs only in the nature of the
placental bases, which are inelastic in Peristrophe and
elastic in Dicliptera. Some workers have tried to apply
other characters (e.g. width of the tertiary bracts) as ge-
neric characters, and this has led to the incorrect clas-
sification of species such as P. angolensis (S .Moore)
K. Balkwill, P. transvaalensis (C.B.Clarke) K. Balkwill
and P. hereroensis (Schinz) K. Balkwill, all of which were
described in the genus Dicliptera. Another species that
has been confused with Dicliptera is P. bivalvis (L.)
Merr., which has been confused with Dicliptera foetida
(Forssk.) Blatt. (Wood et al. 1983). In southern Africa,
the shape and venation of the tertiary bracts, if used si-
multaneously, can act as a guide to the genera. The ter-
tiary bracts of the southern African species of Peristrophe
are either narrow and single-veined, or if broad, then the
veins are pinnate and the secondary veins are reticulate
near the margins (Figure 1), whereas the bracts of Di-
cliptera are 3-, 5- or even 7-veined from the base and
84
Bothalia 26,2(1996)
FIGURE 1. — Inflorescence units of southern African species of Peristrophe , illustrating tertiary bracts. A, P. grandibracteata , Muller 1311
(WIND); B, P hereroensis, De Winter 2356 (PRE),C, P. namibiensis, De Winter & Leistner 5828 (WIND); D, P. paniculata, Goldblatt
1929 (NBG); E, P transvaalensis, Balkwill 764 (J); F, P cliffordii, Balkwill 793 (J); G, P. gillilandiorum, Bruce 58 (K); H, P cernua,
Bulkwill 168 (J); I, P. decorticans, Balkwill 801 (J). Scale bar: 4 mm.
Bothalia 26,2(1996)
85
the secondary veins do not form conspicuous reticulations
near the margins.
Nees (1832, 1847), recognised two sections in Peris-
trophe and subdivided the first of these into two subsec-
tions, but he did not name the sections or subsections.
Since that time, three species of Peristrophe have been
described from Namibia and these do not belong in the
groups described by Nees. Thus, it seems that Nees’s
(1847) first section comprises three groups, but these
three groups are sufficiently different to be recognised
at the rank of section. The species of Peristrophe in
southern Africa belong to two of these groups (referred
to as sections 2 and 3 below). It would be premature
to formally describe these sections here, before a mod-
ern and monographic account of the genus has been
completed.
Section 1: inflorescence of two to seven monochasial
cymes (inflorescence units) umbellately arranged, ‘um-
bels’ sometimes compounded. Tertiary bracts spathulate,
ovate to obovate, with three, five or seven veins from the
base, remaining green when mature. Capsule clavate.
Included species (none southern African): P baphica
(the type of the genus), P. montana Nees and P speciosa
Nees.
Section 2: differs from Section 1 by having only two
inflorescence units in each ‘umbel’; a single primary vein
from the base of the tertiary bracts; tertiary bracts that have
conspicuous secondary veins and become membranous
when mature (Figure 1 A-C); and capsules that are obtrul-
late in profile.
Inflorescence of monochasial cymes (inflorescence
units), usually two umbellately arranged. Tertiary bracts
ovate to widely ovate, 9.1-14.4 x 6.1-13.8 mm, with a
single vein from the base, becoming membranous when
mature. Capsule obtrullate in profile.
Included species (all southern African): P. grandibrac-
teata, P. hereroensis and P namibiensis.
Section 3: differs from Sections 1 and 2 by the narrower,
less conspicuous tertiary bracts, which are lanceolate to
oblanceolate (not ovate to broadly ovate) and narrower
than 4 mm (Figure 1 D-I) and from Section 2 by the tertiary
bracts that are green (not membranous and conspicuously
veined) at maturity and by having clavate (not obtrullate)
capsules.
Inflorescence of monochasial cymes (inflorescence
units), usually 3-4 umbellately arranged, often com-
pounded. Tertiary bracts lanceolate to oblanceolate,
3.0-18.0 x 0.4-3.2 mm, green. Capsule clavate.
Included southern African species: P. cemua, P. clif-
fordii, P. decorticans, P. gillilandiorum, P. paniculata and
P. transvaalensis .
Key to the southern African species of Peristrophe
la Tertiary bracts broader than 5 mm, veins pinnate, secondary veins forming reticulation at bract edges; capsules obtrullate in profile: Section 2:
2a Secondary bracts leaflike, longer than 7 mm; tertiary bracts widely cuneate at base; leaves lanceolate, length : breadth ratio more than
2.4: 1:
3a Young stems appearing white, densely pubescent with eglandular trichomes with enlarged, ornamented terminal cells . . . .1. P. grandibracteata
3b Young stems appearing green, not densely pubescent, trichomes without enlarged terminal cells 2. P. hereroensis
2b Secondary bracts not leaflike, shorter than 7 mm; tertiary bracts reniform or cordate at base; leaves ovate to widely ovate, length :
breadth ratio smaller than 2.4 : 1:
4a Leaf length : width ratio less than 1.75 : 1; tertiary bract length : breadth ratio less than 1.25 : 1; capsules pubescent
3.1. P. namibiensis subsp. namibiensis
4b Leaf length : width ratio greater than 1.75 : 1; tertiary bract length : breadth ratio greater than 1.25 : 1; capsules glabrous
3.2. P. namibiensis subsp. brandbergensis
lb Tertiary bracts narrower than 4 mm, with a midrib only, or if more veins present, then not forming reticulation at edges of bract; capsules
clavate: Section 3:
5a Corolla 6-14 mm long; plant annual, up to 2 m high; rootstock not woody and seldom branched 4. P. paniculata
5b Corolla longer than 14 mm; plant suffruticose perennial, up to 1.2 m high; rootstock woody and often branched:
6a Tertiary bracts longer than 14 mm and length : breadth ratio less than 6.2 : 1 5. P transvaalensis
6b Tertiary bracts shorter than 13 mm, but if longer, then length : breadth ratio greater than 6.2 : 1:
7a Mature stems densely pubescent between ridges; found in the Limpopo River valley:
8a Tertiary bracts up to 5 mm long, length : breadth ratio less than 6:1 6. P. cliffordii
8b Tertiary bracts longer than 7 mm, length : breadth ratio greater than 7:1 7. P gillilandiorum
7b Mature stems glabrous or sparsely pubescent between ridges or not found in the Limpopo River valley:
9a Occurring in Zimbabwe, Botswana and Northern Province; lip in lower position (6.5)— 7.5— 9.2— (10.2) mm long
8. P. decorticans
9b Occurring in KwaZulu-Natal and Eastern Cape; lip in lower position (7.2)— 9.8— 16.5— (22.3) mm long 9 P. cemua
SECTION 2
1. P. grandibracteata Lindau in Botanische Jahr-
biicher 49: 404 (1913); P.G.Mey.: 49 (1968), p. p.; K.Balk-
will et al.: 48 (1988). Type: Namibia, Kuibis, (-DB), Range
613 (SAM!, lecto., designated by Balkwill et al.: 48 (1988);
BOL!).
Diagnostic characters: densely white hairy stems
(Figure 2A) and broad bracts (Figure 1A). Distribution:
southern Namibia, south of 24° latitude and west of 18°
longitude (Figure 3). Habitat: deep sandy soils and dry
rocky slopes in the Namib Desert and Bushy Karoo-
Namib shrubland (White 1983). Flowering time: April to
June. Conservation status: although not highly localised,
P. grandibracteata is known from only five specimens,
86
Bothalia 26,2(1996)
FIGURE 2. — Indumentum on stems of southern African species of Peristrophe. A, P. grandibracteata, Muller 1311 (WIND); B, P.
hereroensis, De Winter 2356 (WIND); C, P. namibiensis, De Winter & Leistner 5828 (PRE); D, P. paniculata, Goldblatt 1929
(NBG); E, P trun.ivaalensis, Balkwill 764 (J); F, P. cliffordii, Balkwill 793 (J); G, P. gillilandiorum, Bruce 58 (K); H, P. cemua,
Balkwill 168 (J); I, P. decorticans, Balkwill 801 (J). Scale bar: 1 mm.
Bothalia 26,2 (1996)
87
FIGURE 3. — Distribution of Peristrophe grandibracteata, ■; P. here-
roensis, ▲.
suggesting it is rare and its conservation status requires
investigation.
2. P. hereroensis (Schinz) K.Balkwill in Getliffe
Norris et al. in South African Journal of Botany 51: 489
(1985); K.Balkwill et al.: 48 (1988). Type: Namibia, Otji-
hua, (-BB), Dinter 459 [Z!, lecto., designated by Balkwill
et al.: 48 (1988), 2 sheets; GRA! K!, SAM!].
Dicliptera hereroensis Schinz in Vierteljahrsschrift der Naturforschen-
den Gesellschaft in Zurich 61: 438 (1916).
Diagnostic characters'. P. hereroensis can be distin-
guished from P. grandibracteata by its green stems with
sparse eglandular hairs (Figure 2B) rather than white
stems with dense ornamented eglandular hairs. It differs
from P. namibiensis by the tertiary bracts that are widely
cuneate at the base (Figure IB) (not reniform or cordate),
secondary bracts that are longer than 7 mm (not shorter
than 6 mm) and by the narrower leaves [length : breadth
ratio greater than (not less than) 2.4 : 1], Distribution:
northern Namibia (Figure 3). Habitat: on various kinds of
soils, usually in the shade of trees of Acacia, in Kalahari
Acacia wooded grassland and deciduous bushland and the
Kalahari/Karoo-Namib transition (White 1983). Flower-
ing time: between November and July, with a peak in
March and April. Conservation status: P. hereroensis is
known from 17 collections from a number of different
localities. It is unlikely to be rare or threatened.
3. P. namibiensis K.Balkwill in Balkwill et al. in
South African Journal of Botany 54: 52 (1988). Type: Na-
mibia, Farm Blasskranz (REH 7), Berghang, (-AC),
Merxmiiller & Giess 28127 (PRE, holo.!; WIND!).
Diagnostic characters: P. namibiensis differs from P.
grandibracteata by its green (not white) stems and from
P. grandibracteata and P. hereroensis by its much smaller
secondary bracts that are not leaflike and by its cordate
tertiary bracts (Figure 1C).
3a. subsp. namibiensis
Diagnostic characters: leaf length : width ratio of less
than 1.75 : 1; tertiary bract length : breadth ratio of less
than 1.25 : 1; and pubescent capsules distinguish subsp.
namibiensis. Distribution: southern half of Namibia (Fig-
ure 4). Habitat: on dolomite koppies (and possibly other
habitats) in the Kalahari/Karoo-Namib transition and
Namib Desert White 1983). Flowering time: April to Au-
gust. Conservation status: this taxon is known from only
five collections, suggesting that its conservation status re-
quires investigation.
3b. subsp. brandbergensis K.Balkwill in Balkwill
et al. in South African Journal of Botany 54: 52 (1988).
Type: Namibia, Outjo Farm OU 516, Sandsteinruecken,
Giess & Barnard 7921 WIND, holo.!; NBG!, PRE!).
Diagnostic characters: leaf length : width ratio greater
than 1.75 : 1; tertiary bract length : breadth ratio greater
than 1.25 : 1; and glabrous capsules distinguish subsp.
brandbergensis. Distribution: northern half of Namibia
(Figure 4). Habitat: Bushy Karoo-Namib shrubland and
Namib Desert White 1983). Flowering time: April to Au-
gust. Conservation status: this taxon is known from nine
collections (some quite recent) from a number of locali-
ties; it is unlikely to be threatened.
SECTION 3
4. Peristrophe paniculata (Forssk.) Brummitt in
Wood et al. in Kew Bulletin 38: 451 (1983). Type: Yemen,
Forsskal 385 [C, designated by Brummitt (Wood et al.
1983), seen on microfiche 38: III. 3^1 at J],
Dianthera paniculata Forssk. (1775).
Dianthera bicalyculata Retz.: 297 (1775 [published in 1776]); Retz.:
10 (1779); Vahl: 6 (1790). Justicia bicalyculata (Retz.) Vahl: 13 (1791);
Willd.: 81 (1797); Vahl: 113 (1804); Roem. & Schult.: 143 (1817); Roxb.:
127 (1820); Roem. & Schult.: 130 (1822); Wall.: 2457 a-g (1830); Roxb.:
126 (1832). Peristrophe bicalyculata (Retz.) Nees: 113 (1832); Nees: 496
(1847); Hook.f. & Benth.: 484 (1849); A.Rich.. 160 (1850); Dalzell &
Gibson: 197 (1861); T.Anderson: 47 (1864); T.Anderson: 521 (1867);
C.B.Clarke: 554 (1885); Lindau: 331 (1895a); Lindau: 371 (1895b); Lin-
dau: 80 (1897); C.B.Clarke in Burkill & Clarke: 242 & 514 (1899);
C.B.Clarke: 85 (1901); Chev.: 501 (1920); Schnell: 20 (1953); Dandy:
28 (1954); Andrews: 185 (1956); Heine: 424 (1963); Cufodontis: 959
(1964); P.G.Mey.: 49 (1968); Agnew: 609 (1974); Solms: 113 & 244
FIGURE 4. — Distribution of Peristrophe namibiensis subsp. namibien-
sis, ■; subsp. brandbergensis, ▲.
88
Bothalia 26,2(1996)
3UIH WEST AfRICA WINDHOEK
SUIDWES-AfRIKA \ ■; , •
HERBARIUM
DUTH WEST AFRIC
SUIDWES-AFRIKA
WINDHOEK
HERBARIUM
South Wojt
ttr othp bk»lfcul>t» (Reli.)
[ticj Um »•
FIGURE 5. — Four specimens of Peristrophe paniculata , showing change in habit over one season. A, 26-02-1963, Giess, Volk & Bleissner
5503 (WIND); B, 25-04-1963, Giess, Volk & Bleissner 6464 (WIND); C, 09-05-1963, Leippert4653 (WIND); D, 05-06-1963, Giess
& Leippert 7317 (WIND). All x 0.35.
Bothalia 26,2(1996)
89
(1867). Type: possibly Koenig (on microfiche of Linnean herbarium, fide
Wood et al. 1983).
Dianthera malabarica L.f.: 85 (1782), nom. illeg. Justicia malabarica
(L.f.) Aiton: 27 (1789), nom. illeg. Type: as for D bicalyculata Retz.
Justicia ligulata Lam.: 632 (1785); Cav.: 52, t. 71 (1791); Lam.: 40,
t. 12, fig. 2 (1791); Lam.: 95 (1811), nom. illeg. Type: as for Dianthera
paniculata Forssk.
Suffruticose annual up to 2 m high with relatively small
tap root system. Leaves ovate to lanceolate, acuminate to
acute, sometimes rounded, attenuate at base, (1 2-) 17^-6
(-65) x 3— 14(— 38) mm, very thin, slightly scabrid above,
soft below, with simple hairs and cystoliths; petiole
0.9-3.9(-36.0) mm. Inflorescence of monochasial cymes
(inflorescence units), (1)2 or 3(4) umbellately arranged,
usually compounded; inflorescence axis (8.0—) 1 1 .8—19.4
(-31.0) mm; longest peduncle of inflorescence units (7.0—)
10.9-1 8. 6(-30.0) mm. Bracts', secondary bracts lanceo-
late, (2.5-)3.5-4.6(-7.8) x (0.2-)0.4-0.6(-1.0) mm, with
short broad-based eglandular trichomes; tertiary bracts
lanceolate, acuminate, larger one (7.0— )9.2— 14.9(— 1 8.0) x
(0.4-)0.6-l.l(-1.5) mm, sparsely strigose, with short-
stalked and sessile glandular trichomes on surface. Calyx:
tube 1 mm deep; lobes lanceolate, 3 mm long, margins
membranous at base, and strigose, with glandular
trichomes more common towards base, outer surface
sparsely strigose, with sessile glandular trichomes, inner
surface sparsely strigose, with appressed eglandular tri-
chomes. Corolla: tube (5.0-)5.2-6.6(-7.0) mm deep, with
medium, straight and curved eglandular trichomes; lip in
lower position elliptic, 3.5-7.2(-7.3) x ( 1.8—) 1 .9— 2.7(— 3.2)
mm; lip in upper position narrowly elliptic, (5.0-)5.2-6.9
(-8.0) x (1.2— )1.3— 1.7(— 1.8) mm; lilac, purple or deep pur-
ple. Stamens: filaments with short, curved, ornamented
eglandular trichomes, white; anther thecae superposed, not
touching, purple. Disc very shallowly cupular, upper edge
crenate with 2 small triangular awns. Gynoecium: stigma
bilobed; style glabrous; ovary ovoid, very hairy and glan-
dular. Fruit (9.0-)9.4-12.3(-14.0) x (2.0-)2.2-2.8(-3.5) mm,
hairy. Seeds discoid, 2.2-2.4(-2.5) mm, rough and tuber-
culate.
The names P. paniculata and P. bicalyculata were both
validly published in publications bearing the date 1775.
They are taxonomic synonyms but P paniculata has pri-
ority (Wood et al. 1983). The species has only been col-
lected between February and August, and most often in
May in southern Africa. The specimens collected in the
earlier months are of younger and smaller plants than
those collected in the later months. This is particularly
noticeable in the series collected in 1963: Giess, Volk &
Bleissner 5503 (26/02/1963); Giess, Volk & Bleissner
6464 (25/04/1963); Leippert 4653 (09/05/1963); Giess &
Leippert 7317 (05/06/1963) (Figure 5). This suggests that
the species is an annual, although it has been said to reach
2 m high (fide notes on Smith 1293 in PRE).
Diagnostic characters: smaller flowers, less than 13.7
mm long, very shallow disc and small simple tap root
system serve to separate P. paniculata from all other
southern African species of Peristrophe. It has often been
confused with P. decorticans and can be separated from
the latter species by the mature bark, which is smooth and
black in P. paniculata and white and peeling in P. decor-
ticans', by the ratio of length of larger tertiary bract to that
of the shorter, which is ( 1 .4 — ) 1 .6— 2.2(— 2.6) : 1 in P. pani-
culata and 1.2— 1.4(— 1 .6) : 1 in P. decorticans', and on the
basis of distribution: P. paniculata grows in Namibia and
in northwestern Botswana; P. decorticans in eastern Bo-
tswana and in Northern Province. The fruits of P. pani-
culata are more densely sericeous than those of any other
species in southern Africa, but the small flowers are the
most reliable character with which to distinguish P. pani-
culata. Distribution: P. paniculata is a widely distributed
species and ranges as far east as India, and as far west
and south as Namibia (Figure 6). Habitat: in southern Af-
rica, it is found in the semi-arid savanna and semidesert
areas. Flowering time: February to August. Conservation
status: this widespread species is well represented in many
herbaria. The first flowers formed are cleistogamous so
that the seed bank is rapidly replenished after seeds ger-
minate. This species is neither rare nor threatened.
5. Peristrophe transvaalensis (C.B. Clarke) K. Balk-
will in Getliffe Norris et al. in South African Journal of
Botany 51: 489 (1985). Type: Northern Province, without
precise locality, Holub s.n. (K, holo.!).
Dicliptera transvaalensis C.B. Clarke: 92 (1901).
Evergreen suffruticose perennial up to 1 m high.
Leaves ovate to narrowly elliptic, acuminate, attenuate at
base, 26 — 4 1 (^42) x 8— 13(— 14) mm, with many multicel-
lular eglandular trichomes and cystoliths; petiole
(1.7-)2.0-4.3(-4.6) mm. Inflorescence of monochasial
cymes (inflorescence units), (1 or 2)3(4) umbellately ar-
ranged, sometimes compounded; inflorescence axis
(4.0-)4.2-7.0(-7.4) mm long; longest peduncle of inflo-
rescence units (2.4-)4. 1-12.9 mm long. Bracts: secondary
bracts 2, free, lanceolate, (4.7— )5. 1—6.6 x 0.6—1 .0(— 1 . 1 )
mm, pubescent, sessile; tertiary bracts oblanceolate, un-
equal, larger one ( 1 3.6— ) 13.8—16. 1 (—1 6.4) x (0.4-)0.6-l.l
(-1.5) mm, almost tomentose, trichomes multicellular and
eglandular. Calyx: tube 1.5 mm deep; lobes, lanceolate,
5.5 mm long, margins membranous and ciliate, with small
glandular trichomes on outer surface. Corolla: tube (7.3-)
7.5-9.0 mm long, sericeous to tomentose; lip in lower
position ovate, (7.1-)8.8(-9.0) x 2.0-2.7 mm; lip in upper
position narrowly elliptic, 7.0-8.4(-8.8) x (2.4-)2.5-3.5
FIGURE 6. — Distribution of Peristrophe paniculata, •; P. transvaalen-
sis, ■; P. cliffordii, ▲.
90
Bothalia 26,2(1996)
(-3.6) mm; light purple with dark purple honey-guides on
white background on lip in upper position. Stamens: fila-
ments with short and long multicellular eglandular
trichomes, white; anther thecae superposed, purple. Disc
shallowly cupular with crenate top and 2 short stubby
awns. Gynoecium: stigma shortly bilobed; style sparsely
strigose; ovary ovoid, sparsely pubescent. Fruit pubescent,
9.0-10.1 x 2. 1-2.5 mm. Seeds 2. 0-2.2 x 1. 8-2.0 mm,
rough and tuberculate.
It was necessary to transfer P transvaalensis from the
genus Dicliptera as the placentae are inelastic at the base.
Diagnostic characters: the closest ally to P transvaalensis
is P. angolensis which has a characteristic inflorescence
structure with many inflorescence units contracted into an
axil, and an occasional inflorescence unit with a very long
peduncle. In contrast, the inflorescence units of P. trans-
vaalensis are not contracted into the leaf axils. The
broader tertiary bracts of these two species serve to sepa-
rate them from the other species of Peristrophe in southern
Africa. P. transvaalensis has a marked vestiture of mul-
ticellular uniseriate eglandular trichomes, which imparts
a grey-green colour to the leaves of living plants; this
feature is very unusual in this section of the genus. Dis-
tribution: Botswana and southwestern Northern Province
(Figure 6). Habitat: interface of Acocks’s (1988) Sourish
Mixed Bushveld and Sour Bushveld [now Mixed
Bushveld (Van Rooyen & Bredenkamp 1996a) and Wa-
terberg Moist Mountain Bushveld (Van Rooyen & Bre-
denkamp 1996b)]. Flowering time: in the field the plants
were beginning to flower in January, and were flowering
profusely when revisited in May, whereas the plants in
cultivation were still in flower in August of the same year.
It appears that the species flowers throughout the year,
but more profusely in winter. Conservation status: P.
transvaalensis is not a common species and is known
from only six gatherings from a relatively restricted area
and populations of the plant are sparse. The conservation
status of this species requires urgent investigation.
6. Peristrophe cliffordii K.Balkwill in Balkwill et
al. in South African Journal of Botany 51: 485 (1985).
Type: Northern Province, East [should be west] of Mes-
sina, on road to Weipe, 2.8 km from Messina-Pontdrift
road, (-BA), Balkwill 793 (NU, holo.!; PRE!).
Diagnostic characters: P cliffordii can be separated
from all other species of Peristrophe in southern Africa
by its much smaller tertiary bracts (Figure IF) that are
covered in rust-coloured glandular and eglandular hairs.
In addition, it, P gillilandiorum and P. transvaalensis have
many grey hairs between the ridges on the stems (Figure
2E) which separates them from P. grandibracteata, which
has dense white hairs on its stem, and all other species,
which have no or very few hairs between the ridges on
the stem. The broader leaves separate this species from P.
gillilandiorum, and the much smaller tertiary bracts sepa-
rate it from P. transvaalensis. Distribution: highly local-
ised near Weipe, west of Messina in the Northern Province
(Figure 6). Habitat: on Kalahari sands amongst Colo-
phospermum mopane in the frost-free Limpopo Valley.
Flowering time: autumn and winter. Conservation status:
this species is extremely rare and localised. Repeated
droughts coupled with browsing do not augur well for it.
7. Peristrophe gillilandiorum K.Balkwill in Balk-
will et al. in South African Journal of Botany 51: 488
(1985). Type: Northern Province, Dongola, Farm Schroda,
(-AB), Bruce 58 (PRE, holo.!; K!).
Diagnostic characters: the long, narrow leaves of this
species are distinctive. For other differences, see under
P. cliffordii. Distribution: very localised in southern Zim-
babwe (on Sentinel Ranch) and Northern Province
(mainly on the Farm Schroda) (Figure 7). Habitat: rocky
ridges and on deep clay amongst Hyphaene coriacea
Gaertn. Flowering time: autumn and winter. Conservation
status: P. gillilandiorum is known from only five speci-
mens and is highly localised. One of the habitats (heavy
clay) is being ploughed and planted to cotton, making this
species both rare and threatened.
8. Peristrophe decorticans K.Balkwill in Balkwill
& Getliffe Norris in South African Journal of Botany 55:
254 (1989). Type: Northern Province, Louis Trichardt
Dist., beside Nl, at gate to Plaas Marius, north of Wyl-
liespoort, K.Balkwill 801 (J, holo.!; E!, K!, NU!, PRE!)
P. kotschyana sensu K.Balkwill: 293 (1985); K.Balkwill et al.: 485,
488 (1985); K.Balkwill et al.: 514-520 (1986), non Nees.
Diagnostic characters: P. decorticans differs from P.
paniculata, with which it has often been confused, by its
perennial (not annual) habit, woody (not herbaceous) stem
bases, white, peeling (not black) bark, strigose (not glan-
dular-pubescent or glabrous) tertiary bracts, larger
[14.7-19.0 (not 6.6-13.7) mm long] flowers and usually
glabrous (not pubescent) capsule. P. decorticans differs
from P. cernua by its white peeling (not green or white,
non-peeling) bark, strigose (not densely glandular) tertiary
bracts and presence of honey-guides on the lip in the
lower position. Distribution: P. decorticans occurs in Bot-
swana, North-West, Northern Province and Mpumalanga
(Figure 7). Habitat: bushveld, usually in the shade of trees.
Flowering time: plants have been collected in flower
throughout the year, with peaks in December, January,
FIGURE 7. — Distribution of Peristrophe gillilandiorum , •; P. decorti-
cans, ■; P. cernua, A.
Bothalia 26,2 (1996)
91
May, July and November. In cultivation, plants flower
sporadically in all seasons, although they produce more
flowers in autumn and early winter. Conservation status:
this species is widespread in southern Africa and is fair-
ly frequent where it occurs; it is neither rare nor threat-
ened.
9. Peristrophe cernua Nees in Linnaea 15: 374
(1841); Hook.: 126 (1840), nom. nud. as Peritrophe cer-
nua; C.B.Clarke: 85 (1901). Type: Eastern Cape, ‘Inter
ffutices ad flumen Zwartkopsrivier, Octobri’ [ = Inter fru-
tices in campis ad flumen ‘Zwartkopsrivier’ prope prae-
dium Pauli Mare, alt. I (Uitenhage), cum Rhytiglossa
ciliata, floret Octobri — in original description], (-CD),
Ecklon Un. It. 556 (GZU, holo.! 2 sheets, STE!).
Peristrophe oblonga Nees: 375 (1841). Rhinacanthus oblonga (Nees)
Nees: 444 (1847). Type: Eastern Cape, sudostlich vom Katberg, Schu-
miberg, Ecklon (S!).
Justicia caulopsila E.Mey.: 137, 195 (1843), nom. nud.
Peristrophe krebsii C.Presl: 94 (1844); C.B.Clarke: 86 (1901). Type:
Eastern Cape, without precise locality, Krebs pi. cap. exs. n. 251 (BR !).
P. caulopsila E.Mey. ex Nees: 498 (1847); C.Presl: 95 (1844), nom.
nud.; T.Anderson: 48 (1864); Lindau: 331 (1895a); C.B.Clarke: 84
(1901). Type: Eastern Cape, Somerset Div., between Zuurberg Range and
Kleinbruintjieshoogte, 2000-2500 ft, (-AD), Drege (K, lecto. ! ; G-DC
seen on microfiche, here designated).
P. natalensis T.Anderson: 48 (1864); C.B.Clarke 85 (1901); J.H.Ross:
325(1972). Type: KwaZulu-Natal, Port Natal, (-CC), Gueinuus (K,
lecto.!, here designated).
Non P. hensii (Lindau) C.B.Clarke in Burkill & Clarke: 243 (1900);
C.B.Clarke: 85 (1901).
Non P. cernua sensu Compton: 557 (1976).
Evergreen suffruticose perennial up to lm high. Leaves
ovate, acuminate, reniform to cuneate and attenuate at
base, ( 1 1— ) 1 8— 37(— 50) x (5— )8— 1 6(— 23) mm, with multi-
cellular eglandular and short-stalked glandular trichomes;
petiole (1 .0 — )1.2— 5.5(— 14.6) mm long. Inflorescence of
monochasial cymes (inflorescence units), (1)2 or 3(4) um-
bellately arranged, often compounded; inflorescence axis
(1 .6— )3.3— 10.8(— 16.0) mm long; longest peduncle of scor-
pioid cymes ( 1 .7— )4.5— 1 1 .6(— 22.7). Bracts: secondary bracts
lanceolate, (2.2-)2.6-4.5(-6.2) x (0.2-)0.3-0.5(-0.6) mm,
pubescent; tertiary bracts lanceolate, unequal, larger one
(4.7-)6.7-12.0(-16.0) x 1 .0— 1.3(— 1.6) mm, with multicellu-
lar eglandular and large and small short-stalked glandular
trichomes. Calyx: tube 1 mm deep; lobes 3.5 mm long, mar-
gins membranous and ciliate, with large and small glandular
trichomes. Corolla: tube (5.5-)8.0-9.8(-12.0) mm deep,
sericeous, with medium-headed glandular trichomes; lip in
lower position ovate, (7.2-)9.8-16.5(-22.3) x (2.4— )2.9-5.6
(-8.6) mm; lip in upper position narrowly elliptic,
(8.0-)10.0-16.2(-21.3) x (1.6-)2.2-3.4(-4.0) mm; purple
with dark purple honey-guides below filaments on lip in
lower position, and on white background on lip in upper
position. Stamens: filaments with short and long, straight and
curved, multicellular, eglandular trichomes, white; anther
thecae superposed, separated, purple. Disc cupular with cre-
nate top and 2 short stubby awns. Gynoecium: stigma
bilobed; style exserted, sparsely strigose; ovary ovoid, with
curved multicellular eglandular trichomes and stalked glan-
dular trichomes. Fruit a short-stalked clavate capsule, with
glandular and eglandular trichomes, (9.4-)10.4-13.1(-14.0)
x ( 1 .5—) 1 .8— 2.3(— 3.0) mm. Seeds (2.0-)2.2-2.7(-2.9) x
(1 .5— )1 .8— 2.2(— 2.3) mm, rough and tuberculate.
Diagnostic characters: P. cernua is closely allied to P.
decorticans as discussed under the latter. P. cernua can
be separated from P. paniculata by the larger flowers;
from P. transvaalensis by the shorter, narrower tertiary
bracts (cf. Figure IE & H), and from P. cliff ordii and P
gillilandiorum by the lack of curved eglandular trichomes
between the ridges on the stem (Balkwill et al. 1986).
Distribution: northern KwaZulu-Natal to Eastern Cape
(Figure 7). Habitat: P. cernua is usually found in valley
bushveld, and the few specimens that are found out of
this veld type are found in coastal forest and thomveld
communities. Flowering time: plants have been collected
in flower throughout the year, but there is a very marked
peak in collections in July and August. Conservation
status: P. cernua is widespread and common where it oc-
curs; it is neither rare nor threatened.
Identification of the type of P. cernua has been con-
fused by the complexities of Ecklon & Zeyher’s collec-
tions and numbering system. Zeyher collected and
distributed material from the Uitenhage area before initi-
ating a collecting pact with Ecklon (Gunn & Codd 1981).
Until then, Ecklon had not collected near Uitenhage and
had distributed his specimens through a botanical ex-
change called Unio Itineraria. Shortly after their pact was
initiated, Ecklon collected at Uitenhage and wrote ‘A list
of plants found in the district of Uitenhage between the
months of July 1829 and February 1830’, in which Jus-
ticia capensis is listed under Family 40 (Ecklon 1830).
Later he and Zeyher collected in the area together and
distributed specimens with either or both their names on
the labels and after their pact lapsed, Zeyher collected
from there again. Their numbering system was not one
with consecutive numbers for consecutive gatherings, but
rather the localities were coded in numbers on the labels.
Because of the coding system, inconsistencies occurred
during the distribution of material so that specimens from
the same gathering may have different data on the labels
and specimens with the same number, locality and col-
lecting data may be from different gatherings (Gunn &
Codd 1981). Some specimens were distributed with per-
sonal numbers, some with combined numbers and some
with locality numbers. It seems that the number 40 (prob-
ably the family number for Acanthaceae) is applied to any
sheet they collected of P. cernua. Nees (1841) does not
cite a herbarium for the type of P. cernua (and cites his
own herbarium as well as others in 1847). The type was
presumably in his own herbarium and distributed to GZU.
There are two sheets at GZU that were collected by Eck-
lon, distributed through the Unio Itineraria and part of a
mixed gathering labelled Justicia capensis, suggesting that
they are part of the type gathering — these sheets are pre-
sumably the holotype. A sheet has been found at^STE,
bearing the number 556 (now crossed out), the name Jus-
ticia capensis , and an English version of the locality on
the sheets at GZU — this sheet is most likely an isotype.
One or both of the sheets presently housed at Kew, must
represent the gathering(s) to which Hooker (1840) referred
when offering Zeyher’s material for sale. Hooker’s listing
of the species antedates Nees’s description, but as it was
not accompanied by a description, diagnosis or precise
locality, it is a nomen nudum. It is uncharacteristic of Nees
not to quote previous references when they existed, and
unlikely that both Nees and Hooker would have inde-
92
Bothalia 26,2(1996)
pendently decided on the same name for the same species,
so that it appears that Hooker must have seen Nees’s
manuscript before it was published, or that Ecklon and/or
Zeyher may have circulated the specimens with a name
which was taken up by Nees. It would, however, have
been characteristic of Nees to cite the source of a name,
even if he had taken it up from a label. One of the sheets
at Kew bears the name Justicia capensis and a locality in
Ecklon’s hand. The sheets at Kew bearing the number 40
are undoubtedly those to which Hooker (1840) referred,
most likely those to which Nees referred in 1847, but not
those to which he referred in 1841. The Kew sheets also
bear the name Justicia capensis and this may indicate that
these sheets were amongst those to which Ecklon (1830)
referred. It is therefore possible that the material at Kew
is part of the type gathering (although it has the wrong
month on the label). There are two Ecklon & Zeyher
specimens and one Zeyher specimen from the type local-
ity at SAM, all with the number 40, but none of these
was collected in October and so are not considered part
of the type gathering.
The size of the tertiary bracts and flowers of this spe-
cies displays clinal variation (Balkwill et al. 1994). The
smaller structures are present in Eastern Cape and the
larger in KwaZulu-Natal. It is likely that this cline ac-
counts for a number of the synonyms of P. cemua. This
species sometimes produces white individuals ( Balkwill
445) and white populations ( Brink 327). As the description
of P. krebsii differs from P. cemua mainly by the white
corolla, it is possible that this name was applied to a white
form of P. cemua. Ross (1972) recorded P. hensii from
KwaZulu-Natal, but this name refers to a tropical species.
ACKNOWLEDGEMENTS
I am grateful to the Foundation for Research Devel-
opment (Core Programme), the University of the Wit-
watersrand (Herbarium Programme) and the Barkers for
funding, and very grateful to the directors and curators of
the quoted herbaria for the loan of herbarium material. I
appreciated permission to work at E, K and PRE and for
use of their libraries. Maps were prepared with a demon-
stration copy of MAPPIT.
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SPECIMENS EXAMINED
(and not quoted in other recent publications)
Abbott 1144 (9) PRU. Acocks 10164, 15970, 17940 (9) PRE. Adam 728
(4) PRE. Adamson D303 (9) PRE.
Balkwill 168, 169, 326, 335, 402, 441, 445, 450, 463, 464, 466, 468,
469, 772 (9); 764, 803 (5) E, NU. Balkwill & Manning 918a (9) E, NU.
Barker 157, 24068 (9) NBG. Bayliss 4825 (9) RUH; 7984, BR1. B. 307
(9) PRE. Benson sub Moss 15705 (9) J. Biggs M620 (4) PRE. Blackbeard
NBF 869/14 (9) BOL. Bolus 12219 (5); 1652, 13888, 32311 (9) BOL.
Borle 314 (4) NBG, PRE, SRGH. Bourquin 144 (9) NU. Brink 240 (9)
GRA; 327, 328 (9) GRA, PRE. Brown 8801 (4) PRE. Burchell 3263 (9)
K.
Chase 2602 (4) SRGH. Codd 7549 (4) PRE, SRGH; 9290 (9) GRA,
NBG, PRE. Comins 1561 (9) PRE; 359 (9) NU. Compton 19744 (9);
326 (4) NBG. Cooper 161 (9) K; 1618 (9) RUH. Craven 956 (4) WIND.
Dahlstrand 535 (9) GRA, J. De Winter & Leistner 5652 (4) PRE, WIND.
Dinter 39, 2909 (4) SAM; 3004 (4) PRE, SAM. Drummond 6035 (4)
PRE; 5560 (4) PRE, SRGH. Drummond & Seagrief 5177 (4) PRE. Du
Toil 2393 (9) PRE. Dyer 4374 (9) PRE.
Ecklon & Zeyher 40 (9) BOL, K, SAM. Edwards 785 (9) NU.
Fanshawe 6829 (4) SRGH. Fisher 36 (9) NU. Flanagan 721 (9) BOL,
NH, PRE, SAM. Forward sub GRA A1611 (9) GRA. Fourcade 2746 (9)
BOL, STE; 5755 (9) NBG, STE.
Gajadhur 39 (9) UDW. Galpin 14806, sub BOL 32312, sub PRE 11553
(9) BOL, PRE; 2044 (9) BOL; 7749, 7805 (9) PRE. Garrett 20 (9) NU.
Gayapersad 34 (9) UDW. Germishuizen 775, 811 (5) PRE. Giess 10377
(4) PRE, WIND; 10480, 15125 (4) WIND. Giess & Leippert 7317 (4)
NBG, WIND. Giess, Volk & Bleissner 6464 (4) WIND; 5503 (4) PRE,
WIND. Gillen 1293 (9) STE; 3804 (5) STE. Goldblatt 1929 (4) NBG,
PRE, WIND. Gonde 92/74 (4) PRE, SRGH. Govender 4 (9) UDW. Green
188 (9) NH. Grice s.n. (9) NU.
Hartley 990 (5) J. Harvey 3597 (9) BOL. Huntley 2 (9) NU, PRE. Hurler
sub RUH 3283 (9) RUH.
Jacobsen 4086 (673) (4) PRE; 51 (9) NU.
Keerath 2 (9) UDW. Khan 13 (9) UDW. Kunhardt 9 (4) SRGH.
Lambrecht 125 (4) PRE. Le Roux 1045 (4) WIND. Leighton sub NBG
48702 (9) NBG P4; Leippert 4653 (4) WIND. Lewis 4564, 4565 (9)
SAM; 4566 (9) PRE, SAM. Liebenberg 7754 (9) PRE. Undstedt 12 (9)
PRE. Long 685 (9) GRA, MPE, PRE.
Marloth 1312 (4) PRE, STE. Meeuse 9644 (5) PRE. Moll 1713 (9) NU,
PRE.
Naidoo 11 (9) UDW. Nappe r 1254 (4) PRE. NH 26839 (9) NH. Nicholson
1760 (9) PRE.
Obermeyer sub Herb. Tvl Mus. 33568 (9) PRE.
Padwa 279 (4) PRE, SRGH. Paterson 2093 (9) BOL. Pearson 2550 (4)
BOL. Phelan 636 (9) NU. Pilland 16571 (9) BOL.
Range 1358 (4) SAM. Rennie 405 (9) BOL. Rodin 1069 (9) BOL, PRE.
Rogers 13179 (4) BOL, NH, PRE.
Schlechter 2545 (9) COI, PRE; 2999 (9) COI. Seydel 1128 (4) PRE.
Sister Frances sub BOL 15707 (9) BOL. Smith 1293 (4) PRE. Story
1284, 2227 (9) GRA, PRE.
Taylor 3516 (9) NBG; 491 (9) PRE; s.n. (9) GRA. Thakersee 5 (9) UDW.
Thoday sub Herb. Mus. Austr.-Afr. 25204 (9) SAM. Thode 2749, sub
STE 7964 (9) STE; A2736 (9) PRE. Thome sub Herb. Musei Austr.-Afr.
35709 (4) SAM. Thornton 644 (4) PRE, SRGH. Tweedie 3854 (4) SRGH.
Van Breda 866 (9) PRE. Van Son sub Tvl Museum 28678 (4) PRE. Van
Wyk 1650 (9) PRE, PLI. Venter 3821 (9) PRE. Vesey-FitzGerald 1209
(4) SRGH.
Wager sub Tvl Mus. 22382 (9) PRE. Ward 4974 (9) NU, PRE, UDW;
6925 (9) UDW; 741 (9) NH. Wells 1340 (9) NU, PRE. West 1238 (9)
PRE. Wild & Drummond 7129 (4) PRE. Wood 11881 (9) BOL, SAM;
609 (9) BOL, K.
Zeyher 40 (9) SAM.
.
■
Bothalia 26,2: 95-123 (1996)
Studies in the Ricciaceae of sub-Saharan Africa: a provisional key to the
currently known species
S.M. PEROLD*
Keywords: key, Riccia, Ricciaceae, sub-Saharan Africa
ABSTRACT
A provisional key to the currently known Ricciaceae species of sub-Saharan Africa is given and is illustrated with
micrographs of the distal and proximal spore faces of each species, wherever possible.
INTRODUCTION
For the following reasons I have deemed it advisable
to publish a provisional key to the sub-Saharan Ric-
ciaceae, prior to an intended revision of the family of the
entire region projected for the not too distant future: 1, it
has been brought to my notice several times that such a
key would be useful to collectors of the family in tropical
Africa; 2, a key to the spores of the southern African
species is available (Perold 1989e) but no comprehensive
key to the southern African species has been published to
date. Such a key was included in my unpublished Ph.D.
thesis (Perold 199 Id) and it will form part of my treatment
of the Marchantiidae for the Cryptogam series of the
Flora of southern Africa projected for 1997.
In my paper, ‘A survey of the Ricciaceae of tropical
Africa’ (Perold 1995), I deliberately omitted such a key
and listed the species alphabetically, as I only intended to
report on what was currently known of the family in sub-
Saharan Africa, as a preliminary to revising the family in
this very undercollected region. In 1994, in an effort to
encourage the collection of more Riccia specimens, I had
submitted requests to all the African delegates present at
the AETFAT congress in Wageningen, The Netherlands,
asking them to please collect Ricciaceae specimens in
their countries and to send duplicates to me, but I have
had no response from them, only from a few European
collectors who have visited Africa in recent years and to
whom I wish to express my sincere gratitude.
Although this key is therefore largely provisional, I
hope that it will serve as an identification aid to collectors
of African Ricciaceae. Spore micrographs have been in-
cluded to facilitate the task; unless otherwise stated, all
specimens cited in the captions of the figures, are held at
PRE.
ARTIFICIAL KEY TO THE GENERA, SUBGENERA, SECTIONS, GROUPS AND SPECIES OF THE
RICCIACEAE IN SUB-SAHARAN AFRICA
Key to the two genera of the Ricciaceae
la Thalli floating or terricolous; assimilation tissue containing large air chambers in several storeys; scales long, pendent, purple ribbons,
but small in land form, margins dentate; oil cells present; gametangia located only along deep central groove Ricciocarpos
(only one species is included here, namely R. mtcins (L.) Corda) (Figure 1A, B)
lb Thalli with very rare exceptions terricolous; assimilation tissue spongy, containing air chambers (subgenus Ricciella), or else compact, consisting
of cell columns enclosing narrow vertical air canals (subgenus Riccia)', scales small to large, imbricate, mostly rounded, margins smooth,
rarely denticulate; oil cells absent; gametangia located along groove or scattered Riccia (all other species are included here)
Key to the taxa of the genus Riccia
la Thalli covered by a dorsal epidermis of mostly thin- walled, generally chlorophyllose cells, very rarely dorsally bearing cellular outgrowths;
air pores fewer in number than in subgenus Riccia, mostly delimited, often ringed by smaller cells, well spaced, frequently becoming
cavernous; assimilation tissue loosely arranged, spongy, unistratose cell plates enclosing large polyhedral air chambers; ventral scales
small and evanescent to occasionally large and persistent, in 1 or 2 ranks when present; habitat mostly mesic, rarely xeric or aquatic;
spores separating at maturity or remaining in tetrads: (lb on p. 97)
2a Dorsal epidermis a single layer of thin-walled, closely joined, flattened cells (very rarely globose, and then somewhat loosely connected),
interrupted by air pores, becoming cavernous over air chambers or not; scales ventral, mostly hyaline and inconspicuous, evanescent:
(2b on p. 97)
* National Botanical Institute, Private Bag X101, Pretoria 0001.
MS. received: 1996-05-30.
96
Bothalia 26,2(1996)
3a Thalli annual or perennial; sometimes in rosettes; always terrestrial; branches not strap-shaped, 2-15 x (1— )3— 6(— 8) mm; often becoming
slightly to markedly cavernous; monoicous or dioicous; sporangia deeply imbedded or bulging somewhat dorsally or ventrally:
4a Spores separating at maturity:
5a Thalli moderately thick and ± opaque, not translucent; branches 2-8 times wider than thick; storage tissue up to ± Vi the thickness
of thallus; spores (50)65-150(-160) pm in diam., polar, tetrahedral, winged, ornamentation reticulate, rarely with vermiculate
ridges (subgenus Ricciella)'.
6a Thalli not inflated but finely to coarsely spongiose; dorsally not deeply grooved or rarely only apically so; often in complete or
partial rosettes; glaucous green to yellow green, sometimes tinged with red or purple (section Spongodes , group ‘Crystallina’):
7a Thalli monoicous; from above air chamber walls visible or not; spores completely or incompletely reticulate:
8a Thalli blue-green; in complete or partial rosettes; from above air chamber walls not clearly visible; without or with tiny
hyaline ventral scales; spores 52-85 pm in diam., light brown, ornamentation on proximal and distal faces similar:
9a Thalli dorsally crystalline and glistening, with rounded cells in loose double tiers, usually obscuring air pores when
fresh, with age becoming spongiose; scales absent or evanescent; ornamentation on 2 spore faces similar, areolae
regular and complete, on distal face 8-10 across, 7.5-10.0 pm wide, walls thin and much higher at nodes, often
with bifid or trifid processes, triradiate mark on proximal face distinct, each facet with up to 20 areolae, 5. 0-7. 5
pm wide R. crystallina (Figure 1C, D)
9b Thalli dorsally vesicular-areolate, air pores not obscured, soon spongiose; ventral scales tiny; ornamentation on 2 spore
faces similar, areolae regular and mostly complete on both faces, distal face with only 5 or 6 areolae across, 12.5-17.5
pm wide, proximal face with up to 13 areolae on each facet, walls with crenulate edges, mostly not higher at the
nodes and lacking bifid processes R. vulcanicola (Figure IE, F)
8b Thalli yellow-green or blue-green, sometimes developing red or purple-red coloration; in rosettes or in gregarious patches;
from above air chamber walls visible, air chambers large or small; small, purple, ventral scales sometimes present;
spores 65-115 pm in diam., red-brown to black or light brown, ornamentation on distal and proximal faces dissimilar:
10a Thalli green to yellow-green, faintly tinged with red at margins; in rosettes; from above walls of large air chambers
visible or air chambers cavernous; scales absent; spores 85-115 pm in diam., red-brown to black, distal face with
thicker, irregularly bi- or trichotomously branching ridges, proximal face with triradiate mark distinct, areolae incom-
plete R. cavernosa (Figure 2A, B)
10b Thalli blue-green; in gregarious patches; from above finely areolate; scales present, purple or hyaline; spores 65-85 pm
in diam., light brown, distal face with 8-10 areolae across, proximal face with triradiate mark absent but with numerous
tiny, shallow areolae R. moenkemeyeri (Figure 2 C, D)
7b Thalli dioicous; heterothallic with smaller male plants; bright green to yellowish green or grey-green; in rosettes; from above
air chamber walls visible, becoming cavernous or hardly so; spore ornamentation foveolate or vermiculate (section Spon-
godes, group ‘Cupuliferae’):
11a Thalli bright green; in medium-sized rosettes; cavernous with age; dorsal pores fairly conspicuous, soon enlarging with
age; spores 95-122 pm in diam., ornamentation foveolate, with small, deep-set areolae, triradiate mark on proximal face
very prominent; distribution restricted to southern Africa R. cupulifera (Figure 2E, F)
lib Thalli grey-green, slightly purple at margins; in small rosettes; dorsal pores slightly enlarging with age; spores 40-65 pm
in diam., ornamentation with ± radiating vermiculate ridges on both faces, triradiate mark distinct; widely distributed
in tropical and northern Africa, Europe and N America, especially at alluvial sites R. frostii (Figure 3A, B)
6b Thalli inflated to rather flat; usually becoming markedly cavernous; dorsally grooved along entire length or only apically;
rarely in rosettes; green to straw-coloured or whitish, very rarely tinged with purple or red (section Spongodes, group
‘Vesiculosa’);
12a Thalli large and very wide, 5.0-15.0 x 3.5— 5.5(— 8.0) mm; when dry, yellowish to straw-coloured or white; spores 100-150
(-160) pm in diam., with 8-12 areolae across distal face, wing , 10 pm wide and thin:
13a Thalli straw-coloured when dry; deeply grooved along entire length; cavernous in older parts only; spores with areolae on
distal face 10-15 pm wide R. bullosa (Figure 3C, D)
13b Thalli white when dry; mostly only apically grooved; honeycomb-pitted dorsally; spores with areolae on both faces mostly
wider than 10-15 pm, often up to 20 pm R. garsidei (Figure 3E, F)
12b Thalli medium-sized and narrower, up to 12 x 2. 5-3.0 mm; when dry, greyish white to yellowish; spores 88-112 pm in
diam., with 5-8 areolae across distal face, wing not thin but 3-5 pm wide:
14a Thalli occasionally in rosettes; branches broadly ligulate to Ungulate; antheridial necks inconspicuous, hyaline; widespread
in summer rainfall areas of southern Africa R. volkii (Figure 4A, B)
14b Thalli never in rosettes; branches somewhat linear; antheridia with conspicuous purple necks; very rare, only known from
Knysna Dist. in the Western Cape R. rubricollis (Figure 4C, D)
5b Thalli very thin and ± translucent; branches 10-15 times wider than thick; storage tissue vestigial; spores 40-65 pm in diam.,
apolar, globose, wingless, ornamentation with numerous small, low truncate spines or papillae (subgenus Leptoriccia) ....
R. membranacea (Figure 5E, F)
4b Spores remaining in tetrads (subgenus Thallocarpus with two species):
15a Thalli in complete or incomplete rosettes, 10-20 mm across; heterothallic, male plants once or twice furcate, very rarely in
incomplete rosettes; branches small, 2.0 x 0.5 mm, occasionally larger; spores joined together by narrow band or ridge into
tetrahedral tetrads, ornamentation with slender spinules up to 5 pm long R. curtisii (Figure 6A, B)
15b Thalli reportedly in rosettes up to 11 mm across; male plants small; spores joined together by wide band into rhomboidal tetrads,
ornamentation with stout spines, 10-15 pm long R. perssonii (Figure 6C, D)
3b Thalli generally annual; mostly not in rosettes; terrestrial or occasionally aquatic; branches usually linear, strap-shaped or ‘ribbon-like’,
15-20 mm long and up to 2 mm wide, occasionally smaller; dorsally often tinged with violet, seldom lacunose; at times forming
apical stolons; sporangia ventrally bulging markedly (section Ricciella ):
16a Thalli monoicous:
17a Thalli sometimes aquatic; strap-shaped or ribbon-like; widespread in mostly summer rainfall areas of southern Africa and in
tropical Africa; scales few, ventral, proximally split into 2, apically single; sporangia oblique; distal spore face with areolar
walls thick and prominent, proximal spore face with distinct triradiate mark R. stricta (Figure 4E, F)
17b Thalli terrestrial; in small rosettes; branches narrow; ventral scales vestigial; dorsally somewhat lacunose proximally; sporangia
vertical; spores with complete, regular areolae on both faces, proximal face lacking triradiate mark; rare in tropical Africa,
mostly in Europe and N America R. huebeneriana* (Figure 5A, B)
16b Thalli dioicous; terrestrial; linear, thin and lax; distribution restricted to winter rainfall area of the Western Cape; sporangia vertical;
distal spore face with areolar walls thin, proximal face with distinct triradiate mark R. purpurascens (Figure 5C, D)
* Perold (1995) misplaced R. huebeneriana in her classification of African Ricciaceae, as she referred it to section Spongodes, group ‘Crystallina’. It
should be placed in section Ricciella on account of its ventrally protruding sporangia.
Bothalia 26,2 (1996)
97
2b Dorsal epidermis not with thin-walled, single-layered flat cells interrupted by air pores:
18a Dorsal epidermis devoid of cellular outgrowths, but with thick-walled cells, lacking chlorophyll; air pores surrounded by superimposed
ring of smaller, thin-walled cells; thallus narrowly grooved, acutely winged; scales rounded, persistent, laxge, reaching thallus
margins; spores single (subgenus Chartacea) R. schelpei (Figure 6E, F)
18b Dorsal epidermis with cellular outgrowths of very tall, hair-like pillars; air pores surrounded by radially arranged wedge-shaped cells;
thallus broadly grooved, obtusely winged; scales triangular, persistent, large, filamentous apices extending above thallus margins;
spores remaining in globular to tetrahedral tetrads (subgenus Pannosae ) R. tomentosa (Figure 7A, B)
lb Thalli covered by a dorsal ‘epithelium’ of echlorophyllose cells in one or several strata; air pores numerous, small and regular intercellular
spaces; assimilation tissue compact, in vertical rows of chlorophyllose cells separated by mostly very narrow interstitial air canals;
ventral scales small to large; habitat often xeric, sometimes mesic; spores separating at maturity (subgenus Riccia ):
19a Epithelial cells closely associated, in one or two layers, generally orientated regularly and in parallel rows running from median groove
to margin; top cells globose, mammillose or pyriform, outer walls (or cells) often collapsing; scales small to large, rounded (section
Riccia ): (19b on p. 99)
20a Thalli with cilia along margins, occasionally also present over sporangia; ventral scales mostly not conspicuous; sometimes flanks
dark purple (group ‘Ciliatae’):
21a Thalli shortly or distinctly winged laterally; medium-sized to large, 7.0-30.0 x 2.5-10.0 mm; cilia short or long; spores straw-coloured
or reddish brown:
22a Thalli shortly winged, medium-sized; branches 10-12 x 4 mm; cilia tapering, white when dry, up to 300(-400) x 30-50 pm at
base, finely granular, absent over sporangia; spores straw-coloured, (95— )1 10—1 15 pm in diam., with wing 10 pm wide, slightly
undulating; distribution restricted to southern Africa; hygrophyte R. natalensis (Figure 7C, D)
22b Thalli distinctly winged, large; branches 7-30 x 2.5-10.0 mm; cilia 100-325 x 75-150 pm at base, hyaline and shiny; spores
reddish brown, (130-)17(M80(-215) pm in diam., with wing 9— 10(— 15) pm wide; widely distributed in the northern hemisphere
R. gougetiana (Figure 7E, F)
21b Thalli not winged laterally, smaller, less than 8.0 x 1.5 mm; cilia hyaline, dry or wet, long or short; spores brown to black, with
or without wing; distribution widespread:
23a Distribution in southern Africa restricted to winter rainfall region of the Western Cape; hygrophyte; branches 3-6 x 1 mm; cilia
generally quite sparse, 250(^100) pm long, finely granular, absent over sporangia; scales small, purple; flanks green; spores
with wing ± 7.5 pm wide R. crozalsii (Figure 8 A, B)
23b Widespread in summer rainfall areas; xerophytes; cilia crowded, dense, variously long, not granular, present over sporangia;
flanks dark purple; spores mostly wingless:
24a Thalli 5.0-6.0 x 0.9-1 .5 mm; cilia straight to slightly flexuose, up to 950 pm long, smooth; spores 100-120 pm in diam.,
ornamentation reticulate, areolar walls heavily encrusted with papillae R. trichocarpa (Figure 8C, D)
24b Thalli small, 1.0-4.0 x 0.6-0.8 mm; cilia arched, up to 300 pm long, channelled and finely striate; spores 80-90 pm in diam.,
ornamentation reticulate, areolar walls mostly smooth R. microciliata (Figure 8E, F)
20b Thallus margins not ciliate, marginal cells enlarged or not; ventral scales small to laige:
25a Thalli medium-sized; margins apically with row of mammillate cells up to 150 pm long; scales small, not extending to thallus
margins, violet to hyaline; very rare (group ‘Mammillatae’) R. mammifera (Figure 9A, B)
25b Thalli small to large; margins glabrous; scales larger, extending to margins or projecting above, hyaline or variously pigmented:
26a Scales not conspicuous, hyaline or partly hyaline; thallus margins hyaline; groove narrow and deep, persistent along most or
whole length of thallus; dorsal epithelium sometimes with some cell walls thickened (group ‘Squamatae’):
27a Dorsal epithelial and subepithelial cells always with markedly thicker walls; thalli light green, appearing almost waxy; margins
not undulate; flanks green; spores polar, dark reddish brown to black, proximal face densely granulate; distribution widespread
(nearly cosmopolitan) but apparently quite rare in tropical Africa R. sorocarpa (Figure 9C, D)
27b Dorsal epithelial cells with walls rarely thicker; thalli glaucous green to bright green; margins undulate; flanks dark red-brown
or violet; spores almost apolar or polar, dull dark brown to nearly black, reticulate:
28a Thalli large; apices rounded; margins usually strongly and irregularly undulate, overhanging in cross section; spores 110-125
pm in diam., triangular-globular, polar, not winged, uniformly reticulate on both faces, ± 7 areolae across distal face,
15-20 pm wide, triradiate mark on proximal face indistinct; plants rare R. nigerica (Figure 9E, F)
28b Thalli medium-sized or smallish; apices tapering to obtusely rounded; margins not strongly undulate, not overhanging in
cross section, but flanks rising ± steeply, to slightly obliquely; spores smaller, 75-105 pm in diam.:
29a Thallus flanks not appearing vertically ‘striped’; spores almost apolar, subglobular, wingless, proximal face without triradiate
mark, ornamentation on 2 faces similar, distal face with 1 0— 1 2(— 1 4) areolae across, 7.5-12.5 pm wide
R. atropurpurea (Figure 10A, B)
29b Thallus flanks often appearing vertically ‘striped’ due to hyaline margins of scales above purple bases; spores polar, triangular-
globular, winged, proximal face with triradiate mark distinct:
30a Thalli medium-sized; branches up to 2.5 mm wide; spores with 2 faces dissimilar, facets on proximal face almost smooth
or with poorly developed areolar walls, distal face with (5)6-8 areolae across the diam., (10— )12— 22 pm wide, but
areolae often incomplete, areolar walls up to 5 pm high R. lanceolata (Figure 10C, D)
30b Thalli smallish; branches 1.0-1. 5 mm wide; repeatedly furcate; spores with 2 faces similar or dissimilar:
31a Spores with 2 faces similar, distal face with 8 or 9 areolae across the diam., 13-15 pm wide, areolar walls sometimes
± incomplete, rather thick and raised at nodes into sharply truncate, plate-like papillae, 3-5 pm high
R. radicosa (Figure 10E, F)
31b Spores with 2 faces dissimilar, distal face with central areolae ± 10 pm wide, decreasing in size toward margins,
proximal face with numerous small areolae, ± 2.5 pm wide, on each facet R. schweinfurthii (Figure 11 A, B)
26b Scales often large and conspicuous, pigmented, with or without hyaline border or entirely hyaline to white; groove various;
dorsal epithelial cell walls not thickened:
32a Scales dark, black or reddish black to deep violet, shiny; thallus size variable; dorsally glaucous green to green or purplish,
rarely brown:
33a Thalli medium-sized to large; in section 2. 5-5.0 times wider than thick; flanks sloping obliquely:
34a Thalli large, up to 15 x 5 mm; margins winged, overhanging; spores subglobular, apolar, ornamentation reticulate with
thin, high areolar walls R. congoana (Figure 1 1C, D)
34b Thalli medium-sized to large, up to 15 x 2(-4) mm; margins attenuate, not overhanging; spores triangular-globular, polar,
areolar walls thick, low:
35a Distribution restricted to winter rainfall regions of Western and Northern Cape; thalli quite fleshy; scales imbricate, clasped
together along midline when dry; spores 90-125 pm in diam., distal face with short spiralling ridges, proximal face
almost smooth, with scattered pores R. limbata (Figure HE, F)
35b Distribution widespread in summer rainfall regions, but not common; thalli mostly thinner; scales imbricate to proximally
more widely spaced; when dry, opposite scales meeting or not, never clasped together; spores 72-82 pm in diam.,
distal face with areolae in more or less concentric rings, proximal face completely or incompletely reticulate
R. angolensis (Figure 12 A, B)
98
Bothalia 26,2(1996)
33b Thalli smallish to medium-sized; in section 1-2 times wider than thick; flanks steeply rising:
36a Thalli medium-sized; bluish green; scales projecting above thallus margins, edge crenate, rather dull black, dry; spores
golden brown, wingless, densely papillate R. okahandjana (Figure 12C, D)
36b Thalli smallish; glaucous green, rust-brown along margins and proximally; scales appressed, not projecting above thallus
margins, edge smooth, shiny dry or wet; spores light brown to dark brown, winged, incompletely reticulate
R. nigrella (Figure 12E, F)
32b Scales other than black, variously coloured or white; thalli small to medium-sized; dorsally green to yellow-green, white or
brownish;
37a Scales brown or various shades of red, pink or violet:
38a Thallus margins and scales brownish yellow; idioblasts (enlarged cells with brown contents) present throughout thallus;
spores vermiculate; species very rare in sub-Saharan Africa R. macrocarpa (Figure 13 A, B)
38b Thallus margins not brown; scales dark red or pink or violet; idioblasts absent; spores reticulate:
39a Thalli very small, 1.5— 2.5(— 3.0) x 0.7-1. 0 mm; bottle-green, tumid; scales regular, appressed, dark red, white-bordered;
spores 60-78 pm in diam., triangular-globular, polar; distribution restricted to Free State and Eastern Cape
R. pottsiana (Figure 13C, D)
39b Thalli medium-sized to robust; bright green or light green to whitish, sometimes tinged with red or violet; scales purple-red,
wine-red or cherry-red to rose-pink, exceptionally not persistent; spores 75— 125(— 140) pm in diam., mostly globular
to subglobular, apolar, rarely triangular-globular and polar; widespread or rare:
40a Thalli with assimilation tissue containing somewhat wider than usual air canals toward margins; if in rosettes these are
irregular; thallus margins parallel or subparallel; spores always apolar:
41a Thalli robust, up to 12 mm long; not in rosettes; scales smallish, pink-red to violet, distant; perennating by apical
stolons; spores globular, apolar, 90—1 10— ( 1 35) pm in diam., areolae 12— 15(— 18) pm wide, lacking spines at nodes
R. discolor (Figure 13E, F)
41b Thalli smallish, up to 5 mm long; in irregular rosettes; scales not persistent; not perennating by stolons; spores globular,
apolar, 80-90(-95) pm in diam., areolae 12-15 pm wide, at nodes truncate spines 4-8 pm high
R. symoensii (Figure 14 A, B)
40b Thalli with assimilation tissue containing narrow air canals throughout; rarely in incomplete rosettes; thallus margins
generally tapering toward apex; spores mostly apolar, only exceptionally polar:
42a Thalli up to 8.5 x 1. 5-2.0 mm, light green; scales purplish red to deep cherry-red, lacking hyaline border and only
slightly projecting above thallus margins, imbricate; spores triangular-globular, polar, both faces ornamented with
tiny areolae R. erubescens (Figure 14C, D)
42b Thalli 6-12 x 1-4 mm; green, turning grey-green, or not, or pale green and white along margins or light bluish green,
becoming violet or brownish grey with age; scales dark red or rose-pink or purple, with or without hyaline border;
spores apolar:
43a Thalli 7.0-12.0 x 1. 0-2.5 mm; bright green, turning grey-green or pale green and white along margins; scales dark
red or rose-pink, with narrow or wide hyaline border:
44a Thalli 7. 0-9.0 x 1.2- 1.6 mm; bright green, turning grey-green; scales dark red, shiny, with narrow hyaline border;
monoicous; spores ruby-red to black, (80-)85-105 (-110) pm in diam., subglobular, (6-)8-10 areolae across
spore face, 10.0-12.5 pm wide, periphery with prominent projections, up to 7.5 pm long, appearing cogwheel-
like in profile *R. runssorensis (Figure 14E, F)
44b Thalli up to 12.0 x 1.0-2. 5 mm; pale green, along margins white; scales wavy, rose-pink with wide hyaline
border; dioicous; spores rare, light brown, periphery with low projections R. rosea (Figure 15A, B)
43b Thalli 6-12 x 2-4 mm; light bluish green, turning violet to brownish grey with age, or green; scales purple-red or
dark red to purple, without hyaline border:
45a Thalli up to 12 x 2-4 mm; light bluish green, becoming violet to brownish grey with age; in section 2-2V2 times
wider than thick; scales purple-red, projecting slightly beyond thallus margins; spores 70-80 pm in diam., dark
red, margin with numerous low, blunt projections R. saharensis (Figure 23C, D)
45b Thalli 6-7 x 2-3 mm; green; in section up to 7 times wider than thick; scales dark red to purple, shiny, inserted
beneath thallus wings and projecting slightly beyond margins; spores 92.5-105.0 pm in diam., dark red to
almost black, marginal papillae ± 5 pm high R. papillispora (Figure 23E, F)
37b Scales predominantly hyaline or white, wavy or appressed; thalli without or with calcium deposits:
46a Thalli and scales lacking calcium deposits:
47a Thalli up to 7.0 x 1.3-1 .8 mm, white-green to blue-green; dioicous; antheridial necks conspicuous, up to 250 pm long;
spores deep red-brown, wingless, ( 1 05—) 1 1 5— 125 pm in diam., with 11 areolae across distal face; very rare species ....
R. somaliensis (Figure 15C, D)
47b Thalli 5-15 x 2-4 mm, light grey-green; monoicous; antheridial necks inconspicuous, 60-85 pm long; spores dark brown,
narrowly winged or wingless, (7 5— )90— 1 20(— 1 25) pm in diam., with up to 10 areolae across distal face; widespread
species R. lamellosa (Figure 15E, F)
46b Thalli and scales with calcium deposits:
48a Scales large, 850-1250 x 500-750 pm, irregularly wavy to frilly, closely imbricate; thalli mostly 8.0-9.0(-12.0) x 1. 5-2.0
(-4.0) mm; apically grooved; dorsally green, turning white and spongy over sporangia:
49a Thalli in rosettes or gregarious; widespread and quite common; spores with 10-12 round to angular areolae across
diam. of distal face R. albolimbata (Figure 16A, B)
49b Thalli not in rosettes; spores with 14-20 small deep areolae across diam. of distal face; quite rare, distribution apparently
restricted to Northern Cape R. albomata (Figure 16C, D)
48b Scales rather smaller, up to 850 x 500 pm, mostly appressed and regular, imbricate; thalli generally slightly smaller,
7.0— 8.0(— 12.0) x 0.7-2.0(-4.0) mm; apically grooved or along almost the entire length; dorsally dull or shiny:
50a Thalli deeply grooved along most of entire length; dioicous; spores apolar or polar:
51a Thalli dorsally dull grey-green, compact; dorsal epithelial cells often rather thick-walled; air pores triangular or qua-
drangular; scales regular, appressed, base purple-grey; spores apolar, wingless; fairly widespread
R. argenteolimbata (Figure 16E, F)
51b Thalli dorsally glistening, light green to green; finely spongy; dorsal epithelial cells thin-walled, not in regular pattern;
scales apically wavy, soon appressed; spores polar, winged; distribution restricted to eastern mountainous regions
of southern Africa R. montana (Figure 17 A, B)
* In Jones’s (1957) opinion, R runssorensis Steph. and R papillispora Steph. are closely allied plants and judging from the portions of the types that
he had seen, they may be identical. I, however, regard them as different species, the scales of R. papillispora lack a hyaline border and its spores have
shorter papillae, as noted by Jones.
Bothalia 26,2(1996)
99
50b Thalli only apically grooved; monoicous; spores polar:
52a Thalli with wide, shallow groove, 8x4 mm, dorsally very heavily encrusted with calcium deposits; air pores large,
regular; scales white; spores finely reticulate R. alboporosa (Figure 17C, D)
52b Thalli narrowly grooved, up to 5 x 1-2 mm, dorsally with fine calcium deposits; air pores small to large, irregular;
scales bicoloured, with deep purple base; spores more coarsely reticulate R. bicolorata (Figure 17E, F)
19b Epithelial cells in free-standing 2-5(6)-celled, uniseriate pillars, irregularly orientated; top cells variously shaped, soon collapsing;
scales small to large, mostly rounded and smooth-margined, rarely triangular and dentate or apically filiform (section Pilifer );
53a Dorsal pillars short, often less than 200 pm long, consisting of 2 or 3(4) cells, mostly wider than long, tapering or not tapering:
54a Dorsal pillars tapering; air canals up to 100 pm wide:
55a Spores wide-winged, wing up to 10 pm wide, elaborately ornamented; very rarely found:
56a Thalli large, up to 10.0 x 2.5-3.8 mm; scales hyaline R. hantamensis (Figure 18A, B)
56b Thalli small, up to 5.0 x 0.9-1. 5(— 2.0) mm; scales red R. alatospora (Figure 18C, D)
55b Spores narrow-winged, wing up to 5 pm wide; ornamentation less elaborate, especially on proximal face, where often reduced
to simple projections and stipplings; widespread R. albovestita (Figure 18E, F)
54b Dorsal pillars not tapering; air canals narrow:
57a Thalli quite large, up to 10 x 3-4 mm; broadly ovate to obovate; in section times wider than thick; flanks sloping obliquely;
distal spore face with 10-14 areolae across diam., sometimes with central papilla or short radiating ridges
R. concava (Figure 19 A, B)
57b Thalli smaller, up to 8 x 1-2 mm; ligulate to ovate; in section as wide as, to twice wider than thick; flanks steep; spores variously
ornamented:
58a Branches frequently simple, long and narrow; spore distal face with 5-7 large, incomplete areolae across diam., often with
central boss R. elongata (Figure 19C, D)
58b Branches mostly several times furcate; spore distal face with more than 7 smaller areolae across diam., lacking central boss:
59a Branches apically keeled to wedge-shaped; margins somewhat tumid; dorsal cell pillars up to 180 pm long; proximal spore
face reticulate:
60a Dorsal pillars with top cell mammillose; distal spore face with ridges generally forming a central cross; distribution restricted
to Western and Northern Cape R. furfuracea (Figure 19E, F)
60b Dorsal pillars with top cell globose; distal spore face with ± 8 angular, irregular areolae across diam.; distribution apparently
restricted to Lesotho R. trachyglossum (Figure 20A, B)
59b Branches apically rounded; margins not tumid; dorsal cell pillars only 70-105 pm long; proximal spore face granulate . . .
R. pulveracea (Figure 20C, D)
53b Dorsal pillars tall, longer than 200 pm and up to ± 450 pm (rarely 1000 pm) long, consisting of (3)4-6 narrow, elongated cells,
(l‘/2-)2-3 (or more) times longer than wide:
61a Dorsal surface of thallus generally somewhat velvety or furry when fresh; emerald-green to lighter green; pillars gradually tapering
to narrower apical cell:
62a Scales rounded, wavy, large, 1500 x 600-900 pm, margin smooth; thallus emerald-green; basal cells of pillars almost equally
long, walls somewhat thickened; spores polar, ornamentation finely or coarsely reticulate R. simii (Figure 20E, F)
62b Scales triangular, very large, up to 1800 pm long; basal cells of pillars variably long, walls not thickened; spores variously
ornamented:
63a Triangular scales with dentate margins; dorsal pillars up to 450 pm long; spores papillose; not rare, but distribution restricted
to the Northern and Western Cape R. villosa (Figure 21 A, B)
63b Triangular scales with filamentous apices; dorsal pillars up to 1000 pm long; spore ornamentation incompletely reticulate; very
rare R. hirsuta (Figure 2 1C, D)
61b Dorsal surface of thallus rarely velvety or furry; steel-grey to bright green or olivaceous green; cell pillars not, or hardly tapering:
64a Dorsally steel-grey; dorsal pillars tall (up to 450 pm long), like slivers of glass, often interlocking over groove; thalli larger, up
to 9 mm long; scales large and billowing, basal cell walls appearing ‘stretched’; proximal spore face incompletely reticulate
and granulate, distal face with a few short, well-spaced, radiating, central ridges R. vitrea (Figure 21E, F)
64b Dorsally olivaceous green or crystalline; dorsal pillars shorter (mostly less than 350 pm long), not interlocking, or if so, only
temporarily toward apex; thalli mostly less than 8 mm long; scales smaller and not billowing; spore ornamentation completely
reticulate on proximal face, not granulate, distal face reticulate or with several, long, thick, radiating ridges:
65a Dried thalli with brown, parchment-like flanks, proximally almost without scales; wet thalli velvety, olivaceous green; distal
spore face with thick radiating ridges R. albomarginata (Figure 22A, B)
65b Dried thalli frequently with somewhat purple flanks; wet thalli rather crystalline, bright green or purplish green:
66a Thalli in section 1 .5 times to nearly twice wider than thick; sides tightly indexed when dry .... R. namaquensis (Figure 22C, D)
66b Thalli in section 2-4 times wider than thick; sides incurved when dry:
67a Cells in dorsal pillars mostly somewhat constricted in middle, ampulla-shaped; spores 90-95(-105) pm in diam., distal face
with several radiating ridges; distribution restricted to eastern mountains in southern Africa R. ampullacea (Figure 22E, F)
67b Cells in dorsal pillars not constricted; spores 70-80(-90) pm in diam., ornamentation with numerous small areolae; distri-
bution restricted to Western Cape R. parvo-areolata (Figure 23A, B)
ACKNOWLEDGEMENTS
I wish to express my sincere thanks to Mrs J. Mul-
venna, NBI, for kindly typing the manuscript and to Dr
O.A. Leistner and Mrs B.A. Momberg for their assistance
and support. The curators of BM, BOL, BR, E, G, H and
PC, Dr T. Bomefeld and Prof. O.H. Volk are sincerely
thanked for the loan of specimens.
SOME LITERATURE REFERENCES TO SUB-SAHARAN
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BERRIE, G.K. 1975. The biology of a West African species of Riccia L.
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FISCHER, Eb. 1993. Taxonomic results of the BRYOTROP Expedition
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FISCHER, Eb. 1995. The genera Ricciocarpos and Riccia (Hepaticae,
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FRAHM, J.-P. & POREMBSKI, S. 1994. Moose von Inselbergen aus
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PEROLD, S.M. & VOLK, O.H. 1988a. Studies in the genus Riccia
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STEPHANI, F. 1888. Hepaticae africanae. Hedwigia 27: 106-113.
STEPHANI, F. 1895. Hepaticae africanae. Botanische Jahrbiicher 20:
299-321.
STEPHANI, F. 1898. Species hepaticarum. Bulletin de THerbier Boissier
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VANDEN BERGHEN, C. 1952. Note sur un Riccia de Mauritanie. Revue
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VANDEN BERGHEN, C. 1965. Hepatiques recolt6es par le Dr J.-J.
Symoens dans la region pdri-Tanganyikaise. Bulletin de la Societe
Royale de Botanique de Belgique 98: 129-174.
VANDEN BERGHEN, C. 1972. H£patiques et anthoc6rot6es. In J.J.
Symoens, Resultats scientifiques de T exploration hydrobiolo-
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Cercle Hydrobiologique de Bruxelles, Bruxelles.
VANDEN BERGHEN, C. 1978. Hepatiques du Shaba. Corrections et
additions. Bulletin du Jardin botanique national de Belgique 48:
367-372.
VOLK, O.H. 1984. Pflanzenvergesellschaftungen mit Riccia-Aiten in
Sudwestafrika (Namibia). Vegetatio 55: 57-64.
VOLK, O.H. & PEROLD, S.M. 1986a. Studies in the genus Riccia
(Marchantiales) from southern Africa. 4. Three endemic species,
R. natalensis, R. microciliata sp. nov. and R. mammifera sp. nov.
Bothalia 16: 169-180.
VOLK, O.H. & PEROLD, S.M. 1986b. Studies in the genus Riccia
(Marchantiales) from southern Africa. 5. R. rosea, a new species.
Bothalia 16: 181-185.
VOLK, O.H., PEROLD, S.M. & BORNEFELD, T. 1988. Studies in the
genus Riccia (Marchantiales) from southern Africa. 10. Two new
white-scaled species of the group ‘Squamatae’: R. argenteolim-
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Bothalia 26,2 (1996)
101
FIGURE 1. — SEM micrographs of spores. A, B, Ricciocarpos natans : A, distal face; B, proximal face. C, D, Riccia
crystallina: C, distal face; D, proximal face. E, F, R. vulcanicola : E, distal face; F, proximal face. A, B, Ward s.n.;
C, D, Duthie 5313 (BOL); E, F, Poes 8068/158. A-D, x BOO; E, F, x 700.
102
Bothalia 26,2(1996)
FIGURE 2. — SEM micrographs of spores. A, B,Riccia cavernosa : A, distal face; B, proximal face. C, D, R moenkemeyeri :
C, distal face; D, proximal face. E, F, R. cupulifera: E, distal face; F, proximal face. A, B, Shearing 17H\ C, D, T.R.
Sim 9072\ E, F, Out hie 7471. A-F, x 800.
Bothalia 26,2 (1996)
103
FIGURE 3. — SEM micrographs of spores. A, B, Ricciafrostii : A, distal face; B, proximal face. C, D, R. bullosa: C, distal
face; D, proximal face. E, F, R. garsidei: E, distal face; F, proximal face. A, B, Monod 10537 (PC); C, Duthie
5486a ; D, S.M. Perold467\ E, F, S.M. Perold536. A, x 800; B, x 900; C, x 600; D-F, x 500.
104
Bothalia 26,2(1996)
FIGURE 4— SEM micrographs of spores. A, B, Riccia votkii : A, distal face; B, proximal face. C, D, R. rubricollis: C,
distal face; D, proximal face. E, F, R. stricta : E, distal face; F, proximal face. A, B, Volk 81/230', C, D, Duthie 5014\
E, F, M. Koekemoer 988. A, B, E, F, x 700; C, D, x 600.
Bothalia 26,2 (1996)
105
FIGURE 5. — SEM micrographs of spores. A, B, Riccia huebeneriana : A, distal face; B, proximal face. C, D, R.
purpurascens: C, distal face; D, proximal face. E, F, R. membranacea: E, distal face; F, proximal face. A, B,
Lubenau-Nestle s.n.\ C, D, MacLoughlin CH4197\ E, F, Jones 672 (BM). A, B, x 750; C, D, x 650; E, F, x 870.
106
Bothalia 26,2(1996)
FIGURE 6.— SEM micrographs of spores. A, B, Riccia curtisii: A, B, tetrads. C, D, R. perssonii: tetrads. E, F, R. schelpei :
E, distal face; F, proximal face. A, B, S.M. Perold 2 730\ C, D, Volk 2059', E, F, CM. van Wyk 2524. A-D, F, x
600; E, x 550.
Bothalia 26,2(1996)
107
FIGURE 7. — SEM micrographs of spores. A, B, Riccia tomentosa: tetrads. C, D, R. natalensis: C, distal face; D, proximal
face. E, F, R. gougetiana: E, distal face; F, proximal face. A, B, S.M. Perold 1495', C, S.M. Perold 307\ D, T.R. Sim
8228', E, F, Volk 82/980 (Herb. Volk). A-D, x 600; E, F, x 400.
108
Bothalia 26,2(1996)
FIGURE 8. — SEM micrographs of spores. A, B, Ricciu crozalsii: A, distal face; B, proximal face. C, D, R. trichocarpa :
C, distal face; D, proximal face. E, F, R. microciliata : E, distal face; F, proximal face. A, Duthie 5436a (BOL); B,
Malherbe & Davies 5373 (BOL); C, D, Toelken 5561 \ E, F, S.M. Perald 102. A-F, x 800.
Bothalia 26,2 (1996)
109
FIGURE 9. — SEM micrographs of spores. A, B, Riccia mammifera : A, distal face; B, proximal face. C, D, R. sorocarpa:
C, distal face; D, proximal face. E, F, R. nigerica: E, distal face; F, proximal face. A, B, S.M. Perold 447\ C, D,
S.W. Arnell 303\ E, F, Jones 1167 (BM). A, B, x 600; C, D, x 800; E, F, x 400.
110
Bothalia 26,2(1996)
FIGURE 1 0, — SEM micrographs of spores. A, B, Riccia atropurpurea: A, distal face; B, proximal face. C, D, R. lanceolata :
C, distal face; D, proximal face. E, F, R. radicosa ; E, distal face; F, proximal face. A, S.M. Perold 782b; B, Volk
84/710 ; C, D, Porembski & Biedinger 1313 ; E, F, Vanderyst s.n. (BR). A, B, x 600; C, D, x 450; E, F, x 500.
Bothalia 26,2(1996)
111
FIGURE 1 1 . — SEM micrographs of spores. A, B, Riccia schweinfurthii : A, distal face; B, proximal face. C, D, R. congoana:
C, distal face; D, proximal face. E, F, R limbata: E, distal face; F, proximal face. A, B, Schweinfurth 1832 (H); C,
D, Volk 00747a ; E, F, Oliver 8858. A-D, x 450; E, F, x 600.
112
Bothalia 26,2(1996)
FIGURE 12.— SEM micrographs of spores. A, B, Riccia angolensis: A, distal face; B, proximal face. C, D, R. okahandjana:
C, distal face; D, proximal face. E, F, R. nigrella : E, distal face: F, proximal face. A, B, S.M. Perold 2466a\ C, S.M.
Perold 267 ; D, Volk 84/707 ; E, F, S.M. Perold 1147. A-D, x 600; E, F, x 800.
Bothalia 26,2 (1996)
113
FIGURE 13. — SEM micrographs of spores. A, B, Riccia macrocarpa: A, distal face; B, proximal face. C, D, R. pottsiana:
C, distal face; D, proximal face. E, F, R. discolor. E, distal face; F, proximal face. A, B, S.M. Perold HH8\ C, D,
Duthie 5463a (BOL); E, F, Bingham 8108. A, B, x 700; C, D, x 800; E, F, x 600.
114
Bothalia 26,2(1996)
FIGURE 14. — SEM micrographs of spores. A, B, Riccia symoensii: A, distal face; B, proximal face. C, D, R. erubescens :
C, distal face; D, proximal face. E, F, R. runssorensis: E, distal face; F, ? proximal face. A, B, Symoens 7075 (BR);
C, D, Townsend 75/1 7; E, S.M. Perold 2466; F, Volk 8 1/1 25c. A, B, x 700; C, D, x 400; E, F, x 600.
Bothalia 26,2 (1996)
115
FIGURE 15. — SEM micrographs of spores. A, B, Riccia rosea : A, distal face; B, proximal face. C, D, R. somaliensis : C,
distal face; D, proximal face. E, F, R. lamellosa; E, distal face, F, proximal face. A, B, S.M. Perold 135a; C, D,
Thulin &. Mohamed 7110 ; E, F, Faure s.n.( E). A, B, x 700; C, D, x 420; E, F, x 550.
116
Bothalia 26,2 (1996)
FIGURE 16. — SEM micrographs of spores. A, B, Riccia albolimbata: A, distal face; B, proximal face. C, D, R. albornata:
C, distal face; D, proximal face. E, F, R. argenteolimbata: E, distal face; F, proximal face. A, Volk HI/921, B,
Stephansen 5393; C, Volk 81/081; D, Smook 6961; E, F, Volk 86/930a. A, B, x 600; C, D, x 700; E, F, x 600.
Bothalia 26,2 (1996)
117
FIGURE 17. — SEM micrographs of spores. A, B, Riccia montana : A, distal face; B, proximal face. C, D, R. alboporosa:
C, distal face; D, proximal face. E, F, R. bicolorata ; E, distal face; F, proximal face. A, B, Van Rooy 3549a ; C, D,
Oliver 8849 ; E, F, Smook 6990a. A, B, x 800; C-F, x 700.
118
Bothalia 26,2 (1996)
FIGURE 1 8. — SEM micrographs of spores. A, B, Riccia hantamensis : A, distal face; B, proximal face. C, D, R. alatospora:
C, distal face; D, proximal face. E, F, R. albovestita : E, distal face; F, proximal face. A, B, S.M. Perold 1830 ; C,
D, Duthie 5004b\ E, F, J.M. Perold 44. A, B, x 800; C, D, x 500; E, F, x 700.
Bothalia 26,2 (1996)
119
FIGURE 19.- — SEM micrographs of spores. A, B, Riccia cuncava : A, distal face; B, proximal face. C, D, R. elongata : C,
distal face; D, proximal face. E, F, R. furfuracea: E, distal face; F, proximal face. A, B, 5.1V. Arnell 30; C, D, S.A7.
Perold 2018\ E, F, Oliver 8957a. A, B, x 700; C-F, x 600.
120
Bothalia 26,2(1996)
FIGURE 20. — SEM micrographs of spores. A, B, Riccia trachyglossunv. A, distal face; B, proximal face. C, D, R.
pulveracea : C, distal face; D, proximal face. E, F, R simii: E, distal face; F, proximal face. A, B, J.M. Perold 34\
C, Oliver 8957a; D, Duthie 5484a ; E, F, Smook 3908. A, B, x 600; C, D, x 700; E, F, x 800.
Bothalia 26,2 (1996)
121
FIGURE 21. — SEM micrographs of spores. A, B, Riccia villosa: A, distal face; B, proximal face. C, D, R. hirsuta: C, distal
face; D, proximal face. E, F, R. vitrea: E, distal face; F, proximal face. A, B, Levyns s.n. \ C, D, Oliver 8040\ E, F,
S.M. Perold 1425. A, B, x 800; C-F, x 600.
122
Bothalia 26,2(1996)
FIGURE 22. — SEM micrographs of spores. A, B Riccia albomarginata: A, distal face; B, proximal face. C, D, R.
namaquensis: C, distal face; D, proximal face. E, F, R. ampullacea: E, distal face; F, proximal face. A, B, S.M.
Perold 2385; C, D, S.M. Perold 1420\ E, F, Van Rooy 3164a. A, B, x 600; C, D, x 800; E, F, x 700.
Bothalia 26,2 (1996)
123
FIGURE 23. — SEM micrographs of spores. A, B, Riccia parvo-areolata: A, distal face; B, proximal face. C, D, R.
saharensis: C, distal face; D, proximal face. E, F, R. papillispora : E, distal face; F, proximal face. A, B, J.M. Perold
24 ; C, D, Chudeau s.n. (G); E, F, Beccari 802 (G). A, B, x 700; C, D, x 560; E, F, x 450.
Bothalia 26,2: 125-140(1996)
FSA contributions 6: Orchidaceae: Holothrix
K.L. IMMELMAN*
1408000 HOLOTHRIX
Holothrix Rich, ex Lindl., The genera and species
of orchidaceous plants: 283 (1835), nom. cons.; Benth. &
Hook.f. 3: 623 (1883); Schltr.: 443 (1898); Schltr.: 12
(1899a); Schltr.: 202 (1899b); Rolfe: 96 (1913); Schelpe:
66 (1966); Schelpe: 5 (1977); R.A.Dyer: 989 (1976); Im-
melman: 455 (1981); J.L.Stewart et al.: 58 (1982). Type:
H. parvifolia Lindl. (1835) nom. illegit., type cons. ( = H.
brevipetala Immelman & Schelpe).
Scopularia Lindl.: t. 1701 (1834)
Monotris Lindl.: t. 1701 (1834)
Tryphia Lindl.: t. 1701 (1834)
Saccidium Lindl.: 302 (1835)
Bucculina Lindl.: 209 (1836)
Deroemera Rchb.f.: 29 (1852); Rendle: 277 (1895) as
Deroemeria.
Terrestrial or lithophytic tuberous herbs. Leaves two,
one sometimes much reduced (or absent?), ovate or or-
bicular, flat on ground, sometimes withered before flow-
ering. Scape arising between two leaves, erect, unbranched,
usually pubescent, with or without bracts. Inflorescence a
spike, usually secund or subsecund. Flowers resupinate
(except in H. majubensis), from 1.5 up to 25 mm long.
Sepals subequal, sometimes connate, green, smaller than
petals. Petals free or partly adnate to lip, green, cream,
white or marked with red or purple, entire or with apex
fimbriate, if entire then apices either thickened and fleshy
or whole petal membranous; lip fleshy or membranous,
entire or divided into three to many lobes, sometimes fim-
briate, coloured as for petals, produced into a spur. Col-
umn usually small, simple and erect, fused basally to lip,
in one species forming an open arch; anther sacs above
stigma; rostellum much reduced. Pollinia two, in anther
sacs, granular, each with a short caudicle and separate
viscidium; viscidia naked, often resting on small flaps of
tissue on either side of column.
Species about 55, distributed from Arabia through east-
ern and West tropical Africa, and the Cameroons, to South
Africa; also in Madagascar, the Comoro Islands and So-
cotra. There are 23 species in southern Africa, with the
greatest concentration in the Western Cape (see Guide for
authors to Bothalia, p. 208: figure 1).
All measurements of lip and petal breadth are taken
across the base of the lobes where these occur, except
where otherwise stated. In undivided petals and lip the
breadth is taken at the broadest point. Length is the total
length of lip or petal. Scape length is also the total length
of the scape, including the flowering portion.
In some of the species, details of flower colour, inflo-
rescence and papillae are given. This has been possible
where living or pickled specimens were seen. It is of in-
terest to note that in all these the anther sacs are usually
a darker and contrasting colour to the rest of the column,
and to the flower as a whole.
Holothrix is derived from the Greek words for ‘com-
pletely hairy’, and refers to the squamous or hispid vege-
tative parts of the type species.
Key to southern African species of Holothrix
la Scape with bracts:
2a Petals entire 19 . H. culveri
2b Petals divided:
3a Flowers dimorphic 20. H. burchellii
3b Rowers not dimorphic:
4a Lip lobes filamentous; Gauteng and Northern Province 21. H. randii
4b Lip lobes thickened; Northern, Western and Eastern Cape:
5a Spur Vi to nearly equal length of lip; scape with a covering of short, fine, somewhat velvety hairs 22. H. schlechteriana
5b Spur 1/5-V4 length of lip; scape usually glabrous but sometimes with a few short hairs at base 23. H. grandiflora
lb Scape without bracts:
6a Petals divided at apex into 3 or 5 lobes:
7a Spur 114-2 times length of lip; petals 5-lobed; Namaqualand (Northern Cape) 12. H. filicomis
7b Spur (4-equal length of lip; petals 3-lobed; Eastern Cape, KwaZulu-Natal, Lesotho, Mpumalanga 13. H. scopularia
6b Petals entire:
8a Petals green or cream, camose at apices:
9a Rowers not resupinate 7. H. majubensis
9b Rowers resupinate:
10a Petals linear; lip lobes short and triangular with obtuse apices 1. H. pilosa
10b Petals broader at centre than at apex; lip lobes various:
11a Lip entire or three-lobed:
12a Spur longer than lip 2. H. longicomu
12b Spur shorter than lip:
* Range and Forage Institute, Private Bag X05, East Lynne 0039, Pretoria.
MS. received: 1990-01-30.
126
Bothalia 26,2(1996)
FIGURE 1. — A-E, Holothrix pilosa, Schelpe s.n.\ A, habit; B, plant with leaf before shrivelling and tubers; C, flower, side view; D, flower, front
view; E, column. F-J, H. villosa var. villosa, Esterhuysen s.n F, habit; G, flower, side view; H, flower, front view; I, lip from above, J,
column. A, E-H, J, drawn by K. Immelman; B-D, I, drawn by G. Condy. Scale bars: A, B, F, 10 mm; C-E, G-J, 1 mm.
Bothalia 26,2 (1996)
127
13a Lip entire 3 . H. exilis
13b Lip 3-lobed:
14a Breadth of central lip lobe at least half its length 4. H. brevipetala
14b Breadth of central lip lobe less than half its length, linear:
15a Lip lobes subequal in length:
16a Hairs on leaves usually broad and flattened (squamules), these rarely restricted to margins, hairs on scape reflexed
and hispid 5. H. cemua
16b Leaves villous, hairs on scape at right angles to scape:
17a Scape slender; inflorescence lax; lip lobes narrowly linear 6a. H. villusa var. villosa
17b Scape short and stout; inflorescence dense; lip lobes broadly linear 6b. H. villosa var. condensata
15b Side lobes of lip half or less than half of length of central lobe:
18a Leaf usually withered at anthesis, with hispid hairs; Drakensberg 8. H. thodei
18b Leaf not usually withered at anthesis, with fine soft hairs, Western and Eastern Cape, mainly lowland 3 . H. exilis
lib Lip lobes 5-7:
19a Gauteng, Free State, Lesotho, KwaZulu-Natal, if Eastern Cape then montane and hairs villous:
20a Petals more than 5 mm long; flowering December to April 10 . H. incurva
20b Petals less than 5 mm long; flowering September and October 11.//. micrantha
19b Western Cape, if Eastern Cape then not montane and hairs squamous or hispid:
21a Spur longer than lip 2. H. longicornu
21b Spur shorter than lip:
22a Base of petals adnate to lip; leaves glabrous 9. H. secunda
22b Petals free from lip; leaves with hairs or squamules:
23a Length of lip lobes usually less than twice the breadth 4. H. brevipetala
23b Lip lobes linear, more than twice as long as broad 5. H. cemua
8b Petals white, membranous at apices:
24a Lip lobes 5:
25a Spur circinnate; leaves uniform in colour; bases of petals adnate to lip 14 . H. parviflora
25b Spur straight; leaves veined with white; petals free from lip 15 . H. orthoceras
24b Lip lobes 7 or more, or indefinite with lip serrate along margin:
26a Lip lobes 7:
27a Lip lobes acute, broadly triangular except for a longer and narrower lobe on either side; petals tapering to an acuminate apex;
leaves veined with white; east of Port Elizabeth 15 . H. orthoceras
27b Lip lobes truncate or obtuse, lobes on either side of central lobe shorter than central lobe; petal apices rounded; leaves uniformly
green; mostly west of Port Elizabeth:
28a Inflorescence condensed at apex of spike; spur short and straight 16 . H. mundii
28b Inflorescence lax; spur curved abruptly forward under lip 17 . H. aspera.
26b Lip lobes more than 7, or lip serrated:
29a Outermost lip lobe on either side longer than central lobes; leaves veined with white; Northern Province to Eastern Cape . . .
15. H. orthoceras
29b Lobes of lip subequal, very short, giving lip a serrated appearance; leaves not veined with white; Eastern Cape
18 . H. macowaniana
1. Holothrix pilosa (Burch, ex Lindl.) Rchb.f. in
Otia botanica hamburgensia: 119 (1881); J.L. Stewart et
al.: 60 (1982). Type: Swellendam Div., on a dry hill east
of Breede River, Burchell 7483 (K!).
Saccidium pilosum Lindl.: 302 (1835).
Leaves large, succulent, upper surface glabrous, rarely
with hairs, undersurface densely pilose with long, fine
hairs, usually withered at anthesis; upper leaf reduced in
size. Scape without bracts, 160-550 mm long, densely
pilose with long, fine hairs, hairs recurved at base of
scape, at right angles to scape at its apex. Inflorescence
secund. Sepals 2.5-5 .0 x 1. 0-2.5 mm, glabrous, edges no-
ticeably paler than bright green centre. Petals entire, car-
nose at apices, narrowly linear, 5.5-10.0 x 0.5-1. 0 mm,
creamy white with a green centre; lip divided into (3— )5— 8
short, broad, obtuse, camose lobes, long and narrow, 5. 5-9.0
x 4.5-9.0, cream with green veins. Spur broadly conical,
straight or slightly curved, 1. 5-5.5 mm long. Anther sacs
yellow. Figure 1A-E.
H. pilosa occurs in the southern Cape region (Figure
2) from Bredasdorp (Western Cape) to Port Elizabeth
(Eastern Cape), and grows in semi-arid, stony localities.
Flowering time: November to March.
The long, linear petals and long narrow lip, which to-
gether form a narrow cylindrical tube, are characteristic
of this species. The inflorescence is strictly secund.
Vouchers: Linder 1704 (BOL); Long 938 (PRE); Muir 1225 (BOL);
Thode A26392 (PRE).
2. Holothrix longicornu G.J. Lewis in Journal of
South African Botany 4: 53 (1938); J.L.Stewart et al.: 60
(1982). Type: Port Elizabeth, Cutting 69 (BOL, holo.!).
Figures 2; 8N.
Leaves with dense, short, fine, retrorse hairs. Scape
without bracts, 65-154 mm long, with dense, short, re-
FIGURE 2. — Distribution of Holothrix pilosa , □; H. longicornu. •.
128
Bothalia 26,2 (1996)
FIGURE 3. — A-F, Holothrix exilis: A, habit; B, flower, side view; C, flower, front view; D, lip with short side lobes, Esterhuysen 21205 ; E, lip
without side lobes, Esterhuysen 18356', F, column. G— J, H. secundu, Schelpe 7900'. G, habit; H, flower, side view; I, flower, front view, J,
column. Drawn by K. Immehnan. Scale bars: A, G, 10 mm; B-F, H-J, 1 mm.
Bothalia 26,2 (1996)
129
curved hairs. Sepals 1. 5-2.5 x 1.0 mm, glabrous, connate
at base. Petals entire, camose at apices, 2.5-3.5 x 0.5
mm; lip camose, divided into 3-5 short, broadly linear
lobes, 2. 5-3.0 x 1.0-1. 5 mm. Spur narrowly conical,
straight, longer than lip, 3-4 mm long.
Holothrix longicomu is probably closely related to H.
cemua (No. 5) and H. brevipetala (No. 4), all three spe-
cies having recurved, hispid hairs on the scape. However,
the short lip lobes (Figure 8N) distinguish it from the for-
mer, and the long, straight spur distinguishes it from both.
Only the type has been seen but, as there are a number
of plants in the type gathering, and they consistently show
these characters, it can be regarded as a species separate
from both the above-mentioned species.
Voucher: the only specimen of this species is the type, which was
collected in October. The habitat is not known.
3. Holothrix exilis Lindi, The genera and species
of orchidaceous plants: 283 (1835); Immelman: 456
(1981); J.L. Stewart et al.: 59, t. 1.1 (1982). Type: in a
walk to the white clay pit, bearing true N-E from our
station at Zoetmelksrivier, Burchell 6738-1 (K! ).
H. exilis var. brachylabris (Sond.) Bolus: t. 14A (1896). H. brachylabris
Sond.: 78 (1847). Type: Uitenhage, Zeyher s.n. (K!)
Leaves densely to very sparsely pilose with long, fine
hairs, small, sometimes withered at anthesis. Scape with-
out bracts, slender, 40-290 mm long, densely to very
sparsely pilose with long, fine hairs at right angles to
scape. Inflorescence usually lax with small flowers. Sepals
0.8-2.5 x 0.5-1. 0 mm, usually glabrous or with a few
long hairs. Petals entire, with carnose apices, 1. 5-4.0 x
0. 3-1.0 mm, creamy green; lip camose, entire or three-
lobed with outer lobes from very short to half as long as
central lobe, 1. 8-3.5 x 0.5-1. 5 mm, creamy green. Spur
slightly curved, 0.8-1. 8 mm. Anther sacs pale yellow-
green. Figure 3A-F.
H. exilis occurs in the coastal regions of the Western
and Eastern Cape from the Saldanha area, the Cape Pen-
insula and Riversdale to Kei Mouth (Figure 4). Flowering
time: October to March.
H. exilis can be distinguished from H. villosa (No. 6)
by the side lobes of the lips which are up to 0.4 times
the length of the central lobe, or absent, while H. villosa
has side lobes (always present) from 0.6 to 1.0 times the
length of the central lobe.
The variety H. exilis var. brachylabris was described
by Bolus on the basis of an unlobed rather than a three-
lobed lip. However, the length of the side lobes varies in
a continuous rather than disjunct manner in the species,
from scarcely discemable to well defined. The variety is
therefore not upheld.
Vouchers: Flanagan 1298 (PRE); Glass 6237 (BOL); Hall 1162
(BOL); Jeppe s.n (PRE).
4. Holothrix brevipetala Immelman & Schelpe in
Immelman in Bothalia 13: 455 (1981); J.L.Stewart et al.:
60 (1982). Type: Humansdorp, koppie above Oudebos,
Jeppe in PRE 33391 (PRE, holo.!; K, iso.!).
H. parvifolia Lindi.: 283 (1835), partly as to part of description (see
also H. cernua. No. 5), non H. parvifolia Lindi. (1836) which is H.
villosa (No. 6).
Holothrix hispidula sensu Schltr.: 443 (1898); Bolus: t. 17 (1913); Bo-
lus: t. 13 (1918) all excl. syn. L.f. (1782) and Thunb. (1794, 1823), non
(L.f.) T.Durand & Schinz which is H. cemua (No. 5).
Leaves densely covered with small squamules or stout
hairs, sometimes withered at anthesis. Scape without
bracts, stout, 60-310 mm long, with stout, hispid, reflexed
hairs. Sepals 1.0-2. 5 mm, densely pilose. Petals entire,
carnose at apices, 2. 5-3. 5 x 0.5-1. 5 mm, yellow-green;
lip and petals sometimes warty, lip camose, oval in outline
with 3-5 short, broad lobes (Figure 8Q), 2.5-3.5 x 2.0-3. 5
mm, yellow-green. Spur slightly curved, 1. 0-2.0 mm.
H. brevipetala occurs in the Western and Eastern Cape
(Figure 4). It grows in sand and shallow soil in rock crev-
ices. Flowering time: can be found in flower from August
to April, usually between October and January.
The squamules or stout hairs on the leaves of this spe-
cies, and the reflexed hairs on the scape, are similar to
those of H. cemua. However, the lip lobes are much shorter
and relatively broader than the linear lobes of H. cemua.
Vouchers: Bruvns 141/75 (PRE); Glass s.n. (34859 in PRE); Schlechter
464 (BOL); Wolley Dod 2339 (BOL).
5. Holothrix cernua (Burm.f.) Schelpe in Orchid
Review 74: 394 (1966); Immelman: 456 (1981); J.L.Ste-
wart et al.: 60, t. 1.13 on p. 63 [not t. 1.13 on p. 62 which
is H. secunda (No. 9)]. Type: Cap. bon. spei, Burmann f.
s.n. (G!).
Orchis cemua Burm.f.: 30 (‘26’) (1768).
Orchis hispidula L.f.: 401 (1782). Holothrix hispidula (L.f.) T.Durand
& Schinz: 5: 70 (1895) Type: Cape of Good Hope, Sparrman 34 (LINN,
microfiche in PRE!).
Orchis hispida Thunb.: 4 (1794); Thunb.: 6 (1823). H. parvifolia Lindi.:
283 (1835) partly excl. part of description which fits H. brevipetala (No.
4). Lectotype: C B S., in arenosis depressis extra Cap, in summitate Taf-
felberg (sic!) et rupibus lateris occidental^ ejusdem montis, Thunberg a
(UPS, microfiche No. 21201 at PRE!).
FIGURE 4. — Distribution of Holothrix exilis, □; and H. brevipetala, •.
130
Bothalia 26,2(1996)
FIGURE 5. — A-D, Holothrix cernua, Linder 1 792 : A, habit; B, flower, side view; C, flower, front view; D, column. E-H, H. orthoceras, Royal Natal
National Park s.n E, habit; F, flower, side view; G, flower, front view; H, column. A-E, H, drawn by K. Immelman; F, G, drawn by G.
Condy. Scale bars: A, E, 10 mm; B-D, F-H, 1 mm.
Bothaiia 26,2 (1996)
131
Monntris secunda Lindl.: 303 (1835) non Orchis secunda Thunb. H.
monotris (Lindl.) Rchb.f.: 119 (1881). Type: between Mossel Bay and
Gouritz River, Burchell 6369 (K!).
H. gracilis Lindl.: 207 (1836); Bolus: 116 (1888). Type: Table Moun-
tain, Drege 1253a (P!).
H. harveyana Lindl.: 206 (1836); Hook.f.: 103 A (1837). Type: Cape
Peninsula, low sandy moist plains, called 'The Flats’ which extend almost
from Table Bay to False Bay (Cape Flats), Harvey s.n. (TCD!).
H. squamulosa Lindl.: 206 (1836); Bolus: t. 18 (1913), t. 11 (1918).
H. squamulosa var. typica Schltr.: 442 (1898). Type: Cape Province,
mountains near De Liefde, Koratra, Drege 1235c (K !).
H. squamulosa var. hirsuta Bolus: 236 (1888). Type: Cape Peninsula,
moist sandy places on Cape Flats, especially after burning; sometimes
on old thatched roofs near Rondebosch, Bolus 7022b (BOL!; K!).
H. squamulosa var. scabra Bolus: 114, t. 23a (1888). Type: Cape Pen-
insula, moist sandy places on Cape Flats, especially after burning; some-
times on old thatched roofs near Rondebosch, Bolus 7022a (BOL!).
H. squamulosa var. glabrata Bolus: t. 18 (1913). Type: Cape Peninsula,
moist sandy places on Cape Flats, especially after burning; sometimes
on old thatched roofs near Rondebosch, Bolus 7022c (BOL!; K!).
Leaves with a dense covering of squamules or small,
stiff hairs, rarely glabrous with hairs on the margins only,
sometimes withered before anthesis. Scape without bracts,
slender or stout, 90-240 mm long, with long, hispid, de-
flexed hairs. Sepals 1.5-3. 0 x 0.5-1. 5 mm, densely hir-
sute. Petals entire, camose at apices, 3.5-7.0 x 0.5-1. 5
mm, cream to lime-green; lip carnose with 3-5 or (6 or
7) lobes (Figure 8D), central lobe the longest, lobes com-
prising J/3 to */2 the length of whole lip, colour as for
petals. Spur curved, 1. 5^4.0 mm long. Anther sacs chest-
nut-brown (Figure 5A-D).
H. cemua is found mainly in the Western Cape, from
the Gifberg to the George area, but extends also along the
southern Cape lowland as far east as Grahamstown in the
Eastern Cape (Figure 6). It grows in sandy or stony places,
often flowering after fire. Flowering time: July to January.
Vouchers: Bolus 11383 (BOL); Galpin 4587 (PRE); Hall 1107 (BOL);
Oliver 4587 (PRE).
6. Holothrix villosa Lindl. in Hooker’s Companion
to the Botanical Magazine 2: 207 (1836).
FIGURE 6. — Distribution of Holothrix cemua.
FIGURE 7. — Distribution of Holothrix villosa var. villosa , •; H. villosa
var. condensata , □; H. majubensis, ▲; and H. thodei, ■
The species is divided into two varieties: var. conden-
sata is generally shorter and stouter, with a denser inflo-
rescence and broader lip lobes than var. villosa.
6a. var. villosa
Bolus: t. 14b (1896), Bolus: t. 14B (1918); Immelman: 456
(1981); J.L. Stewart et al.: 61, t. 1.9a (1982). Type: Groot
Drakenstein and at the foot of Paarl Mountain, under 1000
ft, Drege 1235a (K!).
Orchis hispida Thunb.: 4 (1794); Thunb.: 6 (1823) p.p. quoad spec.
Thunberg (3 (UPS, microfiche no. 21202 at PRE!).
H. parvifolia Lindl.: 207 (1836) non H. parvifolia Lindl. (1835) see
H. brevipetala (No.4) and H. cemua (No. 5); Hook.f.: 103B (1837).
Type: about Cape Town and Wynberg, in the driest and most barren hills
and wastes in a stony and gravelly soil, Harvey s.n. (TCD?).
Leaves with few to many long, straight hairs. Scape
without bracts, 30-365 mm long, densely to sparsely pi-
lose with long, straight hairs at right angles to scape. Se-
pals 1. 0-2.5 x 0.5-1.5 mm, glabrous or with short, straight
hairs. Petals entire, camose at apices, 1 .5 — 4.5 x 0.5- 1.5
mm, cream to yellow-green; lip carnose, divided into three
subequal lobes, 1. 4-4.0 x 0.5-3.0 mm. Spur broadly coni-
cal, curved, 2.0-5. 5 mm long. Column green. Anther sacs
yellow-green. Figure 1F-J.
H. villosa is probably the commonest species of
Holothrix in southern Africa, as well as the most wide-
spread. The typical variety occurs in western Northern
Cape and throughout the Western Cape, and in the western
part of Eastern Cape (Figure 7). It has recently been found
(disjunctly) as far north as Gauteng (Johannesburg) and
the Northern Province. It grows in rock crevices and in
fynbos on hillsides. Flowering time: August to December.
The 3-lobed lip and patent hairs on the scape will dis-
tinguish H. villosa from the occasional specimens of H.
cemua (No. 5) which do not have squamules on the leaves.
Vouchers: Bolus 11638 (BOL); Esterhuysen 22291 (BOL); Galpin 4588
(PRE); Leistner 435 (PRE); Oliver 5058 (PRE); Venter 10276 (PRE).
Bothalia 26,2(1996)
132
FIGURE 8. — A, B, flowers of Holothrix : A, H. mundii, A. Bean s.n., front view; B, H. majubensis, front view. C-R, comparison of lip shape (not
drawn to scale): C, H. fdicornis, Van der We.sthuiz.en s.n. (only part of spur shown); D, H. cernua. Bolus 7022\ E, H villosa var. condensata,
Esterhuysen 294 18', F, H randii, McLaughlin s.n. ; G, H. micrantha, Mottley 2051 ; H, H. schlechteriana, Glass 423', I, H. grandiflora, Bruyns
34/76', J, H. incurva, Compton 21434', K, H. thodei, Wood 5574\ L, H. parviflora, Gordon s.n.', M, R, H. scopularia, Linder 103 L, N, H.
longicornu, Cutting 69, O, H. culveri, Culver 84', P, H. macowaniana, Scully 6204', Q, H. brevipetala, Schelpe s.n. Drawn by G. Condy.
Bothalia 26,2 (1996)
133
6b. var. condensate (Sond.) Immelman in Bothalia
13: 456 (1981); J.L.Stewart et al.: 61, t. 1.9b (1982). Type:
Swellendam, in sand dunes, Mund s.n. (K, lecto.!).
H. condensata Sond.: 76 (1847), Bolus: 36 (1911), Bolus: t. 12 (1918).
H. lithophila Schltr.: 446 (1898). Type: in rock crevices on mountain
above Vogelgat lagoon, Schlechter 9556 (Bt, one flower at K!).
Leaves often broader than long, slightly to densely pi-
lose with long, fine hairs. Scape 85-240 mm long, with
long fine hairs at right angles to scape, without bracts,
usually stout with inflorescence dense. Sepals 1.5-3. 5 x
1. 0-2.5 mm, glabrous to fairly densely pilose with short,
fine hairs, broad, often more than half the length of petals.
Petals entire, camose, 3.5-7.0 x 1.5 mm, green or yel-
lowish green; lip camose, with three broadly linear lobes
^3 to 1/2 length of lip (Figure 8E), lip 3. 5-6. 5 x 1.5-3 .5
mm. Spur broadly conical, curved, 2.0-5. 5 mm long.
H. villosa var. condensata has been found in the west-
ern Northern Cape, the southern Western Cape, and as far
east as Humansdorp in the southern Eastern Cape (Figure
7). It grows in moist areas, and is either lithophytic or
grows in shallow soil in rock crevices. Flowering time:
October to January.
Vouchers: Andreae 843 (PRE); Esterhuysen s.n. (BOL); Leighton 748
(BOL); Pillans 3508 (PRE).
7. Holothrix majubensis C.Archer & R.H. Archer
in South African Journal of Botany 62: 209-211 (1996).
Type: KwaZulu-Natal, Newcastle Dist., Majuba Mtn, sand-
stone cliffs on northern slopes near summit, ± 2 225 m
(2729BD), 7 Jan. 1995, C. Archer & R.H. Archer 2063
(PRE, holo. !; BOL!, NH!).
Leaves, up to 35 x 25 mm, fairly thick-textured, thinly
woolly on margin only. Scape without bracts, up to 55
mm long, with dense to sparse white, woolly hairs. Inflo-
rescence dense, secund; flowers non-resupinate. Sepals ±
2.0 x 1.2 mm, broadly ovate, sparsely hairy. Petals entire,
camose, ± 2.8 x 1.0 mm, white; lip with undivided portion
1.5 x 2.0 mm, white (Figure 8B); lobes 3, ± equal, 1.0 x
0.6 mm. Spur conical, slightly recurved, 1 mm long.
To date the new species is known only from the type
locality (Figure 7), where it is fairly plentiful. Archer &
Archer (1996) expect that it will be found on similar
northern faces of neighbouring mountains in northern
KwaZulu-Natal and southern Mpumalanga. Its habitat is
rather vulnerable to erosion since the plants grow in ex-
posed soil in cracks in vertical sandstone cliffs. Flowering
time: December and January.
The species is well distinct from all other known spe-
cies by its non-resupinate flowers.
In the genus, the non-resupinate flowers of Holothrix
majubensis are the main distinguishing character. It is pos-
tulated that the non-resupination is an adaptation to the
pollinator: due to the inflorescences arching outwards and
downwards, the flowers are placed in the correct position
for pollination. Non-resupinate flowers are also known in
the unrelated Habenaria Willd.
8. Holothrix thodei Rolfe in Dyer, Flora capensis
5,3: 100 (1912); J.L.Stewart et al.: 60 (1982). Type: [Or-
ange] Free State, Witsieshoek, summit of QuaQua Moun-
tains in stony and grassy places, 7500 ft, Thode 48
(BOL!).
Leaves with a dense covering of small, stout, squamous
hairs, withered at anthesis. Scape without bracts, 100-240
mm long, densely covered with stout, scabrid, reflexed
hairs. Sepals 1.5-3. 5 x 0. 5-1.0 mm, with dense short
hairs, usually connate. Petals entire, carnose, 3.5-5.0 x
0.5-1. 5 mm, yellow or yellow-green; lip carnose, with
three lobes (Figure 8K), outer pair of lobes l/5-1/2 the
length of central lobe, 2.5^1. 5 x 1. 5-3.5 mm, colour as
for petals. Spur slightly curved, 0. 5-2.5 mm.
This montane species occurs in Lesotho and the Dra-
kensberg of the Free State and KwaZulu-Natal, with one
record from near Engcobo, Eastern Cape (Figure 7). It is
recorded from basalt-derived soils in rocky grasslands or
crevices in rocks. Flowering time: January and February.
Although it has a flower very similar to H. exilis (No.
3), H. thodei has thick, scabrid hairs (squamules) on the
scape like those of H. cemua. It also has a completely
different distribution.
Vouchers: Jacobs s.n. (BOL); Jacot Guillarmod, Getliffe & Mzamane
32 (PRE); Killick 1304 (PRE); Schelpe 297 (BOL); Trauseld 961 (NU).
9. Holothrix secunda (Thunb.) Rchb.f. in Otia bo-
tanica hamburgensia 2: 119 (1881); Bolus: t. 37 (1911);
J.L.Stewart et al.: 62, t. 1.13 on p. 62 [not 1.13 on p. 63
which is H. cernua (No. 5)] (1982). Type: Thunberg s.n.
(UPS, holo., microfiche no. 21243 at PRE!).
Orchis secunda Thunb.: 4 (1797); Thunb.: 6 (1823).
Tryphia major Sond.: 82 (1847). Type: Brakfontein, 2000-4000 ft, Zey-
her s.n. (K!).
Leaves glabrous, succulent. Scape without bracts,
45-300 mm long, slightly to densely pilose with short,
fine hairs. Inflorescence subsecund. Sepals 1.5-3. 5 x
1. 0-2.0 mm, glabrous. Petals entire, carnose, adnate to lip
at their bases, 2.5-7.0 x 0.5-2.0 mm, creamy white to
yellowish to yellow-green; lip carnose, divided into five
subequal lobes, lobes linear, lip with a few papillae at
entrance to spur, 3. 5-8. 5 x 1. 5-4.0 mm, colour as for
petals. Spur slightly curved, 1 .5 — 4.5 mm. Column cream
and lime-green. Anther sacs deep reddish mauve. Figure
3G-J.
H. secunda is mainly distributed in the western North-
ern Cape and in Western Cape, with a few records from
Eastern Cape (Figure 9). It does not, however, occur on
the Cape Peninsula. It grows in dry areas in the shade of
bushes, on stony soil, in rock crevices and on ledges.
Flowering time: June to October.
The glabrous leaves, and the petals, which are adnate
to the lip at their bases, will distinguish this species from
others having a 5-lobed lip and camose petals. Thunberg
did not designate a type, but there is only one specimen
134
Bothalia 26,2(1996)
FIGURE 9. — Distribution of Holothrix secunda , •; H. incurva, ■; and
H. micrantha, A.
of this species at UPS (21243 on microfiche), which fits
his description and is annotated in his hand.
Vouchers: Acocks 17725 (PRE); Bruyns 1132 (BOL); Compton 4329
(BOL); Muir 2764 (PRE).
10. Holothrix incurva Lindl. in Hooker’s Compan-
ion to the Botanical Magazine 2: 207 (1836); Immelman:
456 (1981); J.L. Stewart et al.: 62 (1982). Type: Stocken-
strom Div., Katberg, Drege 8275a (K!).
H. rupicola Schltr.: 419 (1898); Bolus: t. 40B (1911). Syntypes: Mont-
aux-Sources, summit, in fissures of rocks, Thode 6 (not found); slopes
of Mont-aux-Sources, 8000-9000 ft, Flanagan 11981 (BOL!).
Leaves glabrous or with a few short, scattered hairs, mar-
gins with a dense fringe of short hairs. Scape 72-170 mm
long, densely villous with short, fine hairs at right angles to
the scape, without bracts. Sepals 2.0-4.5 x 1. 5-2.0 mm,
densely villous with fine short hairs. Petals entire, with car-
nose, much-attenuated apices, 5.0-9.5 x 1. 5-1.0 mm, yellow
or greenish yellow; lip camose, with 5 linear lobes (Figure
8J), 3.5-4.5 x 1. 5-2.0 mm, yellow or greenish yellow. Spur
broadly conical, curved, 1.0-1. 5 mm long.
This montane species is distributed in the high-lying
areas of the Eastern Cape, KwaZulu-Natal, the Free State
and Lesotho (Figure 9), and grows on basalt ledges and
rocky crevices. Flowering time: December to April.
Vouchers: Hilliard & Burtt 9803 (NU); Prescott s.n. (BOL); Schelpe
7677 (BOL); Stewart 1947 (NU).
1 1 . Holothrix micrantha Schltr. in Botanische
Jahrbiicher 20, Beihefte 50: 31 (1895); J.L.Stewart et al.:
61 (1982). Type: mountains above Heidelberg, on grassy
cliffs, 5400 ft, Schlechter 3522 (Bf).
Leaves pilose, withered at anthesis. Scape without
bracts, 72-170 mm long, with long, straight hairs. Sepals
1. 5-2.0 x 0.5-1.0 mm, with a few hairs at apices. Petals
entire, with carnose apices, 3. 0-4. 5 x 0. 5-1.0 mm; lip
carnose, divided into five linear, acute lobes (Figure
8G), 3. 5-4. 5 x 1 .5-2.0 mm. Spur broadly conical,
curved, 1.0-1. 5 mm long.
The type, collected at Heidelberg in Gauteng, was not
seen; it was probably destroyed in Berlin. Three other
specimens exist which match the description of H. mi-
crantha: two from Killamey near Johannesburg (PRE!, J!)
the J specimen from a marsh (Figure 9), and one from
Modderfontein (K!). The description is taken from the Kil-
larney specimen. Flowering time: September and October.
This rare species is very similar to H. incurva (No. 10)
but is smaller, has a slightly denser spike, and a different
distribution and flowering time. When more specimens
are found, it may prove to be only a variant of that mon-
tane species.
Voucher: Mottley 2015 (J).
12. Holothrix filicornis Immelman & Schelpe in
Immelman in Bothalia 13: 455 (1981); J.L.Stewart et al.:
64, t. 1.16 (1982). Type: Namaqualand, 23 km west of
Springbok, Acocks 19269 (BOL, holo.!; PRE!, K!).
Leaves glabrous, large. Scape without bracts, 65-260
mm long, glabrous. Sepals ovate, acute, 1. 5-2.0 x 0.5-1. 0
mm, glabrous, green tinged with red. Petals with three
filiform lobes ^2-^/?, as long as petals; petals 3.0-8.0 x
0.5-1. 0 mm, greenish white; lip 4.5-9.0 x 1. 5-2.5 mm,
greenish white, divided into five filiform lobes (Figure
8C), lobes comprising 2/3-3/4 of total lip length. Spur
slightly curved and pendulous, 1.25-2.0 times length of
lip, 7-11 mm long. Ovary twisted, green tinged with red.
H. filicornis occurs in northern Namaqualand (North-
ern Province), growing in the shelter of rocks and shrubs
on mountainsides (Figure 10). Flowering time: June to
September.
This species, together with the montane H. scopularia
(No. 13), is unusual among the species of South African
Holothrix in having divided petals, but no bracts on the
scape. The two species always have three-lobed petals,
instead of the variable number of petal lobes of the other
species with divided petals and a bracteate scope. The
extremely long spur, the different distribution and habitat,
and the less strongly secund spike of H. filicornis, how-
FIGURE 10. — Distribution of Holothrix filicornis , ■; H. scopularia, O;
and H. parviflora, •.
Bothalia 26,2 (1996)
135
ever, make it easy to distinguish this species from H.
scopularia.
Vouchers: Oliver, Tdlken & Venter 652, 702, 823 (PRE); Tdlken 3293
(BOL); Van der Westhuizen s.n. (in liquid preservative in BOL, labelled
H.M.F. 20 and 21); Williamson 2562, 2565 (BOL).
13. Holothrix scopularia (Lindl.) Rchb.f , in Otia
botanica hamburgensia 2: 119 (1881); Schltr.: 21 (1899a)
partly, excl. H. pleistodactylcr, Bolus: 38 (1911); Immel-
man: 456 (1981); J.L.Stewart et al.: 64, t. 1.17 (1982).
Type: Witbergen summit, 6000 ft, Drege 8275b (K! lecto.,
here designated; P!).
Scopularia secunda Lindl.: 207 (1836) non Orchis secunda Thunb.
(1823).
H. multisecta Bolus: 170, 190, t. 7 (1890); T.Durand & Schinz: 71
(1895), as H. multiseta. Type: Stockenstrom, Elandsberg summit, Scully
391 (K!, BOL!).
H. burchellii sensu Kraenzl.: 589 (1899) non Lindl. (1835).
Leaves moderately to densely pilose with long, straight,
slender hairs, often withered at anthesis. Scape without
bracts, 110-340 mm long, densely pilose with long,
straight, slender hairs at right angles to scape. Inflores-
cence strongly secund. Sepals 1. 5-4.0 x 1. 0-2.0 mm,
glabrous or with a few long hairs. Petals three-lobed,
3.0- 11.0 x 0.5-2.5 mm, cream or pinkish cream, some-
times wine-coloured; lip 5-12-lobed (Figure 8M, R),
2.0- 13.0 x 1. 5-4.5 mm, colour as for petals. Spur curved,
1.0- 4.5 mm long.
H. scopularia occurs in the high mountain areas of the
Eastern Cape, KwaZulu-Natal and Lesotho, and also (dis-
junctly) in Mpumalanga (Figure 10). It grows on grassy
slopes and rocky outcrops, and flowers from September
to January.
Vouchers: Dieterlen 1301 (PRE); Galpin s.n. (BOL); Schelpe 7159
(BOL); Sim 1267 (PRE); Stewart 1825 (NU).
14. Holothrix parviflora (Lindl.) Rchb.f., Otia bo-
tanica hamburgensia: 119 (1881); Immelman: 456 (1981);
J.L.Stewart et al.: 63 (1982). Type: Swanepoelspoort
Mountains, 3000-4000 ft, Drege 8276a (K!).
Tryphia parviflora Lindl.: 209 (1836). H. lindleyana var. parviflora
(Lindl.) Rolfe: 106 (1912).
T. secunda Lindl.: 209 (1836) non Orchis secunda Thunb. (1823). H.
lindleyana Rchb.f.: 119 (1881); Bolus: t. 35 (1911). Type: Koratra
(Karatara?), Drege s.n. (K!).
Leaves glabrous, succulent. Scape without bracts, 70-
240 mm high, glabrous. Sepals 1. 5-3.5 x 0.5-1. 5 mm,
glabrous. Petals entire, membranous in texture, adnate to
lip at base, 3.0-6.5 x 0.5-1. 5 mm, pure white or flushed
with purple at base; lip 3.5-9.0 x 0.5-1. 5 mm (breadth
taken at base of three central lobes), pure white or with
pale purple lines and throat, membranous, divided into
five lobes, the outer pair of lobes broadly linear, inner
three shorter, broadly linear to nearly rectangular (Figure
8L). Spur narrowly conical, circinnate, 1.5-5. 5 mm long.
H. parviflora occurs from around the Cape Peninsula
(Western Cape) to Stutterheim (Eastern Cape), then (dis-
junctly) in southern KwaZulu-Natal (Figure 10). It grows
in damp places and on mossy boulders, usually in a shel-
tered position or under bushes and trees, and has also been
recorded from rocks just above high tide (at Knysna).
Flowering time: July to November.
The circinnate spur and the petals which are basally
adnate to the lip, are characteristic of this species.
Vouchers: Bayliss 35/1562 (PRE); Long 1018 (PRE); McLoughlin
s.n. (BOL); Parker 1059 (BOL); Sim 950 (NU).
15. Holothrix orthoceras (Harv.) Rchb.f., Otia bo-
tanica hamburgensia: 119 (1881); Bolus: t. 23 (1911);
J.L.Stewart et al.: 62, t. 1.11 (1982). Type: Howieson’s
Poort, and on the Katberg, Hutton s.n. (TCD!, lecto., here
designated; K!).
Tryphia orthoceras Harv.: 4, t. 105 (1863).
Leaves glabrous, reticulated with white or silver. Scape
without bracts, 60-280 mm long, densely pilose with fine,
short hairs. Sepals 1. 5-3.5 x 0.5-1. 0 mm, glabrous. Petals
entire, membranous, base of each petal rolled into a tube
with its edge hooked into edge of opposite petal, 3.5-7. 5
x 1 .0-2.0 mm, white, apices attenuate; lip papillate at en-
trance to spur, 4.0-8.5 x 2.0-5.5 mm, white with or with-
out purple veins, membranous, divided into five to many
lobes, outer pair of lobes broadly linear and acute, two or
more times as long as triangular central lobes. Spur acute,
straight, 2.5-6. 5 mm long. Anther sacs purple. Figure
5E-H.
This fairly common species occurs in forests in the
Eastern Cape, KwaZulu-Natal, Mpumalanga and the
Northern Province (Figure 11). Flowering time: March to
May, but also occasionally in October and November.
H. orthoceras is similar to the rarer H. macowaniana
(No. 18) but differs in having silver- veined leaves and a
longer outer pair of lobes on the lip. It is also often con-
fused with H. parviflora, but its straight spur, and the tri-
angular rather than rectangular lobes in the centre of the
lip, will distinguish it.
Vouchers: Allsop s.n. (PRE); Bokelmann s.n. (PRE); Botha s.n. (PRE);
Hilliard & Burtt 6519 (NU); McLoughlin 626 (BOL).
FIGURE 1 1 . — Distribution of Holothrix orthoceras, •; H. aspera, ■; H.
mundii, O; H. macowaniana, * ; and H. culveri, ▲.
136
Bothalia 26,2 (1996)
FIGURE 12. — A-D, Holothrix aspera, Schelpe s.n.\ A, habit; B, flower, side view; C, flower, front view; D, column, front view. E-H, H. burchellii:
E, habit, A. Bean 142\ F, flower from base of spike, Seagrief s.n.\ G, flower, V4 way up spike, Seagrief s.n.\ H, column, front view. Drawn
by K. Immelman. Scale bars: A, E, 10 mm; B-D, F-H, 1 mm.
16. Holothrix mundii Sond. in Linnaea 19: 77
(1847); Bolus: t. 13 (1896); Bolus: t. 10 (1918); J.L.Ste-
wart et al.: 60, t. 1.2 (1982). Sytnypes: Swellendam area,
Mund s.n. (K!), and in Worcester area, Winterhoekberge,
4000-5000 ft, among ericas, Zeyher s.n.
Leaves glabrous, small. Scape without bracts, 60-160
mm long, with short, fine, slightly reflexed hairs. Inflo-
rescence condensed (Figure 8A). Sepals 1. 0-2.0 x 0. 3-0.5
mm, glabrous. Petals entire, membranous, 1.5-2. 5 x
0.3-0.5 mm, white; lip membranous, divided into seven
lobes on the same pattern as H. aspera (central lobe and
outer pair longer than other four lobes), papillate at en-
trance to spur, 1.5-3. 5 x 0.5-2.0 mm, white. Spur straight,
0.5-1. 5 mm long. Column very short, green. Anther sacs
bright pinkish mauve.
H. mundii is distributed from the Cape Peninsula
through the southern Western and Eastern Cape as far east
as Port Elizabeth (Figure 11). Flowering time: September
to November.
H. mundii is very similar to H. aspera (No. 17) in the
shape of its lip, but can easily be distinguished by the
condensed inflorescence, as well as by the smaller flowers
and straight spur. It is one of the smallest species in south-
ern Africa, and the only one with a condensed inflores-
cence.
Vouchers: Bolus 4971 (PRE); Esterhuysen 19033 (BOL); Fair s.n.
(BOL); Muir s.n (PRE).
17. Holothrix aspera (Lindl.) Rchb.f, Otia botanica
hamburgensia: 119 (1881); Bolus: 19B (1913); Immel-
man: 456 (1981); J.L.Stewart et al.: 63, t. 1.15 (1982).
Type: Namaqualand, between Mierenkasteel and Zwart-
doomrivier (13 miles south of Garies), Drege 8276b (K!).
Bucculina aspera Lindl.: 209 (1836).
Holothrix confusa Rolfe: 105 (1912). Syntypes: mountainsides about
Clanwilliam, Leipoldt in MacOwan <6 Bolus s.n. (Herb. Norm. Aust.
1757); stony places on Blaauberg, Schlechter 8465 (K!, BOL!); near
Oliphant’s River Mountains, Schlechter 5036 (K!); near Piekenierskloof
and near Modderfontein, Schlechter 5077\ Hex River Valley, Wolley Dod
4054 (K!, BOL!).
Leaves glabrous. Scape without bracts, 30—250 mm
long, nearly glabrous to moderately pubescent with short,
fine hairs. Sepals 1.0-3. 5 x 0.8-2.5 mm, glabrous. Petals
entire, broad, membranous, 3.5-6.5 x 1. 0-4.0 mm, white
Bothalia 26,2(1996)
137
with bright green bases and two broad maroon or purple
stripes; lip membranous, divided into seven unequal lobes,
usually densely papillate at throat, 3. 0-8.5 x 1. 0-6.0 mm,
colour as for petals. Spur very broad, strongly curved and
bent forward under lip, 2.0-5.0 mm long. Column arched
over entrance to spur, lime-green. Anther sacs purple or
maroon. Figure 12A-D.
A species of semi-arid areas, H. aspera occurs in the
Western Cape and in southern Namaqualand, with one
record from the Springbok District, Northern Cape (Figure
11). It is found in sandy or rocky ground or in crevices
in rocks. Flowering time: June to October.
This species cannot be confused with any other from
the Western Cape or the Namaqualand area. The column
differs from the usual state in Holothrix , forming an open
arch across the mouth of the spur. The lip is densely papil-
late under and in front of this arch. The attitude of the
spur is also unique, being doubled up like a penknife un-
der the spur (and also curved), rather than gently curved
or circinnate as in other species.
Vouchers: Acocks 17032 (PRE); Bolus s.n. (BOL); Esterhuysen 3394a
(BOL), 5567 (PRE).
18. Holothrix macowaniana Rchb.f. in Otia botanica
hamburgensia 2: 108 (1881). Type: Bedford Div., Kagaberg,
Weale s.n., sent by MacOwan (BOL, lecto. !; K!).
Leaves glabrous, small. Scape without bracts, 50-60
mm long) glabrous or with short, fine hairs at right angles
to scape. Sepals 1. 5-3.0 x 0.5 mm, glabrous. Petals entire,
membranous, with attenuate apices, 3.0-4.5 x 0.5-1. 5 mm,
white; lip 5. 5-6.5 x 2.5-5. 5 mm, white, membranous, di-
vided into very shallow, broad, triangular lobes, number
of lobes indeterminate (Figure 8P). Spur narrowly conical,
straight, 3.5-6.5 mm long.
This small and probably not very common species has
a restricted distribution in the forests of the southern East-
ern Cape (Figure 11). Flowering time: August to October.
Vouchers: Glass 6204b (BOL); MacOwan s.n. (BOL); Scully s.n.
(20503 in SAM).
19. Holothrix culveri Bolus in Transactions of the
South African Philosophical Society 16: 147 (1905); Im-
melman: 456 (1981); J.L.Stewart et al.: 61 (1982). Type:
near Barberton, Fig Tree Creek, on rocky slopes, 2000 ft,
Sept. 1890, Culver 84 (BOL!).
Deroemera culveri (Bolus) Schltr.: 144 (1907).
H. culveri var. integra Bolus: 147 (1905). Type: Barberton, Culver 84a
(BOL!, K!).
Leaves withered at anthesis, apparently only one present.
Scape with bracts, slender, ± 140 mm long, with a few short
hairs at base. Sepals 2.5 x 0.8 mm, glabrous. Petals entire,
membranous, oblong, 4.0 x 1.5 mm, white; lip membranous,
entire or with small acute lobe on either side of broadly oval
central limb (Figure 80), 3.5 x 3.0 mm, white. Spur straight,
acute, half as long as lip, 1.5 mm long.
Only the type specimen has ever been collected, from
near Barberton, Mpumalanga (Figure 11), on rocky slopes
at 2000 ft. Flowering time: September.
Voucher: type only.
H. culveri is unique among the southern African spe-
cies of Holothrix in having bracts on the scape and entire,
delicate petals. H. squammata (A. Rich.) Rchb.f. from
Uganda and Ethiopia, however, has the same combination
of characters. Bolus states that there were spurs on the
side-sepals of Culver 84a, but this was not seen; possibly
a strip of tissue was tom off the ovary when the sepals
were dissected.
20. Holothrix burchellii (Lindl.) Rchb.f. in Otia bo-
tanica hamburgensia: 119 (1881); Bolus: t. 39 (1911);
J.L.Stewart et al.: 66, t. 1.22 (1982). Type: Zoetmelks
Rivier, Burchell 6709 (K!).
Scopularia burchellii Lindl.: 304 (1835).
Leaves glabrous. Scape with bracts, 150-500 mm long,
with short, deflexed hairs, stout. Inflorescence secund, with
dimorphic flowers. Sepals 2.5-5 .0 x 1. 0-2.5 mm, glabrous.
Petals divided at apex into 5-10 filiform lobes, longer on
upper than lower flowers of spike, petals 3.5-18.0 x 1. 0-3.5
mm, cream-coloured; lip 3.0-9.0 x 1. 5-4.0 mm, cream-col-
oured, divided into 5-13 lobes, lobes longer on upper than
on lower flowers of spike. Spur curved, smaller on upper
than lower flowers, 3. 0-6.0 mm long. Anther sacs pale chest-
nut to tan in colour. Figure 12E-H.
This species occurs in the Western Cape at Piketberg,
and then at Bredasdorp and from there as far east as the
Queenstown and Stutterheim Districts in Eastern Cape
(Figure 13). Flowering time: August to November.
A striking feature of H. burchellii is the dimorphic
flowers, the upper having greatly elongated petal and lip
lobes compared to the lower. The spur and column of the
upper flowers are smaller, but in the same proportion to
the size of the flower (excluding the lobes of lip and pet-
als) as in the lower flowers. The upper flowers are said
to be sterile, but the author has seen them produce cap-
FIGURE 1 3. — Distribution of Holothrix burchellii, •; and H. randii, O.
138
Bothalia 26,2 (1996)
sules with seed in them, and the column and pollinia are
no different in structure from those of the lower flowers.
Also, although the flowers are generally described as
‘dimorphic’ this is not strictly correct, as the transition
from short- to long-lobed flowers is not abrupt, and there
are about four or five intermediate flowers.
Vouchers: Bolus 11382 (BOL); Schelpe 7133 (BOL); Schlieben &
Ellis 12387 (PRE); Sidey 1255 (PRE).
21. Holothrix randii Rendle in Journal of Botany,
British and Foreign 37: 208 (1899); Summerh.: 11 (1968);
J.L.Stewart et al.: 65, t. 1.19 (1982); Kidson & Fletcher:
14 (1984). Type: Rhodesia, Salisbury [Zimbabwe,
Harare], in shady woods, Rand 596 (BM!).
H. reckii Bolus: t. 21 (1913). Type: Transvaal (Gauteng), Koedoespoort,
Reck 1003 (BOL!, PRE!).
Leaves glabrous, withered at anthesis. Scape with
bracts, 170-420 mm long, minutely pilose with short, fine,
sinuous hairs. Sepals 2. 5^4.5 x 1. 5-2.0 mm, glabrous. Pet-
als divided into 7-11 fine, filamentous lobes, 6.0-13.5 x
3.0-5. 0 mm, white; lip divided into 8-15 fine, filamentous
lobes (Figure 8F), 10.0-13.5 x 3.0-6.0 mm, white. Spur
curved to nearly circinnate, 3.5-8.0 mm long.
This species is found at Zebediela (Northern Province)
and in Gauteng in the districts of Pretoria and Johannes-
burg (Figure 13), and occurs also in Tanzania and Zim-
babwe. It grows on grassy slopes and rocky ledges.
Flowering time: September to December.
H. randii is very similar to H. schlechteriana (No. 22)
but differs in the fine, hair-like lobes of the lip and petals,
in its white (not cream or greenish) petals, and in its dis-
tribution. It also tends to have a more strongly curved
spur but this does not hold for all specimens.
Vouchers: Bolus 14082 (BOL); Codd 6162 (PRE); Hewlett s.n.
(PRE); McLoughlin 144 (BOL).
22. Holothrix schlechteriana Schltr. ex Kraenzl. in
Orchidacearum genera et species: 588 (1899); Bolus: t.
20 (1913); J.L.Stewart et al.: 65 (1982). Syntypes: rocky
area near Clarkson, 5000 ft, Schlechter 6015 (K!); Queens-
town Dist., in mountains near Howieson’s Poort, 2000-3000
ft, Mrs Barber s.n (not found).
Leaves glabrous, large. Scape with bracts, stout,
135-700 mm long, moderately to densely pilose with
short, fine, velvety hairs. Sepals 2.0-6.0 x 0.5-2.5 mm,
glabrous. Petals divided at apex into 4-9 filiform lobes,
2.5-10.0 x 1. 0-2.5 mm, green to pale greenish cream to
yellow-ochre; lip divided into 5-1 1 lobes (Figure 8H), col-
our as for petals. Spur curved, half to nearly equal length
of lip, 2.5-5. 5 mm long.
This species occurs in a number of scattered localities.
It has been recorded from Springbok in the Northern
Cape, from Caledon, Montague and Laingsburg, in the
Western Cape and it extends into Eastern Cape as far as
Steynsburg and King William’s Town (Figure 14). It
FIGURE 14. — Distribution of Holothrix schlechteriana, O; and H.
grandiflora, •.
grows in dry areas shaded by rocks or bushes. Flowering
time: October to February.
For characters distinguishing this species from H.
randii (No. 21), see under that species.
Vouchers: Acocks 12071, 16511 (PRE); Ehrens 1962 (PRE); Hardy
& Bayliss 1115 (PRE); Theron 1399 (PRE).
23. Holothrix grandiflora (Sond.) Rchb.f. in Otia
botanica hamburgensia: 119 (1881); J.L.Stewart et al.: 65,
t. 1.21 (1982). Type: Uitenhage, Wiedemann s.n. (not found).
Scopularia grandiflora Sond.: 79 (1847).
Leaves glabrous, large, often withered at anthesis.
Scape with bracts, stout, 190-510 mm long, glabrous or
with a few minute hairs at base. Sepals 4.5-9.0 x 1. 5^1.5
mm, glabrous. Petals divided into 5-9 lobes, 10.0-22.0 x
1.5 mm, white with bases green or pale lilac; lip 12.5-25.5
x 3. 5-6.5 mm (breadth taken at base of flared section),
divided into 13-26 lobes (Figure 81); limb often flared
out into a broad distal section; colour as for petals. Spur
very broadly conical, slightly curved, */5-*/4 length of lip,
3.0-4.5 mm long. Column with an apiculus.
The records for this species are scattered, and come
from the west coast, from Bredasdorp, and from Ladismith
in Western Cape and from Port Elizabeth (the type) in
Eastern Cape (Figure 14). It grows in hollows and crevices
in rocks, under semi-arid conditions. Flowering time:
March.
H. grandiflora , the tallest representative of the genus
in southern Africa, is similar to H. schlechteriana. It can
usually be distinguished by the shorter spur to lip ratio,
the longer petals, and the nearly glabrous scape; it also
generally has a larger number of lip lobes and a flared
distal section of the lip. The flowering time is later, being
in March, while H. schlechteriana flowers mainly from
October to December and occasionally in January and
February.
Bothalia 26,2 (1996)
139
Although the type has not been seen, the description
by Sonder (1847) is detailed and clearly refers to this spe-
cies.
Vouchers: Bruyns 1282 (NBG); Guthrie s.n. (BOL); Hall 3521c (BOL);
Schelpe 7663 (BOL).
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in the collection formed by Mr Drege, at the Cape of Good Hope.
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LINNAEUS, C. (fil.). 1782 (‘1781’)- Supplementum plantarum systema-
tis vegetabilium: 401. Impensi orphanotrophei, Brunsvigae.
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INDEX TO TAXA
Bucculina Lindl., 125
aspera Lindl., 136
Deroemera Rchb.f., 125
culveri (Bolus) Schltr., 137
Deroemeria, 125
Holothrix Rich, ex Lindl., 125
aspera (Lindl.) Rchb.f, 136
brachylabris Sond., 129
brevipetala Immelman & Schelpe, 129
burchellii (Lindl.) Rchb.f., 137
burchellii sensu Kraenzl. non Lindl., 135
cemua (Burm.f.) Schelpe, 129
condensata Sond., 133
confusa Rolfe, 1 36
culveri Bolus, 137
var. Integra Bolus, 137
exilis Lindl., 129
var. brachylabris (Sond.) Bolus, 129
filicomis Immelman & Schelpe, 134
gracilis Lindl., 131
grandiflora (Sond.) Rchb.f, 138
harveyana Lindl., 131
hispidula (L.f.) T.Durand & Schinz, 129
hispidula sensu Schltr., 129
incurva Lindl., 134
lindleyana Rchb.f., 135
var. parviflora (Lindl.) Rolfe, 135
lithophila Schltr., 133
longicomu G.J. Lewis, 127
macowaniana Rchb.f, 137
majubensis C. Archer & R.H. Archer, 133
micrantha Schltr., 134
monotris (Lindl.) Rchb.f., 131
multisecta Bolus, 1 35
mundiiSrW., 136
orthoceras (Harv.) Rchb.f, 135
parviflora (Lindl.) Rchb.f., 135
parvifolia Lindl. (1835), 129
parvifolia Lindl. (1836), 129, 131
pilosa (Burch, ex Lindl.) Rchb.f, 127
randii Rendle, 138
reckii Bolus, 138
rupicola Schltr., 134
schlechteriana Schltr. ex Kraenzl., 138
scopularia (Lindl.) Rchb.f., 135
secunda (Thunb.) Rchb.f, 133
squamulosa Lindl., 131
var. glabrata Bolus, 131
var. hirsuta Bolus, 131
var. scabra Bolus, 131
var. typica Schltr., 131
140
Bothalia 26,2(1996)
thodei Rolfe, 133
villosa Lindl., 131
var. villosa, 131
var. condensata (Sond.) Immelman , 133
Monotris Lindl., 125
secunda Lindl. non Orchis secunda Thunb., 131
Orchis
cernua Burm.f., 129
hispida Thunb. p.p., 129, 131
hispidula L.f., 129
secunda Thunb., 133, 135
Saccidium Lindl., 125
pilosum Lindl., 127
Scopularia Lindl., 125
burchellii Lindl., 137
grandiflora Sond., 138
secunda Lindl., 135
Tryphia Lindl., 125
major Sond., 133
orthoceras Harv., 135
parviflora Lindl., 135
secunda Lindl., 135
Bothalia 26,2: 141-151 (1996)
FSA contributions 7: Verbenaceae ' : Vitex
C. L. BREDENKAMP* and D.J. BOTHA**
Editor’s note: this contribution duplicates most of the synopsis of the genus Vitex published by Bredenkamp & Botha (1993) before the column ‘FSA
contributions’ was initiated in Bothalia. The synopsis lacks species descriptions and can therefore not be accepted as a direct contribution to the FSA.
7186000 VITEX
Vitex L. in Species plantarum 638 (1753) & Genera
plantarum edn 5: 285 (1754); Adans.: 200 (1763); Juss.:
107 (1789); Lour.: 389 (1790); G.Mey.: 218 (1818); Roth:
516 (1821); Endl.: 635 (1836); Schauer: 682 (1847);
Seem.: 190 (1865); Benth. & Hook.: 1135 (1876);
C.B. Clarke: 583 (1885); Briq.: 170 (1894); Baker: 315
(1900); H. Pearson: 211 (1901); Hutch.: 309 (1926);
W.Piep.: 1-91 (1928); Moldenke & A.Moldenke: 32
(1946); Moldenke: 142 (1955a); Moldenke: 186 (1955b);
Moldenke: 13 (1957a); Moldenke: 70 (1957b); Moldenke:
129 (1958a); Moldenke: 197 (1958b); Moldenke: 21
(1961a); Melch.: 435 (1964); Merxm.: 9 (1967); Mold-
enke: 73 (1967a); Moldenke: 222 (1967b); Moldenke: 130
(1973); R.A.Dyer: 520 (1975); Moldenke: 216 (1979a);
Moldenke: 328 (1979b); Verde.: 50 (1992). Type: Medi-
terranean areas and central Asia. V. agnus-castus L. (Farr
et al. 1979).
Mailelou Rheede ex Adans.: 200 (1763). Limia Vand.
in Vand. et al.: 42 (1788). Nephandra Willd.: 8 (1790).
Allasia Lour.: 84 (1790). Tripinna Lour.: 391 (1790).
Chrysomallum Thouars: 8 (1806). Pyrostoma G.Mey.: 219
(1818). Wallrothia Roth: 317 (1821). Ephialis Banks, Sol.
& A.Cunn.: 461 (1838). Psilogyne DC.: 132 (1838). Casaret-
toa Walp.: 91 (1844). Macrostegia Nees: 218 (1847).
Trees or shrubs. Leaves opposite, palmately compound
with 3-5(7) pinnae, margins entire or seldom dentate;
petiolule present or pinnae sessile. Inflorescences terminal,
many-flowered racemes or axillary, determinate com-
pound dichasia. Flowers bisexual, slightly zygomorphic
or zygomorphic. Calyx persistent, enlarging during fruit-
ing stage; tube campanulate, 5-lobed; lobes often reduced.
Corolla with 5 lobes, bilabiate, mauve; lip with yellow
markings at base; tube white, often curved and hairy ex-
ternally as well as internally, infundibular or campanulate.
Androecium : stamens 4, didynamous, epipetalous, ex-
serted; filaments basally broadened and hairy; anthers dor-
sifixed, divergent, extrorse. Gynoecium: ovary superior,
glabrous, scarcely hairy apically, or hairy in upper half,
bicarpellous, 4-locular with a single orthotropous ovule
per locule, with axile placentation; style terete, exserted.
* National Botanical Institute, Private Bag X101, Pretoria 0001.
** National Botanical Institute, Private Bag X7, Claremont 7735, Cape
Town.
MS. received: 1996-05-20.
t Labiatae could be expanded to include the Viticoideae (Cantino 1992:
35).
Fruit a 4-loculed drupe, sessile on a firm, crateriform ca-
lyx, or a nut enveloped by an infundibular chartaceous
calyx. Seeds 1-4, oblong to narrowly obovoid.
The genus Vitex has a world-wide, tropical to subtropical
distribution, and comprises approximately 380 taxa (Verd-
court 1992). Pieper (1928) described 87 taxa from Africa
and divided these into the subgenera Vitex and Holmskiol-
diopsis W.Piep. Of the six southern African representatives
of the subgenus Vitex, Vitex ferruginea Schumach. &
Thonn. subsp. amboniensis (Gtirke) Verde., V angolensis
Giirke, V. harveyana H. Pearson, and V. mombassae Vatke
have a mainly tropical distribution just extending south-
wards into southern Africa ( sensu FSA). Apart from a
few records from Zimbabwe and Mozambique, V. pooara
Corbishley, and V patula E.A.Bruce (subgenus Vitex) appear
to be restricted to southern Africa. All members of the sub-
genus Holmskioldiopsis are endemic to southern Africa.
la Calyx crateriform during fruiting stage (Figure 2), firm; fruit
fleshy (drupe), globose or ellipsoid: Subgenus Vitex:
2a Gynoecium glabrous, or scarcely hairy at apex:
3a Flowers relatively large, 16-18 mm long; young stems
ribbed 1. V. harveyana
3b Flowers relatively small, 5-10 mm long; young stems not
ribbed:
4a Adaxial surface of pinnae scarcely hairy, trichomes firm,
imparting a rough feel to surface; fruit ellipsoid . .
2. V. patula
4b Adaxial surface of pinnae pubescent, velvety; fruit glo-
bose 3. V. pooara
2b Gynoecium velutinous in upper half:
5a Leaves palmate with three pinnae (occasionally five); pin-
nae sessile (petiolulate pinnae exclusively in Angola);
median pinna widely obovate or widely elliptic ....
6. V. mombassae
5b Leaves palmate with five pinnae (exceptionally seven);
pinnae petiolulate; median pinna narrowly ovate, nar-
rowly obovate or elliptic:
6a Adaxial leaf surface glabrous or scarcely hairy on vena-
tion; median pinna 55-142 mm long; petiolule 6-21
mm long 4. V. ferruginea subsp. amboniensis
6b Adaxial leaf surface puberulent; median pinna 26-50
mm long; petiolule absent or up to 5 mm long . . .
5. V. angolensis
lb Calyx campanulate during fruiting stage (Figure 7), chartaceous;
fruit dry (nut), obovoid: Subgenus Holmskioldiopsis:
7a Leaves with both surfaces of lamina macroscopically glabrous;
median pinna narrowly oblong, narrowly elliptic or
lanceolate 7. VT rehmannii
7b Leaves with both surfaces of lamina hairy; median pinna
oblong, elliptic, narrowly ovate or narrowly obovate:
8a Adaxial lamina surface densely velvety with white hairs;
leaves greyish green 8. V. zeyheri
8b Adaxial lamina surface glabrous or pubescent with white
hairs; fresh leaves dark green 9. V. obovata
142
Bothalia 26,2 (1996)
SUBGENUS VITEX
Subgenus Vitex
Euvitex W.Piep.: 29 (1928). According to Art 21(3) of
the International Code (Greuter et al. 1994) Euvitex must
be rejected.
Flowers with total length 6-18 mm, strong tendency
towards zygomorphy. Calyx 5-lobed; lobes often reduced,
0.5-1.5 mm long. Corolla tube 3.8-10.0 mm long; lobes
5, dorsal lobes almost completely fused. Ovary globose
or broadly flask-shaped, glabrous or hairy in upper half.
Fruit a drupe, ellipsoid to globose, 1 0(— 2 1 ) x 7(-21)mm,
sessile on a persistent, firm, crateriform calyx.
Section Axillares Briq. emend. W.Piep. in Botani-
sche Jahrbiicher 62,3, Beiblatt 142: 30 (1928). Type: V.
ferruginea Schumach. & Thonn. Ghana near Aquapim,
Thonning 265 (C, syn., P-JU, isosyn.)
Axillares Briq.: 172 (1894).
1. Vitex harveyana H. Pearson in Flora capensis 5:
212 (1901); W.Piep.: 56 (1928); Moldenke: 40 (1961a);
Compton: 66 (1966); Moldenke: 257 (1971a); Palmer &
Pitman: 1951 (1973); Coates Palgrave: 808 (1977). Type:
eastern region, banks of the higher Tugela River, Gerrard
& McKen 1250 (PRE, lecto.!, fide Bredenkamp & Botha
(1993); K, photo.!).
V. geminata H.Pearson: 213 (1901); W.Piep.: 56 (1928); Moldenke;
257 (1971b). V. harveyana forma geminata (H.Pearson) Moldenke: 329
(1979b). Type: eastern region, Zululand, at the Umlasi River, Gerrard &
McKen 2027 (TCD, holo. photo.!, K!).
V. schlechteri Giirke: 299 (1904); W.Piep.: 56 (1928); Moldenke: 389
(1959); Moldenke: 727 (1971a). Type: Komatipoort, Schlechter 11731
(Z, holo.!; E!, G!, GRA!, K, photo.!; P!, S!, W!).
Shrubs 2-3(-4) m high, many-stemmed with lax
shoots; adventitious roots bome at nodes; young stems
angular, striated, often ribbed; cork grey-brown, finely stri-
ated. Leaves palmately compound with 3-5 pinnae; petiole
(10-)20-22(-39) mm long, adaxially grooved, abaxially con-
vex, glabrous; petiolule (0-)6-7(-10) mm long, glabrous;
median pinna (30-)53-55(-79) x (16-)23-24(-29) mm, el-
liptic to narrowly obovate; margin entire or toothed in
upper half; dark green, surface shiny, waxy, adaxial sur-
face glabrous, abaxial surface sparsely pubescent, glandu-
lar trichomes present. Inflorescence a compound dichasium,
total length (30-)52-55 (-85) mm; length of bracts at first
branch of inflorescence ( 1— )3— 4(— 1 5) mm. Pedicel resupi-
nating. Flowers zygomorphic, total length in lateral view
17 mm. Calyx: tube (4.0-)4.2^1.5(-5.0) mm; lobes re-
duced, 0.5 mm long. Corolla violet; tube 10-12 mm long;
lobes in front view 22 x 17(— 1 8) mm, dorsal lobes folded
backwards parallel to tube. Androecium: longest pair of
stamens 10-11 mm, shortest pair 9 mm long. Ovary: style
14 mm long. Fruit a fleshy drupe, sessile on a firm crateri-
form calyx, ellipsoid (5— )9— 1 0(— 1 2) x (4-)7-8(-9) mm,
black when ripe.
Distribution and habitat: Mpumalanga, especially near
Komatipoort, Swaziland, NE KwaZulu-Natal and Eastern
Cape; also recorded from Angola and Mozambique (Fig-
ure 1). Most frequently grows between rocks in sandy soil
or on river and stream banks. Flowering time: October to
December. Common names: Umbendula (Zulu); three-fin-
ger-leaf, three-finger vitex, whorled finger-leaf (English);
drievingerblaar, krantzvingerblaar, kransvingerblaar (Afri-
kaans). Utilisation: fruit edible from February to May.
Vouchers: Bredenkamp 255 (PUC); Codd 5242 (K, PRE, NH);
Gerstner 4212 (NH); Rogers TM13278 (PRE, NH).
2. Vitex patula E.A.Bruce in Bruce et al. in Bothalia
6: 237 (1951); Moldenke: 44 (1957a); Moldenke: 40
(1961a); Moldenke: 317 (1967d); Moldenke: 258 (1971b);
Palmer & Pitman: 1962 (1973); J.H.Ross: 300 (1972);
Coates Palgrave: 809 (1977). Type: Transvaal [Northern
Province], Zoutpansberg Dist., Kruger National Park,
Dzundweni Hill, 11.5 miles SE of Punda Maria, Codd
5319 (PRE, holo.!).
Multistemmed shrub (1.0-)2.4^4.0(-5.0) m high; cork
grey-brown and striated; young stem hairy. Leaves pal-
mately compound with 3-5 pinnae, frequently 3 pinnae
present, arranged at right angles; petiole (26-)45^49
(-112) mm long, sparsely tawny-pubescent; petiolule
(2-)4-5(-9) mm long, sparsely tawny-pubescent; median
pinna (39-)61-66(-144) x ( 1 7— )3 1— 32(— 60) mm, elliptic
to narrowly obovate, margin entire or toothed in upper
half, dark green, adaxial surface scabrous due to presence
of macroscopical sturdy trichomes, abaxial surface
sparsely hairy; chartaceous. Inflorescence a compound di-
chasium, total length (29-)59-62(-105) mm; length of
bracts at first branch of inflorescence (4-)9-10(-22) mm.
Flowers zygomorphic, total length (5.0-)6.0-7.0(-9.4)
mm. Calyx enlarges during fruiting stage, (3.0-)3. 2-3.4
(-3.8) mm; lobes (0.5-)0.7-0.9 (-1.28) mm long. Corolla:
tube white, lobes mauve; tube (3.8-)4.0-5.0 (-6.0) mm
long; lobes in front view 8.0-8.5 x (6.0-) 6.5-7.0(-8.0)
mm, held at right angles to tube. Androecium: longest pair
of stamens 4.0-4. 8 mm, shortest pair (3.6-)3.7-3.8(-4.0)
mm, not exserted. Ovary glabrous; style (4.9-)5 .0-5.1
(-5.4) mm long. Fruit a fleshy drupe, sessile on a firm
crateriform calyx, ellipsoid, (9.0—) 1 1 .Q— 12.0(— 1 3.0) x (7.0-)
7.5-8.0(-9.0) mm, black when ripe. Figure 2.
Bothalia 26,2 (1996)
143
FIGURE 2. — Vitex patula : A, habit, reduced, height variable, ± x 0.01; B, flowering branch, x 0.5; C, variation in leaf shape, x 0.25; D, fruiting
inflorescence, x 2; E, side view of flower, x 6; F, flower face showing corolla lobes, x 5. Drawn from Bredenkamp 272 and Botha 3969 (PRE).
Artist: Gillian Condy.
144
Bothalia 26,2(1996)
FIGURE 3. — Distribution of Vitex patula in southern Africa.
Ross (1972) and Von Breitenbach & Von Breitenbach
(1995) treat V. patula as a synonym of V. amboniensis
Giirke [see V. ferruginea Schumach. & Thonn. subsp. am-
boniensis (Giirke) Verde, var. amboniensis ]. Morphologi-
cal and anatomical studies have shown that they are
separate taxa (Bredenkamp 1988).
Distribution : NE parts of Northern Province, Venda,
Kruger National Park and N KwaZulu-Natal; also in
Mozambique (Figure 3). Flowering time: end of Novem-
ber to December.
Vouchers: Bredenkamp 247, 248 & 267 (PUC); Gomes & Sousa 3885
(PRE); Pooley 780, (NH); Van der Schijff 3343 (PRE).
3. Vitex pooara Corbishley in Kew Bulletin 10:
333 (1920); I.Verd.: 45 (1938); Moldenke: 89 (1957b);
Moldenke: 388 (1959); Moldenke: 76 (1961b); Moldenke:
258 (1971b); Palmer & Pitman: 1953 (1973); Coates Pal-
grave: 810 (1977). Type: Transvaal [Northern Province],
Modder Nek, Nylstroom, l.B. Pole Evans 19671 (PRE,
holo.!; K, photo.!).
Small to medium-sized trees, to 5 m high; branches
spreading; cork grey-brown, incised, peels in strips; young
stems tawny. Leaves palmately compound with 3-5 pin-
nae; petiole (27-)45-46(-63) mm long, sparsely tawny-
pubescent; petiolule usually absent. Pinnae tawny at base;
median pinna (39-)58-60(-75) x (16-)27-28(-37) mm,
narrowly obovate, margin entire, yellow green to khaki,
both ad- and abaxial surfaces are pubescent, glandular
trichomes present on both surfaces; coriaceous. Inflores-
cence a compound dichasium, total length (34-)54-56
(-84) mm; length of bracts at first branch of inflorescence
(5— )7— 8(— 15) mm. Flowers zygomorphic, total length 6
mm. Calyx: tube 2 mm; lobes 0.5 mm. Corolla: tube
strongly curved, 5 mm long, white; lobes in front view
8.5 x 8.0 mm, held at right angles to tube. Androecium:
longest pair of stamens 4 mm, shortest pair 3.5 mm, not
exserted. Ovary glabrous, sparsely pubescent at apex; style
4.5 mm long. Fruit a fleshy drupe, sessile on a firm
crateriform calyx, spheroid or ellipsoid, (9— ) 14— 15(— 1 8) x
(9—) 1 2— 1 3(— 1 8) mm, black when ripe.
Bredenkamp (1988) has shown that V. pooara and V.
mombassae Vatke are morphologically and taxonomically
clearly defined and cannot be regarded as synonymous
(Coates Palgrave 1977; Von Breitenbach & Von Breiten-
bach 1995). A study of the type material has also shown
that V. pooara is related to the tropical V. isotjensis Gibbs
(Corbishley 1920), but that these taxa are not conspecific.
Distribution: rare in North-West and Northern Prov-
inces and Gauteng, found especially in the vicinities of
Nylstroom, Thabazimbi and Rustenburg (Figure 4); also
recorded for S Zimbabwe. Flowering time: end of No-
vember to December. Common names: pooar (Corbishley
1920); mphoer or mphuru, referring to the characteristic
‘house-bug’ odour ( Theron 2049, PRE); hardekool, poera-
bessie, poera(s)boom, stinkbessie, weeluisbessie (Afri-
kaans); poora-berry, resin-leaf and wild cherry. Utilisation:
Moldenke (1961b) mentions that the wood is not fit for
utilisation. The fruit is eaten by the local people (Verdoom
1938).
Vouchers: Bredenkamp 238 (PUC); Carlson in PRE1481 (PRE); Gal-
pin 11652 (PRE); Westfall 1556 (PRE).
4. Vitex ferruginea Schumach. & Thonn., Be-
skrivelse af Guineiske planter: 62 (1827); Baker: 324, 521
(1900); W.Piep.: 70 (1928); H.Huber in Huber et al.: 447
(1963); Hepper: 130 (1976). Type: Ghana near Aquapim,
Thonning 265 (C, syn., P-JU, isosyn.).
subsp. amboniensis (Giirke) Verde, in Flora of
tropical East Africa: 66 (1992). Type: East Africa, Am-
boni, Holst 2578 (B, holo.f; HBG, iso.!; K, iso., photo.!).
Shrubs 2.5-3.0(-9.0) m high; multistemmed with long
lax shoots; cork grey-brown, incised, peels in strips; young
stems and growing points with tawny indumentum.
Leaves palmately compound with 3-7 pinnae; petiole
(36— )63— 65(— 125) mm long, tawny-pubescent; petiolule
slender, (6—) 11—1 2(— 2 1 ) mm long, tawny-pubescent; me-
dian pinna (54— )80-87(-142) x ( 1 8— )30— 3 1 (—5 8 ) mm, el-
liptic to narrowly ovate, acute or obtuse, apex acute to
acuminate, margin entire or toothed in upper half, yellow
FIGURE 4. — Distribution of Vitex pooara, •; V. ferruginea subsp. am-
boniensis var. amboniensis, □; V. ferruginea subsp. amboniensis
var. amaniensis, O; V. angolensis, ■, in southern Africa.
Bothalia 26,2(1996)
145
to dark green, adaxial surface glabrous to sparsely pubes-
cent, abaxial surface yellowish pubescent, glandular
trichomes present; softly chartaceous. Inflorescence a
compound dichasium, total length ( 16— )3 1— 32(— 50) mm,
tawny-pubescent; length of bracts at first branch of inflo-
rescence (5— )9— 1 0(— 1 5 ) mm. Flowers zygomorphic, total
length 14-15 mm. Calyx : tube (3.0— )3.5 — 4.0(— 5.0) mm
long; lobes 0.5-1. 5 mm long. Corolla : tube white, lobes
and lip mauve, older flowers cream-coloured; tube
strongly curved, (8.0— )8.5—9.5(— 10.0) mm long; lobes in
front view (15— )16— 17(— 19) x 11-12 mm, dorsal lobe
folded backwards, parallel to tube. Androecium : longest
pair of stamens (8.0-)9.0-9.7(-10.0) mm, shortest pair
(6.0-)7.0-7.2(-8.0) mm, not exserted. Ovary velutinous
in upper half; style 6.4-10.0 mm long. Fruit a fleshy
drupe, sessile on a firm crateriform calyx, spheroid or el-
lipsoid ( 1 8— )20— 2 1 (—22) x ( 1 5— )20— 2 1 (—23 ) mm, purplish
black when ripe.
Two varieties can be distinguished:
la Adaxial surface of pinnae glabrous or scarcely hairy; apex
acute, acuminate or characteristically cuspidate
4a var. amboniensis
lb Adaxial surface of pinnae puberulent; apex rounded to acute
and not acuminate 4b. var. amaniensis
4a. var. amboniensis
V. amboniensis Giirke: 340 (1895); Baker: 329 (1900); Sim: 94 (1909);
Moldenke: 204 (1955b); Moldenke: 380 (1959); Moldenke: 28 (1961a);
Dale & Greenway: 539 (1961); Moldenke: 227 (1967b); Moldenke: 496
(1968a); Van der Schijff: 81 (1969); Moldenke: 257 (1971b); Palmer &
Pitman: 1955 (1973); Coates Palgrave: 807 (1977); Moldenke: 361
(1979b).
V. tangensis Giirke: 339 (1895): Baker: 321 (1900); W.Piep.: 68 (1928).
Type: Tanzania Tanga Volkens 92 (B, holo.f).
V. amboniensis Giirke var. schlechteri W.Piep.: 69 (1928); Moldenke:
120 (1949); Moldenke: 206 (1955b); Moldenke: 89 (1967a). Type: East
Africa Lourenfo Marques [Mozambique, Maputo], Schlechter 11715
(BOL, iso.; GRA!).
V. polyantha Baker: 321 (1900). Type: Kenya Mombassa Wakefield
(K, holo.).
V. laevigata Baker. 330 (1900); W.Piep.: 69 (1928). Type: German East
Africa: Usaramo; Dar-es-Salaam [Tanzania] Kirk s.n. (K, holo. photo.!).
V swynnertonii S. Moore: 168 (1911); W.Piep.: 70 (1928). Type:
Mozambique, Gazaland, Swynnerton 1054 (BM, holo.!)
Distinguished from var. amaniensis by the characters
given in the key above.
Distribution and habitat : Venda in Northern Province,
NE Kruger National Park (especially at Nwambiya),
Swaziland and NE KwaZulu-Natal; also Mozambique
(Figure 4). This plant occurs sympatrically with V. patula
in deep sand in sand forests. Flowering time:V. ferruginea
subsp. amboniensis var. amboniensis and V. patula are
sympatric species, but the former flowers in early and the
latter in late December. Fruits ripen in May. Common
names', mufudu (Gir.) (Giirke 1895; Dale & Green way
1961); miahali (Watt & Breyer-Brandwijk 1962); mup-
fumbu-pfumbu (V) (Palmer & Pitman 1972); large-fruited
vitex, Amboni vitex, plum finger-leaf (English); pruim-
vingerblaar, (Afrikaans). Utilisation: Watt & Breyer-
Brandwijk (1962) mentions that the plant is used as a
remedy against snakebite by the Swahili. The fruit is not
edible (Palmer & Pitman 1972).
Vouchers: Balsinhas 242 (PRE); Bredenkamp 268 (PRE, PUQ; Car-
valho 671 (PRE); Codd 5434 (PRE, SRGH); Van Wyk 4015 (PRE, PRU);
Ward 2731 (PRE, NH).
4b. var. amaniensis (W.Piep.) Verde, in Flora of
tropical East Africa: 67 (1992); W.Piep.: 69 (1928); Mol-
denke: 206 (1955b); Moldenke: 29 (1961a). Type: East
Africa, Amani, Institut Amani 221a (B, holo.f).
Distinguished from var. amboniensis by the characters
given in the key above.
Distribution and habitat: only known from the Kruger
National Park (Mpumalanga) at Numbi, and Shabeni Hill
near Pretoriuskop, where the shrub was found at the foot
of the massive rocky summit of the hill (Figure 4). Flow-
ering time: flowers were scarce but could still be found
in the beginning of February.
Vouchers: Bredenkamp 225 (PRE, PRU); Van der Schijff 152 & 740
(PRE, KNP).
5. Vitex angolensis Giirke in Botanische Jahrbiicher
18: 167 (1893); Baker: 325 (1900); Hiem: 835 (1900);
W.Piep.: 70 (1928); Gossw.: 382 (1953); Moldenke: 207
(1955b); Moldenke: 98 (1967a); Moldenke: 227 (1967b).
Type: Angola, Welwitsch 5758 (BM, holo.!; K!, LISC!).
Shrubs 1.5(-3.0) m high, multistemmed and well
branched; cork grey-brown, incised, peels in strips; young
stems and growing points with tawny indumentum.
Leaves palmately compound with 3-7 pinnae; petiole
( 1 3— )34 — 37(— 54) mm long, tawny-pubescent; petiolule
slender, (0-)2-3(-5) mm long, tawny-pubescent; median
pinna (26-)40-41(-55) x (12-)18-20(-24) mm, elliptic to
narrowly obovate, apex acute or obtuse, margin entire or
irregularly toothed in the upper half, yellow to dark green,
ad- and abaxial surfaces yellowish pubescent, glandular
trichomes present; softly chartaceous. Inflorescence a
compound dichasium, total length (29— )3 1— 33(— 4 1 ) mm,
tawny-pubescent; length of bracts at first branch of inflo-
rescence (5.0-)5.8-6.0(-7.0) mm. Flowers zygomorphic,
total lateral length 11 mm. Calyx: tube 4.5-5 .0 mm long;
lobes 1.0-1. 5 mm long. Corolla: tube strongly curved,
9-10 mm long; lobes in front view 12.0(— 13.5) x 9.0 mm,
not folded backwards against tube. Androecium: longest
pair of stamens 8 mm, shortest pair 6 mm, not exserted.
Ovary velutinous in upper half; style 10 mm long. Fruit
a fleshy drupe, sessile on a firm crateriform calyx, sphe-
roid or ellipsoid, (13— )17— 18(— 25) x (9—) 14 — 15(— 23) mm,
purplish black when ripe.
Distribution and habitat: Kuring-Kuru in Namibia, and
E Caprivi (Figure 4); also from Angola, especially at
Huila. This plant and V. mombassae grow sympatrically
in deep sand. Flowering time seems to be the isolating
mechanism between these taxa, as V. angolensis flowers
earlier than V mombassae. Flowering time: December to
mid-January. Fruits ripen between May and June. Com-
mon names: Gossweiler (1953) mentions that the name
muvomba (Ganguela) is used in the Vila da Ponte [Ku-
vango]; according to information obtained from herbarium
specimens, the names mufilo and mumpula are also used.
146
Bothalia 26,2(1996)
Vouchers: Bredenkamp 296 (PUC); Geldenhuys 321 (PRE); Janko-
witz 1334 (PRE, WIND).
6. Vitex mombassae* Vatke in Linnaea 43: 533
(1882); Giirke: 339 (1895); Baker: 326 (1900), pro parte;
Th.Dur. & H.Dur.: 437 (1909); W.Piep.: 66 (1928); Mold-
enke: 120 (1949); Gossw.: 381 (1953); Moldenke: 383
(1959); Dale & Greenway: 597 (1961); Moldenke: 47
(1961a); Moldenke: 12 (1968b); Moldenke: 717 (1971a);
Verde.: 155 (1989); Verde.: 63-64 (1992). Type: Kenya,
prope Mombassa in ora Zanzibarensi [near Mombassa, on
the Zanzibar Coast], Hildebrandt 1972 (B, holo.f; BM,
iso.!, K, iso.!).
V. flavescens Rolfe: 87 (1893); Baker: 321 (1900); Moldenke: 383
(1959); Merxm.: 122 (1967); Moldenke: 717 (1971a). Syntypes:
Malange, S. Marques 8; Angola, Welwitsch 5731 (BM!; K!, P!).
V mechowii Giirke: 167 (1893); Hiem: 834 (1900); Moldenke: 386
(1959); Moldenke: 722 (1971a). Syntypes: Angola, Welwitsch 5731 (BM!,
K!, P!); Malange, Mechow 247 (B, t).
V. goetzei Giirke: 464 (1900); Baker: 521 (1900); Moldenke: 383
(1959); Moldenke: 718 (1971a). Type: Sansibar-Kiistengebiet: in der
Steppe siidlich des Rufidji [Zanzibar coastal region in the steppe south
of the Rufidji], M. Goetze 85 (B, holo.t; K, iso.!).
V. shirensis Baker: 326 (1900); Moldenke: 389 (1959); Moldenke: 727
(1971b). Syntype: Buchanan 231, fide Verdcourt (1992).
V mufiitu De Wild.: 142 (1914); Moldenke: 386 (1959); Moldenke:
723 (1971a). Type: Ober-Katanga, Etoile du Congo [Zaire]. No collector
or number mentioned.
V. mombassae Vatke var. acuminata W.Piep.: 68 (1928); Moldenke:
267 (1967c). Type: Britisch-Ost-Afrika [Kenya] Battiscombe 2 (K,
holo.!).
V. mombassae Vatke var. parviflora (Gibbs) W.Piep.: 68 (1928); Mol-
denke: 717 (1971a). V. flavescens Rolfe var. parviflora Gibbs: 463 (1906);
Gibbs: 459 (1916). Type: Hab. Victoria Falls, veld, Gibbs 135 (BOL,
holo.!; BM!, SAM!).
Shrubs 2-3 m high; multistemmed and well branched;
cork grey-brown, incised, peels in strips; young stems and
growing points with tawny indumentum. Leaves pal-
mately compound with 3-5 pinnae; petiole (23-)48-50
(-88) mm long, tawny-pubescent; petiolule usually absent,
0—1 .5(— 1 1 ) mm long, tawny pubescent; median pinna
(37-)60-65(-108) x (19-)35-40(-65) mm, broadly obo-
vate, base acute to obtuse, apex obtuse, rounded or mucro-
nulate, margin entire, yellow green, ad- and abaxial surfaces
yellowish pubescent, glandular trichomes present, charta-
ceous to coriaceous. Inflorescence a compound dichasium,
total length (3 1— )57— 62(— 125) mm, tawny-pubescent; length
of bracts at first branch of inflorescence (3-)9-10(-22) mm.
Flowers zygomorphic, total length 11 mm. Calyx: tube 4
mm long; lobes 1.5 mm long. Corolla: tube strongly
curved, 8 mm long; lobes in front view 16 x 11 mm,
folded backwards against tube. Androecium: longest pair
of stamens 9 mm, shortest pair 8 mm, exserted. Ovary
velutinous in upper half; style 10 mm long. Fruit a fleshy
drupe, sessile on a firm crateriform calyx, spheroid or el-
lipsoid (8—) 1 8— 20(— 33) x ( 1 0—) 1 5— 1 6(— 3 1 ) mm, purplish
black when ripe.
Pieper (1928) mentioned the occurrence of V. mombas-
sae var. erythrocarpa (Giirke) W.Piep. in Livingstone,
♦Until recently the correct name for this taxon was Vitex payos (Lour.)
Merr. (Verdcourt 1989, 1992). However, since this name has been con-
served with a new type (Nicolson & Brummitt 1994), the correct name is
V. mombassae Vatke.
Zambia. This variety does not seem to extend into south-
ern Africa and no material of it was seen.
Distribution and habitat: N Namibia; also in Angola,
Zimbabwe and Mozambique (Figure 5). V. mombassae
and V. angolensis are sympatric species growing in deep
sand. Flowering time: December to early January. Fruits
ripen in May. Common names: lufuka (Durand & Durand
1909); mfudumaji (Swa.); mfududu (Gir.); mfudukoma
(Dig.); mkufu (San.); mkalijote (Bon.) (Dale & Greenway
1961); muxilo-xilo (Kimbundu) (Gossweiler 1953); mwe-
vumdu mazi ( Taylor 1886, PRE); mushembere and mbu-
kushu (De Winter & Marais 4769, PRE). Utilisation: fruit
is edible.
Vouchers: Bredenkamp 299 (PUC); De Winter & Marais 4796 (PRE,
WIND); Hornby 2342, 2752 (PRE) N Mozambique; Mendonga 869
(LISC) Angola.
SUBGENUS HOLMSKIOLDIOPSIS
Subgenus Holmskioldiopsis W.Piep. in Botanische
Jahrbiicher 62,3, Beiblatt 142: 30 (1928). Type: V. obovata
E. Mey., in valle inter Komga et Kei [in a valley of the
Kei Rivier, near Komga] Drege s.n. (K!, BM!, G!, MO!,
OXF!, P!, S!).
Flowers with total length 6-10 mm, regular. Calyx 5-
lobed, lobes 1. 5-3.5 mm long; tube 3-4 mm long; external
surface glandular pubescent, internal surface glandular.
Corolla: tube straight 6-9 mm long; lobes 5, front view
of lobes 10.0-11.0 x 6.0-6.5 mm, situated at right angles
to tube; tube white, lobes mauve. Androecium didy-
namous, longest pair of stamens 6 mm and shortest pair
5 mm, exserted. Ovary globose or obovoid, hairy and
white glandular in upper half; style 6-9 mm long. Fruit
dry, obovoid, 3x4 mm, sessile in base of a persistent,
chartaceous, campanulate calyx.
7. Vitex rehmannii Giirke in Bulletin de l’Herbier
Boissier 4: 818 (1896); H.Pearson: 214 (1901); Galpin:
23 (1923); W.Piep.: 74 (1928); Hutch.: 335 (1946); Mol-
denke: 122 (1949); Moldenke: 112 (1957b); Moldenke:
FIGURE 5. — Distribution of Vitex mombassae (subgenus Vitex), ■; V.
rehmannii (subgenus Holmskioldiopsis ), •, in southern Africa.
Bothalia 26,2 (1996)
147
388 (1959); Moldenke: 78 (1961b); Moldenke: 34
(1968b); Moldenke: 258 (1971b); Palmer & Pitman: 1957
(1973); Coates Palgrave: 811 (1977). Type: Transvaal
[Northern Province], Makapansberg Mtns, Strydpoort,
Rehmann 5422 (Z, holo.!; BM!, K!).
Trees (2-)3-4(-9) m high; single-stemmed and well
branched; cork dark grey-brown, fissured, rough, tessel-
late; young stems and growing points glabrous or sparsely
white-pubescent. Leaves palmately compound with 3-5
pinnae; petiole (9— )30— 35(— 55) mm long, glabrous or
sparsely white-pubescent; petiolule slender, (0-)5-6(-18)
mm long, sparsely hairy; median pinna (34-)60-65(-135)
x (9—) 1 7— 1 8(— 3 1 ) mm, narrowly oblong, narrowly elliptic
or lanceolate, apex as well as base acute, margin entire
or toothed in upper half; yellow green to khaki; adaxial
surface glabrous to sparsely pubescent, glandular, abaxial
surface sparsely pubescent and glandular; chartaceous. In-
florescence a compound dichasium, total length (35-)45-60
(-95) mm, sparsely white-pubescent; length of bracts at
first branch of inflorescence (l-)9-10(-22) mm. Descrip-
tion of flowers and fruit as for subgenus.
Distribution and habitat : North-West, Gauteng, North-
ern Province (particularly common in the Waterberg area)
and Mpumalanga, Swaziland and KwaZulu-Natal (Figure
5). These plants grow mainly in frost free areas, and a
clear discontinuity is visible in the distribution of this
taxon from the northern provinces to KwaZulu-Natal.
V rehmannii frequently grows on quartzite ridges, on
sandstone and also in deep sand. Flowering time: Decem-
ber to February. Fruits ripen in April. Common names:
mokoele (Sesuto) (Galpin 1923); manohani (Shangaan);
umduli, umluthu (Z); mokwele (NS) (Palmer & Pitman
1972); pipe-stem tree (English); pypsteelboom, vaalbos
(Afrikaans). Utilisation: Galpin (1923) describes the wood
as yellow-grey and of medium hardness as well as weight.
The grain is straight, short and of reasonable strength; the
wood is easy to work with and gives a beautiful finish;
it is not infested by termites after utilisation but it is sel-
dom used due to the availability of stronger wood types.
The local people use it to produce stools. The young stems
are hollow and used to make pipe stems.
Vouchers: Bredenkamp 200, 285 (PRE, PUC); Galpin M.282 (PRE);
Meeuse 9491 (topotype) (MO, PRE, SRGH); Wood 6794 (NH, PRE).
8. Vitex zeyheri Sond. ex Schauer in A.P. De Can-
dolle, Prodromus systematis naturalis regni vegetabilis 11:
639 (1847); Kuntze: 258 (1898); H.Pearson: 216 (1901);
W.Piep.: 74 (1928); Moldenke: 122 (1949); Moldenke:
230 (1958b); Moldenke: 154 (1959); Palmer & Pitman:
1959 (1973); Coates Palgrave: 812 (1977). Type: in ripis
flum. Crocodillorum in inter. Afric. austr. [North-West: on
the banks of the Crocodile River], Burke & Zeyher 73 (K,
holo., photo.!; BM!, PRE!, SAM!).
V. zeyheri Sond. ex Schauer var. brevipes H.Pearson: 216 (1901);
W.Piep.: 74 (1928); Moldenke: 122 (1949); Moldenke: 231 (1958b);
Moldenke: 154 (1959). Type: Transvaal [North-West], Magaliesberg near
Crocodile River, Zeyher 1369 (BOL, holo.!; BM!, G!, K!, OXF!, P!,
SAM!, S!, W!).
V. rehmannii Giirke forma subtomentosa Moldenke: 20 (1954); Mol-
denke: 113 (1957b); Moldenke: 154 (1959); Moldenke: 79 (1961b); Mol-
denke: 34 (1968b); Moldenke 258 (1971b). Type: Transvaal [Gauteng]:
Wonderboom Nature Reserve, Pretoria, Repton 2781 (PRE, holo.!).
Trees (2.5-)3.0-4.0(-6.0) m high; single-stemmed, di-
ameter (120-)200-225(-250) mm; cork dark grey-brown,
fissured, rough, tessellate; young stems and growing
points glabrous or sparsely white-pubescent. Leaves pal-
mately compound with 3-5 pinnae; petiole (10-)25-30
(^41) mm long, white-tomentose; petiolule (0-)2-3(-5)
mm long, white-tomentose; median pinna (36-)55-60
(-78) x ( 1 0— )2 8— 30( — 4 1 ) mm, oblong, elliptic, narrowly
ovate or narrowly obovate, grey-green, base acute, apex
acute, obtuse or rounded, margin entire or toothed in upper
half, ad- and abaxial surfaces white-tomentose and glan-
dular; chartaceous. Inflorescence a compound dichasium,
total length (34-)55-60(-81) mm, white-tomentose; length
of bracts at first branch of inflorescence (4 — ) 11—1 2(— 25 ) mm.
Description of flowers and fruit as for subgenus.
Distribution and habitat: endemic to Gauteng, North-
West and Botswana (Figure 6). These plants grow mainly
in stony soil on south and east facing slopes of the Ma-
galiesberg Mountains and associated ridges. In the Zeerust
District it was found on shale, but it occurs on dolomite
in the vicinity of Pretoria and Marico. Flowering time:
December and January, occasionally up to March. Fruits
ripen in April and May. Common names: mokwele (Tsw)
(Palmer & Pitman 1972); silver pipe-stem tree (English);
vaalpypsteelboom en vaalbos (Afrikaans).
Vouchers: Bredenkamp 91 (PUC); Hutchinson 2927 (GRA, PRE);
Phillips 558 (PRE); Pole Evans H18060 (PRE).
9. Vitex obovata E.Mey., Commentariorum de plan-
tis Africae australioris: 273 (1835); Walp.: 87 (1844);
Schauer: 693 (1847); H.Pearson: 214 (1901); Sim: 287
(1907); W.Piep.: 74 (1928); Moldenke: 23 (1957a); Mol-
denke: 154 (1959); Moldenke: 312 (1967d); Moldenke:
258 (1971b); Palmer & Pitman: 1961 (1973); Coates Pal-
grave: 810 (1977). Type: in valle inter Komga et Key [in
a valley of the Kei River near Komga], Dr'ege s.n. (K,
holo.!; BM!, G!, MO!, OXF!, P!, S!).
V. reflexa H.Pearson: 215 (1901); W.Piep.: 74 (1928); Moldenke: 225
(1958b); Moldenke: 388 (1959); Moldenke: 727 (1971a); Coates Pal-
grave: 812 (1977). V. wilmsii Giirke var. reflexa (H.Pearson) W.Piep.: 74
(1928); Moldenke: 225 (1958b); Moldenke: 122 (1949); Moldenke: 154
& 388 (1959); Moldenke: 258 & 727 (1971a); Palmer & Pitman: 1961
(1973); Coates Palgrave: 812 (1977). Type: Transvaal [Mpumalanga], in
FIGURE 6. — Distribution of Vitex zeyheri, •; V. obovata subsp.
obovata, ■, in southern Africa.
148
Bothalia 26,2(1996)
FIGURE 7. — Vitex obovata subsp. wilmsii : A, habit, reduced, height variable, ± x 0.01 ; B, flowering branch, x 0.5; C, variation in leaf shape, x 0.33;
D, fruiting inflorescence, x 2; E, side view of flower, x 4; F, flower face showing corolla lobes, x 3. Drawn from Bredenkamp 291, 293 and
295 (PRE). Artist; Gillian Condy.
Bothalia 26,2(1996)
149
dongas around Barberton, Galpin 602 (BOL, holo.!; GRA!, PRE!, SAM!,
SRGH!, Z!).
Trees (2.5-)3.0-4.0(-9.0) m high; single stemmed, di-
ameter (120-)220-250(-300) mm; cork dark grey-brown,
fissured, rough, tessellate; young stems and growing
points glabrous or sparsely white-pubescent. Leaves pal-
mately compound with 3-5 pinnae; petiole (14 — )23— 38
(-49) mm long, glabrous of sparsely white-pubescent;
petiolule (0-)4-7(-12) mm long, sparsely pubescent; me-
dian pinna (3 1 .5— )5 1— 74(— 99) x ( 17— )2 1—29(^18) mm, el-
liptic, narrowly ovate or narrowly obovate, yellow-green
to khaki or dark green, base acute, apex acute, shortly
acuminate or rounded, margin entire or toothed in the up-
per half, adaxial surface glabrous, occasionally sparsely
pubescent and glandular, abaxial surface pubescent and
glandular, chartaceous. Inflorescence a compound dicha-
sium, total length (33— )56— 83(— 1 20) mm, white-pubescent;
length of bracts at first branch of inflorescence (4 — )9— 13(— 19)
mm. Description of flowers and fruit as for subgenus.
Two subspecies are distinguished;
la Adaxial leaf surface appearing glabrous due to the lack of
linear trichomes; median pinna ( 1 7— )2 1— 29(— 48) mm
broad 9a. subsp. obovata
lb Adaxial leaf surface pubescent due to the presence of linear
trichomes; median pinna (22— )37— 38(— 70) mm broad . .
9b. subsp. wilmsii
9a. subsp. obovata
Distinguished from subsp. wilmsii by the characters
given in the key above.
Distribution and habitat : Northern Province, Mpuma-
langa, particularly in the vicinity of Barberton, NE KwaZulu-
Natal, Swaziland and Eastern Cape as far south as Komga
and the Kei River, the southernmost record of the genus in
Africa (Figure 6). The habitat is variable as illustrated by the
following records: forms part of the undergrowth in riverine
bush; in grassland against slopes of hills (Krantzkop,
KwaZulu-Natal); in a very dry and stony habitat on shale
(Komga); next to the road on stony red loam (Itala Nature
Reserve); between forest trees (Ubombo Mountain Pass);in
mixed Combretum apiculatum-ve. Id (Jozini); in savanna
(Mbabane); in dense savanna against steep, rocky north fac-
ing slopes (White River); on lime stone (Strydpoort). Flow-
ering time: end of November to January. Fruits ripen in April.
Common names: makosikate (, Miller S254, PRE). Ama-kosi-
kate, manohani, mokoele and pypsteel (Moldenke 1958b);
Cape vitex, white vitex, Kei finger-leaf (English); Keivin-
gerblaar (Afrikaans). Utilisation: wood is seldom used due
to the availability of stronger wood types.
Vouchers: Bredenkamp 208, 261 (topotype) (PRE, PUC); Brown &
Shapiro 347 (PRE); Codd 4734 (PRE, SRGH); Flanagan 578 (PRE);
Gerstner 3706 (NH); Miller S254 (PRE).
9b. subsp. wilmsii (Giirke) C.L. Bredenkamp & D.J.
Botha in South African Journal of Botany 59: 619 (1993).
Types: Transvaal [Mpumalanga]: Bei Lydenburg, Dezem-
ber 1895, fruchtend, Wilms 158 (Z, lecto.!, fide Bre-
denkamp & Botha (1993); BM!, G!, K!, P!).
V. wilmsii Giirke var. wilmsii, Giirke: 76 (1900); H. Pearson: 216 (1901);
W.Piep.: 74 (1928); Worsd.: 501 (1941); Moldenke: 122 (1949); Mold-
FIGURE 8. — Distribution of Vitex obovata subsp. wilmsii in southern
Africa.
enke: 224 (1958b); Moldenke: 54 (1959); Moldenke: 258 (1971b); Pal-
mer & Pitman: 1961 (1973); Coates Palgrave: 812 (1977).
Trees (2— )3— 4(— 9) m high; young stems and growing
points white-tomentose. Leaves palmately compound with
3-5 pinnae; petiole (16-)30-35(-72) mm long, white-pu-
bescent; petiolule (0-)3^f(-13) mm long, white-pubes-
cent; median pinna (43-)73-76(-137) x (22-)35-40(-70)
mm, elliptic or narrowly obovate, dark green, base acute,
apex acute, rounded or mucronate, ad- and abaxial sur-
faces white-pubescent and glandular. Inflorescence a com-
pound dichasium, total length (47— )84— 89(— 1 13) mm,
white-pubescent; length of bracts at first branch of inflo-
rescence (4 — ) 1 0— 1 1 (— 1 7) mm. Description of flowers and
fruit as for subgenus. Figure 7.
Distribution and habitat: endemic to SE parts of the
Northern Province, N parts of Gauteng, Mpumalanga and
Swaziland (Figure 8). The habitat of this plant varies, as is
illustrated by the following records: against north and south
facing slopes on dry stony dolomitic soils (Kaspersnek);
against east facing slopes on stony soil in grassland (Karino);
in riverine bush in the Crocodile River valley (Nelspruit);
on steep west facing slopes in forest (Louw’s Creek). Flow-
ering time: December to January. Fruits ripen in April. Com-
mon names: Wilms’s vitex, umluthu (Z); ama-khosikate (Sw)
(Palmer & Pitman 1972); hairy fmgerleaf, hairy vitex (Eng-
lish); harige vingerblaar (Afrikaans).
Vouchers: Bredenkamp 251, 294 (PRE, PUC); Burn Davy 7289 (BM);
Codd 6756 (PRE, SRGH); Meeuse 9325 (PRE SRGH); Thomcrvft 13 (PRE).
EXCLUDED SPECIES
Vitex guerkeana H. Pearson non Hiem. is a synonym of V. pearsonii
W.Piep. (Pieper 1928), a tropical African species.
Vitex mooiensis H. Pearson is a synonym of Premna mooiensis
(H. Pearson) W.Piep. (Moldenke 1971b).
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Typus Speneri, Berolini.
INDEX
Allasia Lour., 141
Axi llares Briq., 142
Casarettoa Walp., 141
Chrysomallum Thouars, 141
Ephialis Banks, Sol. & A.Cunn., 141
Limia Vand., 141
Macrostegia Nees, 141
Mailelou Rheede ex Adans., 141
Nephandra Willd., 141
Premna mooiensis (H. Pearson) W.Piep., 149
Psilogyne DC., 141
Pyrostoma G.Mey., 141
Tripinna Lour., 141
Vitex L„ 141
subgenus Euvitex W.Piep., 142
subgenus Holmskioldiopsis W.Piep., 146
subgenus Vitex, 142
section Axillares Briq. emend, W.Piep., 142
amboniensis Giirke, 145
var. schlechteri W.Piep., 145
angolensis Giirke, 145
ferruginea Schumach. & Thonn., 144
subsp. amboniensis (Giirke) Verde., 144
var. amboniensis, 145
var. amaniensis (W.Piep.) Verde., 145
var. erythrocarpa (Giirke) W.Piep., 146
flavescens Rolfe, 146
var. parviflora Gibbs, 146
geminata H. Pearson, 142
goetzei Giirke, 146
guerkeana H. Pearson non Hiem, 149
harveyana H. Pearson, 142
forma geminata (H. Pearson) Moldenke, 142
isotjensis Gibbs, 144
laevigata Baker, 145
mechowii Giirke, 146
mombassae Vatke, 146
var. acuminata W.Piep., 146
var. parviflora (Gibbs) W.Piep., 146
mooiensis H. Pearson, 149
mufutu De Wild., 146
obovata E. Mey., 147
subsp. obovata, 149
subsp. wilmsii (Giirke) C.L. Bredenkamp & D.J. Botha, 149
patula E. A. Bruce, 142
payos (Lour.) Merr., 146
pearsonii W.Piep., 149
polyantha Baker, 145
pooara Corbishley, 144
reflexa H. Pearson, 147
rehmannii Giirke, 146
forma subtomentosa Moldenke, 147
schlechteri Giirke, 142
shirensis Baker, 146
swynnertonii S. Moore, 145
tangensis Giirke, 145
wilmsii Giirke
var. reflexa (H. Pearson) W.Piep., 147
var. wilmsii, 149
zeyheri Sond. ex Schauer, 147
var. brevipes H. Pearson, 147
Wallrothia Roth, 141
Hm
.
Bothalia 26,2: 153-158 (1996)
Notes on African plants
VARIOUS AUTHORS
AMARYLLIDACEAE
SHOULD CRINUM FORBESII (LINDL.) SCHULT. & SCHULT.F. BE REINSTATED?
Lehmiller (1992) reinstated the name Crinum forbesii
(Lindl.) Schult. & Schult.f. and criticised Verdoorn’s
(1962) decision to reject it. However, Lehmiller’s long
reasoning misses the real issue.
Verdoorn (1962, 1968) described two new species
of Crinum, C. delagoense I.Verd. and C. paludosum
I.Verd. from Natal and Mozambique, being unable to
determine the identity of Crinum forbesianum Herb.
This name, used by Herbert (1837) and Baker (1881,
1896), was based on Amaryllis forbesii Lindl. Lindley
(1826) described the species from bulbs collected by
John Forbes in Delagoa Bay which flowered at Chiswick
near London. No type specimen or drawing exists. Her-
bert (1837) noted that all Forbes’s bulbs had rotted. Al-
though Herbert referred to an unpublished illustration,
this too is presumably lost.
Verdoorn clearly pondered for many years on the iden-
tity of Crinum forbesii. She (1962, 1966) discussed in
detail why the newly described C. delagoense does not
fit Lindley’s description of Crinum forbesii. In addition
she (1966) indicated that Forbes’s bulbs could have rep-
resented more than one species of Crinum. Verdoorn
(1966, 1968) described the discovery of a new plant by
Mr Gordon McNeil that fitted Lindley’s description better
than C. delagoense. Eventually Verdoorn (1968) described
this plant as C. paludosum. Her (1968, 1973) reasoning
is clear. Crinum forbesii should be regarded as a nomen
incertum since it is impossible to prove beyond reasonable
doubt that McNeil’s discovery represents the same species
of Crinum as discovered by Forbes.
Lehmiller (1992) appears to have misunderstood Ver-
doom and concentrated on the fact that Verdoorn (1973:
45) considered Crinum forbesii (Lindl.) Schult. & Schult.f.
as illegitimate. In fact Verdoorn correctly considered C.
forbesianum Herb, as an illegitimate name and continued
to reject C. forbesii as a nomen confusum. The synonyms
mentioned under C. delagoense by Verdoorn (1973), Ama-
ryllis forbesii var. purpurea Lindl. and Crinum forbe-
sianum var. punicea should also be regarded as nomina
incerta. At least three species of Crinum occur in Mozam-
bique that could well fit Lindley’s and Herbert’s descrip-
tions. These are the very variable C. macowanii Baker,
C. delagoense and C. paludosum. Lehmiller himself (1992)
argues that discrepancies in the above-mentioned descrip-
tions could be attributed to the fact that Forbes’s bulbs
were cultivated under less than optimal conditions in Eng-
land. Incidentally Lehmiller refers to the author of the
new combination, C. forbesii as ‘Schultz’, but it is Schult.
& Schult.f. in Sy sterna vegetabilium 7: 864 (Stafleu &
Cowan 1985; Smith 1990).
The present Code of Botanical Nomenclature (Greuter
et al. 1994) provides new facilities to avoid unnecessary
name changes in order to promote a stabilized nomencla-
ture. Lehmiller’s (1992) argument for reinstating the con-
fused name C. forbesii, 30 years after Verdoorn dealt
satisfactorily with it, is misplaced and in strong contrast
with the aims of the Code.
It remains likely that C. delagoense is conspecific with
C. stuhlmannii Baker from East Africa, perhaps only dis-
tinguishable at subspecific level (Nordal 1977). The latter
name will have priority if detailed studies confirm this
possibility.
REFERENCES
BAKER, J.G. 1881. Crinum forbesianum. Curtis' Botanical Magazine
37: t. 6545.
BAKER, J.G. 1896. Amaryllide*. Flora capensis 6: 171-246. Reeve,
London.
GREUTER, W„ BARRIE, F.R., BURDET, H.M., CHALONER, W.G.,
DEMOULIN, V., HANKSWORTH, D.L., J0RGENSEN, P.M.,
NICOLSON, D.H., SILVA, PC., TREHANE, P. & MCNEILL, J.
1994. International code of botanical nomenclature (Tokyo
Code). Koeltz Scientific Books, Konigstein.
HERBERT, W. 1837. Amaryllidaceae. Ridgway, London.
LEHMILLER, D.J. 1992. Restoration of Crinum forbesii (Lindl.) Schultz
emend. Herbert. Herbertia 48: 86-90.
LINDLEY, J. 1826. Report upon the new or rare plants which have
flowered in the garden of the Horticultural Society at Chiswick,
from its first formation to March 1824. Transactions of the Hor-
ticultural Society of London 6: 62-99.
NORDAL, I. 1977. Revision of the East African taxa of the genus Crinum
(Amaryllidaceae). Norwegian Journal of Botany 24: 179-194.
SMITH, G.F. 1990. The correct author citations of Aloe bowiea and A.
myriacantha (Alooideae). Bothalia 20: 80-82.
STAFLEU, F.A. & COWAN, R.S. 1985. Taxonomic literature. Vol. 5.
Bohn, Scheltema & Holkema, Utrecht.
VERDOORN, I. 1962. Crinum delagoense. The Flowering Plants of
Africa 35: t. 1389.
VERDOORN, I. 1966. Identity of John Forbes’s Crinum. African Wild
Life 20: 197-203.
VERDOORN, I. 1968. Crinum paludosum. The Flowering Plants of
Africa 39: t. 1523.
VERDOORN, I. 1973. The genus Crinum in southern Africa. Bothalia
11:27-52.
R.H. ARCHER* and C. ARCHER*
’National Botanical Institute, Private Bag X101, Pretoria 0001.
MS. received: 1995-12-12.
154
Bothalia 26,2 (1996)
PROTEACEAE
A NEW SPECIES OF SERRURIA FROM THE NORTHERN PART OF THE WESTERN CAPE
This unusual new species was discovered by Mrs Bets
Schlebusch in 1993 on the family property Sewefontein
in the Matsikamma Mountains above VanRhynsdorp in
the northern part of the Western Cape. She showed one
of the populations to Dr Ivor Jardine who submitted a
specimen to the Protea Atlas Project. Further field inves-
tigations were not possible until November 1995 when I
accompanied Tony Rebelo of the Protea Atlas Project to
Sewefontein to examine flowering specimens in their
natural habitat and collect the material on which this ac-
count is based.
Serraria lacunosa Rourke sp. nov., species distinc-
tissima, distinguitur capitulis axillaribus ovoideis, pedun-
culis glabris perlongis (85-140 mm); perianthiis rectis
ante anthesin; stigmatibus clavatis; stylis reflexis post an-
thesin, et dense puberulis basaliter.
Frutex erectus, 0.5-1. 0 m altus, unicaulis. Rami mol-
liter sericei, mox glabrescentes. Folia bipinnata, molliter
sericea, mox glabrescentia, 60-80 x 40-60 mm. Inflores-
centiae pedunculatae, rami floriferi 3-12 inflorescentias
gerentes; pedunculi glabri, graciles, 85-140 mm longi.
FIGURE 1 . — Serruria lacunosa: A, perianth in bud plus subtending floral bract; B, gynoecium; C, perianth at anthesis; D, floral bract with thickened
apex; E, mature leaf, x 0.6; F, perianth after anthesis; G, achene; H, flowering shoot, x 0.6. All from Rourke 2108. Artist: Charmaine Bartman.
Bothalia 26,2 (1996)
155
Nieuwoudtvi 1 li
Vanrhynsdorp
^Piketberg
'Krulenbui
18°00'
FIGURE 2. — Known distribution of
Serruria lacunosa.
Capitula axillaria, solitaria, ovoidea, 20-35 mm in diam.,
floribus 28-35. Bracteae anguste ovatae, 5-6 mm longae,
2 mm latae, glabrae, apicibus recurvatis crassis et cartila-
gineis. Perianthium rectum ante anthesin, 12-14 mm lon-
gum, sparsim sericeum vel glabrum. Stylus rectus,
10-12 mm longus, proximaliter dense puberulus, distaliter
glaber. Stigma atrum, clavatum, 1.5 mm longum.
TYPE. — Western Cape, 3118 (Vanrhynsdorp): Matsi-
kamma Mtn, on Sewefontein Farm in a kloof southwest
of homestead, (-DB), 23-11-1995, Rourke 2108 (NBG,
holo.; BOL, K, MO, NSW, PRE, S, iso.).
Erect, loosely branched shrub 0.5-1. 0 m tall, up to
0.7 m in diam., with a single main stem often branched
near base, up to 20 mm in diam. Bark smooth, greyish
brown. Branches softly sericeous initially, soon glabrous
and reddish flushed, 3-5 mm in diam. Leaves bipinnate,
softly sericeous initially, soon glabrous, 60-80 mm x
40-60 mm, ascending, petiolate; petiole 20-25 mm long.
Leaflets terete, upper surface minutely canaliculate, apices
acute. Flowering shoots bearing 3-12 pedunculate, soli-
tary, axillary inflorescences, opening in centrifugal suc-
cession. Peduncle glabrous, slender, 85-140 mm long
with a few widely spaced lanceolate-acute, glabrous
peduncular bracts. Inflorescence a solitary globose axillary
capitulum, 28-35-flowered, 20-35 mm in diam. Recepta-
cle ellipsoid, 20-25 x 3 mm, with the floral bracts form-
ing a pseudo-involucre at the base. Floral bracts narrowly
ovate, 5-6 x 2 mm, glabrous, apices cartilaginous, thick-
ened, tooth-like, recurved. Perianth straight in bud,
12-14 mm long; tube region 3 mm long, inflated, sparsely
sericeous to glabrous; claws equally recurved at anthesis,
sericeous; limbs elliptic, 2 mm long, sericeous. Anthers
4. Style straight at anthesis, 10-12 mm long, reflexing
abaxially and pointing towards peduncle in post pollina-
tion phase; very densely pubescent, proximally becoming
sparsely pubescent and glabrous distally; pubescence cov-
ering 2/3 of length. Pollen presenter black, clavate-obtuse,
1.5 mm long with a slight annular ring at junction with
style; stigmatic groove terminal. Ovary globose, sharply
differentiated from style, densely lanate. Hypogynous
scales minute, 1 mm long, deltoid. Fruit an ellipsoid, vil-
lous achene 6-7 x 2.5-3.0 mm, beaked terminally, trun-
cate and pedicellate basally with a basal fringe of stout
trichomes (Figure 1).
Diagnostic characters
Distinguished by its solitary, ovoid, axillary capitula
with unusually long (85-140 mm) glabrous peduncles
produced in groups of 3-12 towards the upper half of
each flowering shoot; by the straight perianth and densely
villous styles, pubescent for two thirds of their length
which reflex towards the peduncles through almost 180°
from anthesis to the post pollination phase and by the
clavate pollen presenters.
The specific epithet lacunosa ( = full of pools) alludes
directly to the Matsikamma Mountains where S. lacunosa
occurs. This Koi-Koi name is generally understood to
mean ‘pools of water’ (Nienaber & Raper 1977); it could
also apply to a specific site on Matsikamma, namely the
Farm Sewefontein (Seven Springs), which is the type lo-
cality of the species.
Relationships
Serruria lacunosa is not obviously related to any other
species in the genus although the inflorescence architec-
ture is similar to that in S. reflexa Rourke, namely, a
156
Bothalia 26,2(1996)
FIGURE 3. — Stages in opening of inflorescence in Serruria lacunosa
showing movement of styles until they point downwards towards
ground: A, bud stage; B, most perianths open, styles spreading
but beginning to turn down; C, remaining perianths opening with
styles from two earlier opened perianths pointing downwards,
parallel to peduncle.
pubescent style. The styles are glabrous in all other species
having straight perianths at anthesis.
After the erect, vertically placed styles spring free of
the perianths during the first stages of anthesis when the
pollen presenters are still covered in pollen, they move
through almost 180° until they point downwards towards
the ground. This movement is presumably linked to pol-
lination cues and changes in the receptivity of the stig-
matic surface (Figure 3).
Distribution, habitat and conservation status
This species appears to be very rare and is presently
known only from the type collection. At the time of writ-
ing no more than two populations had been discovered,
one consisting of approximately fifteen plants, the other
of approximately nine plants. Both populations are within
a few hundred metres of each other in undisturbed veld
on Sewefontein Farm at the northern end of the Matsi-
kamma Mountain (Figure 2). Serruria lacunosa is almost
certainly a Matsikamma endemic. While it is probable that
other populations exist in the same general area, it seems
unlikely that this species ranges beyond its specialised
habitat in the northern Matsikamma massif. Several one-
to two- year-old seedlings were observed on bare patches
of soil between the parent plants indicating its ability to
regenerate without the intervention of fire.
The two populations examined are situated at an ele-
vation of about 750 m in a gently sloping gully flanked
by high Nardouw Sandstone Cliffs of the Table Mountain
Group (Kent 1980). Unlike the typically coarse-grained
soils usually derived from sandstone, the soil at this site
is white, very fine-grained and clay-like in consistency.
The associated vegetation consists mainly of dense clumps
of Hypodiscus laevigatus (Kunth) Linder (Restionaceae).
Mean annual rainfall at this site is of the order of 500 mm
(D. Schlebusch pers. comm.).
Flowering apparently extends over a period of at least
four months or longer, peaking between September and
December but continuing until February. Odd open inflo-
rescences have been reported as late as April. This is due
to the slow development of the inflorescence buds which
open successively towards the end of each flowering shoot
in centrifugal order. The inflorescences have no percepti-
ble scent. When the type material was collected in early
summer (November) numerous hairy scarab beetles were
observed clambering over open inflorescences apparently
feeding on pollen (very little nectar is produced in this
species). This particular scarab ( Peritrichia antennata
Schein, Scarabaeidae) is likely to be a seasonally impor-
tant pollinator but other pollinators may be active at other
times during the extended flowering period.
number of solitary axillary capitula with attenuated pedun-
cles grouped towards the apex of a flowering shoot. The
mobile style strongly reflexed above the ovary, is also
comparable to the style in S. reflexa but S. lacunosa dif-
fers sharply from the former species due to its heavily
ACKNOWLEDGEMENTS
Mr and Mrs D. Schlebusch kindly granted permission
to collect specimens of S. lacunosa on their farm. Mrs
Schlebusch also collected fruiting material from which
mature achenes were later obtained.
Bothalia 26,2(1996)
157
REFERENCES
KENT, L.E. (compiler) 1980. Stratigraphy of South Africa. Hand-
book 8. Part 1: Lithostratigraphy of the Republic of South
Africa. South West Africa/Namibia, and the Republics of
Bophuthatswana, Transkei and Venda. Government Printer,
Pretoria.
NIENABER, G.S. & RAPER, P.E. 1977. Toponymica hottentotica (2)
(H-Z). Raad vir Geesteswetenskaplike Navorsing, Pretoria.
J.P. ROURKE*
♦Compton Herbarium, National Botanical Institute, Kirstenbosch, Private
Bag X7, Claremont 7735, Cape Town.
MS. received: 1996-05-09.
CAMPANULACEAE
TRANSFER OF ELEVEN VARIETIES OF LIGHTFOOTIA NOM. ILLEG. TO WAHLENBERGIA
In his recent paper, Lammers (1995) transferred 39
southern African species of Lightfootia nom. illeg., listed
by Welman (1993), to Wahlenbergia nom. cons. Some
Lightfootia species which also occur in tropical Africa,
had previously been transferred to Wahlenbergia by Thulin
(1975). Eleven varieties of Lightfootia , all described by
Adamson (1955), most of them from the winter rainfall
area in South Africa, have not yet been transferred to Wa-
hlenbergia. Although it may prove later that varietal status
in some or all cases is not warranted, it is however, now
necessary to transfer these varieties to Wahlenbergia as
well.
8668000-03075 Wahlenbergia denticulata (Burch.)
A.DC. var. transvaalensis (Adamson) W.G. Welman comb,
nov.
Lightfootia denticulata (Burch.) Sond. var. transvaalensis Adamson:
172 (1955). Holotype: Johannesburg, Moss 9515 (BOL).
8668000-06350 Wahlenbergia longifolia (A.DC.)
Lammers var. corymbosa ( Adamson ) W.G.Welman comb,
nov.
Lightfootia longifolia A.DC. var. corymbosa Adamson: 177 (1955).
Holotype: Cape Peninsula, Klaver Valley, Moss 7499 (BM).
8668000-10730 Wahlenbergia rubens (H.Buek)
Lammers var. brachyphylla (Adamson) W.G.Welman
comb. nov.
Lightfootia rubens H.Buek var. brachyphylla Adamson: 188 (1955).
Holotype: Swellendam, Potteberg, Adamson 4798 (BOL).
8668000-10770 Wahlenbergia rubioides (Banks ex
A.DC.) Lammers var. stokoei (Adamson) W.G.Welman
comb. nov.
Lightfootia rubioides Banks ex A.DC. var. stokoei Adamson: 164
(1955). Holotype: Stokoe s.n. (SAM 63249).
8668000-12360 Wahlenbergia subulata (L’Her.)
Lammers var. conges ta (Adamson) W.G.Welman comb,
nov.
Lightfootia subulata L’Hdr. var. congesta Adamson: 162 (1955). Holo-
type: Swellendam, Infanta, Adamson 4812 (BOL).
8668000-12380 Wahlenbergia subulata (L’Her.)
Lammers var. tenuifolia (Adamson) W.G.Welman comb,
nov.
Lightfootia subulata L’Her. var. tenuifolia Adamson: 161 (1955). Holo-
type: Paarl, Franschhoek Pass, Adamson 4911 (BOL).
8668000-12530 Wahlenbergia tenella (L.f.) Lam-
mers var. palustris ( Adamson ) W.G.Welman comb. nov.
Lightfootia diffusa H.Buek var. palustris Adamson: 193 (1955). Holo-
type: Swellendam, Tradouw Pass, Adamson 4821 (BOL).
8668000-12540 Wahlenbergia tenella (L.f.) Lam-
mers var. stokoei (Adamson) W.G.Welman comb. nov.
Lightfootia diffusa H.Buek var. stokoei Adamson: 194 (1955). Holo-
type: Caledon, Palmiet River, Stokoe 8658 (BOL).
8668000-12570 Wahlenbergia tenerrima (H.Buek)
Lammers var. montana (Adamson) W.G.Welman comb,
nov.
Lightfootia tenella Lodd. var. montana Adamson: 199 (1955). Holo-
type: Swartberg Pass, Adamson 4889 (BOL).
8668000-13080 Wahlenbergia uitenhagensis (H.Buek)
Lammers var. debilis (Sond) W.G.Welman comb. nov.
Lightfootia divaricata H.Buek var. debilis (Sond.) Adamson: 182
(1955). Holotype: Uitenhage, Winterhoek, Zeyher 3131 (S).
8668000-13085 Wahlenbergia uitenhagensis (H.Buek)
Lammers var. filifolia (Adamson) W.G.Welman comb. nov.
Lightfootia divaricata H.Buek vat. filifolia Adamson: 183 (1955). Holo-
type: Uitenhage, Swartkops River, Zeyher s.n. in herb. Sonder (S).
REFERENCES
ADAMSON, R.S. 1955. The South African species of Lightfootia. Jour-
nal of South African Botany 21:1 55-2 1 8.
LAMMERS, T.G. 1995. Transfer of the southern African species of
Lightfootia nom. illeg., to Wahlenbergia (Campanulaceae, Cam-
panuloideae). Taxon 44: 333-339.
THULIN, M. 1975. The genus Wahlenbergia s.l. (Campanulaceae) in
tropical Africa and Madagascar. Symbolae Botanicae Upsaliensis
21: 1-223.
WELMAN, W.G. 1993. Campanulaceae. In T.H. Arnold & B.C. de Wet,
Plants of southern Africa: names and distribution. Memoirs of the
Botanical Survey of South Africa No. 62.
W.G.WELMAN*
* National Botanical Institute, Private Bag X101, Pretoria 0001.
MS. received: 1996-07-05.
158
Bothalia 26,2(1996)
ASPHODELACEAE
GENERIC NUMBERS IN THE SUBFAMILY ALOOIDEAE
In Plants of southern Africa: names and distribution
(Arnold & De Wet 1993), three alooid genera (family As-
phodelaceae subfamily Alooideae) are recognized in the
region. In future publications resulting from the PRECIS
database (Smith et al. 1995), this number will be expanded
to six ( Aloe L., Astroloba Uitewaal, Chortolirion A. Ber-
ger, Gasteria Duval, Haworthia Duval and Poellnitzia
Uitewaal). One of these, the monotypic Poellnitzia , was
recently treated (Smith 1995) in the new series, * lFSA con-
tributions’, in Bothalia. In this paper its generic number
was given as 1028010. Originally (De Dalla Torre &
Harms 1900-1907), the number 1028 was allotted to the
genus Apicra Willd. [non Haw.] (Reid & Glen 1993). For
nomenclatural reasons (Smith et al. 1994) this genus was
renamed Astroloba by Uitewaal (1947). Since Astroloba is
therefore biologically identical to Apicra, it should best be
allotted the first derivative number, namely 1028010. Poell-
nitzia and Astroloba, both previously treated as synonyms
of Haworthia, will in future be upheld as distinct genera (at
least by PRE). To prevent confusion, the full PRECIS cita-
tions for these three haworthioid genera are proposed as:
1028000 Apicra Willd. = Haworthia
1028010 Astroloba Uitewaal
1028020 Poellnitzia Uitewaal
The remaining southern African alooid genera are then,
alphabetically:
1026000 Aloe L.
1029010 Chortolirion A.Berger
1027000 Gasteria Duval
1029000 Haworthia Duval
The only remaining alooid genus which is currently,
but doubtfully upheld, Lomatophyllum Willd., is a Mada-
gascan and Mascarene endemic and does not occur in
continental southern Africa. It has the number 1030 (De
Dalla Torre & Harms 1900-1907).
REFERENCES
ARNOLD, T.H. & DE WET, B.C. (eds) 1993. Plants of southern Africa:
names and distribution. Memoirs of the Botanical Survey of South
Africa No. 62.
DE DALLA TORRE, C.G. & HARMS, H. 1900-1907. Genera Siphono-
gamarum ad Sy sterna Engle rianum conscripta. Gulielmi Engel-
mann, Lipsiae.
REID, C. & GLEN, H.F. 1993. Asphodelaceae (Part B). In T.H. Arnold &
B.C. de Wet, Plants of southern Africa: names and distribution.
Memoirs of the Botanical Survey of South Africa No. 62: 1 37-144.
SMITH, G.F. 1995. FSA contributions 3: Asphodelaceae/Aloaceae,
1028010 Poellnitzia. Bothalia 25: 35, 36.
SMITH, G.F., DAVIS, G.W., DONALDSON, J.S. & ROURKE, J.P.
1995. Research in the National Botanical Institute of South Af-
rica. South African Journal of Science 91: 597-604.
SMITH, G.F., MEYER, N.L. & GLEN, H.F. 1994. Little-known generic
names in the family Aloaceae. South African Journal of Science
90: 489, 490.
UITEWAAL, A.J.A. 1947. Revisie van de nomenklatuur der genera
Haworthia en Apicra. Succulenta SeptVOct. 1947: 51-54.
G.F. SMITH* and A.E. VAN WYK3
* Research Directorate, National Botanical Institute, Private Bag X101,
Pretoria 0001.
1 H.G.W.J. Schweickerdt Herbarium, Department of Botany, University
of Pretoria, Pretoria 0002.
MS. received: 1996-06-18.
Bothalia 26,2: 159-165 (1996)
Exormotheca bulbigena sp. nov. (Hepaticae, Marchantiales) and its relation to
E. holstii in southern Africa
T. BORNEFELD*, O.H. VOLK** and R. WOLF***
Keywords: Exormotheca bulbigena, E. holstii, Hepaticae, Marchantiales, southern Africa
ABSTRACT
A new species Exormotheca bulbigena is described from southern Africa and its relation to E. holstii Steph. is discussed.
Morphologically these species are very similar and can be distinguished only when fertile. The chromosome numbers, however,
n = 32 for E. bulbigena and n = 18 for E. holstii, distinguish sterile living plants.
INTRODUCTION
Stephani (1899), in his Species hepaticarum, described
a new species, Exormotheca holstii , from Muse (Tanza-
nia). Subsequently, Marquand (1930), apparently unaware
of Stephani’s publication, described E. megastomata from
a site near Middelburg, Mpumalanga (Transvaal), South
Africa, as a new species. Finally Amell (1953) published
a new species, E. youngii , from Pilgrim’s Rest, Mpumalanga,
(Transvaal) South Africa, which he himself later placed in
synonymy under E. holstii in 1963. After a morphological
examination of all the respective herbarium samples Perold
(1994) sank E. megastomata under E. holstii.
Chromosome numbers of Exormotheca samples from
the loci classici of Marquand and of Amell and from other
sites in eastern Africa as well, always yielded a chromo-
some number of n = 18. However, Exormotheca plants
collected by Volk at Gaikos and Otjua in Namibia have
a chromosome number of n = 32. Nevertheless, all other
morphological characters of the sterile plants were iden-
tical to those from the eastern sites. A priori, different
chromosome counts cannot be considered as distinctive
on species level, because there are many different species
in the Marchantiales with the same chromosome number.
On the other hand, some species with as many as six
different caryotypes are also reported (Bomefeld 1989;
Fritsch 1982).
By cultivation in a greenhouse, Volk succeeded in
growing fertile plants from material collected in the east-
ern localities and in Namibia. The present study is based
on the examination of living and fertile plants, and only
those specimens are considered for which the chromo-
some number and/or the spores are known. The study of
the sexual organs and their products and the asexual re-
production of Exormotheca has shown that the sinking of
E. megastomata under E. holstii (Perold 1994) is justified.
* Schollergasse 14, Wurzburg, Germany.
** Julius v. Sachs-Institut fur Biowissenschaften, Mittlerer Dallenberg-
weg 64, 97082 Wurzburg, Germany; to whom correspondence should be
addressed.
*** Theodor Boveri-Institut fur Biowissenschaften, Am Hubland, 97074
Wurzburg, Germany.
MS. received: 1996-04-22.
However, an additional new species, E. bulbigena, has to
be established.
MATERIALS AND METHODS
Dry herbarium samples of Exormotheca from southern
Africa were cultivated in a greenhouse on a mixture of
garden mould and sand over a base of peat.
For chromosome counts, thallus tips were fixed, extracted
and stained with orceine as described earlier (Bomefeld
1984). In the present study only samples with known chro-
mosome numbers are considered, although the number of
Exormotheca localities is far greater (Perold 1994).
The localities are listed according to Edwards & Leist-
ner (1971).
For SEM studies the samples were fixed in a mixture
of 70% ethanol/glacial acetic acid/40% formalin = 90/5/5
v/v/v at room temperature for 24 hrs, and then dehydrated
in an acetone series. After critical point drying with CO2
and sputtering with gold, SEM micrographs were taken
with a DSM 962 model by Zeiss.
Exormotheca bulbigena Bomefeld, O.H.Volk &
R.Wolf, sp. nov.
Thallus monoicus. Frons hyalina, usque ad 20 mm
longa, simplex vel furcata, linearis, antica plana, crassa.
Costa maxima, strato antico aequialta, postice valde ro-
tundata, lateribus convexo-adscendentibus. Squamae pos-
ticae magnae, cellulis longissimis (50-600 pm) formatae,
uno latere ad basin grosse lacinulatae, oblique oblongae
acuminatae obtusae. Stratum anticum costae aequialtum,
in fundo fila aggregata gerens. Stomata densissima, altis-
sima, ad 2/3 coalita, tertio supero libera, cylindrica, obtusa,
vertice poro oblongo (± 50 x 140 pm) perforata. Frons
ad apicem stolonum bulbillos pulviniformes ± 2-4 mm
longos et latos 2-3 m altos gerens. Antheridia minuta in
duabus lineis (30 x 40 pm) caudaliter et sagittaliter car-
pocephali, in strato antico, stomatibus contecta, disposita.
Ostiola ± 160 pm lata, e sulco prominentes. Carpocephala
in fundo alveoli nudis, strato antico recedente et ante di-
160
Bothalia 26,2 (1996)
FIGURE 1. — Exormotheca bulbigena: A, thallus with carpocephalum; arrow shows antheridial necks. C, stomata with pores. E, F, bulb: E, dorsal
view, where surface is broken (arrow), spongy interior tissue is visible; F, ventral view, arrow shows remnant of stolon which formed bulb.
E. holstii: B, female thallus. D, stomata with pores near thallus tip. Scale bars: A, B, 2 mm; C, D, 500 pm; E, 1 mm; F, 500 pm.
chotomiam frondis. Complura carpocephala sequentialiter
et aequidistantes (± 3-4 mm) in una frons inserta esse
possunt. Receptacula sessilia globosa, vertice haud porosa
stratoque chlorophyllifero tecta; subtus obconico-angus-
tata, utroque latere involucrata; involucra 2, opposita,
capituli vertice convexo-prominente separata, oblique ad-
scendentia, conchaeformia, antice subcarinata, subtus
apiceque aperta, labiis late hiantibus usque ad basin decur-
rentibus; capsula longius pedicellata, irregulariter quadri-
valvata, valvae rufo-brunneae, maxime incrassatae. Spo-
rae ± 140 pm, papulae distales irregulariter vermiculifor-
mes. Elateres degeneratae (20 pm latae et 50-150 pm
longae) cum anulis et spiralibus brevibus. Chromosoma-
tum numems: n = 32.
TYPE. — Namibia, 2216 (Otjimbingwe) Farm Otjua,
granitic outcrop, (-AA), Volk 85/766 c. fr. (M, holo.; M,
PRE).
Bothalia 26,2 (1996)
161
DISCUSSION
In her study of Exormotheca holstii samples from
southern Africa, Perold (1994) necessarily used dry ma-
terial from various localities collected in different years.
In our studies, samples of E. holstii from Namibia and
Mpumalanga (eastern Transvaal) and of E. bulbigena,
grown under identical conditions, developed carpocephala
(Figure 1A, B). Other characters such as overall size and
shape, shape in cross-section, cell shape of the assimila-
tory tissue, width and height of the stomata [the papillae
on top of (or) side of which stomata are located], shape
of the ventral scales, and colour were all the same. Be-
cause these characters are meticulously described by
Perold (1994) they are not repeated here, where the dif-
ferences between the two species are emphasized. The
only morphological feature which differs in the sterile
thalli are the pores of the stomata; they are more elongate
in E, bulbigena than in E. holstii (Figure 1C, D; Table 1).
Whether or not these relations are affected by ecological
factors in the field is unknown and therefore these differ-
ences should not be overestimated.
The formation of small bulbs is known for E. tuberifera
(Kashyap 1914) and for Corbierella ( = Exormotheca) al-
geriensis Douin & Trabut (1919). Therefore we investi-
gated our cultures for such organs and indeed found these
in E. bulbigena ( inde nomen). They are cushion-shaped,
2-4 mm long and broad, and 2-3 mm thick. The bulbs
consist of a chlorophylliferous, spongy tissue with some
oil cells, somewhat more compact ventrally. When mature
these little bulbs are no longer attached to the thallus and
occur isolated in the ground. When dry they shrivel up
and are easily overlooked in the field. Figure IE & F
shows the dorsal and ventral view of bulbs which were
rehydrated for one day after seven months of desiccation.
On the lower side the remnant of the stolon which formed
the bulb is visible (arrow). Exormotheca holstii also forms
short stolons with a slightly enlarged terminal bud, but
these are not drought tolerant and upon remoistening be-
come covered by mould.
As there appears to be little variation in the caryotype,
the main character for discrimination of sterile plants remains
the chromosome number: n = 32 for Exormotheca bulbigena
and n = 1 8 for E. holstii (Figure 2A, C). Chromosome analy-
sis (Bomefeld 1984) reveals that E. bulbigena is eutetraploid
to a basic number of eight chro- mosomes and E. holstii is
eudiploid to a basic number of nine chromosomes (Figure
2B; D). The difference in the basic numbers is not surprising
when taking into account that for Exormotheca fimbriata
(Brazil) both caryotypes are reported: n = 8 (Jovet-Ast 1976)
and n = 9 (Bomefeld unpubl.).
TABLE 1. — Size relations of stomatal pores of Exormotheca bulbigena
and E. holstii grown simultaneously under identical conditions
(n = 16)
A (<CC<tU
- Ml tt<
c Cj<CC<{c
■> 6> mo
B
■ W<
c
D
FIGURE 2. — Exormotheca bulbigena: A, original arrangement of chromo-
some set; B, chromosomes sorted by types. E. holstii: C, original
arrangement of chromosome set; D, chromosomes sorted by types.
The type of Exormotheca holstii is dioecious; the types
of Marquand (1930) and of Amell (1953) are sterile. In
our cultures out of 12 samples from South Africa only
one contained both sexes; four were female, three were
male, and four remained sterile. When comparing fertile
plants of E. bulbigena and E. holstii the most obvious
difference is the deep sulcus caudal to the carpocephalum
of the monoecious E. bulbigena (Figure 1A) which is
lacking in the female E. holstii plant (Figure IB). Closer
inspection of the sulcus shows the antheridial necks which
are composed of eight rows of cells (Figure 3A). The
whole structure has a diameter of ± 160 pm. The antheri-
dial necks of E. holstii (Figure 3B, D) have a diameter
of only 100 |im. A series of sagittal sections of the car-
pocephalum of £. bulbigena (Figure 4A, B) reveals the
presence of two rows of additional ‘microantheridia’ 40
x 30 pm; the respective values for the main antheridia in
the depth of the sulcus, which are arranged in two parallel
rows, are 350 and 150 pm. These microantheridia are a
unique feature not described as yet for liverworts. The
reinforcing bands in the sporangium wall are very variable
within one sporangium and thus are of no value for dis-
tinguishing between the two species. The spores of both
species are about the same size (120-150 pm), dark brown
to black, anisopolar, without a wing. The ornamentation
of the distal face of E. bulbigena can be described as
‘vermiculate’ (sensu Perold 1989) (Figure 5A). If the
ridges become very short (e.g. Perold 1994, fig. 6A) it
can form a papillate pattern. The corresponding structure
of E. holstii, very broad, indented papillae, could be de-
scribed as ‘polygonal’. Figure 5C shows a spore of the
sample Crosby 1115 from Zoetvlei and its similarity to
that of the type of E. holstii (Figure 5E) proves the cor-
162
Bothalia 26,2 (1996)
FIGURE 3. — Antheridial necks. A, C, E. bulbigena: A, from sulcus, arrow shows broken neck with 8 rows of cells forming the channel. B, D, E.
holstii: B, male thallus with antheridial necks. Scale bars: A, 200 pm; B, 2 mm; C, D, 50 pm.
rectness of the identification by Perold (1994). In the de-
scription of E. holstii, with respect to the papillae of the
spores, Stephani (1899) mentions ‘papulis saepe rostratis’.
In Figure 5E no rostrum-like structures can be detected
on the papillae and it remains unclear to which structures
Stephani refers; elaters stuck to the papillae probably led
to the remark mentioned. The ornamentation of the proxi-
mal face of the spores is rather fine and the triradiate mark
is inconspicuous (Figure 5B, D, F). The differences of the
distal face become visible in another way by dark-field
microscopy where the spores are seen in a bright orange
colour. With this method structures are visible on the
ridges of the ornaments which in E. bulbigena are punc-
tiform or very short grooves, for E. holstii they may be
FIGURE 4. — E. bulbigena: A, sagittal section through carpocephalum; arrow shows site of microantheridia. B, microantheridia. Scale bars: A, 1
mm; B, 100 pm.
Bothalia 26,2 (1996)
163
FIGURE 5. — Spores. A, B, E. bulbigena: A, distal view; B, proximal view. C-F, E. holstii , Crosby 1115, Zoetvlei: C, distal view, D, proximal view;
E, F, type specimen, E, distal view, F, proximal view. Scale bars: A-D, 50 pm; E, F, 50 pm.
described as long, finely branched furrows (Figure 6A,
B). The elaters of both species are degenerate, about 20
pm in diameter and between 50 and 150 pm long, with
rings or incomplete spirals. Summarising all the differ-
ences between E. bulbigena to E. holstii (Table 2) we
consider it obligatory to consider the former as a distinct
species, even though sterile plants are very similar.
Consideration of the climate diagrams (Walter & Lieth
1967) for typical sites of the two species (Figure 7A, B)
suggests that E. bulbigena is better adapted to a hotter
and drier climate than E. holstii. The site of the latter at
Rietfontein in Namibia is near a fountain and thus not
contrary to this suggestion.
The immediate influence of external ecological factors
on physiological activity of these plants is difficult to es-
timate. Both species form droplets of condensed water
inside the stomata which consist of living cells. By cool-
ing down the air within the stomata by evaporation and
164
Bothalia 26,2(1996)
FIGURE 6. — Ornamentation of the distal spore faces as seen by dark-
field microscopy. A, E. bulbigena-, B, E. holstii. Arrows: fine
structure on ridges of ornaments.
TABLE 2. — List of characters differing in Exormotheca bulbigena and
E. holstii
by their sheltered humidity the plants seem to establish a
greenhouse-like microclimate of their own, certainly fa-
vourable for photosynthesis. Figure 8 shows the distribu-
tion of the two species in southern Africa. So far E.
bulbigena has only been found in Namibia and may thus
be considered endemic to this region.
ACKNOWLEDGEMENTS
We wish to thank Mrs C. Gehrig for excellent work
in preparing the SEM samples, Prof. W. Hartung for read-
ing the manuscript, Dr S.M. Perold, NBI, Pretoria, for the
loan of the photographs of the spores of the E. holstii
type, and Mr R. Wacker for the microtomy of the carpo-
cephalum of E. bulbigena.
SPECIMENS EXAMINED
Exormotheca bulbigena, n = 32
NAMIBIA. — 1918 (Grootfontein): Gaikos, on quartzite sand, (-AD),
Volk 81/124 (M, PRE). 2116 (Otjimbingwe): Otjua, granitic outcrop,
(-AA), Volk 84/696, 85/766, 88/030 (M, PRE).
Exormotheca holstii, n = 1 8
NAMIBIA. — 2217 (Windhoek): Rietfontein, Granitzersatz, durch
Sickerwasser zeitweise feucht bis nass, (-CA), Volk 01160 (B=L, PRE).
FIGURE 7. — Climatic diagrams of typical sites. A, in Tsumeb, Namibia
for E. bulbigena', B, in Pilgrim’s Rest, Mpumalanga (Transvaal)
for E. holstii.
FIGURE 8. — Distribution map of E. bulbigena, O; and E. holstii, ■, in
southern Africa. Only samples with known spores and/or chro-
mosome numbers were considered.
NORTHERN PROVINCE.— 2229 (Waterpoort): Lokovhela 793
Farm, Soutpansberg, (-DD), Glen 2650 (PRE).
MPUMALANGA. — 2430 (Pilgrim’s Rest): on R532 to Bourke’s
Luck Potholes ± 1 km before coming to SADF Dog Training Centre, on
dry sandy slope above stream, between grass, (-DB), S.M. Perold 2702
(PRE). 2529 (Witbank): Felsiger Hang liber Elefantenfluss, (-CD), Volk
88/026. 2530 (Lydenburg): S of Lydenburg, Spitskop, drier ledge above
waterfall, (-AB), Perold & Koekemoer 2872 (PRE). 2629 (Bethal): Er-
melo, (-DB), s.n. 2630 (Carolina): Knock Dhu Farm, 13 km SE of Lake
Chrissie on Lothair road, common on rich black loamy soil, in grassland,
(-AD), Germishuizen 2839 (PRE).
GAUTENG. — 2528 (Pretoria): Donkerhoek, 22.5 km E of Pretoria
along Pretoria- Witbank Freeway, just beyond road cutting, seepage area,
(-CD), Germishuizen 5624\ S.M. Perold 2795, 2796.
NORTH-WEST. — 2724 (Taung): Farm Zoetvlei, ± 50 km W of pan,
on higher ground, (-AA), Crosby 1115 (PRE).
REFERENCES
ARNELL, S.W. 1953. A new South African species of Exormotheca.
Transactions of the British Bryological Society 2: 283, 284.
ARNELL, S.W. 1963. Hepaticae of South Africa. Swedish Natural Sci-
ence Council, Stockholm.
BORNEFELD, T. 1984. Chromosomenanalyse der Gattung Riccia L. aus
S- und SW-Afrika und allgemeine Bemerkungen zur Zytogenetik
der Lebermoose. Nova Hedwigia 40: 313-328.
BORNEFELD, T. 1989. The Riccia species of S- and SW-Africa. Chro-
mosome numbers and composition of the chromosome sets. Nova
Hedwigia 48: 371-382.
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DOUIN, C. & TRABUT, L. 1919. Deux Hepatiques peu connues. Revue
generate de Botanique 31: 321-328.
EDWARDS, D. & LEISTNER, O.A. 1971. A degree reference system for
citing biological records in southern Africa. Mitteilungen der
botanischen Staatssammlung, Miinchen 10: 501-509.
FRITSCH, R. 1982. Index to chromosome numbers-Bryophyta. Regnum
vegetabile 108, Bohn, Scheltema & Holkema, Utrecht/Antwer-
pen. Junk, The Hague/Boston.
JOVET-AST, S. 1976. Exormotheca fimbriata (Nees) Lindb. & Amell:
localites nouvelles et caryotype. Revue de Bryologique et Liche-
nologique 42: 785-787.
KASHYAP, S.R. 1914. Morphological and biological notes on new and
little-known West Himalayan liverworts II. New Phytologist 13:
308-323.
MARQUAND, C.V.B. 1930. XXVI. Anew species of Exormotheca from
South Africa. Kew Bulletin 1930: 237-239.
PEROLD, S.M. 1989. Spore-wall ornamentation as an aid in identifying
the southern African species of Riccia (Hepaticae). Journal of the
Hattori Botanical Laboratory 67: 109-201.
PEROLD, S.M. 1994. Studies in the Marchantiales (Hepaticae) from
southern Africa. 5. The genus Exormotheca , E. pustulosa and E.
holstii. Bothalia 24: 15-23.
STEPHANI, F. 1899. Species hepaticarum. Bulletin del'Herbier Boissier
7: 198-225.
WALTER, W. & LIETH, H. 1967. Klimadiagramm Weltatlas. VEB G.
Fischer Verlag, Jena.
Bothalia 26,2: 167-175(1996)
Morphological and biochemical genetic evidence for hybridization in the
genus Centella (Apiaceae), with notes on phylogenetic and taxonomic implica-
tions
M.T.R. SCHUBERT*, B-E. VAN WYK*, F.H. VAN DER BANK** and M. VAN DER BANK*
Keywords: allozyme, Centella, hybridization, morphology, relationships
ABSTRACT
The main aim of this paper is to explore the occurrence of hybridization in the genus Centella. Morphological as well as
genetic characters are investigated to confirm the identity of a putative hybrid between C. triloba and C. macrocarpa. These two
independent data sets, one from enzyme electrophoresis and one from morphology, are compared and interpreted by means of
cladistic analysis. Centella glauca and C. virgata were included in the analysis and the effect of hybridization on cladistics is
demonstrated. Hybridization gives a new perspective on infrageneric relationships within the genus Centella , as it may have
obscured discontinuities between previously discrete inffageneric groups.
INTRODUCTION
The subject of hybrids and hybridization has been cov-
ered extensively by a number of authors (e.g. Stace 1989;
Nason et al. 1992), and the influence of hybridization on
cladistics has also previously been the subject of detailed
analyses (McDade 1990, 1992, 1995). These authors in-
dicated that hybridization significantly affects infrageneric
classification because the limits between taxa may become
obscured by reticulate evolution. The following facts
pointed to hybridization in the genus Centella: 1, lack of
correlation between characters of species; 2, reported oc-
currence of putative hybrids in the herbarium record; and
3, the discovery of a putative hybrid initially thought to
be a new species. Apart from the two populations of parent
species (C. triloba and C. macrocarpa) and their putative
hybrid, one population each of C. virgata and C. glauca
(and an additional population of C. macrocarpa) were in-
cluded in a cladistic study of morphological characters
and enzyme data. A second C. macrocarpa population was
included to compare relationships at the population and
species level. Centella virgata and C. glauca were chosen
because of their obvious close relationship to C. macro-
carpa. Centella macrocarpa from the Swartberg Pass and
C. virgata are reseeders, i.e. plants that are killed by fire
and which can only regenerate from seed after fire,
whereas all the other populations and species are resprout-
ers, i.e. plants that survive fire by coppicing, and have a
slow rate of seed germination. As a result, only a relatively
small number of seedlings are added to the populations
after each fire. The inclusion of reseeders and resprouters
gives the results broader applications in terms of the ef-
fects of fire-survival strategy on genetic variation, and also
on the circumscription of species in the C. virgata group.
* Department of Botany, Rand Afrikaans University, P.O. Box 524, Auck-
land Park, Johannesburg 2006.
** Research Unit for Aquatic and Terrestrial Ecosystems, Rand Afrikaans
University, P.O. Box 524, Auckland Park, Johannesburg 2006.
MS. received: 1995-09-27.
MATERIALS AND METHODS
Herbarium specimens and material preserved in FAA
of Centella triloba , C. macrocarpa and the putative hy-
brid, as well as C. glauca and C. virgata were examined
and the leaves, fruits and inflorescences were drawn under
camera lucida. For microtome sectioning, leaves and ma-
ture fruits preserved in FAA were used. Material was em-
bedded in glycol methacrylate (GMA) according to a
modification of the method of Feder & O’Brien (1968)
as used by Van Wyk & Tilney (1994), who also give the
procedures for ultramicrotome sectioning, staining and
photography.
Voucher specimens
C. triloba (7): 3418 Simonstown: Kogel Bay, (-BD), Schubert & Van
Wyk 15.
C. macrocarpa (4): 3322 Oudtshoom: Swartberg Pass, (-AC),
Schubert & Van Wyk 90.
C. macrocarpa (7): 3418 Simonstown: Kogelberg, (-BD), Schubert
& Van Wyk 53.
C. virgata (5): 3320 Montague: Tradouws Pass, nr Barrydale, (-DC),
Schubert & Van Wyk 67.
C. glauca (5): 3219 Wuppertal: Groot Winterhoek plateau, (-CC),
Schubert & Van Wyk 101.
Putative hybrid (2): 3418 Simonstown: Kogelbaai, Schubert & Van
Wyk 98.
Extraction, sample preparation and gel loading meth-
ods are described in Van der Bank et al. (1995); a Tris-HCl
extraction buffer (pH = 7.5) was used. The supernatant
was absorbed directly onto paper wicks, and twelve per-
cent starch (Sigma: S-4501) gels were used. Genetic in-
terpretation of enzyme banding patterns was based on the
subunit structure and subcellular compartmentalization of
the enzymes (Gottlieb 1981, 1982). Locus nomenclature
followed Harris & Hopkinson (1976), Soltis & Soltis
(1989), Hillis & Moritz (1990) and Shakalee et al. (1990).
Locus abbreviations, monomorphic loci, enzyme commis-
168
Bothalia 26,2(1996)
TABLE 1. — Locus abbreviations, enzyme commission numbers (E.C.No.), optimal buffer systems and pH
* monomorphic loci
TABLE 2. — Allele frequencies of polymorphic loci for six Centella populations: C. triloba (PI); a putative hybrid between C. triloba and C.
macrocarpa (H); C. macrocarpa (resprouter, population from Kogel Bay) (P2); and C. macrocarpa (reseeder, population from Swartberg
Pass) (P3); C. virgata (V); and C. glauca (G). Allele frequencies with significant differences (P<0.5) among populations are marked with an
asterisk
Bothalia 26,2 (1996)
169
sion numbers and buffer system combinations yielding the
best results are listed in Table 1.
Data analysis
We used DISPAN (Ota 1993) for phylogenetic analysis
of allozyme data by using neighbour-joining and bootstrap
methods (1000 replications) and Nei’s (1978) genetic dis-
tance values. The analysis of allozyme data was executed
using BIOSYS-1 (Swofford & Selander 1981). The mor-
phological data as well as the allele frequencies in the
different populations were polarized using C. triloba as
outgroup. It is probably the least derived of all the species
included in the study, judged by the shrubby habit and
the broad, dentate, dorsiventral leaves, which we consider
to be plesiomorphic within the genus. Of these, the dor-
siventral leaves are perhaps the most convincing plesio-
morphy, as this character state occurs in related genera
and in all the basal species of Centella (C. asiatica and
the C. eriantha group). All character states of the outgroup
were polarised as plesiomorphic. Alleles that were absent
from one or more populations, while exhibiting both a
low and a high frequency of occurrence in other popula-
tions, were treated as binary or multistate characters. Al-
lele frequencies without any obvious discontinuities (such
as AAT-1 and ACP-2) were not included in the analysis.
Only those frequencies of which the values were obvi-
ously low or high, with no intermediate values (low taken
as less than 0.3, high taken as 0.5 or more) were polarised.
Again, C. triloba was used as outgroup. The intermediate
states of the GPI A and PGM-1 B alleles were present in
the outgroup, so we were unable to polarize these char-
acters (only the one polarity is shown in Table 3). How-
ever, reversing this polarity had no effect, neither on the
topology nor on the tree lengths or consistency indices,
even when the genetic data were analysed separately (Fig-
ure 3D, E). Table 2 lists the allele frequencies used to
polarize the enzyme characters, as shown in Table 3.
These data were analysed using HENNIG86 (Farris 1988).
Five to ten individuals were studied in each population
except for the hybrid where only two individuals were
present at the locality sampled. Despite the small sample
size on which allele frequencies for the hybrid were based,
we believed that useful results could be obtained because
the observed allele frequencies in the other populations
were generally either very high or very low.
RESULTS
Occurrence of hybrids in the herbarium record
Some established species of Centella may in actual fact
be of hybrid origin, but this is difficult to prove. Only a
few herbarium specimens are possible hybrids. Ester-
huysen 16840, for example, was identified as a hybrid on
the label, and the presence of both parents at the same
locality [Uniondale Dist., Zitzikamma Mountains near
Joubertina (3323 DD)] was noted. Representative speci-
mens of both parents were collected: C. eriantha, Ester-
huysen 16838 (BOL); C. montana, Esterhuysen 16939
(BOL). Another specimen of a possible hybrid between
C. triloba and C. eriantha is Pillans 5894 (BOL), from
the Noordhoek Mountains (3418 AB).
TABLE 3. — Characters and polarization of character states in C. triloba
(PI); a putative hybrid between C. triloba and C. macrocarpa (Yi)',
C. macrocarpa (resprouter, population from Kogel Bay) (P2); C.
macrocarpa (reseeder, population from Swartberg Pass) (P3); C.
virgata (V); and C. glauca (G)
Characters and polarization of character states using C. triloba as out-
group:
1. Leaf type; broad (6-36 mm wide) = 0; narrow (3-5 mm wide) = 1 ;
acicular (approximately 1 mm wide) = 2.
2. Number of teeth on leaf margin: 3(5-11) = 0; l(-3) = 1; 1 = 2.
3. Presence of petiole: distinguishable from lamina = 0; not distinguish-
able from lamina = 1 .
4. Tissue arrangement in leaf: dorsiventral = 0; isobilateral = 1 .
5. Surface sculpturing in fruit: smooth = 0; ribbed = 1; prominently
ribbed = 2.
6. Size of fruit: 3-4 x 3 mm = 0; 4. 0-5. 5 x 4.0 mm = 1.
7. Indumentum of petals: glabrous or villous = 0; glabrous = 1.
8. CAP-1 A: allele present = 0; allele absent = 1.
9. CAP-1 B: allele absent = 2; allele present at low frequencies = 1;
allele present at high frequencies = 0.
10. CAP-1 C: allele absent = 0; allele present = 1.
1 1 . GPI A: allele absent = 0; allele present at low frequencies = 1 ; allele
present at high frequencies = 2.
12. GPI C: allele present at high frequencies = 0; allele present at low
frequencies = 1; allele absent = 2.
13. PER B: allele absent = 0; allele present = 1.
14. PER C: allele present at high frequencies = 0; allele absent = 1; allele
present at high frequencies = 2.
15. PGM-1 A: allele present = 0; allele absent = 1.
16. PGM-1 B: allele absent = 0; allele present at low frequencies = 1;
allele present at high frequencies = 2.
17. PGM-1 C: allele present at high frequencies = 0; allele present at low
frequencies = 1 ; allele absent = 2.
1 8. PGM-2 A: allele present at low frequencies = 0; allele present at high
frequencies = 1 .
19. PGM-2 B: allele absent = 1; allele present = 0.
Hybridization between C. triloba and C. macrocarpa
At Kogel Bay we discovered two morphologically in-
termediate plants in the transitional zone between a popu-
lation of C. macrocarpa and C. triloba. Since the plants
were morphologically intermediate between the only two
Centella species present at this site, we concluded that
they must be hybrids. Supporting evidence for their hybrid
origin is presented below. Different data sets were ana-
lysed to obtain a better understanding of character state
distributions in the study group.
170
Bothalia 26,2(1996)
FIGURE 1 . — Comparison of leaves, fruit and inflorescences of C. triloba, C. hybrid and C. tmcrocarpa. C. triloba'. A1-A5, leaves; Dl, D2, inflorescence; G,
fruit. C. hybrid: B1-B5, leaves; E, inflorescence; H, fruit. C. macrocarpa: C1-C4, leaves; FI, F2, inflorescence; I, fruit. Al, D2, G , Barker 6091: A2,
Adamson 4175', A3, Ecklon & Zeyher s.n:, A4, Compton 13512', A5, Stokoe s.n :, Bl, B2, E, H, Boucher 564', B3-B5, Schubert & Van Wyk 98', Cl,
Esterhuysen 32366: C2, C4, Zeyher 4901: C3, Parker 4262: Dl ,De Vos 1035: FI, I, Compton 17587,¥2, Adamson 4262. Scale bars: 10 mm
Bothalia 26,2(1996)
171
Morphology
Morphological evidence for hybridization was based
on characters of the leaves, petioles and mature fruit of
the parent species and the hybrid. Figure 1A-C shows
that the leaves of the putative hybrid are intermediate be-
tween the parent species in width and in the number of
marginal teeth. The leaves are laminar in C. triloba and
acicular in C. macrocarpa. Based on the outgroup method,
broad leaves as well as the presence of three or more teeth
are polarized as plesiomorphic while narrow and acicular
leaves with few or no teeth respectively are considered
as the apomorphic character states of these multistate
characters (Table 3). Furthermore, in the hybrid and in C.
triloba , a lamina can be distinguished from the petiole (as
in most other Hydrocotyloideae), whereas in C. macro-
carpa, no differentiation between lamina and petiole is
evident. The differentiation of lamina and petiole is con-
sidered plesiomorphic whereas the phyllodinous state of
the leaves of C. macrocarpa is polarized as apomorphic
in Table 3. Transverse sections through the lamina and
the petiole of the two species and the hybrid (Figure 2)
show differences in shape and in the presence of a layer
of palisade cells around the entire circumference of the
leaf (dorsiventral in C. triloba, isobilateral in C. macro-
carpa and the hybrid). In the petiole this palisade layer
is absent. The continuous layer of palisade cells is not as
evident in the lamina of the putative hybrid (Figure 2E)
as in C. macrocarpa (Figure 2F). The leaves of C. triloba
are clearly dorsiventral (Figure 2D). In Table 3 dorsiven-
tral leaves are thus considered plesiomorphic whereas iso-
bilateral leaves are considered apomorphic (see note under
Data analysis). The leaves of C. virgata and C. glauca
are acicular and isobilateral with no differentiation of lam-
ina and petiole. Transverse sections of the petioles and
the lamina of C. triloba, the hybrid and C. macrocarpa,
(Figure 2D, G; E, H; F, I) confirm the intermediacy of
the hybrid because of the intermediate shape of both the
lamina and the petiole. The petiole of C. macrocarpa is
winged, that of the hybrid is slightly winged and the peti-
ole of C. triloba is not winged at all.
Centella macrocarpa and the hybrid have ribbed fruits,
but the ribs on the fruits of C. macrocarpa are more dis-
tinct than those of the hybrid (Figure 2B, C). The fruits
of the putative hybrid are intermediate in shape and are
polarized as intermediate between the plesiomorphic char-
acter state (smooth fruits) of C. triloba and the apomorphic
character state (distinctly ribbed fruits) of C. macrocarpa
(Table 3). Ribbed fruits are exceptionally rare, not only
within Centella, but within the subfamily Hydrocoty-
loideae as a whole. As a result, virtually any choice of
outgroup would lead to the same polarity decision. Fur-
thermore, the fruits of C. macrocarpa are much larger
than those of the other species and the hybrid (Figure
1G-I) and the relatively large size is considered apomor-
phic. Smaller fruits are universal in Centella and the po-
larity decision will therefore not be different for any other
outgroup. The fruits of C. virgata are smaller than those
of C. macrocarpa and less distinctly ribbed, whereas the
fruits of C. glauca are particularly large with well pro-
nounced ribs. Centella triloba may be andromonoecious
FIGURE 2. — Ultramicrotome sections showing anatomy of leaf lamina, petiole and fruit of C. triloba, C. hybrid and C. macrocarpa. C. triloba: A,
fruit; D, leaf lamina; G, petiole. C. hybrid: B, fruit; E, leaf lamina; H, petiole. C. macrocarpa: C, fruit; F, leaf lamina; I, petiole. A, D, G,
Schubert & Van Wyk 15', B, Boucher 564; E, H, Schubert & Van Wyk 98; C, F, I, Schubert & Van Wyk 53. Scale bars: 0.5 mm.
172
Bothalia 26,2(1996)
FIGURE 3. — Cladograms and phenograms showing relationships between some species of Centella. A, cladogram based on morphological data. B,
phenogram, including hybrid, constructed with DISPAN (Ota 1993). C, phylogenetic tree, excluding hybrid, constructed with DISPAN (Ota
1993). D, cladogram, including hybrid, based on genetic data. E, cladogram, excluding hybrid, based on genetic data. F, cladogram based on
morphological and genetic data. Characters and polarization of characters used in construction of cladograms listed in Table 3. Solid square,
apomorphy without homoplasy; open square, apomorphy with reversal higher up; =, convergence; x, reversal.
(male and bisexual flowers on the same plant but no fe-
male flowers) or androdioecious (male and bisexual flow-
ers on different plants and no female flowers) (Figure 1D1,
1D2). The bisexual, functionally female umbellules are
made up of a single fruit while the male umbellules have
three to five flowers of which the petals occasionally are
villous. Centella macrocarpa may be andromonoecious or
androdioecious as well, but instead of the inflorescence of
functionally female plants having many functionally female
umbellules as in C. triloba , it is made up of a single um-
bellule (Figure 1F1). Furthermore, when male and bisex-
ual flowers are present, the inflorescence will bear only
a single functionally female umbellule (Figure 1F2). The
male umbellules have three to five male flowers with
glabrous petals. The hybrid is andromono-ecious (Figure
IE), but only a limited number of specimens is available.
The inflorescence is otherwise compar- able to that of C.
macrocarpa with villous male flowers (Figures 1F1, 1F2).
In Table 3 the glabrous petals of C. virgata, C. glauca
and the two C. macrocarpa populations are polarized as
apomorphic, because of the presence of villous petals in
the outgroup. Centella virgata is andromonoecious and C.
glauca may be androdioecious or andromonoecious.
The morphological data analysis resulted in a partially
resolved cladogram with a length of 1 1 steps and a con-
sistency index of 0.90 (Figure 3A). The cladogram is not
fully resolved as a polytomy occurs between the two C.
macrocarpa populations and C. glauca.
Allozyme data
Genetic variation within species was observed at seven
of the 15 enzyme-coding loci. The choice of five of these
loci, i.e., CAP-1, GPI, PER, PGM-1 and PGM-2, for fur-
ther data analysis was confirmed by their relatively high
fixation index (F-statistic) values (Table 4, the values are
explained in the caption). The alleles that contributed most
to population differences at these loci are represented by
characters 8 to 19 in Table 3 (their Fst values are all
Bothalia 26,2 (1996)
173
TABLE 4. — Summary of F-statistics at all loci. F^ and Fn-are the fixa-
tion indices of individuals relative to the total population and its
subpopulations, respectively. FST measures the amount of differ-
entiation among subpopulations relative to the limiting amount
under complete fixation
above 0.500). Figure 3B shows the dendrogram produced
with DISPAN and the complete set of allozyme data
whereas Figure 3C shows the dendrogram without the hy-
brid. Four cladograms were produced from the characters
(8-19) in Table 3, of which the consensus tree is shown
in Figure 3D. The length of the consensus tree is 23 steps
with a consistency index of 0.73. When the hybrid was
excluded from the data set, the resulting cladogram was
fully resolved, with a length of 21 steps and a consistency
index of 0.80 (Figure 3E). According to McDade (1995),
a higher consistency value is to be expected, since the
removal of a hybrid from the analysis should lead to a
significant reduction in homoplasy.
Table 5 lists the genetic distances between the species
and populations studied. From this information it is evi-
dent that the shortest genetic distance was found between
the hybrid and the C. macrocarpa population from Kogel
Bay. The genetic distance between the two C. macrocarpa
populations is relatively small (0.091) when compared to
the average distance between species (0.218). The species
with the shortest genetic distance are C. macrocarpa and
C. triloba (0.108) and the species with the longest distance
between them are C. macrocarpa and C. virgata (0.350).
The shortest genetic distances are those between C.
triloba, the hybrid, and the two C. macrocarpa popula-
tions. This is what one would expect, given the proposed
hybrid origin. Centella glauca and C. virgata are only
linked into this group at a much greater genetic distance.
Table 5 lists results of the BIOSYS analysis used to
establish the genetic diversity within populations. The ge-
netic diversity within the C. triloba and the C. macrocarpa
(Kogel Bay) populations is by far the greatest (mean het-
erozygosity per locus 14.2%, 15.4%; percentage of loci
polymorphic 40, 40; mean number of alleles per locus
1.4, 1.13 respectively) while that of the C. virgata and
the C. macrocarpa (Swartberg) populations is the smallest
(mean heterozygosity per locus 4.9, 5.0; percentage of loci
polymorphic 13.3, 13.3; mean number of alleles per locus
1.13, 1.13 respectively).
When the morphological and genetic data sets were
combined, the result was a fully resolved cladogram with
a length of 40 steps and a consistency index of 0.70 (Fig-
ure 3F). When the hybrid was removed from this data set,
the tree length reduced to 32 steps and the consistency
index improved to 0.87. This reduction in homoplasy is
again consistent with our assumption of hybridization, fol-
lowing McDade (1995).
Geographical distribution of the putative hybrid and the
parent species
Both C. triloba and C. macrocarpa occur in the West-
ern Cape but C. triloba is limited to the Cape Peninsula
and coastal areas of the Caledon District, whereas C.
macrocarpa is more widespread in the region. The hybrid
is very localized in occurrence. Only two plants were
found at Kogel Bay and both parent species occur at the
same locality (Figure 4). The C. triloba population occurs
on a relatively flat area closer to the sea than the C. macro-
carpa population, which grows on a steep scree slope.
The putative hybrids occurred between the two popula-
tions at the foot of the scree slope.
DISCUSSION
Centella triloba, the hybrid and C. virgata formed mor-
phologically discrete clades with only one overlapping
morphological character between C. virgata and the hy-
brid, namely the presence of ribbed fruit. Centella glauca
and C. macrocarpa are mainly distinguished by their habit
differences and the distinctly glaucous leaves of the for-
mer species (Schubert & Van Wyk 1996). A polytomy
results as these autapomorphic characters were not used
in the cladogram (Figure 3A). The morphological data
showed that the hybrid is intermediate between C. triloba
and the rest of the study group. Characters, such as the
presence of isobilateral leaves in the hybrid, group it with
C. macrocarpa, whereas the similarities with C. triloba
all appear to be symplesiomorphies. McDade (1990) also
found that hybrids do not lead to unresolved cladograms
with high levels of homoplasy, but that they emerge as
TABLE 5. — Genetic variation and differentiation within and between:
C. triloba (PI); a putative hybrid between C. triloba and C.
macrocarpa (H); C. macrocarpa (resprouter, population from
Kogel Bay) (P2); and C. macrocarpa (reseeder, population from
Swartberg Pass) (P3); C. virgata (V); and C. glauca (G)
Nei’s (1978) unbiased genetic distance between: C. triloba (PI); a
putative hybrid between C. triloba and C. macrocarpa (H); C. macro-
carpa (resprouter, population from Kogel Bay) (P2); and C. macrocarpa
(reseeder, population from Swartberg Pass) (P3); C. virgata (V); and C.
glauca (G)
174
Bothalia 26,2 (1996)
FIGURE 4. — The known geographical distribution of C. triloba , O; C.
hybrid (indicated by arrow); and C. macrocarpa, •.
basal to the clade which includes the most derived of the
two parents (Figure 3A, F).
In choosing C. triloba as outgroup, we expected the
topology A in Figure 3. Obviously, the similarities be-
tween the hybrid and C. macrocarpa would be interpreted
as synapomorphies. If we had chosen C. virgata or C.
glauca as outgroup, the reverse would have been true, i.e.
the hybrid would have grouped with C. triloba. According
to McDade (1995), ‘Hybrids may express the derived
states (apomorphies) of both parents, the apomorphies of
one parent, partially derived conditions that are interme-
diate between the parents, or states that are more extreme
than either (autapomorphies)’. If our putative hybrid was
unrelated to the two species with which it was found
growing, we would not have expected any intermediate
characters. The data in Table 3 show the following pattern
of character expression: four characters shared with C.
triloba , seven characters shared with C. macrocarpa, three
intermediate characters and three unique characters. It thus
seems reasonable to conclude that the relatively high
number of intermediate characters support our assumption
of hybrid origin based on field observation. Even if the
two plants found at Kogel Bay eventually turned out to
be a new species (we do not exclude this possibility) we
are arguing that it is of hybrid origin, being derived from
C. triloba and C. macrocarpa.
Allele frequencies calculated at CAP-1 (A, B and C
alleles), PGM-1 (A and B alleles) and PGM-2 (B and C
alleles) grouped the hybrid together with C. macrocarpa,
whereas the allele frequencies recorded for PER (B and
C alleles) coincided for the hybrid and C. triloba. The
larger number of allele frequencies shared with C. macro-
carpa resulted in the hybrid being nested in the polytomy
in Figure 3D. This is again in agreement with the propos-
als of McDade (1995), who suggested that the hybrid
would be placed near to the parent that has the most de-
rived characters. The combined morphological and genetic
data also present supporting evidence for the hypothesized
hybrid origin. When the hybrid was omitted, a fully re-
solved topology resulted, with a substantial improvement
in both the tree length and consistency index (Figure 3E).
The morphology gives only a partially resolved cladogram
(Figure 3A) but the C. glauca/C. macrocarpa polytomy
becomes fully resolved in the combined analysis, with the
genetic data responsible for the improvement. These
analyses are consistent with two generalizations: 1, re-
moval of the hybrid improves the resolution and 2, both
data sets (morphological and genetic) contribute to resolv-
ing different parts of the topology, as is evident in Figure
3A & F. C. virgata is clearly separated from C. macro-
carpa in all the analyses, which gives useful support for
considering it as a distinct species despite the close simi-
larity to some forms of C. macrocarpa (particularly the
reseeding forms). Fire-survival appears to be a ho-
moplasious character which has evolved convergently in
different species.
The C. macrocarpa populations were grouped together
on the final cladogram (Figure 3F). This is interesting
since the Swartberg population is a reseeder and the Kogel
Bay population is a resprouter, and the genetic distance
between them (Table 5) was less (0.091) than between the
C. macrocarpa reseeder population and another closely simi-
lar reseeder species, C. virgata (0.350). Likewise, the dis-
tance of 0.091 is less than the distance between the re-
sprouting C. glauca and the resprouting population of C.
macrocarpa (0.127). Thus the phenomena of reseeding and
resprouting (Schutte et al. 1995) may occur in a single spe-
cies.
There is a possible correlation in the genetic variation
of the reseeder populations and also of the resprouter
populations, i.e. that resprouters are genetically more vari-
able than the reseeder populations. The latter are more
prone to genetic bottlenecks, as was proposed by Schutte
et al. (1995), because presumably only one generation is
present at any given time, while the survival of the re-
sprouters leads to a mixture of numerous generations with-
in the same population. Further studies, including larger
sample sizes, are needed to confirm the relationship be-
tween genetic variation and fire-survival strategy in Cen-
tella.
The demonstration that hybridization occurs between
two species of Centella that are not sister taxa indicates
that a strictly hierarchical infrageneric classification may
be an unobtainable goal. Even the cladistic method may
become problematic, particularly when nodes are weakly
supported [see McDade (1992, 1995) for a detailed dis-
cussion of the impact of hybrids on cladistic analysis].
The assumption of divergent evolution needed for the
cladistic method may thus not be an accurate reflection
of the actual mechanisms of evolution within the genus.
If evolution is indeed reticulate, then the identification of
lineages of ancient hybrid origin becomes problematic
(McDade 1990). Nevertheless, the use of different data
sets in determining phylogenetic relationships between
Centella species, resulted in a closer proximation to the
true phylogeny than would have been possible with either
of the data sets (Shakalee & Whitt 1981; McDade 1995).
The new insight into evolutionary relationships in Cen-
tella provides a possible explanation for the reticulate pat-
tern of character expression in the genus. Adamson (1951)
was unable to create mutually exclusive infrageneric taxa
and coped with the problem by including some species
in more than one series. Roux et al. (1978), Winter &
Van Wyk (1995) and Allison (1995) have used cladistic
methods to study Apiaceae, marking the beginning of a
Bothalia 26,2 (1996)
175
rigorous, empirical approach to classification in this fam-
ily. However, if a hybrid disrupts the pattern of character
expression by inheriting the defining apomorphies of un-
related parents from different sections or series, then it
will not be possible to use cladistics in the normal way
because the method relies on divergence. McDade (1995)
highlighted several new attempts at resolving this prob-
lem.
ACKNOWLEDGEMENTS
We would like to thank Dr P. Tilney for assistance with
the GMA embedding and microtome sectioning. Financial
support from the Foundation for Research and Develop-
ment is gratefully acknowledged.
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Bothalia 26,2: 177-182(1996)
Red Data List of southern African plants. 1. corrections and additions
C. HILTON-TAYLOR*
The recently published Red Data List of southern Af-
rican plants (Hilton-Taylor 1996), by its very nature, is
dynamic and constantly changing. It was therefore pro-
posed that any corrections to the List, such as changes in
names, conservation status and distributions; deletions;
and new additions, should be published annually in
Bothalia to provide users of the List with information as
up-to-date as possible. These corrections and additions
will form a series similar to the contributions to the Flora
of southern Africa ( FSA ) published in Bothalia as ‘FSA
contributions’.
It should be noted that all corrections and comments
given here pertain only to the information published in
the Red Data List (Hilton-Taylor 1996), whereas the ad-
ditions are new records which must be added to the List.
Readers are referred to Hilton-Taylor (1996) for an expla-
nation of the methodology used and for definitions of all
the terms. The old IUCN Red Data Categories have been
used here again, rather than the new categories. A depar-
ture from the methodology described in Hilton-Taylor
(1996) is the use of the new South African provinces. The
abbreviations for the old provinces are, however, given in
brackets so that information is comparable to that in the
published List. Name changes for those taxa in the Red
Data List whose conservation status is not threatened (nt),
neither locally nor globally, are not given here. Changes
in conservation status (hereafter referred to as status) ap-
ply to both the local and global level, unless otherwise
specified. Corrections, additions or deletions of taxa in
the Extinct, Endangered or Vulnerable categories should
also be made in Appendix 2 where necessary.
CORRECTIONS
1. The status of Amphithalea rostrata A.L.Schutte &
B.-E.van Wyk, should be changed from Rare (R) to En-
dangered (E) as it is only known from a small population
near Pearly Beach on the Agulhas Plain.
2. The author citation for Apodolirion lanceolatum
should be (Thunb.) Baker not (L.f.) Benth. (Hilliard &
Burtt 1973).
3. A population of Argyrolobium crinitum (E.Mey.)
Walp. (Edwards 1996) was recently rediscovered (J.H.J.
Vlok pers. comm.). The conservation status of this attrac-
tive shrub should therefore be changed from Extinct (Ex)
to Endangered (E). The population comprises only a
dozen plants along a road verge and approximately 200
individuals in a remnant piece of Renosterveld vegetation
between wheat fields.
4. Argyrolobium sp. nov. ( Drege s.n., BM) has been
formally described by Edwards (1996) as Argyrolobium
* Ecology and Conservation Unit, National Botanical Institute, Private
Bag X7, Claremont 7737, Cape Town.
MS. received: 1996-07-09.
angustissimum (E.Mey.) T.J.Edwards. Its Extinct (Ex)
status is unchanged.
5. Brachystelma discoideum R.A.Dyer was incorrectly
listed as being globally Rare (R) and endemic to the FSA
region. S. Venter (pers. comm.) reports that it also occurs
in Botswana and Zimbabwe. Pending further details, its
global status and status in Botswana should be changed
to Insufficiently Known (K).
6. Ceropegia filiformis (Burch.) Schltr. is now known
to occur on the Great Karas Mountains in southern Na-
mibia where it is fairly common, and there have also been
a number of new collections from the Northern and East-
ern Cape (Bruyns 1995). The conservation status for this
species should therefore be changed from Rare (R) locally
(Cape) and globally to not threatened (nt) and a status of
nt should be listed under Namibia (N). The Vulnerable
(V) status under the Free State (O) remains unchanged as
it is only known from a single collection made in that
province (Bruyns 1995).
7. The status of Coelidium obtusilobum Granby, should
be changed from Rare (R) to Endangered (E) as it is only
known from the type collection made by Elsie Esterhuysen
in 1954. As this area has been fairly well collected this
species may even be Extinct (Ex).
8. Coelidium vlokii A.L.Schutte & B.-E.van Wyk, is
threatened by agricultural activities and its status should
be changed from Rare (R) to Endangered (E).
9. Nordal & Fangan (1994) have shown that Crinum
zeylanicum (L.) L., pro parte, sensu Nordal (1977) should
be regarded as a synonym of Crinum omatum (L.f. ex
Aiton) Bury. Crinum kirkii sensu Verdoorn in Bothalia 11:
38 (1973) non Baker, previously listed as a synonym of
C. zeylanicum (L.) L., should therefore also be listed as
a synonym of C. omatum. The conservation status re-
mains not threatened (nt) as it is a common widespread
taxon.
10. Cyclopia bowieana Harv. is not as scarce as pre-
viously thought and its status should be changed from
Indeterminate (I) to not threatened (nt).
11. Cyclopia burtonii Hofmeyr & E.Phillips (note the
corrected spelling of Hofmeyr) is a scarce endemic from
the upper slopes of the Great Swartberg Mountains. As it
is susceptible to frequent burning, the conservation status
should be changed from Indeterminate (I) to Rare (R).
12. Several attempts to relocate Cyclopia filiformis
Kies, have been unsuccessful; the conservation status
should therefore be changed from Insufficiently Known
(K) to Extinct (Ex) (A.L. Schutte pers. comm.).
13. The status of Cyclopia latifolia DC. should be
changed from Vulnerable (V) to Endangered (E) as it is
known only from two sites, one comprising 23 plants and
the other just a single individual (J.H. de Lange pers. comm.)
14. Cyclopia pubescens Eckl. & Zeyh. is severely threat-
ened by expanding urbanisation and its status should be
changed from Insufficiently Known (K) to Endangered (E).
178
Bothalia 26,2(1996)
15. The conservation status of Echiostachys spicatus
(Burm.f.) Levyns was incorrectly given as Vulnerable (V);
this should be changed to Rare (R) (Hall & Veldhuis 1985).
16. The realisation that plants named as Encephalartos
lebomboensis I.Verd. from the Lebombo Mountain range
in KwaZulu-Natal and Swaziland were incorrectly iden-
tified and represent a separate and undescribed species
(Vorster 1995, 1996e) means that the distribution and
status of E. lebomboensis needs to be corrected. E. le-
bomboensis only occurs in the Pongola River valley of
KwaZulu-Natal (KN) with a disjunct population on the
Lebombo range in Mpumalanga (T) (Vorster 1996e). As
a result of its reduced distribution range and population
size, the conservation status of this species should be up-
graded from Rare (R) to Vulnerable (V) in both provinces
and globally. The conservation status given under Swazi-
land should be deleted.
17. The population of E. lebomboensis in the Pongola
River valley has popularly been referred to as the ‘Piet
Retief’ form (Vorster 1995) and hence the use of Encepha-
lartos sp. nov. ‘Piet Retiefii’ ( Vorster 2977, PRE) in the
Red Data List (Hilton-Taylor 1996). Following the taxo-
nomic changes described above, this ‘name’ must now be
placed into synonymy under E. lebomboensis (Vorster
1995) .
18. Encephalartos sp. nov. ‘Msinga’ ( Vorster 2949,
PRE) was formally described by Vorster (1996c) as
Encephalartos msinganus Vorster. Its conservation status
is still Endangered (E).
19. Encephalartos sp. nov. ‘venitus’ ( Steyn 318, PRE)
was recently described by Vorster (1996d) as Encephalar-
tos venetus Vorster. Unbeknown to Vorster, a manuscript
decribing this taxon as Encephalartos nubimontanus
P.J.H.Hurter, was submitted almost simultaneously for
publication elsewhere (Hurter 1995; Hurter & Claassen
1996) . As E. nubimontanus was effectively and validly
published before E. venetus, it is accepted as the correct
name for the taxon while E. venetus is regarded as a syno-
nym. The conservation status of this taxon remains En-
dangered (E).
20. E.G.H. Oliver (pers. comm.) has found that speci-
mens previously named as Erica parvulisepala H.A. Baker,
match the type of Erica xanthina Guthrie & Bolus, and
as the latter is the older name it must take priority. E.
parvulisepala will therefore be reduced to synonymy un-
der E. xanthina (E.G.H. Oliver pers. comm.). As a result
of this taxonomic change, the conservation status of E.
xanthina must be changed from Insufficiently Known (K)
to Rare (R).
21. A recent study of Euphorbia bamardii A.C. White,
R.A.Dyer & B.Sloane (Knowles & Witkowsi 1996) has
shown that it is not as threatened as previously thought
and that its conservation status should be changed from
Endangered (E) to Vulnerable (V).
22. The distribution of Gladiolus appendiculatus
G.J. Lewis var. appendiculatus should be extended to
KwaZulu-Natal (KN) where it is also Rare (R).
23. The spelling of the specific epithet for Haemanthus
pauculiflorus Snijman & A.E.van Wyk should be cor-
rected to pauculifolius (Snijman & Van Wyk 1993).
24. The spelling of the specific epithet for Haworthia
mcmurtryi C.L.Scott should be changed to macmurtryi in
accordance with ICBN Article 60C.4 (Greuter et al. 1994).
25. The conservation status of Liparia angustifolia
(Eckl. & Zeyh.) A.L.Schutte should be changed from In-
sufficiently Known (K) to Endangered (E). This species
occurs in marshy areas along the coast between the Cape
Peninsula and Hermanus and many populations have been
destroyed by coastal developments.
26. The status of Liparia splendens (Burm.f.) Bos &
de Wit subsp. splendens, should be changed from Insuf-
ficiently Known (K) to Rare (R).
27. Liparia laevigata (L.) Thunb. is confined to the
Cape Peninsula, occurring only in marshy places on Con-
stantiaberg and Table Mountain; its status should be
changed from Insufficiently Known (K) to Rare (R).
28. Liparia myrtifolia Thunb., is now known to have
a wide distribution in the southwestern Cape occurring on
many mountain ranges where it appears to be secure. Its
conservation status should therefore be changed from In-
sufficiently Known (K) to not threatened (nt).
29. Lobostemon sp. nov. ( Buys 432, STE) is now for-
mally described as Lobostemon belliformis Buys (Buys &
Van der Walt 1996). Its conservation status needs to be
changed from Vulnerable (V) to Endangered (E), as only
twenty plants are known to survive in the wild (Buys &
van der Walt 1996).
30. Podalyria burchellii DC. is incorrectly recorded as
occurring in KwaZulu-Natal. Its status remains un-
changed.
31. Podalyria cordata (Thunb.) R.Br., although local-
ised, is not as threatened as previously thought and its
status should be changed from Vulnerable (V) to Rare
(R).
32. Recent attempts to relocate Podalyria microphylla
E.Mey. in its natural habitat, have been unsuccessful, and
its status should therefore be changed from Indeterminate
(I) to Extinct (Ex). Its habitat has largely been replaced
by urbanisation.
33. The <D> which occurs after the author citation for
Podalyria velutina Burch, ex Benth. is a typographical
error which should be deleted.
34. The following entry should be inserted into Appen-
dix 3: Polycarena capitatum Benth. sensu Hall et al.
(1980) see Phyllopodium heterophyllum (L.f) Benth.
35. The <D> which occurs after the author citation for
Pseudopentameris obtusifolia (Hochst.) N.P.Barker, is a
typographical error which should be deleted. The conser-
vation status of this species should also be changed from
Insufficiently Known (K) to Rare (R) (Barker 1995).
36. Investigation by P.V. Bruyns (pers. comm.) into the
identity and distribution of Senecio sarcoides (DC.) C.Jef-
frey, has shown that it is a far more common and wide-
spread species in the Western and Northern Cape (C) than
previously thought. Its conservation status should there-
fore be changed from Insufficiently Known (K) to not
threatened (nt).
37. For Sonderina streyi Merxm., the status of Insuf-
ficiently Known (K) was omitted from under Namibia.
38. Stirtonanthus taylorianus (L.Bolus) B.-E.van Wyk
& A.L.Schutte, is not as threatened as previously thought
and its status should be changed from Vulnerable (V) to
Rare (R). The spelling of the specific epithet of this taxon
was incorrectly given as ‘ tayloriana ’ in Appendix 2.
39. The following entry under Appendix 3 of the List,
Priestleya tomentosa (L.) Druce see Liparia vestita Thunb.
Bothalia 26,2(1996)
179
is incorrect, and should be replaced with the following:
Priestleya tomentosa sensu Hall & Veldhuis (1985) auct.
non (L.) Druce see Xiphotheca fruticosa (L.) A.LSchutte
& B.-E.van Wyk. The entry for L. vestita should be deleted
from Appendix 1 and a new entry for X. fruticosa should
be added. This endemic Western Cape (C) taxon was pre-
viously thought to be Rare (R) but is now better classified
as not threatened (nt) because its distribution has been
greatly expanded.
40. Xiphotheca canescens (Thunb.) A.L.Schutte & B.-
E.van Wyk, should be reclassified from Insufficiently
Known (K) to Vulnerable (V) because of its very restricted
distribution close to an expanding farming area.
41. Xiphotheca elliptica (DC.) A.L.Schutte & B.-E.van
Wyk, previously listed as Indeterminate (I) should be re-
classified as not threatened (nt) as it is a fairly common
high altitude mountain species.
42. Xiphotheca guthriei (L.Bolus) A.L.Schutte & B.-
E.van Wyk, is severely threatened by farming practices
and its status should therefore be changed from Insuffi-
ciently Known (K) to Endangered (E).
43. The status of Xiphotheca lanceolata (E.Mey.) Eckl.
& Zeyh., must be changed from Indeterminate (I) to En-
dangered (E) as it is severely threatened by urban expan-
sion on the Cape Flats.
44. The status of Xiphotheca reflexa (Thunb.) A.L.Schutte
& B.-E.van Wyk, must be changed from not threatened (nt)
to Vulnerable (V) as it only occurs on the Western Cape (C)
lowlands in areas now largely replaced with agricultural
lands and urban areas. Recent searches for this species were
unsuccessful (A.L. Schutte pers. comm.).
45. The status of Xiphotheca tecta (Thunb.) A.L.Schutte
& B.-E.van Wyk, should be changed from Rare (R) to not
threatened (nt) as it has a wide distribution and is relatively
safe.
46. Erica sp. nov. ‘flexistyla' ( Oliver 10558 STE) has
been formally described by Oliver & Oliver (1995) as
Erica flexistyla E.G.H.Oliv. Its Endangered (E) status re-
mains unchanged.
47. Erythrophysa transvaalensis I.Verd. is not endemic
to the FSA region as it also occurs in southwestern Zim-
babwe, it’s rare (R) status remains unchanged.
48. The <D> which occurs after the author citation for
Nemesia micrantha Hiem, is a typographical error which
should be deleted.
49. The <D> which occurs after the author citation for
Nemesia picta Schltr., is a typographical error which should
be deleted.
50. Based on information from C.J. Geldenhuys (pers.
comm.) the status of Ocotea bullata (Burch.) Baill. should
be changed from a question mark (?) under Cape (C) to
not threatened (nt), and therefore its global status should
also be changed from Vulnerable (V) to not threatened
(nt).
51. Ocotea kenyensis (Chiov.) Robyns, is not endemic
to the FSA region and its global status should be changed
from (R) to not threatened (nt).
52. The status of Strelitzia alba (L.f.) Skeels, should
be changed from Indeterminate (I) in the Cape (C) and
globally, to not threatened (nt), (C.J. Geldenhuys pers.
comm.).
53. The global conservation status of Strelitzia juncea
Link, should be Rare (R).
54. The global status of Vlokia ater S.A. Hammer,
should be Rare (R).
ADDITIONS
All taxa listed here are endemic to the FSA region and
the global conservation status is therefore the same as the
local status (province and/or country) unless otherwise in-
dicated. The status of newly recorded taxa not previously
known to occur in the FSA region is not given here, unless
the taxon concerned is already classified as threatened
elsewhere. The conservation status of these taxa will be
evaluated at a later stage once more collecting information
is available. The additions are listed in alphabetical order
by family and genus.
Aizoaceae
1. Delosperma guthriei Lavis, is a Vulnerable (V) spe-
cies confined to coastal areas at Kleinmond and Hermanus
in the Western Cape (C) where it is threatened by coastal
development and alien plant invasions.
2. Drosanthemum austricolum L.Bolus, is an Endan-
gered (E) limestone endemic from the Agulhas Plain,
Western Cape (C) where it is confined to an area densely
infested by alien species.
3. Lampranthus stenus (Haw.) N.E.Br., is a Rare (R)
species recorded from lowland coastal sites from the Cape
Peninsula to Hermanus, Western Cape (C).
Apiaceae
1. Centella temata M.T.R. Schubert & B.-E.van Wyk,
is an Insufficiently Known (K) species known only from
the type collection made by Elsie Esterhuysen in the Ced-
erberg, Western Cape (C) in 1962 (Schubert & Van Wyk
1995).
2. Centella umbellata M.T.R. Schubert & B.-E.van
Wyk, is an Insufficiently Known (K) species known only
from the type collection made by Elsie Esterhuysen on
the mountain slopes above Porterville in the Western Cape
(C) in 1979 (Schubert & Van Wyk 1995).
Asteraceae
1. Hoplophyllum ferox Sond., is a very poorly known
species collected from only two localities in the Western
Cape (C) portion of the Great Karoo last century. Until
further information is obtained, its conservation status is
Indeterminate (I).
2. Metalasia octoflora DC., is a Vulnerable (V) species
confined to Renosterveld fragments on the lowlands along
the west coast of the Western Cape (C).
Euphorbiaceae
Euphorbia versicolores G.Will., is a Rare (R) species
known only from two small colonies in the central
Richtersveld of the Northern Cape (C). Although only ap-
proximately 20 plants in total are known, this species is
not classified as Endangered or Vulnerable at this stage,
as it is not considered to be severely threatened at present.
There are probably other undiscovered populations in this
botanically poorly explored remote, mountainous area
(Williamson 1995).
180
Bothalia 26,2(1996)
Fabaceae
A large number of species in this family have been
added to the Red Data List as a result of a recent detailed
systematic study of the tribes Podalyrieae and Liparieae
(Schutte 1995a):
1 . Amphithalea alba Granby, is a Vulnerable (V) lime-
stone endemic from the Agulhas Plain, Western Cape (C).
2. Amphithalea axillaris Granby, is a Rare (R) species
from the Langeberg and Outeniqua Mountains, Western
Cape (C).
3. Amphithalea biovulata (Bolus) Granby, is a Vulner-
able (V) species recorded mainly from the Agulhas Plain,
Western Cape (C). It is threatened by agricultural activi-
ties.
4. Amphithalea ericifolia (L.) Eckl. & Zeyh. subsp.
erecta Granby, is an Endangered (E) taxon from the west
coast lowlands, Western Cape (C).
5. Amphithalea ericifolia (L.) Eckl. & Zeyh. subsp.
scoparia Granby, is a Rare (R) taxon from the Hottentots
Holland and Riviersonderend Mountains, Western Cape
(C).
6. Amphithalea ericifolia (L.) Eckl. & Zeyh. subsp.
minima Granby, is a Rare (R) highly localised endemic
taxon from the Riviersonderend Mountains, Western Cape
(C).
7. Amphithalea imbricata (L.) Druce, is a Rare (R)
species from the Cape Peninsula and Hottentots Holland
Mountains in the Western Cape (C).
8. Amphithalea oppositifolia L.Bolus, is an Endangered
(E) species from the Kogelberg and Betty’s Bay area,
Western Cape (C).
9. Amphithalea sericea Schltr., is a Vulnerable (V) spe-
cies from the Agulhas Plain, Western Cape (C).
10. Amphithalea speciosa Schltr., is an Endangered (E)
species from the Elim area near Bredasdorp, Western Cape
(C), where it is threatened by agricultural activities.
1 1 . Amphithalea stokoei L.Bolus, is an Endangered (E)
species known only from a single locality on the Hotten-
tots Holland Mountains, Western Cape (C).
12. Ampithalea tomentosa (Thunb.) Granby, is a Vul-
nerable (V) species occurring on the coastal lowlands be-
tween Betty’s Bay and Riversdale, Western Cape (C),
where it is threatened by coastal development and agri-
cultural activities.
13. Amphithalea virgata Eckl. & Zeyh., occurs only in
the Caledon area of the Western Cape (C), where it is
highly Endangered (E) by agricultural activities.
14. Coelidium bowiei Benth., is a Vulnerable (V)
highly localised species from the Houhoek, Caledon and
Kleinmond areas, Western Cape (C).
15. Coelidium flavum Granby, is a poorly known but
apparently Rare (R) species confined to a small area on
the Outeniqua Mountains, Western Cape (C).
16. Coelidium minimum Granby, is a Rare (R) species
known only from the type collection made near Nie-
woudtville, Northern Cape (C). This area has been rela-
tively well collected and it is surprising that no further
collections have been made.
17. Coelidium pageae L.Bolus, is a Rare (R) species
confined to the Montagu area, Western Cape (C).
18. Coelidium purpureum Granby, is a Rare (R) species
known only from the type collection made in the Koue
Bokkeveld Mountains, Western Cape (C).
19. Coelidium spinosum Harv., is a Rare (R) species
recorded only from the Hex River Valley and Touwsrivier
area, Western Cape (C).
20. Cyclopia laxiflora Benth., is known only from a
few old records collected in the Knysna-Plettenberg Bay
area, Western Cape (C). Despite numerous searches for
this species, no traces of it have been found; it is therefore
presumed to be Extinct (Ex).
21. Liparia bonaespei A.L.Schutte, is a poorly known
Rare (R) species from the Hottentots Holland Mountains,
Western Cape (C), where it has only been collected a few
times (Schutte 1995b).
22. Liparia calycina (L.Bolus) A.L.Schutte, is a Rare
(R) species confined to a few localities on the Hottentots
Holland and Klein River Mountains, Western Cape (C).
23. Liparia congesta A.L.Schutte, is a Rare (R) species
known only from a few localities in the Cederberg, West-
ern Cape (C) (Schutte 1995b). Only one plant could be
found during a recent search for this species and it is
probably susceptible to frequent burning (A.L. Schutte
pers. comm.).
24. Liparia genistoides (Lam.) A.L.Schutte, is a very
Rare (R) species known only from two disjunct localities
on the Kamanassie and Kouga Mountains in the Eastern
Cape (C).
25. Liparia graminifolia L., has not been collected
since 1829, when it was found by Ecklon near Simon’s
Town on the Cape Peninsula, Western Cape (C). This spe-
cies probably became Extinct (Ex) as a result of urban
development.
26. Liparia parva Vogel ex Walp., is a Rare (R) species
confined to the Cape Peninsula, Western Cape (C).
27. Liparia rafnioides A.L.Schutte, is a Rare (R) spe-
cies known only from a single locality in the Kogelberg
Mountains, Western Cape (C) (Schutte 1995b).
28. Liparia striata A.L.Schutte, is confined to two hills
near Swellendam-Heidelberg, Western Cape (C), where
it is seriously Endangered (E) by farming activities
(Schutte 1995b).
29. Podalyria argentea (Salisb.) Salisb., is a Rare (R)
species restricted to the Cape Peninsula, Cape Flats and
Hottentots Holland Mountains, Western Cape (C). Many
of the populations on the lower slopes and flats have been
destroyed by urbanisation (A.L. Schutte pers. comm.).
30. Podalyria lanceolata (E.Mey.) Benth., is a Rare
(R) Langeberg endemic, Western Cape (C).
31. Podalyria orbicularis (E.Mey.) Eckl. & Zeyh., is
a Rare (R) species known only from the Swartberg near
Caledon and Genadendal Mountain, Western Cape (C).
32. Podalyria pearsonii E.Phillips, is a Rare (R) spe-
cies recorded only from a few sites on the Bokkeveld
escarpment near Niewoudtville in the Northern Cape and
the Gifberg in the Western Cape (C).
33. Podalyria reticulata Harv., is a Rare (R) species
known only from three old collections made on the Swart-
berg near Caledon, Western Cape (C). This taxon is doubt-
fully distinct from P. orbicularis (see above).
34. Podalyria sericea (Andrews) R.Br., is a Vulnerable
(V) species confined to granite hills on the Cape Peninsula
Bothalia 26,2 (1996)
181
and Cape Flats, Western Cape (C), where it is threatened
by urbanisation.
35. Stirtonanthus chrysanthus (Adamson) B.-E.van
Wyk & A.L.Schutte, is a highly localised Rare (R) en-
demic of the Klein Swartberg Mountains, Western Cape
(C).
36. Stirtonanthus insignis (Compton) B.-E.van Wyk &
A.L.Schutte, is a Vulnerable (V) species, limited to the
mountains around Montagu. Some populations are threat-
ened by farming activities.
37. Xiphotheca cordifolia A.L.Schutte & B.-E.van
Wyk, is a Rare (R) species known only from a few col-
lections made in the Hex River Mountains, Western Cape
(C).
Iridaceae
1. Galaxia parva Goldblatt, is a Vulnerable (V) species
confined to a single locality on the Agulhas Plain near
Bredasdorp, Western Cape (C) (Goldblatt 1984).
2. Gladiolus abbreviatus Andrews, is a Vulnerable (V)
species from the Western Cape (C). It was once fairly
widespread but is now limited to remnant Renosterveld
patches and road verges (J.C. Manning pers. comm.).
3. Moraea regalis Goldblatt & J.C.Manning, is a Rare
(R) species known only from a single locality at De Rust
in the Little Karoo, Western Cape (C) (Goldblatt & Man-
ning 1995).
Poaceae
Pseudopentameris brachyphylla (Stapf) Conert, is a
Rare (R) species from the Betty’s Bay area of the Western
Cape (C), which although not presently under threat, is
‘potentially threatened’ because of its narrow habitat re-
quirements (Barker 1995).
Orchidaceae
Following a systematic revision of Disa draconis (L.f.)
Sw. by Johnson & Linder (1995), this Western Cape (C)
species needs to be added to the Red Data List as Endan-
gered (E). One small population is protected at Rondevlei,
a pocket-sized nature reserve, while the remainder are all
under severe threat from urban and agricultural develop-
ment (Johnson & Linder 1995).
Zamiaceae
1. Encephalartos hirsutus P.J.H. Hurter, is a Vulnerable
(V) species known only from three small, widely sepa-
rated populations in the Northern Province (T) (Hurter &
Glen 1996).
2. Encephalartos aplanatus Vorster, is a critically En-
dangered (E) species which occurs in Swaziland (S); only
a single population is known, comprising just a few scat-
tered individuals (Vorster 1996a).
3. Encephalartos brevifoliolatus Vorster, is an Endan-
gered (E) species known only from five individuals widely
scattered over a few square kilometers in the Northern
Province (T) (Vorster 1996b).
4. Encephalartos senticosus Vorster, is a Rare (R) spe-
cies occurring along the Lebombo Mountain range in
northern KwaZulu-Natal (KN) and Swaziland (S) (Vorster
1996e). This species is not endemic to the FSA region, as
it also occurs in southern Mozambique where it is con-
sidered to be Vulnerable (V) (Bandeira et al. 1996). Large
numbers of this species have been removed from the wild
for horticultural purposes in the last 50 years (Vorster
1996e) and if this trend continues, its status will have to
be changed to Vulnerable (V).
SUMMARY
As a result of the changes in status and the additions
to the Red Data List given above, the statistics on the
number of taxa in each threatened category need to be
changed [see Table 2 of the Red Data List, Hilton-Taylor
(1996)]. Sixty taxa are now presumed to be Extinct (Ex),
270 are Endangered (E), 432 Vulnerable (V), 1436 Rare
(R), 375 Indeterminate (I) and 905 Insufficiently Known
(K). The numbers in all the categories, except for the last
two, have increased. Twelve taxa were removed from the
List and 55 were added, giving a new total of 3 478 glob-
ally threatened taxa in the FSA region.
ACKNOWLEDGEMENTS
The author wishes to thank J.B.P. Beyers, P.V. Bruyns,
J.H. de Lange, T.J. Edwards, C.J. Geldenhuys, J.C. Man-
ning, E.G.H. Oliver, A.L. Schutte, S. Venter, J.H.J. Vlok
and P. Vorster for the provision of new data and for their
comments on the corrections and additions to the Red
Data List.
REFERENCES
BANDEIRA, S O., MARCONI, L. & BARBOSA, F. 1996. Preliminary
studies of threatened plants of Mozambique. In L.J.G. van der
Maesen, X.M. van der Burgt & J.M. van Medenbach de Rooy, The
biodiversity of African plants : 306-309. Kluwer Academic Publi-
shers, Dordrecht.
BARKER, N.P. 1995. A systematic study of the genus Pseudopentameris
(Arundinoideae: Poaceae). Bothalia 25: 141-148.
BRUYNS, P.V. 1995. New records and new species of Asclepiadaceae
from Namibia. Bothalia 25: 155-172.
BUYS, M.H. & VAN DER WALT, J.J.A. 1996. A new species of Lo-
bostemon section Grandiflori (Boraginaceae) from South Africa.
South African Journal of Botany 62: 31-35.
EDWARDS, T.J. 1996. Rare or extinct species of Argyrolobium. Bothalia
26: 42-46.
GOLDBLATT, P. 1984. New species of Galaxia (Iridaceae) and notes on
cytology and evolution in the genus. Annals of the Missouri
Botanical Garden 71: 1082-1087.
GOLDBLATT, P. & MANNING, J.C. 1995. New species of southern
African Moraea (Iridaceae-Iridoideae), and the reduction of
Rheome. Novon 5: 262-269.
GREUTER, W. et al. 1994. International code of botanical nomenclature
(Tokyo Code). Regnum Vegetabile 131.
HALL, A.V., DE WINTER, M„ DE WINTER, B. & OOSTERHOUT,
S.A.M. 1980. Threatened plants of southern Africa. South African
National Scientific Programmes Report No. 45. CSIR, Pretoria.
HALL, A.V. & VELDHUIS, H. A. 1985. South African Red Data Book:
plants — Fynbos and Karoo Biome. South African National Scien-
tific Programmes Report No. 117. CSIR, Pretoria.
HILLIARD, O.M. & BURTT, B.L. 1973. Notes on some plants of south-
ern Africa chiefly from Natal: III. Notes from the Royal Botanic
Garden, Edinburgh 32: 303-387.
HILTON-TAYLOR, C. 1996. Red Data List of southern African plants.
Strelitzia 4. National Botanical Institute, Pretoria.
HURTER, P.J.H. 1995. Two newly described cycads from Africa. Phy-
tologia 78: 409-416.
HURTER, P.J.H. & CLAASSEN, I. 1996. Encephalartos nubimontanus
Hurter. Encephalartos 45: 4-11.
182
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HURTER, RJ.H. & GLEN, H.F. 1996. Encephalartos hirsutus (Zami-
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JOHNSON, S.D. & LINDER, H.P. 1995. Systematics and evolution of
the Disa draconis complex (Orchidaceae). Botanical Journal of
the Linnean Society 118: 289-307.
KNOWLES, L. & WITKOWSKI, E.T.F. 1996. Conservation biology of
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VORSTER, P. 1996c. Encephalartos msinganus (Zamiaceae): a new
species from KwaZulu-Natal. South African Journal of Botany
62: 67-70.
VORSTER, P. 1996d. Encephalartos venetus (Zamiaceae): a new species
from the Northern Province. South African Journal of Botany 62:
71-75.
VORSTER, P. 1996e. Encephalartos senticosus (Zamiaceae): a new spe-
cies from northern KwaZulu-Natal and Swaziland. South African
Journal of Botany 62: 76-79.
WILLIAMSON, G. 1995. Euphorbia versicolores, a new species from
the northwestern Cape, South Africa. Cactus and Succulent Jour-
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Bothalia 26,2: 183-184(1996)
Book Reviews
DYNAMICS OF WEED POPULATIONS by R.D. Cousens & M. Mor-
timer. 1995. Cambridge University Press , The Edinburgh Bldg,
Shaftsbury Rd, Cambridge CB2 2RU, UK. Pp. 332. Hard cover: ISBN
0 521 49649 7, price £50.00, US$79.95. Soft cover: ISBN 0 521 49969
0, price £17.95, US$29.95.
The study of weed dynamics in agricultural situations has for centu-
ries been largely directed at solving problems (how can the weeds be
killed?). Large-scale problems associated with the invasive spread of alien
plants into natural ecosystems is a much more recent phenomenon, and
agricultural weed scientists have paid little attention to the study of the
dynamics of such invasions. The study of invasions of natural systems
has been the domain of ecologists who see the phenomenon as a special
case of succession with considerable potential for shedding light on the
forces that structure communities. With a few exceptions, studies of such
invasions have paid little attention to advances in ‘classical’ weed science.
Recent synthesis volumes in these disciplines have further contributed to
the widening chasm. The authors of this timely volume set out to integrate
recent advances from the two related fields. They believe there is a need
to ‘establish a framework of interlocking components, which together
help us to predict and to manage the size of populations’.
In attempting this ambitious task, Cousens & Mortimer explore the
dynamics of geographic range expansion (at various scales); dispersal
within and between populations; processes involved in the regulation of
population density; the intrinsic and extrinsic dynamics of population
density; and the spatial dynamics of weed populations. The penultimate
chapter is devoted to the evolution of herbicide resistance. I particularly
enjoyed the section on the dynamics of geographic range expansion.
Here, the authors point out the very real problem of recognizing phases
in the invasion process which bedevil attempts at generalization and
prediction. Extensive coverage is given to the well-studied invasion of
large parts of North America by Bromus tectorum. They re-analyze data
on the spread of this grass species to show how difficult it is to distinguish
exponential increase from a two-phase increase incorporating a lag phase.
Another chapter I found very useful was the one on mathematical models
of population density. I can think of no better introduction on the topic.
The case studies are fairly heavily based in favour of Australian, UK
and, to a lesser extent. North American examples. Herbaceous plants are
also given much better coverage than woody plants. Despite this, the
examples that are chosen cover a very wide range of situations, and
everyone with an interest in weeds will find points that relate to their
particular interest. As a plant ecologist working mainly on the ecology
of woody plants invading natural vegetation, I certainly found a great
deal of interest in this book. I unreservedly recommend it to all plant
population ecologists and other researchers with an interest in manipu-
lating plant populations. The volume would also be ideal as the founda-
tion for a post-graduate course on the subject.
The book is a pleasure to read; it is very well written, the concepts
are clearly developed, and there is a good balance between text, figures
and tables. I found no typographical errors.
D M. RICHARDSON*
* Institute for Plant Conservation, Botany Dept, University of Cape
Town, 7700 Rondebosch, Cape Town.
THE ANTHER: FORM, FUNCTION AND PHYLOGENY, edited by
W.G. D’Arcy and R.C. Keating. 1996. Cambridge University Press, The
Edinburgh Building, Shaftesbury Road, Cambridge CB2 2RU, UK. Pp.
xii + 351, 267 illustrations (line drawings, cladograms and photomicro-
graphs), 13 tables. Size: 253 x 193 mm. ISBN 0 521 48063 9. Price:
hard cover, £55.00, US$80.00.
A comprehensive work on the male reproductive structures of flow-
ering plants has for many years been overdue. This book with its focus
on the anther as the fundamental unit of the male floral organ, provides
an authoritative summation of the present state of knowledge and theo-
ries, scattered in botanical literature, about the stamen of extinct and
extant angiosperms. Essentially, the book is derived from papers read at
the 1993 International Botanical Congress in Yokohama, Japan, but ad-
ditional chapters have been added to include, inter alia, a bibliography
(Chapter 13) which lists 1 347 references, indexed by subject and plant
family, relating to stamen morphology and anatomy.
Although the first chapter is basically intended to outline the contents
of the book and to act as a primer for understanding the terms and concepts
used to describe stamen structure and function, a formidable amount of
detailed information is presented concisely, but in an easily readable style.
Summaries of classical and current theories on stamen design, arrangement,
structure, function and evolution are interwoven with data on morphological
and anatomical characters and are related to modes of pollen delivery and
pollinator behavioural patterns. Importandy, the introductory chapter stresses
the necessity of solid primary observations, both in the field and the labo-
ratory, for the collection of new information and the alignment and testing
of facts and theories about stamens.
Keating’s overview (Chapter 12) of the literature dealing with diverse
techniques for studying the anther is a valuable contribution to the vol-
ume. Initially, methods for preparing specimens intended for serial sec-
tioning and light and electron microscopical studies are discussed and
the advantages of hand sectioning, various wet mount techniques, smears
and clearing agents are pointed out. Numerous recendy developed meth-
ods to elucidate the biochemical and genetic basis behind the known
structural complexity in developing and in mature anthers are referred
to and the well-chosen references contribute more valuable information.
This review of methodology could, in combination with D’ Arcy’s outiine,
serve as an excellent textbook on the angiosperm stamen for biologists
at the non-specialist level.
For specialists in the subdisciplinary fields of palaeobotany, taxon-
omy, comparative morphology, anatomy and embryology, this beautifully
illustrated book offers much useful and interesting information and an-
swers to questions which otherwise would require searching through
many reference works. The mystery of the origin(s) of flowering plants
is explored by using the results of cladistic analyses (Chapter 3) or evi-
dence from the fossil record (Chapter 2). Since the stamen has been
recognized as a model organ for addressing basic and applied problems
concerning plant cell biology, morphogenesis, breeding and evolution,
fundamental questions such as: ‘Are stamens and carpels homologous?’
are considered and critically evaluated from divergent perspectives.
Anatomists and embryologists who are not particularly inclined towards
phylogenetic studies, will appreciate contributions dealing with hetero-
chrony, resorption tissue (the calcium oxalate package) and the diversity
of endothecium patterns in anthers, while pollination ecologists will wel-
come Peter Bemardt’s review on the role of the anther in zoophily.
The volume is elegant in style, has a good standard of finish and the
format makes it a delight to handle. Errors in the text and references are
few and insignificant. The illustrations are superb, informative and not
a recycling of old ones, seen many times before. Literature cited in each
chapter is listed separately by contributors and shows that the book is
up to date: of the nearly 1 300 references included in Chapters 1-12 and
ranging from 22-226 per chapter, 40% were published later than 1985,
while less than 30% are older than 1970.
In conclusion, I must stress that the title of the book is extremely
misleading. Although this is the first-ever authoritative work on angio-
sperm stamens, it is also a comprehensive source of references on floral
biology and will, in time, become indispensible to both students and
professional biologists. The price easily puts it in the realm of constitu-
tional purchasing and the book should be available in all botanical li-
184
Bothalia 26,2(1996)
braries to stimulate and enhance research in numerous fields of basic
and applied biology.
E.M.A. STEYN*
* Research Directorate, National Botanical Institute, Private Bag X101,
Pretoria 0001.
TERRESTRIAL ORCHIDS: FROM SEED TO MYCOTROPHIC
PLANT, by HANNE N. RASMUSSEN. 1995. Cambridge University
Press , The Pitt Building, Trumpington St., Cambridge CB2 1RP, England.
Pp. 444. ISBN: 0 521 45165 5. Price: hard cover: £45.00, US$64.95.
There is an imbalance in our general interest in orchids, as epiphytic
species are preferred to terrestrial ones by most people, botanists as well
as amateurs. Probably the main reason is that (unlike most terrestrial
orchids) epiphytic species have frequently showy flowers and are fairly
easy to cultivate. Epiphytic orchids are also the larger group, and it is
estimated that approximately 70% of orchid species grow as epiphytes.
Consequently, we know a great deal more about the biology of the epi-
phytic orchids than about that of the terrestrial species.
Nevertheless, the biology of terrestrial orchids has, for a variety of
reasons, fascinated some botanists and plant lovers for centuries, but sev-
eral aspects of it are still poorly understood. Many terrestrial species are
rather rare and unpredictable in their appearance which is largely due to
their heavy dependency on the associated mycorrhiza — sometimes thriv-
ing underground for several years and then suddenly emerging from the
soil with a flowering shoot — and this sporadic nature enhances their ap-
peal to the botanically interested community. The life cycle of terrestrial
orchids has been something of a mystery for centuries because of their
underground stage which is difficult to study. Especially, the germination
of the minute seeds is still incompletely known. Due to their mycotrophy
the cultivation of terrestrial species is difficult for the most part, but as
a consequence, maintaining them in culture has become a new challenge
for the advanced grower. Especially at a time when many terrestrial or-
chids are becoming increasingly rare and their conservation is rapidly
becoming a high priority, a better understanding of their germination and
growth requirements is vital.
In view of our incomplete knowledge Hanne Rasmussen’s compre-
hensive book on the life cycle and the fungal associations of terrestrial
orchids is a most welcome addition to the orchid literature. This book
is the first that deals extensively with the aspect of the biology of the
terrestrial orchids since the work of Burgeff in the first half of the century
(Burgeff 1911, 1936, 1954). The present book deals with the holarctic
species only. The first few chapters explain the properties of their dust-
like seeds, the germination requirements, germination processes, associ-
ated mycorrhizal fungi, as well as the development of the seedling to
the mature plant. Ecological and evolutionary consequences of the or-
chid-mycorrhiza symbiosis are also outlined. A chapter on substrates and
the propagation with relation to mycorrhizal fungi describes our present
state of knowledge in this field, and will be particularly useful for orchid
growers. The life cycle of the holarctic species of 36 genera of terrestrial
orchids is discussed in detail in a subsequent chapter. The genera are
mostly European, but a few North American and Asian genera have also
been included. Throughout the text the available literature on these as-
pects is reviewed, and is supplemented by personal observations which
the author has made in the field as well as in laboratories, both in Mary-
land, USA, and in Copenhagen, Denmark. Obviously several aspects are
still insufficiently understood, and such gaps in our knowledge are
pointed out by the author, and it is hoped that future students will be
able to solve some of these problems. Appendix A lists the nutrient sub-
strates mentioned in the text, and Appendix B is a list of scientific names
and synonyms. An extensive list of references and an index conclude the
book.
The text is clearly and concisely written, and is illustrated by black
and white line drawings and photographs. There are very few errors, like
genus Hammerbya instead of Hammarbya (p. 135) or Cypripedium cal-
ceolus var. parviflorus and var. parviflora instead of var. parviflorum (p.
241). Considering the abundance of terrestrial orchids in southern Africa,
the general chapters of this book are certainly relevant to our region
although it has not been established yet if, and how far, the data obtained
in orchids from the northern hemisphere can also be applied to southern
hemisphere species. Australian and South African terrestrial orchids are
occasionally mentioned in the text, but a slight limitation of the new
book to us south of the equator is its geographical range. The publica-
tion— particularly the description of the life cycle of various genera — will
certainly be of greater interest to readers in the northern hemisphere or
at least to those with a strong interest in orchids from that hemisphere.
On the whole this new book can be widely recommended, both for
botanists and horticulturalists. Being invaluable for our understanding of
the biology of the terrestrial orchids, it will also appeal to conservationists
who will find, e.g. information on species suitable for re-introduction, or
information on germination and growth requirements of certain rare spe-
cies.
REFERENCES
BURGEFF, H. 1911. Die Anzucht tropischer Orchideen aus Samen. Jena.
BURGEFF, H. 1936. Samenkeimung der Orchideen und Entwicklung
ihrer Keimpflanzen. Jena.
BURGEFF, H. 1954. Samenkeimung und Kultur europaischer Erdor-
chideen, nebst Versuchen zu ihrer Verbreitung. Gustav Fischer
Verlag, Stuttgart.
H. KURZWEIL*
* Compton Herbarium, National Botanical Institute, Private Bag X7,
Claremont 7735, Cape Town.
Bothalia 26,2: 185-199(1996)
National Botanical Institute South Africa: list of staff and publications,
31st May 1996
Compiler: B.A. Momberg
CHIEF DIRECTORATE
CAPE TOWN
Huntley, Prof. B J. M.Sc. Chief Director
Stafford, Ms D.L. Senior Secretary
Davidson, D.C. Head: Graphic Services
Mantlana, M. Deputy Director: Public Affairs
Mirkin, Ms Y.A. Senior Typist/Receptionist
COMMUNICATION AND PROMOTIONS— CAPE TOWN
Coningsby, Ms N.J. B .Journalism(Hons.). Communications Officer
Palmer, Ms G. Events co-ordinator
GRAPHIC SERVICES— CAPE TOWN
Davidson, D.C. B.A. (Hons.), PRISA Final Dipl.
Loedolff, Mrs J. B.Sc.(Ind.Tech.). Photographer/Illustrator
Sims, W.D. Graphics Assistant
ADMINISTRATION DIRECTORATE
|~CAPE TOWN j
Jordaan, A.W. Adv.Dip.B. & A. Director: Personnel and Finance
Comelissen, Mrs A.M. Secretary to the Board
Hughes, W.S.G. Head: Finance
Van Zyl, J.M. Head: Personnel
186
Bothalia 26,2(1996)
FINANCE— CAPE TOWN
Hughes, W.S.G. B.Com., C.A.(SA). Deputy Director
Armitage, Mrs C.S. Senior Accountant Roman, D.C. Accounts Clerk: cashier
Kriel, Ms C.M.M. Senior Accounts Clerk: debtors Simon, Ms P.I. Assistant Accountant: creditors
Manjo, C. Accounts Clerk: finance Traut, G.D. Chief Accounts Clerk: salaries
Neuwirth, Ms E.V. Accountant: salaries Yeomen, Ms I.N. Senior Accounts Clerk: creditors
Petersen, R.E. Senior Accounts Clerk: salaries
PERSONNEL— CAPE TOWN
Van Zyl, J.M. M.Econ. (Industrial Psychology). Assistant Director: Personnel
Barnard, Mrs A. Assistant Training Officer
Crowie, A.C. Courier
Dollie, Mrs NJ. Administration Aid
Haupt, Ms C.S. Administration Aid
Mulder, Mrs G.P. Senior Typist
Nicholas, Mrs W.L. Photocopy Machine Operator
Oelofse, Ms V.S. Senior Personnel Clerk
Paulse, Mrs D.W. Registration Clerk
Staal, P.B. Senior Personnel Practitioner: labour relations
Woodman, Ms L.R. Telephonist/Receptionist
EDUCATION & INFORMATION DIRECTORATE
CAPE TOWN
Low, A.B. M.Sc. Deputy Director: Education & Information
Ashwell, Ms A.N. Head: Education
Lovric, Mrs N. Typist/secretary
EDUCATION
CAPE TOWN
Ashwell, Ms A.N. M.Ed. Assistant Director
Constable, M.D. Bus Driver Sibikile, Mrs A. Administration Aid. Gold Fields Centre
Cupido, Ms M. Administrative Assistant (contract worker)
Huet, Ms H. Senior Administration Clerk. Gold Fields Centre Slattery, Mrs C.M. B.Sc.(Hons.), H.D.E. Education Officer
Mkefe, T.J. SPTD, Education Outreach Officer Tyokolo, Ms S.E. SPTD, Education Outreach Officer
WITWATERSRAND
Moteka, N.D. PTC, Education Outreach Officer
Van der Westhuizen, Mrs S. M.Sc. Education Officer
Vlok, Mrs S. B.Com. Administrative Assistant
PRETORIA
Symonds, Ms A.M. N.D. (Nature Cons.), H.D.E. Education Officer
INTERPRETATION— PRETORIA
Joffe, Mrs H. B.Sc. Garden Utilization Officer
Bothalia 26,2 (1996)
187
GARDENS DIRECTORATE
CAPE TOWN
Botha, D.J. D.Sc. Director: Gardens
Behr, Ms C.M. Curator: Harold Porter NBG
Britz, R.M. Curator: Free State NBG
Carstens, Mrs M.M. Secr.Dip. Senior Secretary
Chaplin, P.J. Curator: Witwatersrand NBG
Engelbrecht, B. Head: Support Services
Heilgendorff, J.P. Curator: Pretoria NBG
Kluge, J.P. Curator: Lowveld NBG
Oliver, I.B. Curator: Karoo NBG
Tarr, B.B. Curator: Natal NBG
Winter, J.H.S. Curator: Kirstenbosch NBG
PLANNING, MAINTENANCE & DEVELOPMENT— CAPE TOWN
Linde, D.C. N.T.C.III(Technician and Inspector of Works). M.S.A.I.D. Cert. Estate Agency.
Control Works Inspector
Arendse, D.S. Artisan’s Assistant
Manasse, S.P. Dip. (Masonry). Senior Artisan
Peck, W.I. Artisan’s Assistant
HAROLD PORTER NBG— BETTY’S BAY
Behr, Ms C.M. B. Sc. (Hons.). Control Research Technician
Abrahams, F. Groundsman II Jacobs, L.H.J. Groundsman II
Abrahamse, S.J. General Foreman Marinus, E.M. Groundsman I
Arendse, Ms M. Groundswoman I Martinka, A.G. Groundsman II
Bezuidenhout, Mrs H.M. Senior Administration Clerk October, Mrs R.P. Groundswoman I
Forrester, Ms J.A. N.T.C.III(Hort.). Chief Research Technician Rex, G. Groundsman II
Harper, C.C. Driver Van Coller, A.E. Groundsman I
Hendricks, C.F. Groundsman I Visser, C.J.J. Groundsman II
KAROO NBG— WORCESTER
Oliver, I.B. N.D.(Hort.)(PRA). Chief Research Technician
Ashworth, Mrs E.H. Senior Administration Clerk
Dlikilili, N.S. Groundsman II
Kecela, S. Groundsman I
Kumeke, A.M. Groundsman I
Made, M.A. Groundsman I
Madumane, M. Tractor driver
Makubalo, F.N. Nurseryman
Makubalo, J. Groundsman I
Maquanda, D. Groundsman II
Monisi, M.M. Groundsman I
Mpayipeli, W.W. Groundsman I
Mpeke, Mrs E.N. Groundswoman I
Mtetwa, A.M. General Foreman
Nkoloti, P.M. Groundsman I
Qumba, G.A. Groundsman I
Radebe, M.B. Groundsman II
Sibeko, L.A. Groundsman II
Sibozo, N.E. Driver
Simane, D.K. Senior Foreman
Theoha, P.P. Groundsman II
Townsend, D.J. N.D.(Hort). Senior Research Technician
Tuswa, M.H. Groundsman I
Viljoen, D.M. N.D.(Hort.). Senior Research Technician
Yekiso, S.M. Groundsman I
188
Bothalia 26,2 (1996)
KIRSTENBOSCH NBG— CAPE TOWN
Winter, J.H.S. N.D.(Hort.). Assistant Director
CENTRAL
Albu, Ms K. Principal typist
Anderson, D.L. Groundsman II. Workshop
Berman, R.C. Groundsman II. Gateman. Security
Engelbrecht, Mrs L.D. Administration Clerk. Plant re-
cording
Fredericks, Miss N.C.E. Information Officer
Geduldt, D.C. Engraver. Plant labels
Grace, T. Senior Storeman
Haynie, R.D. Groundsman II. Workshop
Jacobs, A.P. Information Officer
Jacobs, F.H. Storeman
Jansen, K.J. Senior Driver
Jaques, R.F. Research Technician. Plant utilization
Karelse, P.B. Tractor driver
Labuschagne (nee Malan), Mrs C.E. B. Sc. (Hons.). Edu-
cation Officer
Lewis, N. I. Security II
Marent, Mrs H.C. Senior Administration Clerk. Plant
recording
McLean, N.S. Groundsman II. Workshop
Newman, W. Driver. Workshop
Petersen, F.J. Driver
Petersen, P.B. Driver
Prins, F.B. Security I
Rudolph, A. Security II
Solomons, T.C. Senior General Foreman. Security
Trautman, C.E. Artisan/Supervisor. Workshop
Williams, G.C. Groundsman II. Gateman. Security
Williams, W.P. Principal General Foreman. Drivers
Woodward, Ms Y.J. Senior Administration Clerk
GARDEN
Abrahams, J.E. Groundsman I. General garden
Adams, H. Groundsman I. New development
Adonis, A. Senior General Foreman. Dell/Ericas
Adonis, J.J.M. Groundsman II. Dell/Ericas
Arendse, D.P. Groundsman I. Proteas
August, T. Groundsman I. New development
Balabala, L. Groundsman I. Dell/Ericas
Beck, G.D. Groundsman I. New development
Bowler, M.A. Groundsman II. Annuals
Crowie, R.W. Senior General Foreman. General garden
Cupido, J. Groundsman I. New development
Davids, S. General Foreman. Trees/shrubs
De Kock, Mrs M.E. Grounds woman I. Proteas
Fienies, C. Groundsman I. Trees/shrubs
Hendricks, S. Groundsman I. Proteas
Jacobs, D.R. Groundsman I. Proteas
Jansen, W. Groundsman I. Annuals
Josephus, D.N. Groundsman I. General garden
Julie, V. Groundsman I. Proteas
Loft, G.E. Groundsman II. Proteas
Lukas, K. Groundsman I. General garden
Manuel, D.R. Groundsman I. Annuals
McKlein, M.L. Groundsman I. General garden
McKlein, P. Groundsman I. Dell/Ericas
Mitchells, G. Chief General Foreman. New development
Morris, J.N.M. Groundsman I. Proteas
Petersen, A.D. General Foreman. Dell/Ericas
Petersen, V.E.M. Groundsman I. Dell/Ericas
Philander, N. Groundsman I. New development
Phillips, R. Groundsman I. General garden
Pietersen, J. Groundsman II. New development
Plaatjies, S.D. Groundsman I. Proteas
Ruiters, M. Groundsman II. New development
Sampson, R. Groundsman I. General garden
Smith, A.E. Groundsman II. General garden
Solomons, J. Groundsman I. Annuals
Solomons, K.J. Groundsman II. Proteas
Tait, A. Scholar
Van der Westhuizen, A.J. Senior General Foreman. Proteas
Van Rooy, K. General Foreman. Annuals
Van Schalkwyk, J. Groundsman I. Trees/shrubs
Van Wyk, F. Groundsman I. Glass house
Van Wyk, N. Groundsman II. Dell/Ericas
Williams, M.L.J. Groundsman II. New development
ESTATE
Le Roux, PH. Chief Research Technician. Estate Manager
Abrahams, M. Groundsman II. Estate
Adams, J. Groundsman I. Estate
Adonis, S.J. General Foreman. Aliens
Alexander, A. Groundsman I. Estate
Andrews, M.M. Groundsman II. Estate
Baadjies, I. Groundsman II. Estate
Bezuidenhout, A.K. Groundsman I. General maintenance
Bouwers, G.G. Groundsman I. Construction
Button, J. Groundsman I. Estate
Claasen, F. Groundsman I. Aliens
Geswind, A.J. Groundsman II. Lawnmowers
Grootboom, C.J. Groundsman I. Construction
Hope, C. General Foreman. Construction
Isaacs, M. Groundsman I. Estate
Jaceni, L.C. Groundsman I. Aliens
Jackson, P. Groundsman II. Lawnmowers
Jacobs, M.D. Groundsman I. Estate
Jaftha, R. Groundsman I. General maintenance
Botha] ia 26,2 (1996)
189
Julius, W.J. Groundsman I. Aliens
Kayster, GJ. General Foreman. Construction
Kuscus, G.W. General Foreman. General maintenance
Louw, Q. Groundsman I. Estate
Matthews, N. Groundsman I. Aliens
Matthews, I.N. Senior General Foreman. Estate
McLean, A. Groundsman II. Aliens
McLean, N. Groundsman II. Aliens
Petersen, J. Groundsman II. Aliens
Petersen, N.H. Groundsman I. Construction
Plaatjies, M.P. Groundsman I. Estate
Plaatjies, P.J. Groundsman I. Aliens
Reed, T.W. Groundsman II. Lawnmowers
Rhode, W.C. Groundsman I. Estate
Sampson, J.J. Tractor driver. Lawnmowers
Smith, R. Groundsman I. Estate
Solomons, G. Groundsman I. Aliens
Solomons, S. Groundsman I. Construction
Stanfield, J.R. Groundsman I. Aliens
Van Gusling, E.J. Senior General Foreman. Lawnmowers
NURSERY
Powrie, Ms F.J. B. Sc. (Hons.), N.D.(Hort.). Chief Research Technician. Head: Nursery
Adams, G. Groundsman II. Nursery
Apolis, A. Groundsman I. Succulents
Bennet, Ms J.I. Student (contract worker)
Beck, J.A. Groundsman I. Sales
Brown, B.M. Groundsman I. Nursery
Carrol, R.R. Groundsman I. Sales
Davids, M.I. General Foreman. Bulbs
Davids, N. Groundsman I. Sales
Dorfling, Ms C.C. Student (contract worker)
Duncan, G.D. N.D.(Hort.). Chief Research Technician
Francis, J. Groundsman I. Nursery
Hitchcock, A.N. N.H.D.(Hort-). Chief Research Techni-
cian. Sales
Jacobs, C.W. Groundsman I. Nursery
Jacobs, H.C. General Foreman. Sales
Jamieson, Mrs H.G. N.D. (Parks & Recreation). Principal
Research Technician. Restio & Asparagus collec-
tions
Jenkins, A. Groundsman I. Succulents
Jodamus, Ms N.L. Student (bursary)
Kettledas, P.G. Groundsman I. Nursery
King, O. Groundsman I. Nursery
Kotze, F.G. N.T.C.III(Hort.). Chief Research Technician.
Erica & Protea collections
Lawrence, E. Groundsman II. Sales
Lewin, T.B. Groundsman II. Nursery
March, A. Student (bursary)
Marthinus, E. Groundsman II. Succulents
Mathys, Ms S. Student (contract worker)
Matthews, Z. Groundsman I. Sales
Oliver, Y. Groundsman I. Sales
Sardien,T.P. General Foreman. Succulents
Sauls, C.J. Groundsman II. Nursery
Sauls, K. Groundsman I. Nursery
Shaide, Ms A. Student (bursary)
Szabo, Mrs B.J. Research Technician. Glass house
Tamboer, J.S. Senior General Foreman. Nursery
Van der Walt, Mrs L.E. N.D.(Hort.). Senior Research
Technician. Herbaceous collections
Van Jaarsveld, E.J. M.Sc., N.D.(Hort). Chief Research Tech-
nician
Young, B.A. Student (contract worker)
SEED ROOM AND TISSUE CULTURE
Notten, Miss A.L. Research Technician. Supervisor: Seed room
August, C. Groundsman II. Seed room Manuel, I.P. Senior General Foreman. Seed room
Crowie, Mrs U.M. Administration Aid II. Seed room Smith, D. Groundsman II. Seed room
Goliath, L. Administration Aid I. Seed room Van Rooyen, Miss S. Administration Aid I. Seed room
Crous, H.T. Senior Research Technician. Tissue culture
Picane, S. Administration Aid II. Tissue culture
LOWVELD NBG— NELSPRUIT
Kluge, J.P. B.Sc.(Hons.), T.H.O.D. Chief Research Technician
Froneman, W.C. N.D. (Nature Conservation & Manage-
ment), N.D. (Parks & Recreation Admin.), N.T.C.III
(Hort.). Principal Research Technician
Hurter, P.J.H. B. Sc. (Hons.). Principal Research Techni-
cian
Khoza, D.E. Groundsman I.
Khoza, F.D. Gardener
Khumalo, N.S. Groundsman I.
Khumalo, S.S. Tractor Driver
Lekhuleni, M.P. Groundsman I
Lekhuleni, W. Groundsman I
Magagula, K.E. Groundsman II
Magagula, N.R. Groundsman I
Mahlahlubane, F.J. Groundsman II
Makamo, Mrs J.E. Groundswoman II
Makhubela, B.J. Groundsman I
Maqungo, D.M. Groundsman I
Maqungo, Miss V.L.B. Groundswoman II
190
Bothalia 26,2(1996)
Maseko, N.E. Groundsman I
Mazibuko, F.E. Groundsman I
Mdhluli, M.B. Groundsman I
Mdluli, M.E. Groundsman II
Mkhatshwa, Mrs N.S. Groundswoman II
Mteto, E.M. Groundsman I
Muswili, K.J. General Foreman
Ndlovu, D. Groundsman II
Ndlovu, L.D. Groundsman II
Ngomane, S. Groundsman I
Ngqani, Mrs L.S. Groundswoman I
Ngwengoma, P.N. Driver
Ngwenya, P.S. Groundsman II
Ngwenyama, K.A. Groundsman I
Nkosi, Mrs P.B. Groundswoman I
Nkosi, Mrs S.L. Groundswoman II
Shabangu, M.E. Driver
Shabangu, S.L. Groundsman I
Shabangu, W.N. Groundsman I
Shawa, P. Groundsman I
Shawe, S.A. General Foreman
Sibule, B.F. Groundsman II
Sibure, W.F. Groundsman I
Sigudla, B.M. Groundsman II
Soka, M.P. Groundsman I
Thabethe, S.S. Groundsman I
Tsela, T.K. Groundsman I
Van der Walt, Mrs G.A. Senior Administration Clerk
NATAL NBG— PIETERMARITZBURG
Tarr, B.B. N.D. (Parks & Recreation Admin.), Advanced Dip.(Adult Education).
Chief Research Technician
Busani, M.A. Driver
Dlungwane, R. General Foreman
Gabuza, A. Groundsman I
Gates, Ms J.E. N.D. (Parks & Recreation Admin.), N.D.
(Hort.) Intermed.D. (Marketing Management). Princi-
pal Horticulturist. Kniphofia, forest spp.
Hlela, C. Groundsman I
Mbanjwa, Z. Groundsman I
Mbense, A. Groundsman I. Machine operator
Mdluli, K. Groundsman I
Mncwabe, Ms A. Groundswoman I
Mncwabe, P. Groundsman I
Mpangase, A. Groundsman I
Mpulo, D.H. Groundsman I
Mthalane, A. Groundsman I
Mtolo, C. Groundsman II. Team leader
Nkabini, A. Groundsman I
Nzakwe, W. Groundsman I
Radebe, A. Groundsman I
Sokhela, P. Groundsman I
Van der Merwe, Ms M.E.H. Senior Administration
Clerk
Zimu, J. Groundsman I
Zimu, S. Groundsman I
Zondi, A. Groundsman I
Zondo, Z. Groundsman I
Zuma, J. Groundsman I
Zuma, Ms K. Groundswoman I
FREE STATE NBG— BLOEMFONTEIN
Britz, R.M. N.D. (Forestry). Chief Research Technician
)
Dweba, K.P. Groundsman I. Entrance
Eysele, Mrs J.P. Senior Administration Clerk
Lekhetho, M.J. Groundsman II. Maintenance
Lekhetho, T.S. Groundsman I. Nursery
Lumley, M.J. Principal Research Technician. Nursery
Mabula, S.K. Groundsman I. Maintenance
Masheane, M.D. Tractor driver
Mbolekwa, G.M. Groundsman II. Nursery
Mbolekwa, L.M. Groundsman II. Rhus, display
Mohokare, L.J. Groundsman II. Maintenance
Moima, T.J. Groundsman I. Braai area, maintenance
Mopeli, M.J. Groundsman I. Bulb area
Moticoe, Mrs M.A. Groundswoman I. Nursery
Motse, V.D. Groundsman I. General garden
Mpeli, PA. Groundsman I. Nursery
Nakanyane, R.B. General Foreman
Nakedi, N.J. Driver
Nonjinge, S.H.B. Principal Research Technician
Olifant, D.M. Groundsman I. Kiosk area
Radithlare, Mrs E.M. Groundswoman I. Offices & nursery
Sebolai, Mrs C.L. Groundswoman I. Seed room nursery
Sebolai, P.R.A.N. Groundsman II. Tools maintenance
Semenyane, T.D. Groundsman I. Maintenance
Thaele, Mrs M.E. Food services Aid I
PRETORIA NBG
Heilgendorff, J.P. H.N.D.(Hort-). Chief Research Technician
Baloyi, F.R. Groundsman I
Baloyi, J.K. Groundsman I
Baloyi, S.J. Driver/Operator
Baloyi, S.M. Dip.(IBM) Specialized Auxiliary Services
Officer
Chipi, S. Security Assistant
Chuma, S.J. Security Assistant
Dry, D.H. N.D. (Hort.). Chief Research Technician. Plant
sales, records and gate control. Technical papers on
horticulture and plants
Bothalia 26,2(1996)
191
Kemp, J. Groundsman I
Keyter, B.A. Senior Security Officer
Klapwijk, N.A. N.D.(Hort.), N.D. (Plant Prod.), N.D.(Die-
sel Fitting). Chief Research Technician. Planning and
development. Index Nursery, New Plant Company
Kutama, B.T. Groundsman I
Letsoalo, H.M. Groundsman I
Mabasa, J.R. Security Assistant
Machika, S.M. Groundsman I
Mahlangu, J.J. Groundsman I
Makafula, W.P. Groundsman I
Makena, J.T. Groundsman I
Makena, S.M. Driver/Operator
Makena, S.N. General Foreman
Makgopo, C.K. Groundsman I
Makhafola, L.W. Groundsman I
Makhubela, D. General Foreman
Makhubela, P.K. Groundsman I
Makoeng, P.T. Groundsman I
Makola, J.M. Groundsman I
Makola, L.M. Groundsman II. Tractor driver
Makua, E.G. Groundsman I
Malewa, D.K. Groundsman I
Malobola, L.J. Groundsman I
Malobola, M. Groundsman II
Maluleke, J.M. Security
Mametja, A.M. Groundsman I
Mariri, J.N. Factotum
Marule, P.M. Groundsman II. Tractor driver
Masango, M.G. General Foreman
Mathabathe, D.S. Groundsman I
Matlala, S.M. Groundsman I
Matshika, S.P. Groundsman I
Mmakujwana, J.K. Groundsman I
Mnyangeni, L.D. Groundsman I
Modisha, D.M. Groundsman I
Mogoru, F.M. Groundsman I
Mogoru, S.C. Groundsman I
Mohale, F.R. General Foreman
Mohale, J.N. Groundsman I
Mokawe, R.N. Groundsman I
Molefe, J.R. Groundsman II
Molokomme, F.K. Groundsman I
Molomo, S.E. Groundsman I
Mononyane, J.B. Groundsman I
Motshweni, V.V. Groundsman I
Mudau, R.T. Groundsman I
Muhali, B.M. Groundsman I
Ngobeni, R.T. Groundsman I
Nkoana, J.M. Groundsman I
Nkwana, F.N. Driver/Operator
Noko, J.M. Research Assistant
Noku, A.Y. Groundsman II. Tractor driver
Ramakgaphola, A.M. Groundsman I
Ramatsetse, P.M. Security Assistant
Rampopana, A.M. Groundsman I
Schnackenberg, Ms H. Senior Provisioning Administra-
tion Clerk
Sete, L.B. General Foreman
Shilubane, E. Storeman Assistant
Shirindi, J.R. Groundsman I
Shirindi, S.M. Groundsman I
Sithole, J.D. Groundsman I
Strydom, D.J.F. N.T.C.ni(Hort.), N.D.(Parks & Rec. Manage-
ment). Chief Research Technician. Maintenance cycads
Swartz, Ms P. M.Sc. Senior Horticulturist. Scientific and
horticultural curation of living collections; garden
development; seedbank of endangered plants and
succulents; Madagascan plants
Tefu, P.R. Groundsman I
Tloubatla, J.L. Groundsman I
Tolo, P.K. Groundsman I
WITWATERS RAND NBG— WILROPARK
Chaplin, P.J. N.T.C.Dip.(Hort.). Chief Research Technician
Hankey, A.J. N.D.(Hort.). Senior Research Technician
Head, Mrs S.E. Senior Administration Clerk
Khedzi, K.P. Groundsman I. Nursery
Lukhwa, N.A. Groundsman I. Garden
Luvhimbi, T.S. Groundsman I. Garden
Majamane, Z.E. Groundsman I. Garden
Mamosebo, M.A. Groundsman I. Garden
Manjati, Mrs N.L. Groundswoman n. Clerical Assistant
Manyikana, T.M. Groundsman I. Garden
Matsea, M.W. Groundsman I. Garden
Mbulaheni, N.P. Groundsman I. Garden
Mulibana, N.S. Groundsman I. Machine operator
Mmola, Ms B.E. Groundswoman I. Cleaner
Moteka, N.D. Outreach Education Officer
Ndou, A.P. Groundsman I. Garden
Ndou, M.W. Groundsman I. Machine operator
Ndwambi, N.W. Groundsman I. Garden
Ndzondo, Ms G.P. Groundswoman I. Cleaner
Nedambale, M.P. Groundsman II. Nursery
Nemalili, M.E. Driver
Nemalili, A.S. Groundsman II. Driver
Nekhavhambe, S.P. Groundsman I. Garden
Nenungwi, M.S. Groundsman I. Nursery
Rammela, N.N. Groundsman I. Machine operator
Randima, M. Groundsman I. Garden
Raphalalani, V.S. Groundsman I. Nursery
Steel, Miss B.S. N.D.(Nature Conservation), N.D.(Parks &
Recreation Admin.), Dip.(Joumalism). Senior Research
Technician. Nursery, succulent and herb gardens, plant
records
Tebeile, Ms Z.M. Groundswoman II. Clerical Assistant
Thupe, G.G. Groundsman I
Tshisikule, G.M. Groundsman I. Garden
Turner, MS S.L. B. Sc. (Hons.). Horticulturist
Van der Westhuizen, Mrs S. M.Sc. Environmental Educa-
tionalist
Vlok, Mrs S. Administration Assistant
192
Bothalia 26,2(1996)
RESEARCH DIRECTORATE
PRETORIA |
Smith, G.F. Ph.D., F.L.S. Director: Research
Wolfson, Mrs M.M. Ph.D. Deputy Director
Smit, Mrs A.C. Personal Secretary
Steyn, Mrs E.M.A. Ph.D. Principal Scientist. Embryology
PLANT SYSTEMATICS SUBDIRECTORATE
| PRETORIA 1
Smith, G.F. Ph.D., F.L.S. Systematics of succulents and rosulate, petaloid monocots
Arnold, T.H. Head: Data Management (Pretoria)
Koekemoer, Miss M. Curator: National Herbarium (Pretoria
Rourke, Dr J.P. Curator: Compton Herbarium (Cape Town)
Williams, Ms R. Curator: Natal Herbarium (Durban)
COMPTON HERBARIUM— CAPE TOWN
Rourke, J.P. Ph.D., F.L.S., F.R.S.S.Af. Assistant Director. Systematics of southern African
Proteaceae, Stilbaceae
Beyers, Mrs J.B.P. M.Sc. Scientist. Assistant Curator: Col-
lections. Taxonomy of the Gnidieae (Thymelaeaceae)
Cupido, Mrs C.S. Specialized Auxiliary Services Officer
Davidse, Mrs. E. Specialized Auxiliary Services Officer
Fellingham, Mrs A.C. B.Sc. Scientific Officer. Taxonomy
of Cliffortia (Rosaceae)
Foster, Mrs S.E. Principal Typist
Kurzweil, H. Ph.D. Scientist. Systematics of southern African
terrestrial orchids
Leith, Mrs J. Senior Administration Clerk
Manning, J.C. Ph.D. Specialist Scientist. Systematics of
Iridaceae and Orchidaceae; anatomy
Oliver, E.G.H. M.Sc. Principal Scientist. Taxonomy of the
Ericoideae (Ericaceae)
Paterson- Jones, Mrs D.A. (nee Snijman) Ph.D. Principal
Scientist. Systematics of Amaryllidaceae; cladistics
Roux, J.P. N.T.C.(Hort-), F.L.S., M.Sc. Principal Scientist.
Systematics of Pteridophyta
Steiner, K.E. Ph.D. Specialist Scientist. Systematics of
Scrophulariaceae and evolutionary interactions be-
tween oil-secreting flowers and oil-collecting bees
NATAL HERBARIUM— DURBAN
Williams, Ms R. B.Sc. (Hons.), H.D.E. Chief Scientific Officer.
Crouch, N.R. Ph.D. Scientist. Ethnobotanist
Mbonambi, B.M. Groundsman II. Gardener
Ngwenya, A.M. Principal Specialized Auxiliary Services
Officer. Herbarium Assistant. Plant identification, plant
information
Noble, Mrs H-E. Senior Administration Clerk
Nzimande, S.B. Groundsman II. Caretaker
Sikhakhane, T.B. Senior Specialized Auxiliary Services
Officer. Herbarium Assistant. Plant identifications,
herbarium services
Singh, Ms Y. B.Sc. (Hons.), H.E.D. Senior Scientific Officer.
Taxonomy of Zantedeschia, plant identifications
Bothalia 26,2 (1996)
193
NATIONAL HERBARIUM— PRETORIA
Koekemoer, Miss M. M.Sc. Assistant Director. Herbarium management. Taxonomy of Poaceae, Asteraceae:
Disparago, Stoebe, Amphiglossa, Elytropappus, Pterothrix and Bryomorphe
Bredenkamp, Mrs C.L. M.Sc. Chief Scientific Officer. Assistant Curator: Public relations.
Taxonomy of Vitex, Phylica, Rhamnaceae, Sterculiaceae and other related families
Germishuizen, G. M.Sc. Principal Scientist. Assistant Curator: Finances. Taxonomy of Poly-
gonaceae, Fabaceae, Loranthaceae, Viscaceae
Herman, P.P.J. M.Sc. Principal Scientist. Assistant Curator: Personnel. Taxonomy of Asteraceae,
Flora of northern provinces
Heymann, Mrs M.Z. T.E.Dip., B.A.(Education & History). Principal Specialized Auxiliary
Services Officer. Assistant Curator: Services, loans, gifts and exchanges
Anderson, H.M. Ph.D. Principal Scientist. Palaeobotany,
palaeogeography
Anderson, J.M. Ph.D. Specialist Scientist. Palaeobotany,
palaeogeography
Archer, R.H. Ph.D. Scientist. Taxonomy of mainly Celas-
traceae, Euphorbiaceae
Archer (nee Reid) Mrs C. M.Sc. Senior Scientist. Taxon-
omy of Cyperaceae, Restionaceae, Orchidaceae
Burgoyne, Ms PM. M.Sc. Senior Scientific Officer. Plant
identifications co-ordinator
Cloete, Mrs M. Dip. (Typing). Senior Provisioning Clerk.
Specimen label typist
Dreyer, Miss L.L. M.Sc. Scientist. Taxonomy of Gera-
niaceae, Crassulaceae, Oxalidaceae
Fish, Mrs L. B.Sc. Chief Scientific Officer. Taxonomy of
Poaceae. Plant collecting programme; supervising
mounters
Glen, H.F. Ph.D. Senior Scientist. Taxonomy of trees and
succulents, especially Aloe, herbarium for cultivated
plants, and botanical collectors
Glen, Mrs R.P. M.Sc. Senior Scientific Officer. Taxonomy
of ferns, water plants
Hoare (nee Victor), Mrs J.E. B.Sc. (Hons.), H.Dip.Journ.
Scientific Officer. Taxonomy of Rutaceae, Ascle-
piadaceae
Jordaan, Mrs M. M.Sc. Chief Scientific Officer. Taxon-
omy of Casuarinaceae — Connaraceae, Maytenus
Kgaditsi, W.T. Specialized Auxiliary Services Officer. Moun-
ter, general assistant in cultivated plants section
Lephaka, M.G. Specialized Auxiliary Services Officer.
Parcelling, pressing and general assistance
Makgakga, M.C. Senior Specialized Auxiliary Services
Officer. Herbarium assistant, Wing B
Makgakga, K.S. Specialized Auxiliary Services Officer.
Mounter of vascular plants
Makwarela, A. B.Sc. Scientific Officer
Marinus, Mrs E. Principal Specialized Auxiliary Services
Officer. Herbarium assistant
Masombuka, Ms A. Specialized Auxiliary Services Officer.
Herbarium assistant
Meyer, J.J. N.D. (Teaching). Scientific Officer. Herbarium
assistant, Wing C
Netnou, Ms C. B.Sc. Scientific Officer
Nkoana, S. Scientific Officer
Perold, Mrs S.M. Ph.D. Taxonomy of Ricciaceae, Hepaticae
(contract worker)
Phahla, T.J. Specialized Auxiliary Services Officer. Moun-
ter of bryophytes and vascular plants
Ready, Mrs J.A. N.D.(Hort.). Senior Specialized Auxiliary
Services Officer. Herbarium assistant, Wing D
Retief, Miss E. M.Sc. Senior Scientist. Pollen studies of
Boraginaceae. Taxonomy of Boraginaceae, Verben-
aceae, Lamiaceae, Asteraceae, Rubiaceae
Riddles, L. B.Sc. Scientific Officer
Sebothoma, P. Specialized Auxiliary Services Officer. Herba-
rium assistant
Smithies, Mrs S.J. M.Sc. Senior Scientific Officer. Taxon-
omy of Scrophulariaceae, Selaginaceae, Lobeliaceae
Steyn, Ms C.C. Principal Auxiliary Services Officer. Anat-
omy, palynology
Van Rooy, J. M.Sc. Senior Scientist. Taxonomy and bio-
geography of mosses; supervising bryophyte moun-
ters
Veldman, Mrs J.M. Senior Provisioning Clerk. Herbarium
administration
Welman, Miss W.G. M.Sc. Senior Scientist. Taxonomy of
Convolvulaceae, Solanaceae, Cucurbitaceae, Campa-
nulaceae, Asteraceae, Acanthaceae
DATA MANAGEMENT— PRETORIA
Arnold, T.H. M.Sc. Assistant Director. Computer application especially in taxonomy
Botha, Mrs A.G. Principal Specialized Auxiliary Services
Officer. Secretary
De Wet, Mrs B.C. B.Sc.(Computer Science), B.A., H.D.L.S.
Principal Datametrician
Evenwel, Mrs E. Specialized Auxiliary Services Officer.
Quality control
Harris, Mrs B.J. Specialized Auxiliary Services Officer.
Encoding, quality control
Joubert, Mrs M.A.E. Senior Data Typist
Kema, Ms N. Student (contract worker)
Stodart, Ms M. Student (contract worker)
Van Staden, Ms M. N.D. (Computer Data Processing).
Assistant Datametrician
194
Bothalia 26,2(1996)
PRETORIA
Wolfson, Mrs M.M. Ph.D. Deputy Director. Physiology/Ecophysiology of Poaceae, carbon uptake metabolism,
allocation in response to environmental and management stress
Clapperton, Mrs S. Typist
Du Plessis, Mrs E. Acting Head: Publications
Du Plessis, Mrs. H. Head: Research Herbarium Support Services
Liebenberg, Mrs E.J.L. Head: Administration
Potgieter, Mrs. E. Senior Librarian
PUBLICATIONS— PRETORIA
Du Plessis, Mrs E. B. Sc. (Hons.), S.E.D. Technical editor. Editing, translating, layout
Bopape, Mrs M.D. Senior Administration Clerk. Book-
shop
Brink, Mrs S.S. Dip. (Typing). Chief Typesetter. Typeset-
ting, layout, word processing
Leistner, O.A. D.Sc., F.L.S. Editing (contract worker)
Mapheza, T.P. Senior Storeman. Bookshop
Momberg, Mrs B.A. B.Sc. (Entomology & Zoology). Tech-
nical editor. Editing, layout
Mulvenna, Mrs J.M. Dip. (Typing). Typist. Word process-
ing, typsetting (contract worker)
RESEARCH HERBARIUM SUPPORT SERVICES— PRETORIA
Du Plessis, Mrs H. M.Sc. Senior Scientist. Cytogenetics.
Condy, Ms G.S. M.A. Senior Industrial Technician. Botanical artist
Romanoswki, Mfs A.J. Dip. (Photography). Senior Industrial Technician
(Photography). Scientific photographer
MARY GUNN LIBRARY— PRETORIA
Potgieter, Mrs E. B. Libr. Senior Librarian
Fourie, Mrs A. B.Libr. Assistant librarian (contract worker)
Louw, Ms A. Library assistant (contract worker)
Van der Walt, Ms A.M.E. Cataloguer (contract worker)
ADMINISTRATION— PRETORIA
Liebenberg, Mrs E.J.L. M.Sc. Chief Scientific Officer. Cytotaxonomy
Gotzel, Mrs A. Senior Telecom. Operator
Kama, Mrs P.B. Administration Aid I
Kekana, Mrs M.R. Administration Aid I
Khumalo, N.P Principal General Foreman. Supervisor:
Office services
Koehne, Mrs R.W.R. Senior Registration Clerk
Makgobola, Mrs M.R. Administration Aid I
Maphuta, Mrs M.S. Administration Aid I
Martin, Ms M.A. Senior Administration Clerk
Phaala, M.C. Administration Aid I
Smuts, Mrs W.E. Administration Officer. Personnel
Tloubatla, J.M. Courier/Photocopy Machine Operator
Venter, W.A. N2(Technical). Principal General Foreman.
Maintenance
Bothalia 26,2(1996)
195
ECOLOGY AND CONSERVATION SUBDIRECTORATE
CAPE TOWN]
Rutherford, M.C. Ph.D., Dip.(Datamet.). Deputy Director
Davis, G.W. Ph.D. Assistant Director. Head: Ecology liaison. Stress and disturbance ecology
Nanni, Ms I. B.Sc., H.E.D. Head: Research Support Services
Reynolds, Ms P.Y. B.A., H.D.L.S., B.Proc. Senior Librarian
Bosenberg, J. de Wet B.Sc.(Hons.). Principal Scientific
Officer. Cycad biology and conservation
Botha, P.A. N.H.D.(Hort). Senior Scientific Officer. Tissue
culture research, seed biology
Brown, N.A.C. Ph.D. Specialist Scientist. Seed biology
research, plant growth regulators
De Lange, J.H. B.Sc.(Hort.), M.Sc. (Plant Physiology),
D.Sc.(Agric.), Ph.D.(Bot.). Senior Specialist Scien-
tist. Ecology, tissue culture, horticulture
Donaldson, J.S. M.Sc. (Entomology), PhD. (Zoology). As-
sistant Director. Cycad biology, plant/insect interac-
tions, conservation biology
Hilton-Taylor, C. B.Sc. (Hons. )(Biological Sci.). Scientist.
Threatened plants, biodiversity of arid regions
Hoffman, M.T. Ph.D. Senior Scientist. Disturbance and
historical ecology
Hunter, Ms D.A. Senior Administration Clerk. Secretary
Jacobs, E.C. Specialized Auxiliary Services Officer. Labo-
ratory Assistant. Cyc/op/a/Honeybush tea project
McDonald, D.J. Ph.D. Principal Scientist. Mountain vege-
tation, biogeography, conservation
Midgley, G.F. M.Sc. Principal Scientist. Plant stress physi-
ology/ecology
Musil, C.F. Ph.D. Specialist Scientist. Plant stress physi-
ology/ecology
O’Callaghan, M.G. Ph.D. Principal Scientist. Plant-climate
interactions
Petersen, Ms A. Scientific Officer. Disturbance and his-
torical ecology (contract worker)
Powrie, L.W. M.Sc. Chief Scientific Officer. Ecological
databases: computer co-ordination
Rebelo, A.G. Ph.D.(Zoology). Senior Scientist. Conserva-
tion biology, biogeography
Wand, S.J.E. M.Sc.(Agric.). Scientist. Plant stress physi-
ology/ecology
RESEARCH SUPPORT SERVICES— CAPE TOWN
Nanni, Ms I. B.Sc., H.E.D. Chief Scientific Officer
Boonzaier, I. Groundsman I
Bowler, Mrs M. Laboratory & Office Aid II
De Witt, D.M. Principal Specialized Auxiliary Services
Officer. Scientific Assistant
Jagger, B.W. Principal Specialized Auxiliary Services Of-
ficer. Scientific Assistant
Parenzee, Ms H.A. Senior Administrative Assistant
Snyders, S.G. Specialized Auxiliary Services Officer. Gene-
ral Assistant
HARRY MOLTENO LIBRARY— CAPE TOWN
Reynolds, Ms P.Y. B.A., H.D.L.S., B.Proc., B. Bibl.(Hons.). Senior Librarian
Edwards, Mrs J. A.L.A.(UK). Cataloguer (contract worker)
Maqetuka, Ms N. Student (contract worker)
196
PUBLICATIONS BY THE STAFF
(1st April 1995-3 1st March 1996)
Bothalia 26,2(1996)
ANDERSON, H.M., HILLER, N. & GESS, R.W. 1995. Archaeopteris
(Progymnospermopsida) from the Devonian of southern Africa.
Botanical Journal of the Linnean Society 117: 305-320.
ARCHER, R. & CONDY, G. 1995. Elaeodendron croceum. Flowering
Plants of Africa 54: 57-62, t. 2112.
ARCHER, R.H. & VAN WYK, A.E. 1995. Proposal to conserve the name
Cassine barbara L. with a conserved type. Taxon 44: 435, 436.
ASHWELL, A.N. 1995. What role education? In A.B. Low & F.E. Jones,
The sustainable use and management of Renosterveld remnants
in the Cape Floristic Kingdom. Proceedings of a symposium.
FCC Report 1994/5 Flora Conservation Committee, Botanical
Society of South Africa, Kirstenbosch, Cape Town: 5-8.
BAXTER, J. & CROUCH, N. 1996. Gerrardanthus tomentosus: a rare
member of the pumpkin family from KwaZulu-Natal. PlantLife
14: 11, 12.
BEYERS, J.B.P. 1995. A new species of Trichogyne from Namaqualand
(Asteraceae). Bothalia 25: 107-109.
BEYERS, J.B.P. & VAN DER WALT, J.J.A. 1995. The generic delimita-
tion of Lachnaea and Cryptadenia (Thymelaeaceae). Bothalia
25: 65-85.
BRITS, G.J., CUTTING, J.G.M., BROWN, N.A.C. & VAN STADEN,
J. 1995. Environmental and hormonal regulation of seed dor-
mancy and germination in Cape fynbos Leucospermum R.Br.
(Proteaceae) species. A working model. Plant Growth Regulation
17: 181-193.
BROWN, N.A.C. 1995a. Conservation through cultivation. Plant-de-
rived smoke and seed germination. African Wildlife, May-Oct.: 29.
BROWN, N.A.C. 1995b. Kirstenbosch Instant Smoke-Plus Seed Primer.
Hortagro 2,1: 19.
BROWN, N.A.C. 1995c. Where there’s smoke, there’s seed. The Austra-
lian Garden Journal, Autumn: 48-50.
BROWN, N.A.C. & BOTHA, PA. 1995. List of species in which
treatment with smoke or aqueous smoke extract has been shown
to give improved germination. Veld & Flora 81: 93.
BROWN, N.A.C., BOTHA, P.A. & PROSCH, D. 1995. Where there’s
smoke. Garden 120: 402-405. Royal Horticultural Society, London.
BURROWS, J.E. & CROUCH, N.R. 1995. New distribution records of
South African pteridophytes (Pteridophyta). Bothalia 23:
236-238.
CAIRNCROSS, B„ ANDERSON, J.M. & ANDERSON, H.M. 1995.
Palaeoecology of the Triassic Molteno Formation, Karoo Basin,
South Africa— sedimentological and palaeoecological evidence.
South African Journal of Geology 98: 452-478.
CHESSELET, P., MOSSMER, M. & SMITH, G.F. 1995. Research
priorities in the succulent plant family Mesembryanthemaceae
Fenzl. South African Journal of Science 91: 197-209.
CROUCH, N. & TARR, B. 1995. And all because the lady loves...
Crocosmia. Veld & Flora 8 1 : 79, 80.
DAVIS, G.W. (ed.) 1995a. Bulletin of the Southern African Institute of
Ecologists and Environmental Scientists 14,1.
DAVIS, G.W. 1995b. Degradation for the better? Bulletin of the Southern
African Institute of Ecologists and Environmental Scientists 14,2.
DAVIS, G.W. 1995c. Manipulating the medium. Bulletin of the Southern
African Institute of Ecologists and Environmental Scientists 14,3: 1 .
DAVIS, G.W. 1995d. International concern with functional aspects of
biodiversity. South African Journal of Science 9 1 : 6 1 , 62.
DAVIS, G.W. & SOMERS, M. (eds) 1995. Conservation of southern
Africa’s wild living resources: exploitation, sustainability and
ethics. Proceedings of a one day symposium, May 1995, hosted
by the Southern African Institute of Ecologists and Environ-
mental Scientists, the South African Wildlife Management Asso-
ciation, and the Marine Science Society of Southern Africa.
DEAN, W.R.J., HOFFMAN, M.T., KERLEY, G.I.H. & MILTON, S.J.
1995. Desertification in developed countries: in search of the
silver bullet. South African Journal of Science 91: 213-215.
DEAN, W.R.J., HOFFMAN, M.T., MEADOWS, M E. & MILTON, S.J.
1995. Desertification in the semi-arid Karoo, South Africa: re-
view and reassessment. Journal of Arid Environments 30: 247-264.
DELANGE, J.H. 1995. Saadlose borne: dieoplossing virbosbou? Arbor,
October: 12.
DONALDSON, J.S. 1995a. An integrated effort to save South African
cycads. Plant Talk 2: 13.
DONALDSON, J.S. 1995b. (ed.) Cycad conservation in South Africa:
issues, priorities and actions. Cycad Society of South Africa,
Pretoria.
DONALDSON, J.S. 1995c. Understanding cycad life histories: an essen-
tial basis for successful conservation. In J.S. Donaldson, Cycad
conservation in South Africa: issues, priorities and actions, March
1995: 8-13. Cycad Society of South Africa, Pretoria.
DONALDSON, J.S. 1995d. Matched photographs as a means of assess-
ing changes in cycad distribution and abundance. In J.S. Donald-
son, Cycad conservation in South Africa: issues, priorities and
actions, March 1995: 20-23. Cycad Society of South Africa,
Pretoria.
DONALDSON, J.S. 1995e. The Winterberg cycad — surviving against
the odds. Veld & Flora 81: 36-39.
DONALDSON, J.S. & BOSENBERG, J. DE W. 1995a. Life history and
host range of the Leopard magpie moth, Zerenopsis leopardina
Felder (Lepidoptera: Geometridae). African Entomology 3:
103-110.
DONALDSON, J.S. & BOSENBERG, J. DE W. 1995b. Duration of
developmental stages of the male gametophyte of Encephalartos
altensteinii in the interval between pollination and fertilization.
In P.J. Vorster, Proceedings of the Third International Congress
on Cycad Biology: 415-422. Cycad Society of South Africa,
Stellenbosch.
DONALDSON, J.S., NANNI, I. & BOSENBERG, J. DE W. 1995. The
role of insects in pollination of the African cycad Encephalartos
cycadifolius (Zamiaceae). In P.J. Vorster, Proceedings of the
Third International Congress on Cycad Biology: 423-434. Cycad
Society of South Africa, Stellenbosch.
DREYER, L.L., MARAIS, E.M. & VAN DER WALT, J.J.A. 1995. A
subspecific division of Pelargonium reniforme Curt. (Gera-
niaceae). South African Journal of Botany 61 : 325-330.
DRY, D.H. 1995a. A promise to keep. The S.A. Nurseryman 14,1: 29.
DRY, D.H. 1995b. Die kat en saffraan. The S.A. Nurseryman 14,2: 22,
23.
DRY, D.H. 1996. Trees of the year for 1996. Local Government Digest,
January 1996: 46-50.
DUNCAN, G.D. 1995a. A visit to California. IBSA Bulletin 43: 24, 25.
FELLINGHAM, A.C. 1995. Anew species of Cliffortia from the Swart-
berg (Rosaceae). Bothalia 25: 104-107.
FELLINGHAM, A.C. & MEYER, N.L. 1995. New combinations and a
complete list of Asparagus species in southern Africa (Aspara-
gaceae). Bothalia 25: 205-209.
FINDLAY, R.F. & GLEN, H.F. 1995. Erythrophysa transvaalensis — the
Transvaal red balloon. Trees in South Africa 45: 40.
GERMISHUIZEN, G. & CONDY, G. 1995. Crotalaria monophylla.
Flowering Plants of Africa 54: 54-56, t. 2111.
GLEN, H.F. 1995a. Ken Cunliff ’s Library. Trees in South Africa 44: 8, 9.
GLEN, H.F. 1995b. Greyia sutherlandii. Trees in South Africa 44: 17, 18.
GLEN, H.F. 1995c. Venison marinade with juniper berries. Trees in South
Africa 44: 29.
GLEN, H.F. 1995d. Avocado pear poisoning: a warning. Trees in South
Africa 44: 30.
GLEN, H.F. 1995e. Plants threatened by Man’s agricultural activities.
Trees in South Africa 44: 34—42.
GLEN, H.F. 1995f. Peddiea africana. Trees in South Africa 44: 43, 44.
GLEN, H.F. 1995g. Bobby Broomberg. Trees in South Africa 44: 51.
GLEN, H.F. 1995h. Foreword to: Olitzky, S. The would-be Desert Queen.
Symbol, Pretoria [‘Green’ children’s storybook.]
Bothalia 26,2 (1996)
197
GLEN, H.F. 1995i. Light in the darkness: towards a guide to exotic trees
grown in southern Africa. Trees in South Africa 45: 44-50.
GLEN, H.F. 1995j. Notes towards a southern African Garden Flora I:
Introduction, Nyssaceae, Comaceae and Bixaceae. Trees in South
Africa 45: 51-60.
GLEN, H.F. 1995k. Contribution: notes on distribution maps published
in southern Africa. In J. Lundqvist, Index holmiensis VIII. Swed-
ish Museum of Natural History, Stockholm.
GLEN, H.F., CARR, D. & SUMAR, D. 1995. What is a Christmas tree?
Trees in South Africa 45: 33-36.
GLEN, H.F. & HARDY, D.S. 1995. Aloe section Anguialoe and the
problem of Aloe spicata L.f. (Aloaceae). Haseltonia 3: 92-103.
GLEN, H.F., LAVRANOS, J.J. & CONDY, G. 1995. Anagallis monelli.
Flowering Plants of Africa 43: 64—67, t. 2113.
GLEN, H.F. & Members of the Tree Society of southern Africa 1995.
Identification of some Highveld trees in winter. Trees in South
Africa 44: 22-29.
GLEN, H.F. & SMITH, G.F. 1995. Notes on the typification of some
species of Aloe (Asphodelaceae/ Aloaceae). Bothalia 25: 37-42.
GLEN, H.F., SMITH, G.F. & HARDY, D.S. 1995. Typification of Aloe
species described by B.H. Groenewald (Asphodelaceae/ Alo-
aceae). Bothalia 25: 97-99.
GLEN, H.F. & WELMAN, W.G. 1 995a. Friedrich von Breitenbach OMS
(1916-1995). Trees in South Africa XLV.l & 2: 11-17.
GLEN, H.F. & WELMAN, W.G. 1995b. Friedrich von Breitenbach OMS
(1916-1995). Forum Botanicum 32,2: 11.
GLEN, H.F. & WELMAN, W.G. 1995c. Obituary: Friedrich von Breiten-
bach OMS (1916-1995). Bothalia 25: 260-264.
GOLDBLATT, P. & MANNING, J.C. 1995a. What is Gladiolus cruen-
tusl Veld & Flora 81:4-7.
GOLDBLATT, P. & MANNING, J.C. 1995b. New species of the southern
African genus Geissorhiza (Iridaceae: Ixioideae). Novon 5:
156-161.
GOLDBLATT, P. & MANNING, J.C. 1995c. New species of southern
African Moraea (Iridaceae: Iridoideae), and the reduction of
Rheome. Novon 5: 262-269.
GOLDBLATT, P & MANNING, J.C. 1995d. Phylogeny and speciation
in Lapeirousia subgenus Lapeirousia (Iridaceae: Ixioideae). An-
nals of the Missouri Botanical Garden 82: 517-534.
GOLDBLATT, P. & MANNING, J.C. 1995e. Phylogeny of the African
genera Anomatheca and Freesia (Iridaceae: Ixioideae), and a new
genus Xenoscapa. Systematic Botany 20: 161-178.
GOLDBLATT, P & MANNING, J.C. 1996. Aristeas and beetle pollina-
tion. Veld & Flora 82: 17-19.
GOLDBLATT, P, MANNING, J.C. & BERNHARDT, P. 1995. Pollina-
tion biology of Lapeirousia subgenus Lapeirousia (Iridaceae) in
southern Africa: floral divergence and adaptation for long-
tongued fly pollination. Annals of Missouri Botanical Garden 82:
518-534.
HERMAN, P.P.J. 1995a. A new species in the genus Clerodendrum
(Verbenaceae). Bothalia 25: 100-102.
HERMAN, P.P.J. 1995b. A note on Ursinia brachyloba (Asteraceae).
Bothalia 25: 244.
HERMAN, P.P.J. 1995c. A new combination in Cotula (Asteraceae).
Bothalia 25: 244, 245.
HICKEY, B. & VAN JAARSVELD, E. 1995. Propagation of three rare
endangered vygies. British Cactus and Succulent Journal 13: 4.
HILTON-TAYLOR, C. 1996. Threatened Ericaceae in southern Africa.
Yearbook of the Heather Society 1996: 7-16.
HOBBS, R.E., DAVIS, G.W. & KEELEY, J.E. 1995. Mediterranean-type
ecosystems. In: V.H. Heywood, Global biodiversity assessment :
366-370. Cambridge University Press, Cambridge.
HOFFMAN, M.T. 1995a. Review: ‘Conservation, reclamation and graz-
ing in the northern Negev: contradictory or complementary con-
cepts?’, by Avi Perevolotsky. Pastoral Development Network
Paper 38b, July: 10-13.
HOFFMAN, M.T. 1995b. Desertification of the Karoo, South Africa: a
review of the recent research and current status of the debate.
Proceedings of the National Research and Development Work-
shop on the Assessment and Monitoring of Desertification in
South Africa , Premos Conference Centre, Pretoria, 10-12 Octo-
ber 1995: 21-36. Department of Agriculture: Directorate of Re-
source Conservation, Pretoria.
HOFFMAN, M.T. 1995c. Ecology and rural development. South African
Institute of Ecologists and Environmental Scientists Bulletin
14,2: 28.
HOFFMAN, M.T. 1995d. Environmental history and the desertification
of the Karoo, South Africa. Giomale Botanico Italiano 129,1:
261-273.
HOFFMAN, M.T. 1996. The impact of the SADF on the Riemvasmaak
environment between 1974-1994 and an outline of a monitoring
programme for the region. Proceedings of conference Reclaiming
the land: from defence to development , Johannesburg, 13-14
March 1996. Group for Environmental Monitoring (GEM), Jo-
hannesburg.
HOFFMAN, M.T., BOND, W.J. & STOCK, W.D. 1995a. Desertification
of the eastern Karoo, South Africa: conflicting palaeoecological,
historical and soil isotopic evidence. Environmental Monitoring
and Assessment 37: 159-177.
HOFFMAN, M.T., BOND, W.J. & STOCK, W.D. 1995b. Desertification
of the eastern Karoo, South Africa: conflicting palaeoecological,
historical and soil isotopic evidence. In D.A. Mouat & C.F.
Hutchinson, Desertification in Developed Countries: 159-177.
Kluwer Academic Publishers, Dordrecht.
HURTER, P.J.H. 1995a. Ex situ cycad conservation in the Lowveld
National Botanical Garden. In P. Vorster, Proceedings of the Third
International Conference on Cycad Biology: 27. Cycad Society
of South Africa, Stellenbosch.
HURTER, P.J.H. 1995b. Two newly described cycads from Africa.
Phytologia 78: 409-416.
HURTER, P.J.H. & CLAASSEN, M.I. 1996. Focus on Encephalartos
nubimontanus Hurter. Encephalartos 45: 4-11.
HURTER, P.J.H. & GLEN, H.F. 1995. Encephalartos equatorialis (Za-
miaceae): a newly described species from tropical Africa. South
African Journal of Botany 61: 226-229.
HURTER, P.J.H. & GLEN, H.F. 1996. Encephalartos hirsutus (Za-
miaceae): a newly described species from South Africa. South
African Journal of Botany 62: 46—48.
HURTER, P.J.H., GLEN, H.F. & CLAASSEN, M.I. 1995. Focus on
Encephalartos equatorialis Hurter. Encephalartos 44: 4-9.
JAMIESON, H. & BROWN, N.A.C. 1995a. Out of Africa: new restios
with garden potential. The Australian Garden Journal 1 5, 1 : 39—42.
JAMIESON, H. & BROWN, N.A.C. 1995b. Restios with garden poten-
tial. Pacific Horticulture 56,2: 36-41.
JAMIESON, H„ HITCHCOCK, A. & BROWN, N.A.C. 1995a. New
restios available at the Botanical Society Garden Shop. Veld &
Flora 81: 92.
JAMIESON, H„ HITCHCOCK, A. & BROWN, N.A.C. 1995b. Growing
restios. Veld & Flora 81: 129-131.
JOFFE, P. 1996. Dassie Trail, Pretoria National Botanical Garden.
Environmental Resource Guide: NBI Garden Trail Guide 1.
National Botanical Institute, Pretoria.
JOHNSON, S.D. & STEINER, K.E. 1995. Long proboscid fly pollina-
tion of two orchids in the Cape Drakensberg Mountains, South
Africa. Plant Systematics and Evolution 195: 169-176.
KURZWEIL, H. 1995. Floral morphology and ontogeny of Satyriinae
(Orchidaceae). Flora 190: 1-20.
KURZWEIL, H. 1996. Floral morphology and ontogeny of Satyriinae
(Orchidaceae). Flora 191: 9-28.
KURZWEIL, H. & SMITH, G.F. 1995. The succulent orchids of southern
Africa. Aloe 32: 40-43.
KURZWEIL, H„ LINDER, HP, STERN, W.L. & PRIDGEON, A. 1995.
Comparative vegetative anatomy and classification of Diseae
(Orchidaceae). Botanical Journal of the Linnean Society 117:
171-220.
LAVRANOS, J.J., CRAIB, C.L. & CONDY, G. 1995. Pelargonium
hystrix. Flowering Plants of Africa 54: 42-46, t. 2109.
LEISTNER, O.A., SMITH, G.F. & GLEN, H.F. 1995. Notes on Podo-
carpus in southern Africa and Madagascar (Podocarpaceae).
Bothalia 25: 233-236.
LINCOLN, THALIA, ROURKE, J„ DUNCAN, G.D. & VAN JAARS-
VELD, E. 1996. Flowers of southern Africa. The Sappi Collec-
tion Calendar.
198
Bothalia 26,2(1996)
LINDER, H.P. & KURZWEIL, H. 1995. Taxonomic notes on the African
Orchidoideae (Orchidaceae): a new genus and combination.
Willdenowia 25: 229-234.
LOW, A.B. & REBELO, A.G. 1996. The vegetation types of South Africa,
Lesotho and Swaziland. Edina Griffiths Publishers.
MANDER, M„ MANDER, J„ CROUCH, N„ MCKEAN, S. &
NICHOLS, G. 1996. Growing and knowing muthi plants. Share-
Net Publications, Howick.
MANNING, J.C. 1006. Diversity of endothecial thickenings in the
Angiosperms. In W. Darcy & R.C. Keating, The anther: 136-158.
MANNING, J.C. & GOLDBLATT, P. 1995. Cupid comes in many guises.
Veld & Flora 81: 50-53.
MARAIS, E.M. & CONDY, G. 1995. Pelargonium punctatum. Flower-
ing Plants of Africa 54: 36-40, t. 2108.
MCDONALD, D.J. & COWLING, R.M. 1995. Towards a profile of an
endemic Mountain Fynbos flora: implications for conservation.
Biological Conservation 72: 1-12.
MCDONALD, D.J., JURITZ, J.M., COWLING, R.M. & KNOTTEN-
BELT, W.J. 1995. Modelling the biological aspects of local ende-
mism in South African Fynbos. Plant Systematics and Evolution
195: 137-147.
MIDGLEY, G.F., STOCK, W.D. & JURITZ, J.M. 1995. The effects of
elevated CO2 on Cape Fynbos species adapted to soils of different
nutrient status: nutrient- and CO2- responsiveness. GCTE Con-
ference Proceedings, Journal of Biogeography 22: 185-191.
MILTON, S.J., DEAN, W.R.J., HOFFMAN, M.T. & KERLEY, G.I.H.
1995. Why can’t developed countries control desertification?
Bulletin of the Grassland Society of Southern Africa 6: 15-17.
MOSSMER, M„ SMITH, G.F., VAN WYK, B-E. & CONDY, G. 1995.
Haworthia fasciata forma fasciata. Flowering Plants of Africa
54: 88-93, t. 2118.
MUSEL, C.F. 1994/1995. Ultraviolet-B irradiation of seeds affects pho-
tochemical and reproductive performance of the arid-environ-
ment ephemeral Dimorphotheca pluvialis. Environmental and
Experimental Botany 34: 371-378.
MUSIL, C.F. 1995. Differential effect of elevated ultraviolet-B radiation
on the photochemical and reproductive performance of dicotyle-
donous and monocotyledonous arid-environment ephemerals.
Plant, Cell and Environment 18: 844—854.
NGWENYA, M.A., SINGH, Y„ CROUCH, N.R., SCOTT-SHAW, C.R.
& SIKHAKHANE, T.B. 1995. Brachystelma natalense: redis-
covered and redescribed. Aloe 32: 44, 45.
OLIVER, E.G.H. 1995. A new species of Erica from the Western Cape
(Ericaceae). Bothalia 25: 242-244.
OLIVER, E.G.H. 1996. The position of Bruckenthalia versus Erica.
Yearbook of the Heather Society 1996: 6.
OLIVER, E.G.H. & OLIVER, I.M. 1995. Studies in the Ericoideae
(Ericaceae). XVI. Six new species of Erica from the Western
Cape, South Africa. Bothalia 25: 87-95.
OLIVER, E.G.H. & OLIVER, I.M. 1996. Studies in the Ericaceae
(Ericoideae). XX. Arare new species of Erica from South Africa.
Yearbook of the Heather Society 1996: 1-5.
PATERSON-JONES, C. & SNIJMAN, D A. 1995. Dramatically differ-
ent: winter rainfall Amaryllids. Africa-Environment & Wildlife
4,2: 25-30.
PEROLD, S.M. 1995a. Studies in the Marchantiaies (Hepaticae) from
southern Africa. 8. The genus Plagiochasma (Aytoniaceae: Ay-
tonioideae) and six local taxa. Bothalia 25: 13-29.
PEROLD, S.M. 1995b. Studies in the Marchantiaies (Hepaticae) from
southern Africa. 9. The genus Marchantia and its five local
species. Bothalia 25: 183-204.
PEROLD, S.M. 1995c. The taxonomic history of the Ricciaceae
(1937-1995) and a classification of sub-Saharan Ricciae.
Bothalia 25: 211-231.
PEROLD, S.M. 1995d. Fruiting Lunularia cruciata , now also reported
from southern Africa (Lunulariaceae). Bothalia 25: 239-242.
PEROLD, S.M. 1995e. A survey of the Ricciaceae of tropical Africa.
Fragmenta Floristica et Geobotanica 40: 53-91.
REBELO, A.G. (ed.). 1995a. Protea Atlas Newsletter 27.
REBELO, A.G. 1995b. SASOL: Proteas. A field guide to the proteas of
southern Africa. Femwood Press, in association with the National
Botanical Institute.
REBELO, A.G. 1995c. Proteas of the summer rainfall region. Environ-
mental Resource Guide: Pocket field guide 5. EKKO, Protea
Atlas Project, National Botanical Institute and the Botanical
Society of South Africa.
REBELO, A.G. 1995d. Algorithms for biodiversity reserve configura-
tions. 1993-94 Yearbook of the Wissenschaftskolleg zu Berlin:
138-140.
REBELO, A.G. 1995e. Five years of Protea atlassing. Veld & Flora 81:
124-127.
REBELO, A.G. & REBELO, P. (eds). 1995a. Protea Atlas Newsletter 28.
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REBELO, A.G. & REBELO, P. (eds). 1995c. Protea Atlas Newsletter 30.
RETIEF, E„ NICHOLAS, A. & BAIJNATH, H. 1995. The psocid
Liposcelis bostrychophilus Badonnel (Psocoptera: Liposceli-
dae): an occasional herbarium pest. Bothalia 25: 247-253.
ROURKE, J.P. 1995a. Obituary: Hedley Brian Rycroft. Transactions of
the Royal Society of South Africa 50: 103-105.
ROURKE.J.P. 1995b. Obituary: Winsome (Buddy) Barker(1907-1994).
Bothalia 25: 255-259.
ROURKE, J.P. & WARD-HILHORST, E. 1995. Serruria meisneriana.
Flowering Plants of Africa 54: 48-52, t. 21 10.
ROUX, J.P. 1995. Systematic studies in the genus Mohria (Pteridophyta:
Anemiaceae). VI. Taxonomic review. Bothalia 25: 1-12.
RUTHERFORD, M.C., O’CALLAGHAN, M„ HURFORD, J.L.,
POWRIE, L.W., SCHULZE, R.E., KUNZ, R.P., DAVIS, G.W.,
HOFFMAN, M.T. & MACK, F. 1995. Realized niche spaces and
functional types: a framework for prediction of compositional
change. Journal of Biogeography 22: 523-531.
RUTHERFORD, M.C., O’CALLAGHAN, M„ HURFORD, J.L.,
POWRIE, L.W. & SCHULZE, R.E. 1995. Modelling survival of
plant populations with differing mobility and sensitivity to cli-
mate change over real and simulated terrain types. In P. Binning,
H. Bridgeman & B. Williams. International Congress of Model-
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SAJEVA, M„ CARIMI, F. & VAN JAARSVELD, E.J. 1995. In vitro
propagation of Aloe polyphylla Schonland ex Pillans (Aloaceae).
Giornale Botanico Italiano 129, 2: 61.
SAMWAYS, M.J., STORK, N.E., CRACRAFT, J„ EELEY, H.A.C.,
FOSTER, M„ LUND, C. & HILTON-TAYLOR, C. 1995. Scales,
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SINGH, Y„ VAN WYK, A.E. & BAIJNATH, H. 1995. Know your arums:
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SLATTERY, C.M. 1995. One more step along the road ... towards an
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cum 32,1: 9.
SMITH, G.F. 1995e. The distribution, uses and history of Agave in
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. i . .
Bothalie 26,2: 201-210(1996)
Guide for authors to Bothalia
This guide is updated when necessary and includes an
index. Important points and latest additions appear in
bold type.
Bothalia is named in honour of General Louis Botha,
first Premier and Minister of Agriculture of the Union of
South Africa. This house journal of the National Botanical
Institute, Pretoria, is devoted to the furtherance of botanical
science. The main fields covered are taxonomy, ecology,
anatomy and cytology. Two parts of the journal and an index
to contents, authors and subjects are published annually.
1 Editorial policy
1.1 Bothalia welcomes original papers dealing with flora
and vegetation of southern Africa and related subjects.
Full-length papers and short notes, as well as book re-
views, are accepted.
1.2 Page charges: As stated in our notification included
in volume 23,1 (May 1993), MSS submitted for publica-
tion in Bothalia after the 10th June 1993 are subject to
payment of page charges of R 1 25,00 per printed page, VAT
included. The following are exempt from these charges:
1, NBI members; 2, persons/institutions who have been
granted exemption by the Executive Committee of the
NBI; 3, authors of contributions requested by the Editor;
4, contributors to the column ‘FSA contributions’. The
Editor’s decision on the number of pages is final. An
invoice will be sent to the author, who must arrange for
payment as soon as possible to NBI, Publications section,
Private Bag X101, Pretoria 0001.
1.3 Articles are assessed by referees, both local and over-
seas. Authors are welcome to suggest possible referees to
judge their work. Authors are responsible for the factual
correctness of their contributions. Bothalia maintains an
editorial board (see title page) to ensure that international
standards are upheld.
2 Requirements for a manuscript
2.1 The original manuscript should be typed on one side
of A4-size paper, double-spaced throughout (including
abstract, tables, captions to figures, literature references,
etc.) and have a margin of at least 30 mm all round. Three
photocopies (photocopied on both sides of the paper to
reduce weight for postage) of all items, including text,
illustrations, tables and lists should be submitted, and the
author should retain a complete set of copies. If the article
was generated on a computer, a copy of the diskette should
be submitted with the final (accepted) version (see 3).
2.2 Papers should conform to the general style and layout
of recent issues of Bothalia (from volume 26 onwards).
2.3 Material should be presented in the following se-
quence: Title page with title, name(s) of author(s), key-
words, abstract (and information that should be placed in
a footnote on the title page, such as address(es) of author(s)
and mention of granting agencies.
2.4 The sequence continues with Introduction and aims,
Material and methods, Results, Interpretation (Discus-
sion), Specimens examined (in revisions and mono-
graphs), Acknowledgements, References, Index of names
(recommended for revisions dealing with more than about
15 species), Tables, Captions for figures and figures. In the
case of short notes and book reviews, keywords and ab-
stract are superfluous.
2.5 All pages must be numbered consecutively beginning
with the title page to those with references, tables and
captions to figures.
2.6 For notes on the use of hyphens and dashes see 3.10
to 3.12.
2.7 Special character: use your own word or code that is
unique and self-explanatory, enclosed between ANGLE
BRACKETS, e.g. <mu>m for pm. Please supply us with
a list of the codes.
3 Requirements for diskettes/stiffies
(to be submitted only with final/accepted version)
3.1 data must be IBM compatible and written in ASCII,
or in Word for Windows from Windows 1; Word for
MS-DOS from MSWord 3; WordPerfect 5 for DOS
only; Windows Write 3 onwards.
3.2 the original printout of the diskette should be sup-
plied in double line spacing.
3.3 tables need not be placed on the diskette — a clearly
laid out hard copy is adequate.
3.4 use a non-breaking space to keep two elements to-
gether on the same line, e.g. 3 500.
3.5 do not justify lines.
3.6 do not break words, except hyphenated words.
3.7 all lines, headings, keys, etc., should start flush at the
margin, therefore no indentations, footnotes or tabs of
any kind.
3.8 in Word and WordPerfect, italics and bold should be
used where necessary.
3.9 paragraphs and headings are delineated by a carriage
return (ENTER) but no indentation.
3.10 a hyphen is designated as one dash, with no space
between the letter and the dash, e.g. ovate-lanceolate. See
also 17.6.
202
Bothalia 26,2 (1966)
3.11 an N-dash is typed as three hyphens with no space
between the letter and the hyphen, e.g. 2 — 5 mm (typeset,
it looks like this, 2-5 mm).
3.12 an M-dash is typed as two hyphens with no space
between the letter and the hyphen, e.g. computers - - what
a blessing! (typeset, it looks like this, computers — what).
3.13 do not use a double space between words, after
commas, full stops, colons, semicolons or exclamation
marks.
3.14 use lower case x as times sign, with one space on
either side of the x, e.g. 2x3 mm.
3.15 use single (not double) opening and closing quotes,
e.g. the so-called ‘stiffy’ refers to a rigid diskette.
3.16 keys — put only three leader dots before number and
name of taxon (with one space before and one space after
the first and last dot), regardless of how far or near the word
is from the right margin, e.g. ... 1. R. ovata (see 13.18).
4 Author(s)
When there are several authors the covering letter
should indicate clearly which of them is responsible for
correspondence and, if possible, telephonically available
while the article is being processed. The contact address
and telephone number should be mentioned if they differ
from those given on the letterhead.
5 Title
The title should be as concise and as informative as
possible. In articles dealing with taxonomy or closely re-
lated subjects the family of the taxon under discussion
(see also 13.2) should be mentioned in brackets but
author citations should be omitted from plant names
(see also 13.6).
6 Keywords
Up to 10 keywords (or index terms) should be provided
in English in alphabetical sequence. The following points
should be borne in mind when selecting keywords:
6. 1 keywords should be unambiguous, internationally ac-
ceptable words and not recently coined little-known
words.
6.2 they should be in a noun form and verbs should be
avoided.
6.3 they should not consist of an adjective alone; adjec-
tives should be combined with nouns.
6.4 they should not contain prepositions.
6.5 the singular form should be used for processes and
properties, e.g. evaporation.
6.6 the plural form should be used for physical objects,
e.g. augers.
6.7 location (province and/or country); taxa (species, ge-
nus, family) and vegetation type (community, veld type,
biome) should be used as keywords.
6.8 keywords should be selected hierarchically where
possible, e.g. both family and species should be included.
6.9 they should include terms used in the title.
6.10 they should answer the following questions:
6. 10. 1 what is the active concept in the document (activity,
operation or process).
6.10.2 what is the passive concept or object of the active
process (item on which the activity, operation or process
takes place).
6. 10.3 what is the means of accomplishment or how is the
active concept achieved (technique, method, apparatus,
operation or process).
6.10.4 what is the environment in which the active con-
cept takes place (medium, location).
6.10.5 what are the independent (controlled) and depend-
ent variables?
6.11 questions 6.10.1 to 6.10.3 should preferably also be
answered in the title.
7 Abstract
7.1 Abstracts of no more than 200 words should be pro-
vided. Abstracts are of great importance and should con-
vey the essence of the article.
7.2 They should refer to the geographical area concerned
and, in taxonomic articles, mention the number of taxa
treated. They should not contain information not appear-
ing in the article.
7.3 In articles dealing with taxonomy or closely related
subjects all taxa from the rank of genus downwards should
be accompanied by their author citations (see also 13.6).
7.4 Names of new taxa and new combinations should not
be italicized. If the article deals with too many taxa, only
the important ones should be mentioned.
8 Table of contents
A table of contents should be given for all articles
longer than about 15 typed pages, unless they follow the
strict format of a taxonomic revision.
Bothalia 26,2(1996)
203
9 Acknowledgements
Acknowledgements should be kept to the minimum
compatible with the requirements of courtesy. Please give
all the initials of the person(s) you are thanking.
10 Literature references
In text
10. 1 Literature references in the text should be cited as
follows: ‘Jones & Smith (1986) stated...’, or ‘...(Jones &
Smith 1986)’ or (Ellis 1988: 67) when giving a reference
simply as authority for a statement. For treatment of litera-
ture references in taxonomic papers see 14.
10.2 When more than two authors are involved in the
paper use the name of the first author followed by et al.
10.3 When referring to more than one literature reference,
they should be arranged chronologically and separated by
a semicolon, e.g. (Nixon 1940; Davis 1976; Anon. 1981,
1984).
10.4 Titles of books and names of journals should prefer-
ably not be mentioned in the text. If there is good reason
for doing so, they should be treated as described in 10.12
& 10.13.
10.5 Personal communications are given only in the text,
not in the list of references. Please add the person’s full
initials to identify the person more positively, e.g. C.
Boucher pers. comm.
In References at end of article
10.6 References of the same author are arranged in
chronological sequence.
10.7 Where two or more references by the same author
are listed in succession, the author’s name is repeated with
every reference, except in an obituary, where the name of
the deceased in the list of publications (not in the refer-
ences) is replaced by an N-dash.
10.8 All publications referred to in the text, including
those mentioned in full in the treatment of correct names
in taxonomic papers, but no others, and no personal com-
munications, are listed at the end of the manuscript under
the heading References.
10.9 The references are arranged alphabetically accord-
ing to authors and chronologically under each author, with
a, b, c, etc. added to the year, if the author has published
more than one work in a year. This sequence is retained
when used in the text, irrespective of the chronology.
10. 10 If an author has published both on his own and as a
senior author with others, the solo publications are listed
first and after that, in strict alphabetical sequence, those
published with one or more other authors.
10.11 Author names are typed in capitals.
10.12 Titles of journals and of books are written out in
full and are italicized as follows: Transactions of the
Linnean Society of London 5: 171-217, or Biology and
ecology of weeds: 24.
10.13 Titles of books should be given as in Taxonomic
literature, edn 2 by Stafleu & Cowan and names of jour-
nals as in the latest edition of World list of scientific
periodicals.
10.14 Examples of references:
Collective book or Flora
BROWN, N.E. 1909. Asclepiadaceae. In W.T. Thiselton-Dyer, Flora ca-
pensis 6,2: 518-1036. Reeve, London.
CUNNINGHAM, A.B. 1994. Combining skills: participatory approaches
in biodiversity conservation. In B.J. Huntley, Botanical diversity in south-
ern Africa. Strelitzia 1: 149-167. National Botanical Institute, Pretoria.
Book
DU TOIT, A.L. 1966. Geology of South Africa , 3rd edn, S.M. Haughton
(ed ). Oliver & Boyd, London.
HUTCHINSON, J. 1946. A botanist in southern Africa. Gawthom, Lon-
don.
Journal
DAVIS, G. 1988. Description of a proteoid-restioid stand in Mesic Moun-
tain Fynbos of the southwestern Cape and some aspects of its ecology.
Bothalia 18: 279-287.
SMOOK, L. & GIBBS RUSSELL, G.E. 1985. Poaceae. Memoirs of the
Botanical Survey of South Africa No. 51: 45-70.
STEBBINS, G.L. Jr 1952. Aridity as a stimulus to plant evolution. Ameri-
can Naturalist 86: 35 — 44.
In press, in preparation
TAYLOR, H.C. in press. A reconnaissance of the vegetation of Rooiberg
State Forest. Technical Bulletin, Department of Forestry.
VOGEL, J.C. 1982. The age of the the Kuiseb river silt terrace at Homeb.
Palaeoecology of Africa 15. In press.
WEISSER, P.J., GARLAND, J.F. & DREWS, B.K. in prep. Dune ad-
vancement 1937-1977 and preliminary vegetation succession chronology
at Mlalazi Nature Reserve, Natal, South Africa. Bothalia.
Thesis
KRUGER, F.J. 1974. The physiography and plant communities of the
Jakkalsrivier Catchment. M.Sc. (Forestry) thesis. University of Stellen-
bosch.
MUNDAY, J. 1980. The genus Monechma Hochst. (Acanthaceae tribe
Justiciae) in southern Africa. M.Sc. thesis, University of the Witwaters-
rand, Johannesburg.
Miscellaneous paper, report, unpublished article, techni-
cal note, congress proceedings
ANON, no date. Eetbare plante van die Wolkberg. Botanical Research
Unit, Grahamstown. Unpublished.
BAWDEN, M.G. & CARROL, D.M. 1968. The land resources of Le-
sotho. Land Resources Study No. 3, Land Resources Division, Director-
ate of Overseas Surveys, Tolworth.
BOUCHER, C. 1981. Contributions of the Botanical Research Institute.
204
Bothalia 26,2(1966)
In A.E.F. Heydom, Proceedings of workshop research in Cape estuaries :
105-107. National Research Institute for Oceanology, CSIR, Stellenbosch.
NATIONAL BUILDING RESEARCH INSTITUTE 1959. Report of the
committee on the protection of building timbers in South Africa against
termites, woodboring beetles and fungi, 2nd edn. CSIR Research Report
No. 169.
11 Tables
11.1 Each table should be presented on a separate sheet
and be assigned an Arabic numeral, i.e. the first table
mentioned in the text is marked ‘Table 1 ’ .
11.2 In the captions of tables the word ‘TABLE’ is written
in capital letters. See recent numbers of Bothalia for the
format required.
11.3 Avoid vertical lines, if at all possible. Tables can often
be reduced in width by interchanging primary horizontal
and vertical heads.
12 Figures
12. 1 Figures should be planned to fit, after reduction, into
a width of either 80, 118 or 165 mm, with a maximum
vertical length of 230 mm. Allow space for the caption in
the case of figures that will occupy a whole page.
12.2 Line drawings, including graphs and diagrams,
should be twice the size of the final reproduction and
should be in jet-black Indian ink, preferably on fine Felix
Schoeller parole or similar paper, 200 gsm, or tracing film.
Lines should be bold enough and letters/symbols large
enough to stand reduction.
12.3 Photographs should be of excellent quality on glossy
paper with clear detail and moderate contrast, and they
should be the same size as required in the journal.
12.4 Photograph mosaics should be submitted complete,
the component photographs mounted neatly on a white
flexible card base (can be curved around drum of
scanner) leaving a narrow gap of uniform width (2 mm)
between each print. Note that grouping photographs of
markedly divergent contrast results in poor reproductions.
12.5 Lettering and numbering on all figures should be
done in letraset, stencilling or a comparable method. If
symbols are to be placed on a dark background it is
recommended that black symbols are used on a small
white disk ± 7 mm in diameter and placed in the lower
left hand corner of the relevant photo.
12.6 If several illustrations are treated as components of
a single composite figure they should be designated by
capital letters.
12.7 Note that the word ‘Figure’ should be written out in
full, both in the text and the captions and should begin with
a capital ‘F’(but see 14.7 for taxonomic papers).
12.8 In the text the figure reference is then written as in the
following example: ‘The stamens (Figure 4A, B, C) are...’
12.9 In captions, ‘FIGURE’ is written in capital letters.
Magnification of figures should be given for the size as
submitted.
1 2. 1 0 Scale bars or scale lines should be used on figures.
12.11 In figures accompanying taxonomic papers, vou-
cher specimens should be given in the relevant caption.
12.12 Figures are numbered consecutively with Arabic
numerals in the order they are referred to in the text.
These numbers, as well as the author’s name and an
indication of the top of the figure, must be written in soft
pencil on the back of all figures.
12.13 Captions of figures must not be pasted under the
photograph or drawing.
12. 14 Authors should indicate in pencil in the text where
they would like the figures to appear.
12.15 Authors wishing to have the originals of figures
returned must inform the editor in the original covering
letter and must mark each original ‘To be returned to
author’.
12.16 Authors wishing to use illustrations already pub-
lished must obtain written permission before submitting
the manuscript and inform the editor of this fact.
12.17 Captions for figures should be collected together
and typed at the end of the MS and headed Captions for
figures.
12.18 It is strongly recommended that taxonomic articles
include dot maps as figures to show the distribution of
taxa. The dots used must be large enough to stand reduc-
tion to 80 mm (recommended size: letraset 5 mm diame-
ter). No open diamonds or open triangles should be
used.
12. 19 Blank distribution maps of southern Africa, Africa
and the world are available from the Bookshop, NBI
Pretoria.
13 Text
13.1 As a rule, authors should use the names (but not of
all authors of plant names — see 13.6) as listed by T.H.
Arnold & B.C. de Wet (eds) in Memoirs of the Botanical
Survey of South Africa No. 62.
13.2 Names of genera and infrageneric taxa are usually
italicized, with the author citation (where relevant; see
1 3.6) not italicized. Exceptions include names of new taxa
in the abstract, correct names given in the synopsis or in
paragraphs on species excluded from a given supraspecific
group in taxonomic articles; in checklists and in indices,
where the position is reversed, correct names are not
italicized and synonyms are italicized.
13.3 Names above generic level are not italicized.
Bothalia 26,2(1996)
205
1 3.4 In articles dealing with taxonomy and closely related
subjects the complete scientific name of a plant (with
author citation) should be given at the first mention in the
text. The generic name should be abbreviated to the initial
thereafter, except where intervening references to other
genera with the same initial could cause confusion (see
16.6).
13.5 In normal text, Latin words are italicized, but in the
synopsis of a species, Latin words such as nom. nud. are
not italicized (see 14.3, 16.4, 17.9).
13.6 In accordance with Garnock- Jones & Webb (1996)
in Taxon 45: 285, 286, authors of plant names are not to be
added to plant names except in taxonomic papers. Names
of authors of plant names should agree with the list pub-
lished by the Royal Botanic Gardens, Kew, entitled,
Authors of plant names, edited by R.K. Brummitt & C.E.
Powell (1992).
13.7 Modern authors not included in the list should use
their full name and initials when publishing new plant
names. Other author names not in the list should be in
agreement with the recommendations of the Code.
13.8 Names of authors of publications are written out in
full except in the synonymy in taxonomic articles where
they are treated like names of authors of plant names.
13.9 Names of plant collectors are italicized whenever
they are linked to the number of a specimen. The collec-
tion number is also italicized, e.g. Acocks 14407.
13.10 Surnames beginning with ‘De’, ‘Du’ or ‘Van’ begin
with a capial letter unless preceded by an initial.
13.11 For measurements use only units of the Interna-
tional System of Units (SI). In taxonomic papers only
mm and m, should be used; in ecological papers cm or
m should be used.
13.12 The use of ‘±‘ is preferred to c. or ca (see 17.7).
13.13 Numbers ‘one’ to ‘nine’ are spelled out in normal
text, and from 10 onwards they are written in Arabic
numerals.
13.14 In descriptions of plants, numerals are used
throughout. Write 2.CM-.5 (not 2 — 4.5). When counting
members write 2 or 3 (not 2-3), but 2^1.
13.15 Abbreviations should be used sparingly but con-
sistently. No full stops are placed after abbreviations end-
ing with the last letter of the full word (e.g. edition = edn;
editor = ed.); after units of measure; after compass direc-
tions; after herbarium designations; after countries, e.g.
USA and after well-known institutions, e.g. CSIR.
13.16 Apart from multi-access keys, indented keys should
be used with couplets numbered la-lb, 2a-2b, etc. (with-
out full stops thereafter).
13.17 Keys consisting of a single couplet have no num-
bering.
13.18 Manuscripts of keys should be presented as in the
following example:
la Leaves closely arranged on an elongated stem; a sub-
merged aquatic with only the capitula exserted ... lb. E.
setaceum var. pumilum
lb Leaves in basal rosettes; stems suppressed; small marsh
plants, ruderals or rarely aquatics:
2a Annuals, small, fast-growing pioneers, dying when the
habitat dries up; capitula without coarse white setae; re-
ceptacles cylindrical:
3a Anthers white ... 2. E. cinereum
3b Anthers black ... 3. E. nigrum
2b Perennials, more robust plants; capitula sparsely to
densely covered with short setae:
13.19 Herbarium voucher specimens should be referred
to wherever possible, not only in taxonomic articles.
14 Species treatment in taxonomic papers
14.1 The procedure to be followed is illustrated in the
example (17.9), which should be referred to, because not
all steps are described in full detail.
14.2 The correct name (not italicized) is to be followed
by its author citation (italicized) and the full literature
reference, with the name of the publication written out in
full (not italicized).
14.3 Thereafter all literature references, including those
of the synonyms, should only reflect author, page and year
of publication, e.g. C.E. Hubb. in Kew Bulletin 15: 307
(1960); Boris et al.: 14 (1966); Boris: 89 (1967); Sims: t.
38 (1977); Sims: 67(1980).
14.4 The description and the discussion should consist of
paragraphs commencing, where possible, with italicized
leader words such as flowering time, diagnostic charac-
ters, distribution and habitat.
14.5 When more than one species of a given genus is dealt
with in a paper, the correct name of each species should
be prefixed by a sequential number followed by a full stop.
Infraspecific taxa are marked with small letters, e.g. lb.,
12c., etc.
14.6 Names of authors are written as in 13.6, irrespective
of whether the person in question is cited as the author of
a plant name or of a publication.
14.7 The word ‘figure’ is written as ‘fig.’, and ‘t.’ is used
for both ‘plate’ and ‘tablet’ (but see 12.7 for normal text).
14.8 Literature references providing good illustrations of
the species in question may be cited in a paragraph com-
mencing with the word leones followed by a colon. This
paragraph is given after the last paragraph of the syn-
onymy, see 17.9.
206
Bothalia 26,2(1966)
15 Citation of specimens
15.1 "type specimen in synopsis: the following should be
given (if available): country (if not in RSA), province, grid
reference (at least for new taxa), locality as given by
original collector, modem equivalent of collecting locality
in square brackets (if relevant, e.g. Port Natal [now Dur-
ban]), quarter-degree square, date of collection (op-
tional), collector’s name and collecting number (both
italicized).
15.2 The abbreviation s.n. ( sine numero) is given after the
name of a collector who usually assigned numbers to his
collections but did not do so in the specimen in question.
The herbaria in which the relevant type(s) are housed are
indicated by means of the abbreviations given in the latest
edition of Index Herbariorum.
15.3 The holotype (holo.) and its location are mentioned
first, followed by a semicolon, the other herbaria are
arranged alphabetically, separated by commas.
15.4 Authors should indicate by means of an exclamation
mark (!) which of the types have been personally exam-
ined.
15.5 If only a photograph or microfiche was seen, write
as follows: Anon. 422 (X, holo.-BOL, photo.!).
15.6 Lectotypes or neotypes should be chosen for correct
names without a holotype. It is not necessary to lectotypify
synonyms.
15.7 When a lectotype or a neotype are newly chosen, this
should be indicated by using the phrase ‘here designated’
(see 17.9). If reference is made to a previously selected
lectotype or neotype, the name of the designating author
and the literature reference should be given. In cases where
no type was cited, and none has subsequently been nomi-
nated, this may be stated as ‘not designated’.
15.8 In brief papers mentioning only a few species and a
few cited specimens the specimens should be arranged
according to the grid reference system: Provinces/ coun-
tries (typed in capitals) should be cited in the following
order: Namibia, Botswana, Northern Province (previously
Northern Transvaal), North-West (previously northeastern
Cape and southwestern Transvaal), Gauteng (previously
PWV), Mpumalanga (previously Eastern Transvaal), Free
State (previously Orange Free State), Swaziland, KwaZulu-
Natal, Lesotho, and Northern Cape, Western Cape and
Eastern Cape (see Figure 1, p. 208).
15.9 Grid references should be cited in numerical se-
quence.
15.10 Locality records for specimens should preferably
be given to within a quarter-degree square. Records from
the same one-degree square are given in alphabetical or-
der, i.e (-AC) precedes (-AD), etc. Records from the same
quarter-degree square are arranged alphabetically accord-
ing to the collectors’ names; the quarter-degree references
must be repeated for each specimen cited.
15.11 The relevant international code of the herbaria in
which a collection was seen should be given in brackets
after the collection number; the codes are separated by
commas. The following example will explain the proce-
dure:
KWAZULU-NATAL. — 2731 (Louwsburg): 16 km E of Nongoma, (-DD),
Pelser 354 (BM, K, PRE); near Dwarsrand, Van der Merwe 4789 (BOL,
M). 2829 (Hanismith): near Groothoek, (-AB), Smith 234\ Koffiefontein,
(-AB), Taylor 720 (PRE); Cathedral Peak Forest Station, (-CC), Marriot
74 (KMG); Wilgerfontein, Roux 426. Grid ref. unknown: Sterkstroom,
Strydom 12 (NBG).
15.12 For records from outside southern Africa authors
should use degree squares without names, e.g.:
KENYA. — 0136: Nairobi plains beyond race course, Napier 485.
15.13 Monographs and revisions: in the case of all major
works of this nature it is assumed that the author has
investigated the relevant material in all major herbaria and
that he has provided the specimens seen with determinavit
labels. It is assumed further that the author has submitted
distribution maps for all relevant taxa and that the distri-
bution has been described briefly in words in the text.
Under the heading ‘Vouchers’ no more than five speci-
mens should be cited, indicating merely the collector and
the collector’s number (both italicized). Specimens are
alphabetically arranged according to collector’s name. If
more than one specimen by the same collector is cited,
they are arranged numerically and separated by a comma.
The purpose of the cited specimens is not to indicate
distribution but to convey the author’s concept of the taxon
in question.
15.14 The herbaria in which the specimens are housed are
indicated by means of the abbreviation given in the latest
edition of Index Herbariorum. They are given between
brackets, arranged alphabetically and separated by com-
mas behind every specimen as in the following example:
Vouchers: T.H. Arnold 64 (PRE); Fisher 840 (NH, NU, PRE); Flanagan
831 (GRA, PRE), 840 (NH, PRE); Marloth 4926 (PRE, STE); Schelpe
6161, 6163, 6405 (BOL); Schlechter 4451 (BM, BOL, GRA, K, PRE).
15.15 If long lists of specimens are given, they must be
listed together before Acknowledgements under the head-
ing Specimens examined. They are arranged alphabeti-
cally by the collector’s name and then numerically for each
collector. The species is indicated in brackets by the
number that was assigned to it in the text and any infra-
specific taxa by a small letter. If more than one genus is
dealt with in a given article, the first species of the first
genus mentioned is indicated as 1.1. This is followed by
the international herbarium designation. Note that the
name of the collector and the collection number are itali-
cized:
Acocks 12497 (2.1b) BM, K, PRE; 14724 (1.13a) BOL, K, P. Archer
1507 (1.4) BM, G. Burchell 2847 (2.8c) MB, K. Burman 2401 (3.3)
MO, S. B.L. Burn 789 (2.6) B, KMG, STE.
1 6 Synonyms
16. 1 In a monograph or a revision covering all of southern
Africa, all synonyms based on types of southern African
origin, or used in southern African literature, should be
included.
Bothalia 26,2(1996)
207
16.2 Illegitimate names are designated by nom. illeg. after
the reference, followed by non with the author and date, if
there is an earlier homonym.
16.3 Nomina nuda (nom. nud.) and invalidly published
names are excluded unless there is a special reason to cite
them, for example if they have been used in prominent
publications.
16.4 In normal text Latin words are italicized, but in the
synopsis of a species Latin words such as nom. nud., et al.
are not italicized (see 13.5, 14.3, 17.9).
16.5 Synonyms should be arranged chronologically into
groups of nomenclatural synonyms, i.e. synonyms based
on the same type, and the groups should be arranged
chronologically by basionyms, except for the basionym of
the correct name which is dealt with in the paragraph
directly after that of the correct name.
16.6 When a generic name is repeated in a given synony-
my it should be abbreviated to the initial, except where
intervening references to other genera with the same initial
could cause confusion (see 13.4).
17 Description and example of species treatment
17.1 Descriptions of all taxa of higher plants should,
where possible, follow the sequence: Habit; sexuality;
underground parts (if relevant). Indumentum (if it can be
easily described for the whole plant). Stems/branches.
Bark. Leaves : arrangement, petiole absent/present, pubes-
cence; blade: shape, size, apex, base, margin; midrib:
above/ below, texture, colour; petiole; stipules. Inflores-
cence: type, shape, position; bracts/bracteoles. Flowers :
shape, sex. Receptacle. Calyx. Corolla. Disc. Androecium.
Gynoecium. Fruit. Seeds. Chromosome number. Conser-
vation status. Figure (word written out in full) number.
17.2 As a rule, shape should be given before measure-
ments.
17.3 In general, if an organ has more than one of the parts
being described, use the plural, otherwise use the singular,
for example, petals of a flower but blade of a leaf.
17.4 Language must be as concise as possible, using
participles instead of verbs.
17.5 Dimension ranges should be cited as in 17.9.
17.6 Care must be exercised in the use of dashes and
hyphens. A hyphen is a short stroke joining two syllables
of a word, e.g. ovate-lanceolate or sea-green, with no space
between the letter and the stroke. An N-dash (en) is a
longer stroke commonly used instead of the word ‘to’
between numerals, ‘2-5 mm long’ (do not use it between
words but rather use the word ‘to’, e.g. ‘ovate to lanceo-
late’); it is produced by typing three hyphens next to each
other. An M-dash (em) is a stroke longer than an N-dash
and is used variously, e.g. in front of a subspecific epithet
instead of the full species name; it is produced by typing
two hyphens next to one another.
17.7 The use of ‘±’ is preferred to c. or ca when describ-
ing shape, measurements, dimensions, etc. (see 13.12).
17.8 The decimal point replaces the comma in all units
of measurement, e.g. leaves 1.0-1. 5 mm long.
17.9 Example:
1. Englerophytum magalismontanum ( Sond. ) T.D.Perm. The genera
of Sapotaceae: 252 (1991). Type: Gauteng, Magaliesbeig, Zeyher 1849 (S,
holo.-BOL, photo.!).
Bequaertiodendron magalismontanum (Sond.) Heine & Hemsl.: 307
(1960); Codd: 72 (1964); Elsdon: 75 (1980).
Chrysophyllum magalismontanum Sond.: 721 (1850); Harv.: 812
(1867); Engl.: 434 (1904); Bottmar: 34 (1919). Zeyherella magalismon-
tana (Sond.) Aubrdv. & Pellegr.: 105 (1958); Justin: 97 (1973).
Chrysophyllum argyrophyllum Hiem: 721 (1850); Engl.: 43 (1904).
Boivinella argyrophylla (Hiem) Aubr6v. & Pellegr.: 37 (1958); Justin et
al.: 98 (1973). Types: Angola, Welwitsch 4828 (BM!, lecto., here desig-
nated; PRE!); Angola, Welwitsch 4872 (BM!).
Chrysophyllum wilmsii Engl.: 4, t. 16 (1904); Masonet: 77 (1923);
Woodson: 244 (1937). Boivinella wilmsii (Engl.) Aubrtiv. & Pellegr.: 39
(1958); Justin: 99 (1973). Type: Mpumalanga, Magoebaskloof, Wilms
1812 [B, holo.f; K!, P!, lecto., designated by Aubrev. & Pellegr.: 38
(1958), PRE!, S!, W!, Z!].
Bequaertiodendron fruticosa De Wild.: 37 (1923), non Bonpl.: 590
(1823); D.Bakker: 167 (1929); H.Fr.: 302 (1938); Davy: 640 (1954);
Breytenbach: 117 (1959); Clausen: 720 (1968); Palmer: 34 (1969). Type:
Mpumalanga, Tzaneen Dist., Granville 3665 (K, holo.!; G!, P!, PRE!,
S!).
B. fragrans auct. non Oldemann: Glover: 149, t. 19 (1915); Henkel:
226 (1934); Stapelton: 6 (1954).
leones: Harv.: 812 (1867); Henkel: t. 84 (1934?); Codd: 73 (1964);
Palmer: 35 (1969).
Woody perennial; main branches up to 0.4 m long, erect
or decumbent, grey woolly-felted, leafy. Leaves linear to
oblanceolate, 3— 10(— 23) x 1.0-1.5(-4.0) mm, obtuse, base
broad, half-clasping. Heads heterogamous, campanulate, 7-8
x 5 mm, solitary, sessile at tip of axillary shoots; involucral
bracts in 5 or 6 series, inner exceeding flowers, tips sub-
opaque, white, very acute. Receptacle nearly smooth. Flow-
ers ± 23-30, 7-11 male, 16-21 bisexual, yellow, tipped pink.
Achenes ± 0.75 mm long, elliptic. Pappus bristles very
many, equalling corolla, scabridulous. Chromosome number.
2n = 22. Figure 23B.
18 New taxa
18.1 The name of a new taxon must be accompanied by
at least a Latin diagnosis. Authors should not provide
full-length Latin descriptions unless they have the re-
quired expertise in Latin at their disposal.
18.2 It is recommended that descriptions of new taxa be
accompanied by a good illustration (line drawing or pho-
tograph) and a distribution map.
18.3 Example:
109. Helichrysum jubilatum Hilliard, sp. nov. H. alsi-
noidei DC. affinis, sed foliis ellipticis (nec spatulatis), in-
florescentiis compositis a foliis non circumcinctis, floribus
femineis numero quasi dimidium hermaphroditorum
208
Bothalia 26,2 (1966)
aequantibus (nec capitulis homogamis vel floribus
femineis 1-3 tantum) distinguitur.
Herba annua e basi ramosa; caules erecti vel decum-
bentes, 100-250 mm longi, tenuiter albo-lanati, remote fo-
liati. Folia plerumque 8-30 x 5-15 mm, sub capitulis
minora, elliptica vel oblanceolata, obtusa vel acuta, mucro-
nata, basi semi-amplexicauli, utrinque cano-lanato- arach-
noidea. Capitula heterogama, campanulata, 3.5^4.0 x 2.5
mm, pro parte maxima in paniculas cymosas terminates
aggregata; capitula subterminalia interdum solitaria vel
2 vel 3 ad apices ramulorum nudorum ad 30 mm lon-
gorum. Bracteae involucrales 5-seriatae, gradatae, ex-
teriores pellucidae, pallide stramineae, dorso lanatae,
seriebus duabus interioribus subaequalibus et flores quasi
aequantibus, apicibus obtusis opacis niveis vix radian-
tibus. Receptaculum fere laeve. Flores ± 35-41. Ache-
nia 0.75 mm longa, pilis myxogenis praedita. Pappi setae
multae, corollam aequantes, apicibus scabridis, basibus
non cohaerentibus.
TYPE. — Northern Cape, 2817 (Vioolsdrif): Richters-
veld, (-CC), ± 5 miles E of Lekkersing on road to Stink-
fontein, kloof in hill south of road, annual, disc whitish,
7-11-1962, Nordenstam 1823 (S, holo.; E, NH, PRE).
19 New provinces of South Africa (Oct. 1996)
FIGURE 1. — 1, Western Cape; 2, Eastern Cape; 3, Northern Cape; 4,
Free State (previously Orange Free State); 5, KwaZulu-Natal; 6,
North-West (previously northeastern Cape and southwestern
Transvaal); 7, Gauteng (previously PWV); 8, Mpumalanga (pre-
viously Eastern Transvaal); 9, Northern Province (previously
Northern Transvaal).
20 Proofs
Only page proofs are normally sent to authors. They
should be corrected in red ink and be returned to the editor
as soon as possible.
21 Reprints
Authors receive 100 reprints free. If there is more than
one author, this number will have to be shared between them.
22 Documents consulted
Guides to authors of the following publications were
made use of in the compilation of the present guide: An-
nals of the Missouri Botanic Garden, Botanical Journal
of the Linnean Society, Flora of Australia, Smithsonian
Contributions to Botany, South African Journal of Botany
(including instructions to authors of taxonomic papers),
South African Journal of Science.
23 Address of editor
Manuscripts should be submitted to: The Editor,
Bothalia, National Botanical Institute, Private Bag X101,
Pretoria 0001.
24 FSA contributions
24.1 Figures and text must conform to Bothalia format.
24.2 These articles will be considered as a full contribu-
tion to the Flora of southern Africa and will be listed as
published in the ‘Plan of Flora of southern Africa' , which
appears in all issues of the FSA series.
INDEX
abbreviation, 13.4, 13.5, 13.12, 13.15, 14.7, 15.2, 15.14, 16.2, 16.3, 16.4,
16.6
abstract, 2.1, 2.3, 7, 13.2
acknowledgements, 9
address of
authors, 2.3, 4
editor, 23
alphabetical, 6, 10.3, 10.9, 10.10, 15.3, 15.10, 15.13, 15.14, 15.15
Arabic numerals, 11.1, 12.12, 13.3
ARNOLD, T.H. & DE WET, B.C. (eds) 1993. Plants of southern Africa:
names and distribution. Memoirs of the Botanical Survey of South
Africa No. 62, 13.1
ASCII, 3.1
author(s), 1, 2.1, 4, 10.14, 12.14
address, 2.3, 4
citation, 5, 7.3, 13.2, 13.4, 13.6, 14.2
first, 10.2
names, 2.3, 10.3, 10.7, 10.9, 10.11, 12.12, 13.7, 13.8, 14.3, 14.6, 15.7,
16.2
names of plant names, 5, 7.3, 13.6, 13.7, 13.8
senior, 10.10
book reviews, 1, 2.4
books, 10.4, 10.12, 10.13, 10.14
Bothalia, 1, 2.2, 11.2, 22
brief taxonomic articles, 15.8
BRUMMITT, R.K. & POWELL, C.E. (eds) 1992. Authors of plant names.
Royal Botanic Gardens, Kew, 13.6
c„ 13.2, 17.7
ca, 13.2, 17.7
Cape, 15.8, 18.3, 19
capitals, 11.2, 12.6, 12.9, 14.2, 15.8
captions, 2.1, 2.4, 2.5, 11.2, 12.7, 12.9, 12.11, 12.13, 12.17
checklist, 13.2
chromosome number, 17.1, 17.9
chronological sequence, 10.3, 10.6, 10.9, 16.5
citation
author, 5, 7.3, 13.2, 13.4, 13.6, 14.2
literature, 14.4
of specimens, 15
cm, 13.11
Bothalia 26,2 (1996)
209
collection
date, 15.1
number, 13.9, 15.1, 15.2, 15.11, 15.13, 15.15
collective book, 10.15
collector, 13.9, 15.1, 15.2, 15.10, 15.13, 15.15
colon, 3.13
comma, 3.13, 15.13
compass directions, 13.15
composite figure, 12.6
congress proceedings, 10.14
contents, 8
correspondence, 4
countries, 6.7, 15.8
decimal point, 17.8
description and example of species treatment, 17
diagrams, 12.2
discussion, 2.4, 14.4
diskette, 1, 3, 3.4
distribution maps, 12.18, 12.19, 15.13, 18.2
documents consulted, 22
DOS, 3.1
dot maps, 12.18, 12.19, 15.13, 18.2
double
line spacing, 3.2
space, 2.1, 3.13
drawing paper, 12.2
drawings, 12.2
Eastern Transvaal, see Mpumalanga
edition, 13.15
editor, 13.15, 22
editorial
board, 1
policy, 1
et al., 10.2, 13.5, 14.3, 17.9
example of
new taxa, 18.3
species treatment, 17.9
exclamation mark, 3.13, 15.4
family name, 5, 6.7
fig., 14.7
figure(s), 12, 14.7, 17.1
reduction of, 12.1, 12.2, 12.18
returned, 12.15
first author, 10.2
flora, 1, 10.14
Flora of southern Africa, 24
footnote, 2.3, 3.7
Free State (previously Orange Free State), 15.8, 19
FSA contributions, 24
full stop, 3.13, 13.15, 13.16, 14.5
GARNOCK-JONES, RJ. & WEBB, C.J 1996. The requirement to cite
authors of plant names in botanical journals. Taxon 45: 285, 286,
13.6
Gauteng (previously PWV), 15.8, 17.9, 19
genera, 13.2
generic name, 13.3, 13.4, 16.6
geographical area, 7.2
granting agencies, 2.3
graphs, 12.2
grid reference system, 15.1, 15.8, 15.9, 15.11
headings, 3.7, 3.9
sequence of, 2.3, 2.4
herbaria, 15.2, 15.3, 15.11, 15.13, 15.14
herbarium
code, 15.11
designations, 13.15, 15.15
voucher specimens, 12.12, 13.19
here designated, 15.7, 17.9
holo., 15.5, 17.9, 18.3
holotype, 15.3, 15.6
homonym, 16.2
hyphenated words, 2.6
hyphen, 3.10-3.12, 17.6
IBM compatible, 3.1
icones, 10.2, 17.9
illegitimate names (nom. illeg.), 16.2
illustrations, 12.4, 12.6, 12.16, 14.8
previously published, 12.16
indentations, 3.7, 3.9
Index Herbariorum, 15.2, 15.14
index of names, 2.4
infrageneric taxa, 13.2
initials, 9, 10.5, 13.7
in prep., 10.14
in preparation, 10.14
in press, 10.14
International
Code of Botanical Nomenclature, 13.7
System of Units (SI), 13.11
invalidly published names, 16.3
italics, 7.4, 10.12, 13.2, 13.3, 13.5, 13.9, 14.2, 15.1, 15.13, 15.15
journals, 10.4, 10.12, 10.14
names of, 10.1, 10.13
justify, 3.5
keys, 3.7, 2.16, 13.16, 13.17, 13.18
keywords, 2.3, 2.4, 6
KwaZulu-Natal (previously Natal), 15.8, 19
Latin, 13.5, 15.2, 16.2, 16.3, 16.4
descriptions, 18.1
layout, 2.2
lecto., 15.6, 15.7, 17.9
lectotype, 15.6, 15.7, 17.9
letraset, 12.5, 12.18
lettering, 12.5
line
drawings, 12.2, 18.2
spacing, 3.4, 3.9
literature
citations, 14.4
references, 2.1, 10, 10.7
within synonymy, 10.7, 14.8
localities outside southern Africa, 15.12
locality, 15.1, 15.10
m, 13.11
magnification of figures, 12.4, 12.9
manuscript
language, 1, 17.4
requirements, 2
map, distribution, dot, 12.18, 12.19, 15.13, 18.2, 19
M-dash, 3.12, 17.6
mm, 13.11
maigin, 2.1, 3.7, 3.16, 17.1
material, 2.3, 2.4
measurements, 13.11, 17.2, 17.7, 17.8
methods, 2.4, 6.10.3
microfiche, 15.5
miscellaneous paper, 10.14
monograph, 2.4, 15.13, 16.1
Mpumalanga (previously Eastern Transvaal), 15.8, 19
MSWord, 3.1, 3.8
name(s)
collector’s, 15.10
illegitimate, 16.2
invalidly published, 16.3
of authorfs), 2.3, 10.7, 10.9, 10.11, 13.7, 13.8, 14.6
of authors of plant names, 5, 13.1, 13.2, 13.6, 14.6
of publications, 13.8
Natal, see KwaZulu-Natal, 15.8, 19
N-dash, 3.11, 17.6
neotype, 15.6, 15.7
new
combinations, 7.4
provinces of South Africa (Oct. 1996), 15.8, 19
taxa, 7.4, 13.2, 13.7, 15.7, 18
nom. illeg., 16.2
nom. nud., 13.5, 16.3, 16.4
non-breaking space, 3.4
Northern Province, see Northern Transvaal, 15.8, 19
North-West, 15.8, 19
notes, 1, 2.4, 10.14
technical, 10.14
number
chromosome, 17.1, 17.9
page, 3.2
numbering, 13.13
figures, 12.5, 12.12, 17.1
keys, 13.16, 13.17
pages, 2.5, 13.4
taxa, 3.16, 7.2, 13.4, 14.5, 15.15
numerals, Arabic, 11.1, 12.12, 13.3
210
Bothalia 26,2 (1966)
Orange Free State, 15.8, 19
page charges, 1.2
PC diskettes, 3
pers. comm., 10.5, 10.8
personal communications (pers. comm.), 10.5, 10.8
photocopies, 2.1
photograph, 12.3, 12.4, 12.13, 15.5, 18.2
mosaic, 12.4
plant
collectors, 13.9
name, 5, 13.4, 13.6, 13.7, 13.8, 14.6
plate (t.), 14.7
prepositions, 6.4
proceedings, 10.14
proofs, 20
provinces, 6.7, 15.1, 15.8
of South Africa, 15.8, 19
publications, 10.8, 13.8, 14.3
name of, 14.2
solo, 10.10
year of, 10.9, 14.3
PWV, see Gauteng, 15.8, 19
quarter-degree squares, 15.1, 15.10
quotes, 3.15
reduction of figures, 12.1, 12.2, 12.18
referees, 1
reference, 2.4, 10.6, 10.7, 10.9, 10.14
figure, 12.8
grid, 15.1, 15.8, 15.9, 15.11
list, 10.5, 10.8, 10.9
literature, 2.1, 10, 10.7
report, 10.14
reprints, 21
requirements for
diskette, 3
manuscript, 2
results, 2.4
revision, 2.4, 8, 15.13, 16.1
scale bar, 12.10
semicolon, 3.13, 10.3, 15.3, 15.13
senior author, 10.10
sequence of headings, 2.3, 2.4
short notes, 1, 2.4
special character, 2.7
species treatment in taxonomic papers, 14
specimens examined, 2.4, 15.5
square brackets, 15.1, 17.9
STAFLEU, F.A. & COWAN, R.S. 1976-1988. Taxonomic literature. Vols
1-7, 10.13
stiffy/stiffies, 3
surnames, 13.10
symbols, 12.5
synopsis, 13.2, 13.5, 15.1, 16.4
synonymy, 10.7, 13.8, 14.4, 14.8, 16.6
t„ 14.3, 14.7, 17.9
table, 2.1, 2.4, 2.5, 3.3, 11
of contents, 8
tablet (t.), 14.7
tabs, 3.7
taxa
name of, 3.16, 5, 7.4, 10.8, 13.2, 13.3
new, 7.4, 13.2, 13.7, 15.7, 18
numbering of, 3.16, 7.2, 13.4, 14.5, 15.15
taxonomic
articles/papers, 7.2, 10.8, 12.11, 12.18, 13.2, 13.6, 13.8, 14
revision, 8
taxonomy, 5, 7.3, 13.4, 15.8
technical note, 10.15
text, 2.1, 10.1, 10.4, 10.5, 10.8, 11.1, 12.7, 12.8, 12.12, 12.14, 13, 15.13,
15.15, 16.4
thesis, 10.15
times sign, 3.14
title, 2.3, 5, 6.9, 6.11
of books, 10.4, 10.12, 10.13, 10.14
of journals, 10.4, 10.12, 10.13, 10.14
page, 1, 2.3, 2.5
Transvaal, 15.8, 17.9, 19
type, 15.2, 15.4, 15.7, 16.1, 16.6, 17.9
here designated, 15.7, 17.9
not designated, 15.7
specimen, 15.1
units of measure, 13.11, 13.15
unpublished article, 10.14
vouchers, 15.13, 15.14
voucher specimens, 12.11, 13.19
Windows Write, 3.1
Word for Windows, 3.1
WordPerfect, 3.1, 3.8
World list of scientific periodicals, 10.13
year of publication, 10.9, 14.3
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Volume 26,2
October 1996
CONTENTS
1. A synopsis of Peristrophe (Acanthaceae) in southern Africa. K. BALKWILL 83
2. Studies in the Ricciaceae of sub-Saharan Africa. S.M. PEROLD 95
3. FSA contributions 6: Orchidaceae: Holothrix. K.L. IMMELMAN 125
4. FSA contributions 7: Verbenaceaet : Vitex. C.L. BREDENKAMP and D.J. BOTHA 141
5. Notes on African plants:
Amaryllidaceae. Should Crinum forbesii (Lindl.) Schult. & Shult.f. be reinstated? R.H.
ARCHER and C. ARCHER 153
Asphodelaceae. Generic numbers in the subfamily Alooideae. G.F. SMITH and A.E. VAN
WYK 158
Campanulaceae. Transfer of eleven varieties of Lishtfootia nom. illeg. to Wahlenbereia. W.G.
WELMAN 157
Proteaceae. A new species of Serruria from the northern part of the Western Cape. J.P. ROURKE 1 54
6. Exormotheca bulbigena sp. nov. (Hepaticae, Marchantiales) and its relationship to E. holstii in
southern Africa. T. BORNEFELD, O.H. VOLK and R. WOLF 159
7. Morphological and biochemical genetic evidence for hybridization in the genus Centella
(Apiaceae), with notes on phylogenetic and taxonomic implications. M.T.R. SCHUBERT, B-E.
VAN WYK, H.F. VAN DER BANK and M. VAN DER BANK 167
8. Red Data List of southern African plants. 1. Corrections and additions. C. HILTON- TAYLOR . 177
9. Book reviews 183
10. National Botanical Institute: list of staff and publications, 31st May 1996. Compiler: B.A. MOM-
BERG 185
11. Guide for authors to Bothalia 201
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