NASIONALE BOTAN”
iNSTITUUT
Private Baa X 101 PRETORIA Ouu
m -06- 2 ?
Privaatsak X101 PRBTCM'A 0_
NATiONA1
.i./'.TiTl ITS
»\1 ;0
MAnnnn,V.NN LIBRARY
c_ 0000017978
B°od^ rifan National
Biodiversity Institute
BOTHALIA
VOL. 19
Published by the
Botanical Research Institute, Pretoria
Department of Agriculture and Water Supply
Gepubliseer deur die
Navorsingsinstituut vir Plantkunde, Pretoria
Departement van Landbou en Watervoorsiening
Edited by— Onder redaksie van
O.A. LEISTNER
Index prepared by— Indeks voorberei deur
B.A. MOMBERG
Digitized by the Internet Archive
in 2016
https://archive.org/details/bothaliavolume1919unse
CONTENTS — INHOUD
Page/Bladsy Date/Datum
No. 1 1-150 May/Mei 1989
No. 2 151-320 Oct./Okt. 1989
Page
Bladsy
ANDERSON, J.M. Review: The Banksia Atlas, ed. by Anne Taylor and Stephen Hopper 142
BARKER, N.P. The caryopsis surface of Pentameris and Pseudopentameris (Arundinoideae, Poaceae)
revisited 134
BRUSSE, F. A new species of Inezia (Anthemideae) from the north-eastern Transvaal (Asteraceae) ... 27
BRUSSE, F. A new Phymaspermum (Anthemideae) species from dolomite areas of the Wolkberg
(Asteraceae) 29
BRUSSE, F. A new species of Fuscidea (Lichenes) from the Cape Fold Mountains (Fuscideaceae) .... 35
BRUSSE, F. A new species of Maronea (Lichenes) from the Drakensberg (Fuscideaceae) 36
BURROWS, J.E. New taxa, combinations and records of Pteridophyta from southern and central Africa 167
CARR, J.D. Review: Trees and shrubs of the Witwatersrand, Magaliesberg and Pilanesberg, by J. van Gogh
and J.M. Anderson 319
CARR, J.D. & RETIEF, E. A new species of Combretum from Natal (Combretaceae) 38
DEALL, G.B. & BACKER, A.P The vegetation ecology of the Eastern Transvaal Escarpment in the Sabie
area. 3. Annotated checklist 91
DjtALL, G.B. , SCHEEPERS, J.C. & SCHUTZ, C.J. The vegetation ecology of the Eastern Transvaal
Escarpment in the Sabie area. 1. Physical environment 53
DEALL, G.B., THERON, G.K. & WESTFALL, R.H. The vegetation ecology of the Eastern Transvaal
Escarpment in the Sabie area. 2. Floristic classification 69
DEALL, G.B. & WESTFALL, R.H. Improving the resolution of floristic/habitat pattern correlations on
phytosociological tables 263
DE WET, B.C., GIBBS RUSSELL, G.E., GERMISHUIZEN, G., SCHRJRE, B.D., JORDAAN, M.,
PIENAAR, B.J., WELMAN, W.G., REID, C., VAN WYK, C.M., FISH, L., IMMELMAN, K.L.,
VAN ROOY, J., GLEN, H.F. & BARKER, N.P New taxa, new records and name changes for southern
African plants 275
ELLIS, R.P. Leaf anatomy of the South African Danthonieae (Poaceae). XVIII. Centropodia mossamedensis 41
ELLIS, R.P Leaf anatomy of the South African Danthonieae (Poaceae). XIX. The genus Prionanthium 217
FOURIE, D.M.C. Obituary: Inez Clare Verdoorn (1896-1989) 313
GERMISHUIZEN, G. Oxygonum altissimum, a new species from central Somalia (Polygonaceae) 210
GERMISHUIZEN, G. , KOK, P.D.F. & ROBBERTSE, PJ. Polygonum hydropiper in southern Africa
(Polygonaceae) 211
GERMISHUIZEN, G., ROBBERTSE, PJ. & KOK., P.D.F. The genera Polygonum and Bilderdykia
(Polygonaceae) in southern Africa: morphology and taxonomic value of the ocrea and fruit 175
GIBBS RUSSELL, G.E. & ELLIS, R.P Taxonomy and leaf anatomy of the genus Ehrharta (Poaceae) in
southern Africa: the Ramosa group 189
HARTMANN, H.E.K. & DEHN, M. A re-examination of the genus Amphibolia (Mesembryanthemaceae) 179
HENDERSON, L. Invasive alien woody plants of Natal and the north-eastern Orange Free State 237
HUTCHINGS, ANNE. A survey and analysis of traditional medicinal plants as used by the Zulu, Xhosa
and Sotho Ill
HUTCHINGS, A. Observations on plant usage in Xhosa and Zulu medicine 225
IMMELMAN, K.L. Studies in the southern African species of Justicia and Siphonoglossa (Acanthaceae):
palynology 151
IMMELMAN, K.L. Siphonoglossa and Aulojusticia in southern Africa (Acanthaceae) 209
JOHNSON, C.T. & HUTCHINGS, A. A contribution to the pteridophyte flora of Transkei 183
KILLICK, D.J.B. Two nomenclatural problems involving Article 63 133
KILLICK, D.J.B. Review: Southern African botanical literature 1600—1988 SABLIT, compiled by A.S.
Kerkham 141
PANAGOS, M.D. Review: Combretaceae in southern Africa, by J.D. Carr 141
PANAGOS, M.D., BRITZ, PJ. & WESTFALL, R.H. Plant collecting apparatus for taxonomic and ecological
studies. 5. A gas drier for field drying of plant specimens 270
PANAGOS, M.D. & WESTFALL, R.H. Plant collecting apparatus for taxonomic and ecological studies.
1. A lightweight plastic plant press for on-site specimen pressing 266
PANAGOS, M.D. & WESTFALL, R.H. Plant collecting apparatus for taxonomic and ecological studies.
4. Drier-transporters for plant presses 269
PECKHAM, G.D. & VAN JAARSVELD, F.A. New botanical perspectives on the origin of the Raphia
palms at Mtunzini 213
PEROLD, S.M. Studies in the genus Riccia (Marchantiales) from southern Africa. 11. Riccia montana and
R. alboporosa, a further two new white-scaled species of the group ‘Squamatae’ 9
PEROLD, S.M. Studies in the genus Riccia (Marchantiales) from southern Africa. 12. Riccia albolimbata
and the status of R. albosquamata, white-scaled species originally described by Arnell 17
PEROLD, S.M. Studies in the genus Riccia (Marchantiales) from southern Africa. 13. A new species, R.
hantamensis, in section Pilifer and a new record for R. alatospora 157
PEROLD, S.M. Studies in the genus Riccia (Marchantiales) from southern Africa. 14. R. concava section
Pilifer 161
POYNTON, R.J. Review: Flora of Australia (Vol. 19), Myrtaceae — Eucalyptus, Angophora, by G.M. Chip-
pendale 319
SANCHEZ, EVANGELINA, ARRIAGA, MIRTA O. & ELLIS, ROGER P. Kranz distinctive cells in the
culm of Arundinella (Arundinelleae; Panicoideae; Poaceae) 45
SCHRIRE, B.D. Obituary: Rudolf Georg Strey (1907-1988) 137
SPIES, J.J., SAAYMAN, E.J.L., VOGES, S.P. & DAVIDSE, G. Chromosome studies on African plants.
9. Chromosome numbers in Ehrharta (Poaceae: Ehrharteae) 125
VAN WYK, B-E. Studies in the genus Lotononis (Crotalarieae, Fabaceae). 2. Three new species of the
section Telina from the Cape Province 1
VAN WYK, B-E. Studies in the genus Lotononis (Crotalarieae, Fabaceae). 5. A new species of the L.
involucrata group (section Polylobium ) from the north-western Cape Province 7
VAN WYK, B-E. The identity of Lotononis elongata (Crotalarieae) 32
WELLS, M.J. Review: The gardener’s labyrinth, by Thomas Hill, ed. by Richard Mabey 320
WESTFALL, R.H. Plant collecting apparatus for taxonomic and ecological studies. 2. Coldat: a field data
capture program for collector’s data and herbarium labels 267
WESTFALL, R.H., BRITZ, P.J. & PANAGOS, M.D. Plant collecting apparatus for taxonomic and ecological
studies. 3. A new top-loading plant press for off-site specimen pressing 268
WESTFALL, R.H. & PANAGOS, M.D. Plant collecting apparatus for taxonomic and ecological studies.
6. A transportable map cabinet for vehicle and office use 272
WESTFALL, R.H., PANAGOS, M.D. & VAN STADEN, J.M. Plant collecting apparatus for taxonomic
and ecological studies. 7. A transportable camping kitchen for vehicle use 273
WILLIAMS, R. Turraea pulchella rediscovered (Meliaceae) 31
Bothalia 19,1: 1-5(1989)
Studies in the genus Lotononis (Crotalarieae, Fabaceae). 2. Three
new species of the section Telina from the Cape Province
B-E. VAN WYK*
Keywords: Cape Province, Fabaceae, Lotononis section Telina, new taxa
ABSTRACT
Three new species of the section Telina (E. Mey.) Benth. of Lotononis (DC.) Eckl. & Zeyh. are described; L.
azureoides B-E. van Wyk, L. gracilifolia B-E. van Wyk and L. lamprifolia B-E. van Wyk. These species appear to be
very rare and are known from only a few localities in marginal fynbos areas of the south-western and southern Cape.
UITTREKSEL
Drie nuwe soorte van die seksie Telina (E. Mey.) Benth. van Lotononis (DC.) Eckl. & Zeyh. word beskryf: L.
azureoides B-E. van Wyk, L. gracilifolia B-E. van Wyk en L. lamprifolia B-E. van Wyk. Hierdie soorte skyn baie
skaars te wees en is bekend van slegs enkele lokaliteite in marginale fynbosgebiede van die Suidwes- en Suid-Kaap.
INTRODUCTION
The section Telina (E. Mey.) Benth. of Lotononis
(DC.) Eckl. & Zeyh. comprises a group of prostrate or
procumbent suffrutices easily recognized by their large,
usually solitary flowers that are borne on long slender
peduncles. The very large standard petal is a particularly
useful diagnostic character.
Meyer (1836) included this group as one of three sec-
tions in his genus Telina, namely the section Chasmo-
neae. When Bentham (1843) changed the status of
Telina to a section of Lotononis, he excluded the sec-
tions Cytisoides and Brachypodae and referred both of
these to the section Krebsia (Eckl. & Zeyh.) Benth.
Chasmoneae E. Mey. and Telina (E. Mey.) Benth. are
therefore synonymous. Most of the species which Eck-
lon & Zeyher (1836) included in their concept of Lotono-
nis belong to this section.
The section as circumscribed by Diimmer (1913) is
not a natural group. Polhill (1973) and Van Wyk (1987)
respectively, transferred Lotononis bracteata Benth. to
Pear sonia Diimmer and L. magnistipulata Diimmer to
Argyrolobium Eckl. & Zeyh. Some more changes are
necessary, but these will be motivated elsewhere (Van
Wyk in prep.). My own concept of the group does not
include L. minor Diimmer & Jennings, L. macrocarpa
Eckl. & Zeyh., L. solitudinis Diimmer and L. marlothii
Engl. It does however include the three new species that
are described below.
Lotononis azureoides B-E. van Wyk, sp. nov., L.
azureae Eckl. & Zeyh. similis, sed habitu denso ramo-
sissimo, foliolis conduplicatis valde recurvatis, stipulis
anguste lanceolatis, inflorescentiis subterminalibus (in
speciebus omnibus aliis Telinae foliis oppositis) atque
vexillo pro ratione parvo carinam aequanti (vexillum
quam carina valde longius in speciebus omnibus aliis
Telinae ) differt.
TYPE. — Cape, 3222 (Beaufort West): Beaufort West,
Karoo National Park, at Blouput on steep rocky and
* Department of Botany, Rand Afrikaans University, P.O. Box 524,
Johannesburg 2000.
MS. received: 1988.03.27.
bouldery sandstone slope (— BC), 2.11.1984, Bengis
442 (PRE, charta 1 , holo.; PRE, charta 2, iso.).
Procumbent, densely branched shrublet, ± 0,3 m
wide. Branches smooth, glabrous; twigs densely strigil-
lose, conspicuously white at the ends. Leaves digitately
trifoliolate; petiole (2-) 3-4 (-7) mm long; leaflets
conduplicate, strongly recurved, obovate, (2-) 4-6
(—8) x (1—) 2—3 (—5) mm, abaxially strigillose,
adaxially glabrous. Stipules consistently present, single
at each node, elliptic-oblong to narrowly lanceolate,
2—3 (—5) mm long. Inflorescences terminal or subter-
minal on lateral branches, slender, long-pedunculate,
25-40 mm long, invariably single-flowered; bracts
small, up to 1 mm long; bracteoles absent. Flowers
12- 14 mm long, deep blue; pedicel 2—3 mm long. Ca-
lyx 7-8 mm long, with the upper and lateral lobes on
either side fused higher up in pairs, minutely strigillose.
Standard broadly ovate to orbicular, 11-13 mm long,
with a line of hairs dorsaily along the middle, deep blue
with yellow at the base. Wing petals oblanceolate,
slightly longer than the keel; apex obliquely truncate;
sculpturing upper basal and upper left central, in 4 rows
of inter- and mtracostal lunae. Keel petals semicircular,
obtuse, auriculate and pocketed near the base. Anthers
dimorphic. Pistil 12—14 mm long; ovary linear, 8—10
mm long. Fruit (slightly immature) 20 x 4-5 mm,
laterally much inflated, lower suture sunken, upper
suture verrucose, surface wrinkled, glabrous. Seed
unknown. Figure 1.
This species is similar to L. azurea Eckl. & Zeyh. but
differs in the dense and much branched habit, the con-
duplicate and strongly recurved leaflets, the narrowly
lanceolate stipules, die subterminal inflorescences (leaf-
opposed in all other species of Telina ) and the relatively
small standard petal, which is as long as the keel (stan-
dard petal much longer than the keel in all other species
of Telina ). The flower structure is reminiscent or some
species in the section Polylobium (Eckl. & Zeyh.)
Benth., but the slender single-flowered inflorescence
and the shape of the calyx and wing petals are typical of
other species in the section Telina.
L. azureoides is known only from two collections
from the Nuweveld Mountains at Beaufort West (Figure
2). It was found on a rocky south-eastern slope in grassy
dwarf shrubland at an altitude of 1 600 m. The specific
2
Bothalia 19,1 (1989)
FIGURE 1. — Lotononis azureoides.
A, flowering branch, showing
the slender peduncles and con-
duplicate, recurved leaflets; Bl,
B2, leaves with leaflets opened
out, showing the single stipule
and sparsely strigillose vesti-
ture: Bl, adaxial view, B2,
abaxial view; C, calyx opened
out, with the upper lobes to the
left, vestiture not shown; D,
standard petal; E, wing petal; F,
keel petal; G, pistil; H, fruit
(slightly immature), showing
the wrinkled surface and verru-
cose upper suture; I, androe-
cium; J, bract (all from Bengis
442). Scales in mm.
FIGURE 2.- The known geographical distribution of Lotononis azureoides. ^ ; L. gracilifolia, and L. lamprifolia, ®
Bothalia 19,1 (1989)
3
epithet suggests a similarity with L. azurea Eckl. &
Zeyh. which it superficially resembles, but the affinities
of the species are not clear. More material is needed for a
detailed study. Available evidence indicates that the
Telina- type flower and inflorescence have evolved more
than once from different lines in the sections Aulacinthus
(E. Mey.) Benth. and Polylobium. The distinction
between the latter two sections and indeed also the sec-
tion Telina is becoming more and more obscured. Some
newly discovered species such as L. azureoides have
combinations of those characters previously used diag-
nostically for each of the three sections.
CAPE. — 3222 (Beaufort West): Nuweveld Mountain, Mountain
view, roadside near FM tower (-AB), D. Shearing 2.2.85 (private
herbarium on the farm Layton, Fraserburg District); Beaufort West,
Karoo National Park, at Blouput on steep rocky and bouldery
sandstone slope ( — BC), 2.11.1984, Bengis 442 (PRE, holo.; PRE,
iso.).
Lotononis gracilifolia B-E. van Wyk, sp. nov., L.
argenteae Eckl. & Zeyh. et speciebus aliis sectionis
Telinae similis, sed distinctissima foliolis gracilibus
acicularibus, stipulis inconspicuis, lobis calycis subula-
tis. L. dissitinodi B-E. van Wyk (sectionis Aulacin-
thus ) valde similis sed habitu nano suffrutescente, folio-
lis valde angustioribus, stipulis inconspicuis, pedunculis
longioribus et lobis calycis angustissimis differt.
TYPE. — Cape, 3320 (Montagu): Laingsburg District,
Tweedside (-AB), 27.9.1951, Barker 7482 (NBG,
charta2, holo.; NBG, charta 1, iso.).
Dwarf suffrutescent perennial up to 0,15 m tall.
Branches procumbent from a central rootstock; sparsely
branched and sparsely leafy; young twigs densely silky-
sericeous. Leaves digitately trifoliolate, very slender,
densely silky-sericeous; petiole up to 22 mm long; leaf-
lets acicular, variable in length, (2-) 7— 15 (-25) mm
long, slightly conduplicate, almost terete. Stipules
inconspicuous, caducous, rarely present, up to 0,5 mm
long when present. Inflorescences subterminal or leaf-
opposed on short lateral branches, 1 -flowered, rarely 2-
flowered; peduncle slender, variable in length, (3—)
20—50 (—60) mm long; bract small, oblong, up to 2 mm
long; bracteoles absent. Flowers large, up to 18 mm
long, yellow; pedicel 2—4 mm long. Calyx 10—13 mm
long; lobes long, subulate, with the lateral ones on either
side fused higher up in pairs, densely sericeous. Stan-
dard very large, broadly ovate to orbicular, 18—20 mm
FIGURE 3. — Lotononis gracilifolia.
A, habit, showing the pro-
cumbent flowering branches,
acicular leaves and slender
peduncles; Bl, leaf in adaxial
view, with leaflets opened out,
B2, B3, leaves in abaxial view,
showing variation in size, leaf-
lets not opened out; C, calyx
opened out with upper lobes to
the left, showing fusion of the
lateral lobes; D, standard petal;
E, wing petal; F, keel petal; G,
pistil; H, androecium; II, 12,
bracts; J, petiole base with vesti-
ture partly removed to show the
single, inconspicuous stipule
(all from Barker 7482 ). Scales
in mm.
4
Bothalia 19,1 (1989)
long, with hairs dorsally along the middle. Wing petals
as long or slightly longer than the keel, oblanceolate, ±
15 mm long; apex obliquely obtuse; sculpturing upper
basal, in 3—4 rows of mostly intercostal lunae. Keel
petals semicircular, obtuse, auriculate and pocketed
near base. Anthers dimorphic. Pistil long; ovary linear,
± 10 mm long. Fruit and seed unknown. Figure 3.
This species is similar to L. argentea Eckl. & Zeyh.
and other species of the section Telina, but it is very
distinct in the acicular leaflets, the inconspicuous
stipules and the subulate calyx lobes. It is remarkably
similar to L. dissitinodis B-E. van Wyk (section Aula-
cinthus ), but differs from this species in the dwarf and
suffrutescent habit, the much narrower leaflets, the
inconspicuous stipules, the longer peduncles and the
very narrow calyx lobes.
L. gracilifolia appears to be a very rare species and is
only known from the type collection (Figure 2). I have so
far been unable to locate the plant. The slender, acicular
leaves are a useful diagnostic character and the species is
unlikely to be confused with any other. This very dis-
tinctive character has suggested the specific epithet.
CAPE. — 3320 (Montagu): Tweedside, Laingsburg (-AB),
27.9. 195 1 , Barker 7482 (NBG, holo. ; NBG, iso.).
Lotononis lamprifolia B-E. van Wyk, sp. nov. , L.
argenteae valde affinis, sed habitu diffusiore, stipulis
obovatis foliolis valdc similibus (in L. argentea lineari-
bus vel lanceolatis), floribus flavis (in L. argentea cae-
ruleis) et vexillo depresso ovato (in L. argentea late
ovato vel orbiculato; etiam L. acuminatae Eckl. &
Zeyh. similis, sed ab hac specie indumento dense sericeo
et petiolis valde longioribus differt.
TYPE. — Cape, 3321 (Ladismith): Riversdale and
Swellendam Districts, between Muiskraal and
Lemoenshoek (— CC), 6.8.1951, Barker 7374 (NBG,
holo.).
Dwarf suffrutescent perennial ± 0,2 m tall. Branches
sparse, woody at the base; young twigs densely seri-
ceous. Leaves digitately trifoliolate, densely and silvery
sericeous; petiole relatively thick, variable in length,
3—4 (—10) mm long, leaflets oblanceolate to obovate,
(3-) 4-5 (-8) mm long, slightly conduplicate, thick in
texture. Stipules present or absent; when present closely
resembling the leaflets in shape, size and vestiture,
usually single to each node, strongly persistent. Inflores-
cences leaf-opposed on short lateral branches, in-
variably single-flowered; peduncle slender, 18-35 mm
FIGURE 4. — Lotononis lamprifolia.
A, habit; Bl, B2, B3, leaves:
Bl, abaxial view, showing the
single stipule and sericeous ves-
titure; B2, adaxial view, show-
ing paired stipules; B3, adaxial
view, stipules absent; C, calyx
opened out, upper lobes to the
left, showing the very slight
fusion of the lateral lobes; D,
standard petal (note the shape);
E, wing petal; F, keel petal; G,
pistil; H, androecium; II, 12,
bracts (all from Barker 7374 ).
Scales in mm.
Bothalia 19,1 (1989)
5
long; bracts small, oblong, ± 1 mm long; bracteoles
absent. Flowers 10—12 mm long, yellow; pedicel 1—2
mm long. Calyx 6-7 mm long; lobes triangular, with
the lateral ones on either side fused only slightly higher
up in pairs, densely sericeous. Standard large, broadly
depressed ovate, ± 12 mm long, with hairs dorsally
along the middle. Wing petals longer than the keel,
broadly oblanceolate, ± 10 mm long; apex obtuse,
rounded; sculpturing upper basal and upper left central,
in 4— 5 rows of mostly intercostal lunae. Keel petals
semicircular, obtuse, auriculate and pocketed near base.
Anthers dimorphic. Pistil short; ovary linear, ± 7 mm
long. Fruit and seed unknown. Figure 4.
This species is closely related to L. argentea Eckl. &
Zeyh. but differs in the more diffuse habit, the obovate
stipules which closely resemble the leaflets (linear to
lanceolate in L. argentea ), the yellow flowers (blue in
L. argentea ) and the depressed ovate standard (widely
ovate to orbicular in L. argentea ). It is also similar to L.
acuminata Eckl. & Zeyh. but differs from this species
in the densely sericeous vestiture and the much longer
petioles.
L. lamprifolia is only known from a single specimen
collected along the northern foothills of the Langeberg
near Barrydale (Figure 2). The shining and silky appear-
ance of the leaflets is very distinct (as in L. argentea ),
hence the specific epithet. The latter species is also very
poorly represented in southern African herbaria and col-
lectors are requested to look out for more complete mate-
rial. Fruiting material in particular is required for a more
detailed study of the full range of variation in these two
closely related species and to verify the diagnostic
features of the new species.
CAPE. — 3321 (Ladismith): Riversdale and Swellen-
dam Districts, between Muiskraal and Lemoenshoek
(-CC), 6.8. 1951 , Barker 7374 (NBG, holo.)
ACKNOWLEDGEMENTS
I wish to thank Dr H. F. Glen (Botanical Research
Institute, Pretoria) for the Latin translations and the
Directors and staff of the cited herbaria for the loan of
specimens. The taxonomic study of Lotononis is a regis-
tered Ph. D. project at the University of Cape Town.
REFERENCES
BENTHAM, G. 1843. Enumeration of Leguminosae, indigenous to
southern Asia, and central and southern Africa. Hooker’s, London
Journal of Botany 2:504—613.
DUMMER, R. A. 1913. A synopsis of the species of Lotononis, Eckl.
& Zeyh. , and Pleiospora Harv. Transactions of the Royal Society
of South Africa 3:275—335.
ECKLON, C. F. & ZEYHER, C. 1836. Enumeratio plantarum Africae
Australis extratropicae 2: 176-178. Perthes & Besser, Ham-
burg.
MEYER, E. H. F. 1836. Commentariorum de plantis Africae Austra-
lians 1,1: 67 —70. Regiomonti, Leipzig.
POLHILL, R. M. 1973. A revision of Pearsonia (Leguminosae -Papi-
lionoideae). Kew Bulletin 29: 383—410.
VAN WYK, B-E. 1987. Taxonomic notes on Argyrolobium variopile
(Fabaceae), and the status of Lotononis magnistipulata. South
African Journal of Botany 53: 395—397.
Bothalia 19,1:7-8(1989)
Studies in the genus Lotononis (Crotalarieae, Fabaceae). 5. A new
species of the L. involucrata group (section Polylobium) from the
north-western Cape Province
B-E. VAN WYK*
Keywords: Cape Province, Fabaceae, Lotononis section Polylobium, new taxon
ABSTRACT
A new species of the section Polylobium (Eckl. & Zeyh.) Benth. of Lotononis (DC.) Eckl. & Zeyh. is described,
namely L. racemiflora B-E. van Wyk. The species, known only from a single collection near Clanwilliam in the north-
western Cape, is closely related to L. involucrata (Berg.) Benth. and L. angustifolia (E. Mey.) Steud.
UITTREKSEL
’n Nuwe soort van die seksie Polylobium (Eckl. & Zeyh.) Benth. van Lotononis (DC.) Eckl. & Zeyh. word beskryf,
naamlik, L. racemiflora B-E. van Wyk. Die soort, wat slegs bekend is van ’n enkele versameling naby Clanwilliam in die
Noord-wes Kaap, is naverwant aan L. involucrata (Berg.) Benth. en L. angustifolia (E. Mey.) Steud.
INTRODUCTION
Lotononis involucrata (Berg.) Benth. and related
species differ from other species of the section Polylo-
bium (Eckl. & Zeyh.) Benth. in the subterranean caudex
from which flowering shoots develop annually. Stipules
are paired or absent and never single or markedly dimor-
phic as in other species. The umbellate inflorescence in
all the species which Diimmer (1913) included in Polylo-
bium was used as a diagnostic character for the section.
The new species described below is morphologically
intermediate between L. involucrata (Berg.) Benth. and
L. angustifolia (E. Mey.) Steud. and is obviously very
closely related to these species. However, the inflores-
cence is a true raceme. Inflorescence structure is there-
fore no longer a useful diagnostic character for distin-
guishing the section Polylobium. It may indeed be
argued that Polylobium sensu lato is an artificial group
which has resulted from the excessive weighting of inflo-
rescence structure as a diagnostic character.
Lotononis racemiflora B-E. van Wyk, sp. nov., L.
angustifoliae (E. Mey.) Steud. valde similis, sed inflo-
rescentia multiflora racemosa (in L. angustifolia pauci-
flora umbellata vel subumbellata), basi stipulae semicor-
dato, indumento densiore piloso differt; etiam L. involu-
crata (Berg.) Benth. similis, sed ab hac specie inflores-
centia racemosa, foliolorum amplitudine texturaque
(quam in L. involucrata maioribus coriaceoribus), stipu-
larum amplitudine formaque, vexillo alisque multo
maioribus differt.
TYPE. — Cape, 3218 (Clanwilliam): Clanwilliam Dis-
trict, Bokwater, W of Clanwilliam (— BB), 28.10.1948,
Acocks 15171 (PRE, holo.; K, iso.).
Suffrutescent herb, with annual flowering branches
* Department of Botany, Rand Afrikaans University, P.O. Box 524,
Johannesburg 2000.
MS. received: 1988.05.25.
from a woody root. Branches procumbent, up to 0,3 m
long, thick and rigid, densely leafy, more sparsely so
towards the base, densely pilose. Leaves digitately tri-
foliolate, densely and softly pilose; petiole (3—) 8-12
(—14) mm long; leaflets linear to narrowly elliptical,
(4-) 10—18 (—22) x 2—2,5 mm, with soft pilose hairs
on both surfaces. Stipules similar to the leaflets, consis-
tently present, paired at each node, ovate to lanceolate,
(3—) 8 — 10 ( — 12) mm long; base distinctly semicordate;
apex obtuse to acute. Inflorescences terminal on primary
and secondary branches, racemose, lax, 90—140 mm
long, up to 18-flowered; peduncle 28-50 mm long,
densely and softly pilose; rachis up to 95 mm long,
bracts conspicuous, thickly textured, ovate-cymbiform,
up to 8 x 3 mm, pilose on both surfaces; bracteoles
absent. Flowers large, 14-16 mm long, yellow; pedicel
3-5 mm long. Calyx 12—14 mm long, lobes narrowly
acuminate, with the lateral ones on either side fused
higher up in pairs, lanately pilose. Standard large,
suborbicular, 15 mm long, longitudinally striated, base
broadly cordate. Wing petals broadly obovate, much
longer than the keel; apex obliquely rounded; sculpturing
upper basal and upper central, in 7-8 rows of mostly
intercostal lunae and lamellae. Keel petals small, semi-
circular, 11-12 mm long, auriculate and pocketed near
the base; apex acute. Anthers dimorphic. Pistil 11-12
mm long; ovary oblong-linear, 6-7 mm long, pub-
escent. Fruit (immature) oblong-linear, twice as long as
the calyx, much inflated laterally, sparsely pubescent,
with evenly spaced warty protuberances along the upper
suture. Seed unknown. Figure 1 .
L. racemiflora is closely related to L. angustifolia but
differs in the many-flowered racemose inflorescence
(few-flowered and umbellate or subumbellate in L. an-
gustifolia), the semicordate stipule base and the more
densely pilose vestiture. It is also similar to L. involu-
crata, but differs from this species in the racemose inflo-
rescence, the size and texture of the leaflets (larger and
more coriaceous than in L. involucrata), the size and
shape of the stipules and the much larger standard and
wing petals.
8
Bothalia 19,1 (1989)
FIGURE 1. — Lotononis racemiflora .
A, habit, showing a flowering
branch, the persistant root and
the long, racemose inflor-
escence. Bl, B2, B3, leaves and
stipules, showing the variation
in size and shape and the pilose
vestiture: Bl, abaxial view of a
mature leaf taken from a basal
node, B2, adaxial view (note the
vestiture), B3, abaxial view. C,
calyx opened out, with the up-
per lobes to the left, vestiture
not shown; D, standard petal; E,
wing petal; F, keel petal, show-
ing the small size and pointed
apex'; G, pistil; FI, immature
fruit in lateral view, showing the
verrucose upper suture; II, 12,
13, long anther, carinal anther
and short anther respectively; J,
bract. All from Acocks 15171 .
Scales in mm.
FIGURE 2.— The known geographical distribution of Lotononis race-
miflora.
The new species may be confused with L. angustifolia
and L. involucrata when not in flower, but the inflor-
escence structure is quite different. It is known from a
single collection near Clanwilliam in the north-western
Cape, where it was found on an old land in Fynbos-
Strandveld vegetation. Figure 2.
CAPE. — 3218 (Clanwilliam): Clanwilliam District, Bokwater, W of
Clanwilliam ( -BB), 28. 10. 1948, Acocks 15171 (PRE, holo.; K. iso.).
ACKNOWLEDGEMENTS
I wish to thank Dr H. F. Glen (Botanical Research
Institute, Pretoria) for the Latin translation. The taxo-
nomic study of Lotononis is a registered Ph. D. project at
the University of Cape Town.
REFERENCE
DUMMER, R. A. 1913. A synopsis of the species of Lotononis , Eckl.
& Zeyh. , and Pleiospora Harv. Transactions of the Royal Society
of South Africa 3: 275-335.
Bothalia 19,1:9-16(1989)
Studies in the genus Riccia (Marchantiales) from southern Africa. 11.
Riccia montana and R. alboporosa, a further two new white-scaled
species of the group ‘Squamatae’
S. M. PEROLD*
Keywords: air pores, anatomy, calcium salts, distribution, endemic, rare, reticulation, scales, spore ornamentation
ABSTRACT
Another two new endemic species of the subgenus Riccia, section Riccia. group 'Squamatae', are described: R.
montana and R. alboporosa. The distribution of R. montana is apparently restricted to high altitudes in the Drakensberg
and Witteberg Mountain ranges. The species is characterized by ligulate branches, finely spongy dorsal surface and hyaline
to white, calcium-encrusted scales. R. alboporosa is found in Namaqualand, but it is rare. It can be recognized by the
distinctly porous appearance of the dorsal surface due to the presence of large, ± regularly spaced air pores, which are
encircled by six or seven radially arranged, wedge-shaped cells that become white on drying, hence the specific epithet.
UITTREKSEL
Nog twee nuwe endemiese spesies van die subgenus Riccia. seksie Riccia. groep 'Squamatae', word beskryf: R.
montana en R. alboporosa. Die verspreiding van R montana is blykbaar beperk tot hooggelee gebiede in die Drakens-
berg- en Witteberggebiede. Die spesie word gekenmerk deur lintvormige vertakkings, fyn, sponserige dorsale oppervlak en
hialiene tot wit, kalkbedekte skubbe. R. alboporosa kom voor in Namakwaland maardit is skaars. Dit kan erken word aan
die duidelik poreuse voorkoms van die dorsale oppervlak, veroorsaak deur groot, ± reelmatig verspreide lugporiee wat
omsluit word deur 'n ring van ses of sewe radiaal gerangskikte, wig-vormige selle wat wit word met uitdroging, vandaar die
spesifieke epiteton.
1 . Riccia montana Per old, sp. nov.
Thallus dioicus, perennis; subviridis vel viridis, in
sicco albidus vel flavidus; subtiliter spongiosus; margini-
bus reflexis vel inflexis saepe labia duo secus ramos
formantibus. Frons usque ad 8 mm longa, (1,5—)
1,7— 2,0 (—2,5) mm lata, 0,6—0,75 mm crassa, 2,5-3-
plo latiora quam crassa, lobis ligulatis, dorsaliter pro-
funde sulcatis. Squamae apicem versus undulatae, mox
ad latera appressae, vix margines thalli superantes,
hyalinae vel calcii depositis tectae. Sporae 70,0—85,0
pm diametro, alis ± 5 pm latis, grosse reticulatae, 7-8
areolis trans diametrum, parietibus areolarum crassis.
Chromosomatum numerus n = 9.
TYPE. — Cape Province, 3027 (Lady Grey): Witte-
berg Mountains, basalt cliffs at top of Jouberts Pass, 10
km E of Lady Grey, eastern aspect, alpine heath-grass-
land (— CB), Van Rooy 2712 (PRE, holo.), with Bryum
alpinum Huds. ex With.
Thallus dioicous (Figure 1A, B), perennial, gre-
garious, not in rosettes, medium-sized; branches apically
symmetrically or asymmetrically furcate, frequently
with short lateral branching more proximally, medium to
widely divergent, up to 8 mm long, segments 3—5 mm
long, (1,5— )1, 7— 2,0(— 2,5) mm wide, 0,6-0,75 mm
thick, i.e. ± 214 to 3 times wider than thick, ligulate,
apex rounded to subacute, emarginate, deeply grooved
on dorsal surface (Figure 2A), proximally ± flat to
slightly concave (Figure 1D1-6); dorsally light green to
* Botanical Research Institute, Department of Agriculture and Water
Supply, Private Bag X 101 , Pretoria 000 1 , RSA.
Ms. received: 1988.04.24.
green, finely spongy and glistening; margins acute,
flanks almost vertical distally to sloping somewhat obli-
quely outward and upward proximally, green; ventral
surface rounded, green; when dry, dorsally white to yel-
lowish, margins indexed or more usually re flexed along
the edges (Figure 1C), generally forming two lips in
proximal parts of branches. Anatomy: dorsal epithelium
unistratose, hyaline, bulging upper walls of cells covered
with fine deposit of calcium salts (Figure 2C), 20-30 x
35x50 pm, cell width somewhat irregular, sometimes
single cells spanning l)4-2(-3) subdorsal cells (Figure
IE), soon collapsing; air pores often only partly aligned
with air canals below (Figures IF, 2C), large, 20-45
pm across, wider towards thallus margins, (3-)
4-5(-6)-sided (Figure 2D); assimilation tissue (chlo-
renchyma) consisting of vertical columns of 6- 10 cells,
± 50-65 x (37 — )42 — 50 jLtm, air canals in between
assimilation cell columns ± 50 (-65) pm wide (Figure
IE, G); storage tissue occupying lower 14 or more of the
thickness of thallus, cells tightly packed. Rhizoids hya-
line, some smooth, others tuberculate, up to 25 pm
wide, arising from ventral epidermis of thallus and base
of scales. Scales wavy at apex, soon appressed to
flanks, imbricate, hyaline or whitened with calcium de-
posits, sometimes flecked with red toward base, hardly
exceeding thallus margins (Figures 1H; 2B), 850 x 500
pm, cells in body of scale 4-6-sided, 50-85(-90) x
± 40 pm, smaller at margin, ± 30 x 50 pm, cell walls
mostly bulging. Antheridia with hyaline or white necks
± 160 pm long, projecting from small, shallow pits on
either side of dorsal groove (Figure 1A). Archegonia
purple-necked, scattered along groove in female plants.
Sporangia proximal, single or 2 adjacent, each con-
taining about 450 spores, large, ± 750 /Ltm wide, bulg-
ing conspicuously dorsally (Figure IB), overlying thal-
10
Bothalia 19,1 (1989)
FIGURE I Riccia moniana. Structure of thalius and scales. A, fresh male thallus; B, fresh female thallus with sporangia; C, dry female
thallus; D1 -6, transverse sections of thallus branch at different distances from apex to basal parts; E, transverse section through dorsal
epithelium and chlorenchyma; F, epithelial cells (solid lines), and air pores (hatched lines) from above, with subdorsal cells (broken lines)
and air canals (stippled); G, horizontal section through chlorenchyma, with air canals stippled; H, scale. A-C, Van Rooy 3046\ D1 -6,
Van Rooy 2712 ; E. Oliver 8354, F ,G, Perold 3 /; H, Van Rooy 2718. A-H drawn by J. Kimpton. Scale baron A-D = 1 mm’ E-G =
50 fim: H = 100 gm.
Bothalia 19,1 (1989)
II
FIGURE 2. — Riccia montana. Struc-
ture of thallus and cells. A, dor-
sal view of thallus, apically
grooved; B, scales at apical
margin; C, intact dorsal cells
with calcium deposits, air pores;
C, collapsed dorsal cells with
wide air pores. A-D, Oliver
8354, SEM micrographs. Scale
bar = 50 gm. All SEM and LM
micrographs by S. M. Perold.
FIGURE 3. — Riccia montana.
Spores. A, proximal face; B,
proximal face viewed from side;
C, distal face; D, distal face,
side view; E, areolae, wing and
pore on distal face, much
enlarged; F, distal face. A, C,
D, F, Van Rooy 2712\ B, Glen
1728', E, Van Rooy 2718. A-E,
SEM micrographs; F, LM
micrograph. Scale bar - 50
Atm; diameter of spore on F = ±
100 fjLm.
12
Bothalia 19,1 (1989)
lus tissue not turning white, but shrinking and disinte-
grating. Spores (70,0— )75, 0—80, 0(— 85,0) pm in diam-
eter, brown, semi-transparent, triangular-globular,
polar, wing ± 5 /u,m wide, wider at perforated angles,
margin somewhat wavy (Figure 3 A, C), finely eroded,
crenulate, ornamentation on both faces completely or
incompletely coarsely reticulate; distal face with 7 or 8
rounded to angular areolae across diameter, ±7,5 pm
wide (Figure 3C, F), walls thick and high, ± 5 pm tall,
extending partly onto wing, with raised papillae at areo-
lar nodes; proximal face with triradiate mark distinct to
less clearly defined, areolae often incomplete, irregu-
larly ridged (Figure 3B), or with complete, angular areo-
lae, raised at nodes (Figure 3A). Chromosome number n
= 9 (T. Bomefeld pers. comm.).
Riccia montana grows on black, humus-rich soil,
overlying rocky outcrops, often near seepages, and in
association with moss species e.g. Bryum alpinum Huds.
ex With, and with other Riccia spp.
In the white-scaled species of the group ‘Squamatae’,
section Riccia, subgenus Riccia, R. montana and R.
argenteolimbata (Volk et al. 1988) are the only two
dioicous species. R. montana differs from the other Ric-
cia species in this group by the finely spongy texture of
the dorsal surface of the thallus, by the ligulate branches
with lip-like, re flexed margins along the proximal parts
when dry, by the coarsely reticulate, polar spores and by
its distribution which is restricted to the eastern mountain
ranges. It is somewhat similar to the other white-scaled
species, R. albolimbata S. Amell, R. albornata Volk
& Perold, R. argenteolimbata Volk & Perold, and R.
alboporosa sp. nov. in the group ‘Squamatae’, but it
never turns chalk- white over the sporangia as does R.
albolimbata. R. albolimbata and R. albornata gener-
ally have much larger, wavy, hyaline (or white) scales
that extend above the thallus margins and broad, not
ligulate branches. The ornamentation of their spores is
also markedly different. R. argenteolimbata Volk &
Perold, has a more compact thallus, stiff white scales
and apolar spores, whereas R. alboporosa has a more
coarsely spongy thallus, is dorsally puffy when dry and
thickly covered with calcium salts, has inconspicuous
scales and more finely reticulate polar spores. R. mon-
tana was originally recognized as a new species by the
late Prof. E. A. C. L. E. Schelpe of BOL, from one of
his gatherings at the Sentinel, Drakensberg in 1946.
Unfortunately the spores were not fully mature, but re-
cently, more collections of the same species with mature
sporangia have been made, also in mountainous regions
2 000— 3 000 m above sea level.
The specific epithet, montana, refers to the moun-
tainous localities, where all the collections have so far
been made. Its distribution area does not appear to over-
lap with that of any of the other white-scaled species,
being confined to high altitudes in the Drakensberg of
Natal and Lesotho and the Witteberg Mountains of the
north-eastern Cape Province (Figure 4).
SPECIMENS EXAMINED
NATAL. — 2828 (Bethlehem): Drakensberg, Sentinel, 9 500 It
(— CB), Schelpe s.n. (BOL).
LESOTHO 2828 (Bethlehem): 6 km from Oxbow Lodge to
Mokhotlong, basalt outcrops (-DC), Van Rooy 3045, 3046 (PRE).
2927 (Maseru): Blue Mountain Pass, 19 km from Maseru on road to
FIGURE 4. — Map showing distribution of R. montana, 9 and R.
alboporosa, H in southern Africa.
Thaba Tseka, calcrete soil on seepage bank ( — BC), J . M. Perold 31
(PRE). 2929 (Underberg): Sani River bank, ± 7 km from Sani Top,
along road to Mokhotlong (— CA), Van Rooy 3446 (PRE); Sani Top,
mountain slopes W of Border Post (— CB), Van Rooy, 3540, 3566
(PRE); Sani flats, 2 km from Sani Border Post to Mokhotlong (-CB),
Van Rooy 3702 (PRE); Sehlabathebe Nat. Park, on black gritty soil
bordering stream, 2 300—2 500 m ( — CC), Jacot Guillarmod, Getliffe
&Mzamane60 (PRE).
CAPE. — 3027 (Lady Grey): Witteberg Mountains, basalt cliffs at
top of Jouberts Pass, 10 km E of Lady Grey (— CB), Van Rooy 2718
(PRE); ibid. Van Rooy 2712 (see type); Ben MacDhui, above Rhodes,
wet loamy plateau S of main peak, very short grassland ( — DD), Oliver
8354 (PRE). 3028 (Matatiel): 20 km from Rhodes, up Naudesnek
towards Maclear(— CA/CC), Glen 1728 (PRE).
2. Riccia alboporosa Perold, sp. nov.
Thallus monoicus, perennis, in vivo flavido-viridis,
dense calcii depositis tectus, poribus magnis regularibus;
in sicco albidus, inflatus, marginibus solum apicaliter
inflexis. Frons usque ad 7,0 mm longa, l,8-3,5(-4,0)
mm lata, 0,8— 1,2 mm crassa, 2— 3-plo latior quam
crassa, obtuse cuneata vel late ovata, sulco dorsali lato.
Squamae inconspicuae, aegre ad margines thalli exten-
sae, calcio tectae. Sporae 75,0-88,0 pm diametro,
polares, ala ± 5 pm lata, tenue reticulata, 11 — 13 areolis
trans diametrum superficiei distalis. Chromosomatum
numerus n = 10.
TYPE.— Cape Province, 3119 (Calvinia): NE of
Nieuwoudtville, Groothoek, at Soetlandsfontein River
(—AD), Oliver 8854 (PRE, holo.), on sandy/clay flats
alongside river, in rock crevices and on ledges, asso-
ciated with Riccia albornata Volk & Perold and small
moss species, Bryum argenteum Hedw.
Thallus monoicous, perennial, gregarious or single,
not in rosettes; once (Figure 5A) or twice symmetrically
or occasionally asymmetrically furcate (Figure 5B),
branches medium to widely divergent, bluntly wedge-
shaped to broadly ovate (Figure 6A), up to 7 mm long,
1,8— 3,5 (—4,0) mm wide and 0,8 — 1,2 mm thick, i.e.
2-3 times wider than thick; apex rounded, emarginate;
dorsally bright yellowish green with ± regularly spaced,
large, conspicuous air pores encircled by lighter
coloured cells; apically deeply grooved by wide sulcus
(Figure 6B), proximally somewhat concave to nearly flat
(Figure 5D1 -5); ventrally green, gently rounded; flanks
Bothalia 19,1 (1989)
13
FIGURE 5. — Riccia alboporosa. Structure of thallusand scales. A, B, fresh thalli; C, dry thallus; Dl-5. transverse sections of thallus branch
at different distances from apex to basal parts: E, transverse section through dorsal epithelium and chlorenchyma; F. epithelial cells (solid
lines) and air pores (hatched) from above, with subdorsal cells (broken lines) and air canals (stippled); G, scale. A. B, E, S. M. Perold
1775: C, D, E, G, Oliver 8854. A-G drawn by J. Kimpton. Scale baron A-D = 1 mm; E. F. = 50/am:G = 100 /am.
14
Bothalia 19,1 (1989)
FIGURE 6. — Riccia alboporosa.
Structure of thallus and cells. A,
dorsal view of thallus; B, wide
dorsal groove at apex; C, scales
near apex; D, air pore with sur-
rounding cells heavily encrusted
with calcium crystals; E, air
pores and dorsal cells; F, dorsal
cells (lightly dusted with cal-
cium crystals) encircling air
pores, air canals and subdorsal
cells faintly visible beneath.
A-C, E, S. M. Perold 1775; D,
F, Oliver 8854. A-E, SEM
micrographs; F, LM micro-
graph. Scale bar = 50 /urn.
sloping obliquely outward and upward to raised blunt
margins (Figure 5D1-4); when dry, greenish white to
white, somewhat puffy, slightly concave, margins erect
to indexed (Figure 5C) or apically clasped together,
revealing regular white, appressed ventral scales.
Anatomy: dorsal epithelium unistratose, cells hyaline,
upper walls rounded near and in dorsal groove, but more
laterally soon collapsing (Figure 5E) and becoming
heavily encrusted with thick layer of calcium salts
(Figure 6D), generally 6 or 7 cells wedge-shaped from
above, 60-75 gm long ± 50 /im wide at broadest part,
radially arranged around each air pore (Figures 5F; 6E,
F), which leads to wider air canal below; assimilation
tissue (chlorenchyma) on section occupying about / the
thickness of the thallus, ± 400 /xm thick, topmost cells
generally somewhat thicker- walled, 20-25(-35) X
30—35 /Am, often 2 under each dorsal cell, soon losing
their chloroplasts as overlying cell layer collapses; air
canals 50—80 /xm wide (Figure 5D), enclosed by
columns of rectangular cells ± 50 x 37-45 /xm; storage
tissue occupying lower 'A of thickness of thallus, cells
50-75 /xm wide. Rhizoids hyaline, some smooth, others
tuberculate, up to ± 25 /xm wide, very long, arising
from ventral epidermal cells and base of scales. Scales
mostly inconspicuous, 550 x 350 /xm, hardly extending
to thallus margins, appressed, imbricate (Figure 6C),
rounded, smooth-edged, heavily encrusted with calcium
salts, especially on exposed distal parts of scales, cells
mostly 6-sided in body of scale, ± 85 x 37 /xm (Figure
5G), marginally smaller, brick-shaped. Antheridia with
hyaline necks, scattered along groove. Arche gonia with
purple necks. Sporangia single near base (Figure 5B),
or crowded in groups along middle of branch (Figure
5A), bulging dorsally, ± 800 /xm wide, containing ±
400—500 spores. Spores (75,0) 80,0-85,0 (-88,0)
/xm in diameter, yellow-brown, semi-transparent, trian-
gular-globular, polar (Figure 7A, E), wing ± 5 /xm
wide, slightly wider at perforated angles, margin mostly
smooth; ornamentation reticulate: distal face with 11-13
areolae across diameter (Figure 7B, C, F), ± 5 /xm
wide, toward centre somewhat larger and with thicker,
higher walls, ridges extending onto wing, radial ones
generally more pronounced than those across (Figure
7D), slightly raised at nodes; proximal face with trira-
diate mark ± distinct (Figure 7A, E), each facet with up
to ± 50 small, round areolae ± 3 /xm wide, some adja-
cent ones confluent, ridges low (Figure 7A, E). Chromo-
some number n = 10 (Bomefeld pers. comm.).
Bothalia 19,1 (1989)
15
FIGURE 7. — Riccia alboporosa.
Spores. A, E, proximal face; B,
C, F, distal face; D, enlarged
view of areolae and margin on
distal face. A-F, Oliver 8849.
A-D, SEM micrographs; E, F,
LM micrographs. Scale bar =
50 /am; diameter of spore on E,
F = ± 100 /am.
Riccia alboporosa is known from only two localities
in the drier area of the north-west Cape Province, with an
annual winter rainfall of less than 200 mm at altitudes of
750—850 m above sea level. It grows on fine yellow-
brown, sandy or slightly brackish soil overlying tillite
rocks, in association with other Riccia species, e.g. R.
albornata Volk & Perold and with the small moss spec-
ies Bryum argenteum Hedw. and with Aloina bifrons
(De Not.) Delgadillo.
It is easily recognized by a feature alluded to in the
specific epithet: the thallus has numerous large, widely
but ± regularly spaced air pores encircled by dorsal epi-
thelial cells, part of which form the roof over the air
canals and on drying, rapidly become white, before the
remainder of these cells, supported by subdorsal cells,
do. The dorsal surface becomes heavily encrusted with
calcium salts, which, in field-grown specimens, must be
removed with dilute hydrochloric acid, before the cell
outlines can be distinguished. As seen from above, each
hyaline dorsal cell is in contact with an air pore, which
necessitates placing this species in the subgenus Riccia
(Volk 1983), although the air canals are wider than is
generally encountered in this subgenus. It differs from
the other white-scaled species by the inconspicuousness
of its scales, by the puffy appearance of the dorsal sur-
face in the dry plant and by the finely reticulated spores.
SPECIMENS EXAMINED
CAPE. — 3119 (Calvinia): S of Loeriesfontein, Skietnes Kloof, E of
Slagberg, rocky ledges, facing South (-AB), Oliver 8849 (PRE); NE
of Nieuwoudtville, Groothoek, at Soetlandsfontein River (—AD), S.
M. Perold 1772, 1775, 2317 (PRE).
ACKNOWLEDGEMENTS
The author wishes to thank Dr habil. T. Bomefeld,
Am Reelein 1, D8706, Hochberg, Germany, for the
chromosome counts and Dr H. F. Glen, BRI, for trans-
lating the diagnoses. Sincere thanks are also due to: Prof,
(emer.) Dr O. H. Volk of Wurzburg University for
advice during preparation of the first draft of the text; the
late Prof. E. A. Schelpe for the loan of his specimen;
Messrs E. G. H. Oliver, Stellenbosch Botanical Unit,
J. van Rooy, BRI and J. Perold, Department of Agricul-
ture and Water Affairs, for collecting specimens, often
in difficult terrain.
16
Bothalia 19,1 (1989)
REFERENCES
ARNELL. S. 1957. Hepaticae collected in South West Africa by Prof.
Dr O. H. Volk. Mitteilungen a us der Botanischen Staatssam-
mlung, Miinchen 16: 262—272.
ARNELL. S. 1963. Hepaticae of South Africa, pp. 411. Swedish
National Scientific Research Council. Stockholm.
BORNEFELD, T. 1984. Chromosomenanalyse der Gattung Riccia von
Sud- und SW-Afrika und allgemeine Bemerkungen zur Zytogene-
tik der Lebermoose. Nova Hedwigia 40: 313—328.
PEROLD, S. M. 1989. Studies in the genus Riccia (Marchantiales)
from southern Africa. 12. Riccia albolimbata and the status of R.
albosquamata, white-scaled species originally described by
Amell. Bothalia 19: 17-25.
VOLK, O. H. 1983. Vorschlag fur eine Neugliederang der Gattung
Riccia L. Mitteilungen aus der Botanischen Staatssammlung,
Miinchen 19:453—465.
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. argenteo-
limbata and R. albornata. Bothalia 18:155-163.
Bothalia 19,1: 17-25(1989)
Studies in the genus Riccia (Marchantiales) from southern Africa. 12.
Riccia albolimbata and the status of R . albosquamata, white-scaled
species originally described by Arnell
S. M. PEROLD*
Keywords: Arnell, Marchantiales, Riccia albolimbata, R. albosquamata, white scales
ABSTRACT
Amell's (1957, 1963) inadequate descriptions, poor drawings, mistakes in the text and key. as well as mixed collec-
tions, have all contributed to the confusion regarding the identity of his species R albolimbata and R albosquamata.
Volk's collections (Arnell 1957) of the latter consist mostly of two white-scaled species in the same packet: one is R.
albolimbata and the other is R. argenteolimbata Volk & Perold (Volk et al. 1988). Evidently Arnell used characters of
both plants to describe R. albosquamata. although the type specimen, Volk 452. consists of only R albolimbata (plus
fragments of R. atropurpurea Sim and R. trichocarpa Howe). R. albosquamata is accordingly to be regarded as a
taxonomic synonym of R. albolimbata.
UITTREKSEL
Arnell se (1957, 1963) onvoldoende beskrywings. swak tekeninge, foute in die teks en sleutel. sowel as gemengde
versamelings, het almal bygedra tot die verwarring omtrent die identiteit van sy spesies R. albolimbata en R. albosqua-
mata. Volk se versamelings (Arnell 1957) van laasgenoemde, bestaan meestal uit twee spesies met wit skubbe in dieselfde
pakkie: die een is R. albolimbata en die ander een R. argenteolimbata Volk & Perold (Volk et al. 1988). Blykbaar het
Arnell kenmerke van beide plante gebruik om R. albosquamata te beskryf. alhoewel die tipe-eksemplaar. Volk 452.
alleenlik bestaan uit R. albolimbata (plus fragmente van R. atropurpurea Sim en R. trichocarpa Howe). R. albosqua-
mata word derhalwe beskou as ’n taksonomiese sinoniem van R. albolimbata.
1 . Riccia albolimbata S. Arnell, Mitteilungen aus
der Botanischen Staatssammlung, Miinchen: 264 (1957);
1963:25.
TYPE.— SWA/Namibia, 2117 (Windhoek): Farm
Voigtland, bei Windhoek, gegen Ondekaremba, Kalk-
boden (— AB), 1956.02.12. Volk 11419 (PRE-CH
4232) (M!, holo., PRE!). 2217 (Windhoek): Binsen-
heimkamp (—CD), Volk 11080 (M!, PRE!, para.).
Thallus monoicous, perennial, in rosettes 15—20 mm
across, or in crowded gregarious patches; medium-sized;
furcate or bi-furcate, branches ± symmetrical or one
branch smaller, diversely divergent, ovate to oblong, up
to 8,0 mm long, (1,5-) 1,8-2, 2 (-3,0) mm broad,
2-2,5 times wider than thick; apex rounded, emarginate
(Figures 1A; 2A); sulcus narrow and deep distally, dis-
appearing in proximal parts, which are ± flat or slightly
concave (Figure 1D1-5); dorsally green and shiny, be-
coming white and spongy over sporangia; margins acute
to subacute, flanks generally sloping obliquely upward
and outward, green or brown to dark red; ventral surface
flat to slightly rounded (Figure 1C, 1D1-5), green;
when dry, margins inflexed with wavy hyaline/white
scales covering most of dorsal surface (Figure IB).
Anatomy of thallus: cells of dorsal epithelium emerg-
ing apically from groove in regular rows, hyaline, thin-
walled, unistratose, dome-shaped or globose (Figures
IE; 2C), ± 45 x ± 50 /am, each cell usually with a
single corresponding column of assimilation cells
* Botanical Research Institute. Department of Agriculture and Water
Supply, Private Bag X 101 . Pretoria 000 1 . RSA.
MS. received: 1988.04.24.
beneath, bulging upper walls sprinkled with calcium
carbonate granules, cells soon collapsing, especially
toward margins and more proximally (Figures IE; 2D);
air pores 4— 5(— 6)-angled, leading to air canals below
(Figure IF); assimilation tissue (chlorenchyma) about Vi
the thickness of thallus, cells short -rectangular, ± 50 x
40-45 pm, in columns of 6-8 (- 10), enclosing 4—5-
sided air canals (Figures IF; 2E) which widen toward
sides of branches (Figure 2B); storage tissue occupying
lower Vi of thallus, cells angular, size ± 55 gm.
Rhizoids mostly smooth, some tuberculate, 15-20 /u.m
wide. Scales hyaline to white, base often flecked with
brown or dark red, closely imbricate, undulating, large,
800 — 900 (-1200) x ± 600 part (Figure 1G), extending
± 150 p m above margin of thallus, rounded, edge
mostly smooth, cells hexagonal to oblong-hexagonal in
body of scale (Figures 1H; 2F), 55-100 x 35-55 gm,
marginally smaller, ± 25-40 x 30—40 pm, surface of
cells often encrusted with calcium carbonate deposits,
cell walls generally free of crystals. Antheridia with
thick hyaline necks becoming white and thread-like, in
one or two rows along middle of lobe. Arche gonia with
purple necks. Sporangia with about 300—450 spores
each, overlying tissue turning white and spongy, disinte-
grating soon and leaving several capsules exposed along
the longitudinal hollow. Spores 82-95 (-105) gm in
diameter, yellow-brown to dark brown, semi-transparent
to opaque, triangular-globular, polar, with wing narrow,
3, 0-5,0 pm wide, often with pores at marginal angles,
margin crenulate or finely eroded (Figure 3A); distal face
with ornamentation quite variable, generally (7—)
10—12 angular to round areolae across (Figure 3B, C,
D), 5,0—7, 5(— 10,0) pm wide, areolar walls varying
from thin to thick, with raised papillae at nodes (Figure
18
Bothalia 19,1 (1989)
FIGURE 1. — Riccia albolimbala Amell. Morphology and anatomy. A, fresh thallus; B, dry thallus; C, ventral view of thallus; Dl-5,
transverse sections of branch at different distances from the apex to older proximal part, 5. M. Perold 1380\ E, enlargement of transverse
section through intact dorsal epithelial cells, capped with calcium crystals and showing narrow air canals in assimilation tissue; on right
collapsed dorsal epithelial cells toward margin and larger air canals between columns of cells; F, epithelial cells with calcium crystals, air
pores (hatched) overlying 4-5-sided air canals (dotted), as seen from above, S. M. Perold 398\ G, scale; H, enlarged scale cells with
calcium crystals, S. M. Perold 803. Drawings by J. Kimpton. Scale bar: A-D = 1 mm; E, F, H = 50 /urn; G = 100 ftm.
Bothalia 19,1 (1989)
19
FIGURE 2. — Riccia albolimbata
Amell. Morphology and anat-
omy. A, dorsal view of apex
and groove; B, large air pores
near margin; C, globose dorsal
cells intact at groove; D, col-
lapsed dorsal cells around air
pores; E, dorsal cells and air
pores, 5. M. Perold 398'. F,
scale cells, Volk 11419', G, scale
cells, Volk 452. A-D, SEM
micrographs; E-G, LM (light
microscope) micrographs. Scale
bar on A-G = 100 /ixm. All
SEM and LM micrographs by
S. M. Perold.
''3s-
3F), frequently only a few of the areolae over central
area complete, and short, irregular ridges radiating out-
wards from central areolae and extending onto wing;
proximal face with triradiate mark distinct and generally
incompletely areolate, walls often thinning out or anasto-
mosing to form irregularly branching ridges (Figure 3A,
E). Chromosome number n = 12 (Volk 81/204 p.p. ); 16
(Volk 81/160); (Volk 811231b as R. albosquamata); 24
(Volk 81/ 164) (Bomefeld 1984).
Different chromosome patterns within the same Riccia
species, due to differential multiplication of individual
chromosomes, were reported by Bomefeld (1984) and
termed ‘nothopolyploidy’. Multiple chromosome num-
bers in some species (such as R. albolimbata ) render
chromosome numbers unsuitable as a diagnostic charac-
ter (Volk etal. 1988).
R. albolimbata is widely distributed in the summer
rainfall area of southern Africa and has been collected in
South West Africa/Namibia and in Transvaal, Orange
Free State and north-eastern Cape Province. It is
apparently quite rare in Natal and central Cape and has
not been found in the winter rainfall area of the north-
western, western and south-western Cape (Figure 5).
20
Bothalia 19,1 (1989)
FIGURE 3. — Riccia albolimbata
Amell. Spores. A, proximal
face; B, distal face; C, side view
of distal face; D, distal face,
Volk 11419, holotype, (spores
from disintegrated sporangium
at base of thallus); E, proximal
face; F, distal face, Volk 1 1080,
paratype. A— C, E, F, SEM
micrographs; D, LM (light
microscope) micrograph. Scale
bar on A— C, E, F = 50 /urn;
diameter of spore on D. ± 100
jam.
It often grows on rocky outcrops, on shallow, calcrete
soil at an alkaline pH, and on loamy soil between tufts of
grass, sometimes in association with other Riccia
species, e.g. R. atropurpurea Sim, R. okahandjana S.
Amell, R. trichocarpa Howe, R. argenteolimbata Volk
& Perold and rarely with Marchantia spp.
Amell (1957, 1963) reported R. albolimbata to be
dioicous, but it is definitely monoicous (Table 1). There
are other inaccuracies in his description as well: with
branches 7x2 mm, it is not truly ‘minor’, but medium-
sized; the dorsal colour of the thallus is green when
fresh, not ‘pale green’; the scale cells are not ‘almost
quadratic’, except occasionally at the margins: in the
body of the scale they are oblong-hexagonal (he also
described the cell shape as ‘generally hexagonal’ (Amell
1957, at the bottom of p. 266), mistakenly referring to it
as R. albomarginata, as explained below), and their cell
sizes at 55-100 x 35-55 gm are usually somewhat
larger than the 30 x 30-40 x 60 /aid he reported; the
scale cell walls are hardly thinner than in R. albosqua-
mata [in both, the vertical walls are frequently visible
(Figures 1 G, H and 2F, G which illustrate only R. albo-
limbata)]. The surfaces of the scale cells are often
encrusted with calcium carbonate, but he did not men-
tion it, only referring to it under R. albosquamata. In R.
albolimbata spores, he described the wing as incom-
plete, but it could have been partly folded inwards or
partly obscured, as his illustrations (Amell 1957, Figure
2; 1963, Figure 14) suggest. Although there are
generally papillae arising from the areolar nodes, he
stated that they were without papillae; his illustrations
show ± 12 areolae across the distal face, but it has been
found that the ornamentation shows a good deal of
variability, with the areolae often incompletely sepa-
rated, and hence fewer in number (Table 2). The inner
face, however, clearly has ‘irregular, thin lamellae’ as
described by him.
In his key to the Riccia species (pp. 13 — 16), R. albo-
limbata has been left out and R. albomarginata is listed
twice, at Nos 7 and 1 1 . On p. 14, at No. 11,/?. albomar-
ginata must be replaced by R. albolimbata, and also on
p. 25, as he is comparing R. albosquamata with R.
albolimbata (and not with R. albomarginata !). In the
couplet (i) and (ii) on p. 14, part of the first sentence
must be transposed, to correspond with his information
in the text, viz. ‘with a deep and sharp furrow’ belongs
with the information about R. albolimbata and ‘slightly
concave’ refers to his description of the dorsal surface of
TABLE 1. — Comparison of Arnell’s data on Riccia albolimbata and R. albosquamata. Observations and/or comments on type and other specimens by author
Bothalia 19,1 (1989)
- *
c Z
o
O Os
E >
3 ^
*o ***
ed ^
-c Jx
£ 'o
„ +■* >>
TD Uh ^
<L> <U c
2 -2P o
x -C o
" s §
■si £
S £
E o
OJj £
o£ £
c E
5 Z
— 1 ON
E ©
E o
m [g
o 5
o. £
3 E
•= .a
X 01)
o c
> 00
° c;
o -H.
c -S
•n t
« c •a
o o <u
ns =3
2 ° ^
; i-i
•g o "
.5 u IS
o Ef
2 a. 2
Coo
g m c
no
^ ia
55
* x
^ o
o o
toS
o io I
CD 00 O
-O ^ etf
03 c
E ^ o
3 £ §P
S ^ x
>- 0)
cu .E *c
£
o ^
o Jf o
“ « a
a. E -cs
3
Q. 3 •—
— 3 o;
E J5
5 ° s
^ .E o
o on o
1 E^i
► 00
2 .5 -o'
A T3
2 ^
Q o
- j>s ~
o ja
h ^ ~
i '> c
►-J > 0)
o £
S* o ‘q
E
3 O -O
’ rs ^
! T3
.2 > E
E-S
2 Q
E ■*
E s
2 E |
o. ON NO
3 o'
03 0C Q
S' S
*o .2" '
>» £ £
< m E
8 x
- o
E 5
® a. °«
3
x 5
o « 2
® o O
ao c
3 o « o E
Q £
04 =
O co Q E
3 3
Cl C/3
« 0XjZ3
— X) £ 03
u 2 .3 -5
■S CQ 00 < o
a 5
2 3
almost quadratic, 30 X 30 to deposits cells large, 50 X 70 mtti, thick -walled, lumen white, heavily encrusted with overlying calcium
40 X 60 /um ± filled of (sic) fine granules deposits, perhaps due to much higher calcium
content in substrate
22
Bothalia 19,1 (1989)
TABLE 2.— Comparison of measurements of spores from the type specimens of Riccia albolimbata ( Volk 11419) and R. albo
squamata ( Volk 452: only the white-scaled portions). A minimum of 10 spores examined from each sporangium
R. albosquamata. It has now, however, become clear
that the dorsal surface of R. albolimbata is only apically
deeply grooved, and flat to somewhat concave proxi-
mally, whereas that of the other white-scaled species in
these Volk collections, R. argenteolimbata (Volk et al.
1988), is deeply grooved along the entire length of the
thallus.
Besides the types, there are fortunately several other
fairly good specimens of R. albolimbata with spores
[Volk 11080 (M!, PRE!), 11401 (Ml), 11705 (PRE!)
and 1 1946 (M!, PRE!)], identified by Amell, to enable
one to form a clear concept of the characters of this
species. Volk 11967 (PRE!), is possibly a large plant of
R. atropurpurea, judging by the spore ornamentation,
but the thalli are broken up and it is difficult to make a
definite decision about its identity. In notes found with
some of Duthie’s collections from Fauresmith and Mid-
delburg (Cape) and now identified as R. albolimbata ,
she referred to them as the ‘Doomberg’ species, thus
clearly indicating that she recognized them as belonging
to a distinct species. Regrettably, she made no attempt to
describe it.
R. albolimbata is closely similar to R. albornata
Volk & Perold (Volk et al. 1988), but the former often
grows in rosettes, the dorsal tissue covering the
sporangia turns chalk-white and spongy, its scales are
somewhat smaller and the spore ornamentation is
coarser. Furthermore, the distribution ranges of the two
species do not appear to overlap, except for two
localities at Middelburg and Britstown, in the central
Cape, and a doubtful one in northern South West Africa/
Namibia.
R. albida Sull., described by Frye & Clark (1937) as
having a white, spongiose dorsal surface (which they
called ‘calcified’), differs in other respects from R. albo-
limbata by being a small plant, 2—4 mm long and 1,1
mm wide; the ventral scales are minute and the spores
nearly smooth. Na-Thalang (1980) regarded R. austinii
Steph., R. albolimbata S. Amell, R. albosquamata S.
Amell and R. albomarginata Bisch. as species closely
related to R. lamellosa Raddi. However, R. albomargi-
nata belongs to section Pilifer Volk, which is endemic
to South Africa and is characterized by a dorsal epithe-
lium of loose cell pillars (Volk 1983). R. lamellosa [=
R. austinii (Muller 1954)], is a larger plant with thallus
lobes up to 20 mm long and with a somewhat different
spore wing and spore ornamentation (Jovet-Ast 1986),
neither has it been recorded from southern Africa.
2. Riccia albosquamata S. Arnell, Mitteilungen
aus der Botanischen Staatssammlung, Miinchen: 266
(1957); 1963:25.
TYPE. — SWA/Namibia, Damaraland, 1918 (Groot-
fontein): Neitsas, am Rande einer Kalkpfanne (-CA),
1956 Volk 452 p.p. (M!, holo.). 2017 (Waterberg):
OTJ147 (-CA), Volk 881 p.p. (M!, PRE!, para.).
Amell (1957, 1963) recognized two species, R. albo-
limbata and R. albosquamata on the basis of the fol-
lowing (see also Table 1):
(i) the monoicism or dioicism of the plants;
(ii) the so-called differences in the colour, size and
branching of the thalli;
(iii) differences in the shape of the dorsal groove, mar-
gins and flanks;
(iv) the thin-walled subspherical or cubic-shaped dor-
sal epithelial cells;
(v) the hyaline or white-spotted scales with,
respectively, thin or thick cell walls;
(vi) the different ‘texture’ of the spores, i.e. ± 12-14
areolae across the diameter, forming a reticulum
without papillae, in his spore drawing [Table I,
Figure 2 (1957), Figure 14 (1963)], as opposed to
7-8 areolae across the spore diameter [Table II,
Figure 1 (1957), Figure 15 (1963)], with ‘pro-
cesses of the reticulum (projecting) as spines in the
wing’.
The holotype collection of R. albosquamata, Volk
452 , is a mixed gathering which also contains R. tricho-
carpa Howe and R. atropurpurea Sim and several
branches of a white-scaled species. Detailed investiga-
tions of the white-scaled material resulted in the follow-
ing findings: the plant appears to be monoicous, as R.
albolimbata was also found to be, and the branching,
shape and size of the lobes fall within the normal range
for R. albolimbata. The material is, of course, no longer
fresh and green and dorsally it has turned whitish. The
groove is only distinct apically, flattening out and
becoming slightly concave proximally. The margins are
subacute, the flanks are steeper and less sloping than
Bothalia 19,1 (1989)
23
FIGURE 4. — Riccia albosquamata,
Volk 452, holotype. Spores. A,
proximal face; B, distal face
(spores from sporangium in
larger thallus); C, proximal
face; D, distal face (spores from
sporangium at base of thallus);
E, distal face (spores from
small, loose thallus); F, distal
face (spores from small thallus).
A-F, SEM micrographs. Scale
baron A-F = 50 gm.
usual and the ventral surface is convex. The dorsal epi-
thelial cells have collapsed, but the pores are 4-5-
angled, the latter not commented on by Amell. The
scales are white with a reddish purple base, imbricate,
large, 850-1000 x ± 550 pm, and extend above the
margin of the thallus; the cells are ± 60—75 x 35 /tm
(Amell reported 70 x 50 pm), calcium carbonate
granules cover the cell surfaces and do not fill the cell
lumens as stated by Amell. The cell walls are not truly
thicker either (Figure 2G). The spores were found to be
70-100 /Am in diameter (Table 2) (Amell reported
70-80 /xm), yellow-brown to dark brown, triangular-
globular, polar, with the wing (3 — )5( — 7) /Am wide,
sometimes with radial folds, pores at the marginal angles
and the margin crenulate (Figure 4A); distal face with
8-10 (-12) areolae across (Figure 4B, D, E, F), (5,0-)
7,5-12,5 /Am wide and often incomplete; sometimes the
branched ridges over the centre of the face are more
prominent (Figure 4B, F), as also seen in Amell’ s figure.
The smaller number of slightly wider areolae Amell
described and illustrated for R. albosquamata spores,
namely 7 or 8 as opposed to ± 12 he illustrated for R.
albolimbata spores, are due to several areolae being in-
completely separated and becoming confluent. Papillae
project from the areolar nodes as in R. albolimbata
spores. Amell made no mention of the proximal face,
but with thin, irregular ridges and a distinct triradiate
mark (Figure 4A, C), it is indistinguishable from that of
R. albolimbata.
I therefore conclude that the white-scaled branches
included in the type gathering of R. albosquamata repre-
sent R. albolimbata. I hereby sink R. albosquamata S.
Amell under R. albolimbata S. Amell, because the type
collection of R. albolimbata represents only one species
and the description refers to only one species, whereas
R. albosquamata is based on a mixed type collection
and its description is based on different species.
The identification of the other specimens which Amell
placed under R. albosquamata are as follows; Volk 453
(M, PRE!): the PRE specimen contains only R. tricho-
carpa; Volk 881 (paratype) (M!, PRE!): R. albolimbata
and R. argenteolimbata Volk & Perold; Volk 883 (M!,
PRE!): R. albolimbata , R. argenteolimbata and R. atro-
purpurea Sim; Volk 11906 (M!): R. argenteolimbata, R.
okahandjana S. Amell , R. trichocarpa and R.atropur-
purea; Volk 12744 (M!, PRE!): R. albolimbata and an
unidentified Riccia species. None of these white-scaled
24
Bothalia 19,1 (1989)
FIGURE 5. — Distribution map of R. albolimbata in southern Africa.
specimens, except for the unidentified fragment of Volk
12744 and Volk 452, have spores.
It will be noticed that three of the above specimens
also contain R. argenteolimbata Volk & Perold in the
mixed collections. It is evident that Amell based his
description of R. albosquamata on the two white-scaled
species, R. albolimbata and R. argenteolimbata, using
characters from both. The references to ‘thin and scale-
like’ margins, perpendicular sides, white scales with
pale purple bases ‘resembling the scales of R. limbata in
shape and size’ and presumably the ‘cubic’ dorsal cells,
indicate that he referred to the R. argenteolimbata part
of the collections, whereas the ‘concave dorsal face’ of
the thallus and the spores with a wing (i.e. polar), are R.
albolimbata characters; R. argenteolimbata has a long,
sharp dorsal groove and the spores are wingless and
apolar, its thallus is compact and the dorsal air pores are
mostly triangular, but he did not note this. To give
Amell the credit due to him, he obviously recognized
that there were two white-scaled species present, but he
failed to distinguish clearly between them. As concluded
above, R. albosquamata is a taxonomic synonym of R.
albolimbata ; R. argenteolimbata has been described as
a new species (Volk et al. 1988).
SPECIMENS EXAMINED
SWA/NAMIBIA. — 1917 (Tsumeb): Farm Kumkauas (-CA), E.
Relief 1459 (PRE). 1918 (Grootfontein): Ossa, on dolomite (-AC),
Volk 81-146 (M, PRE); Gaikos (-AVolk 84-703 (M, PRE); Neitsas
(-BC), Volk 452 (M). 2017 (Waterberg): Oros 98 ( — AA), Hoffmann
PRE-CH 4516 (PRE); OTJ 147 (-CA), Volk 881 p.p .,883 p.p. (M,
PRE); Wilhelmstal (-CD), Volk 84-717, 84-721 (M, PRE). 2116
(Okahandja): Erichfelde, auf Kalk (-DA), Volk 11946 (M, PRE).
2118 (Steinhausen): Gobabis, Farm Sturmfeld (-DB), Toelken 5558
(PRE). 2216 (Otjimbingwe): OM 37 Otjua ( — AA), Volk 81-1 15 (M,
PRE). 2217 (Windhoek): Rietfontein, on calcareous crust (-CD), Volk
81-265 (M, PRE); Binsenheimkamp (—CD), Volk 11080 (M, PRE);
Wilhelmstal (-CD), Volk 84-717 (M, PRE). 2316 (Nauchas): Farm
Naos, on calcareous crust (- BA), Volk 81-200 (M, PRE). 2317 (Re-
hoboth): Gravenstein (-BC), Volk 11705 (M, PRE). 2416 (Malta-
hohe): MAL98(-DD), Volk 01254a (M). 25 16 (Helmeringhausen):
Duwisib (-BC), Volk 12744 p.p. (M. PRE).
TRANSVAAL.— 2228 (Maasstroom): Alldays, 55 km W of, on
calcrete soil ( — DA), S. M. Perold 770 (PRE); Gregory Halt ( — DA),
S. M. Perold 737 (PRE); 38 km W of Alldays (-DB), S. M. Perold
759 (PRE); Bulkop, on calcrete soil (-DC), S. M. Perold 793-795
(PRE). 2229 (Waterpoort): Chasa (?), Limpopo River, precise locality
unknown, Stephansen 5393 (BOL); 4 km W of Alldays, Farm Bavaria
(-CA), S. M. Perold 733 (PRE); Wylie's Poort (-DD), S. M. Perold
803 (PRE). 2327 (Ellisras): Villa Nora, 29 km NW of Farm Fransch-
hoek ( — BD), Smook4231 (PRE). 2329 (Pietersburg): Vivo, 15 km N
of (-AB), S. M. Perold 725 (PRE); Dendron, 25 km S of ( — AD),
S. M. Perold 719 (PRE). 2330 (Tzaneen): Lebowa Ga-Modjadji
( — AD). Glen 1400, 1404 (PRE). 2428 (Nylstroom): between Groen-
vallei and Roedtan, Farm Zoetkoppies ( — DB), S. M. Perold 339
(PRE). 2430 (Pilgrim's Rest): Phalaborwa, Farm Parsons 155, next to
Olifants River (— BB), Venter 12197 (PRE). 2527 (Rustenburg): 14
km N of Rustenburg, on calcareous soil (-CA), 5. M. Perold 222, 228
(PRE); Maanhaarrand, on soil at streamside (-CD). S. M. Perold 454
(PRE). 2530 (Lydenburg): Sudwala, on earth bank ( — BC), S. M.
Perold 398 (PRE). 2531 (Komatipoort): Kaapmuiden, Kaapse plateau
(-CB), Vogel T136 (Mainz). 2627 (Potchefstroom): Wonderfontein-
spruit, 6 km N of Carltonville (-AD), S. M. Perold 1026 (PRE);
Gerhardminnebron (— CA), Ubbink 1156 (PUC); Venterskroon
( — CA), Ubbink 1291 (PUC). 2725 (Bloemhof): Wolmaransstad,
Farm Leeufontein (-BB), A. E. van Wyk5753 p.p. (PRE).
NATAL. — 2730 (Vryheid): Burgers Pass (— CB), S. M. Perold 699
(PRE).
O.F.S. — 2726 (Odendaalsrus): Odendaalsrus (-DC), Smook6583a\
6584 p.p. 2727 (Kroonstad): Heilbron, at stream, N of town ( — BD).
5. M. Perold 1369 (PRE). 2728 (Frankfort): Wonderfonteinspruit, 40
km from Bethlehem on road to Lindley (— CC), S. M. Perold 1365
p.p.; 1366 (PRE). 2825 (BoshoD: Farm Goedehoop (-CA), Volk 81-
204 p.p., 81-210 (M, PRE). 2827 (Senekal): Allemanskraal, plateau
near caravan park ( — AC), Volk 81-041 , 81-231 p.p., 81-214 p.p.,
84-653 (M, PRE). 2925 (Jagersfontein): Fauresmith (— CB). Duthie
5441C, 5445, 5449 (BOL). 2926 (Bloemfontein): Bloemfontein
( — AA), Duthie 5507, 5519 (BOL); Eagle's Nest (-AA), Geo. Potts
CH 1010, CH 1036b (PRE); Henrici CH 3741 p.p. (PRE); Volk
81/289 p.p. (M, PRE). 3025 (Colesberg); H. F. Verwoerd Dam,
watercourse (— CB), 5. M. Perold 950, 951 (PRE).
CAPE. — 2624 (Vryburg): 8 km E of Vryburg (-DD), 5. M. Perold
1380 (PRE). 2724 (Taung): Reivilo, Farm Sebete Tsapitse 899
(-AD), Venter 12457 (PRE). 2823 (Griekwastad): precise locality
unknown, Wilman 5517 (BOL). 3023 (Britstown) Britstown ( — DA),
Duthie 5469 (BOL). 3125 (Steynsburg): Middelburg, Doomberg
(-AC), Duthie 5 1 10, 5438 (BOL).
ACKNOWLEDGEMENTS
Prof, (emer.) Dr O. H. Volk of Wurzburg University
is thanked most sincerely for his generous donation of
specimens to PRE, and for the loan of the holotype of R.
albosquamata, Volk 452, and specimens from his per-
sonal herbarium, as well as many fruitful discussions.
Thanks are also due to the Curators of Botanische Staats-
sammlung, Munich, and the Bolus Herbarium, Uni-
versity of Cape Town, for the loan of specimens and to
Mr Bob Watkins, Wirsam Scientific and Precision
Equipment (Pty) Ltd, Johannesburg, for soil analyses.
REFERENCES
ARNELL, S. 1957. Hepaticae collected in South West Africa by Prof.
Dr O. H. Volk. Mitteilungen aus der Botanische n Staats-
sammlung, Miinchen 16: 262—272.
ARNELL, S. 1963. Hepaticae of South Africa, pp. 411. Swedish
National Scientific Research Council, Stockholm.
BORNEFELD, T. 1984. Chromosomenanalyse derGattung Riccia von
Sud- und SW-Afrika und allgemeine Bemerkungen zurZytogene-
tik der Lebermoose. Nova Hedtvigia 40: 313—328.
FRYE. T. C. & CLARK. L. 1937. Hepaticae of North America. Uni-
versity of Washington Publications in Biology 6: 1-162.
HOWE, M. A. 1899. The Hepaticae and Anthocerotes of California.
Memoirs of the Torres Botanical Club 7: 13—33.
JOVET-AST, S. 1986. Les Riccia de la Region Mediterraneenne.
Cryptogamie, Bryologie, Lichenologie 7: 283-431 .
Bothalia 19,1 (1989)
25
MULLER. K. 1954. Die Lebermoose Europas. In L. Rabenhorst.
Kr\ptogamen-Flora. edn 3, Band 6, 1: 1—756. Leipzig.
NA-THALANG, O. 1980. A revision of the genus Riccia (Hepaticae)
in Australia. Brunonia 3: 61 - 140.
PEROLD, S. M. 1989. Studies in the genus Riccia (Marchantiales)
from southern Africa. 1 1. Riccia montana and R. alboporosa, a
further two new, white-scaled species of the group ‘Squamatae'.
Bothalia 19:9-16.
VOLK. O. H. 1983. Vorschlag fur eine Neugliederung der Gattung
Riccia L. Mitteilwigen aus der Botanischen Staatssammlung.
Munchen 19: 345—465.
VOLK. O. H. 1984. Beitrage zur Kenntnis der Marchantiales in
Sudwest-Afrika (Namibia). Nova Hedwigia 39: 117-143.
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. argenteo-
limbata and R. albornata. Bothalia 18:155—163.
Bothalia 19,1: 27-40(1989)
Notes on African plants
VARIOUS AUTHORS
ASTERACEAE
A NEW SPECIES OF INEZ! A ( ANTHEMIDEAE) FROM THE NORTH-EASTERN TRANSVAAL
Inezia speciosa Brusse, sp. nov.
Herba perennis, erecta, usque ad 450 mm alta (Figure
1). Caulis principalis simplex vel ad basim biramosus,
erectus, 2-4 mm crassus, in scapum productus, hir-
sutus, parce foliatus, ad subbasim multiramosus. Caules
secundarii subbasales, multi, simplices vel sub-
simplices, dense foliati, hirsuti, ascendentes, subaeque
alti terminantes. Folia usque ad 20 mm longa, sat dense
hirsuta, superioribus simplicibus, inferioribus imparipin-
natisectis, usque ad trijugatis, segmentis principalibus
cuneatis vel linearibus, segmentis terminalibus lineari-
lanceolatis, marginibus involutis, apicibus glabris, scle-
roticis, saepe fuscis. Folia axillaria in axillis foliorum
pinnatisectorum vulgaria, simplicia, filiformia vel
lineari-lanceolata, uni- vel bigeminata, pari secundo
semper breviore. Inflorescentia simplex. Scapus
solitarius, ex caule principali productus, hirsutus, per-
parce foliatus. Capitulum solitarium, radiatum, ligulis
exclusis usque ad 25 mm diametro, plus quam 100-flo-
rum. Bracteae involucri 3— 4-seriatae, subaequales,
anguste-lanceolatae, exteme rufae hirsutae, interne gla-
brae, 1 — 1,5 mm latae, 6—9 mm longae, exteriores
interioribus angustiores. Receptaculum nudum. Flores
ligulati circa 25—30, feminei sed steriles. Ligulae al-
bae, quam bracteas multo longiores, simplices vel pro-
funde trilobatae, parce pubescentes, glandibus bicellulo-
sis superficiaribus; cellulae paginae superioris breviter
botuliformes, radiatim leviter striatae; tubus brevis, circa
1 mm longus. Androecium nullum. Stylus 1,7— 1,8 x
0,16-0,23 mm, ad apicem versus biramosus, ramis
0,46-0,54 mm longis, truncatis, base parum dilatata.
Nectarium nullum. Ovarium non bene evolutum, glan-
dibus bicellulosis, superficiaribus. Ovulum redactum.
Pappus nullus. Flores disci hermaphroditi, fertiles,
plus quam 100, 4,2— 5,7 mm longi. Corolla lutea,
2, 5-3,0 mm longa, glandibus bicellulosis superficiari-
bus, anguste infundibuliformis vel cylindrica, tetralo-
bata; lobi 0,4— 0,5 mm longi, quum maturi patens;
margines incrassati, cellulis elongatis protrudentibus,
parietibus crassis et longitudine striatis; cellulae paginae
interioris transverse striatae. Stamina quatema, tubo
corollae supra basin sed infra medium affixa. Fila-
mentum breve, 0,44—0,55 mm longum; collum fila-
menti 0,29—0,33 x 0,10— 0,1 1 mm; cellulae in sectione
longitudinali 6— 8-seriatae; thecae 0,96—1,11 mm
longae; appendix apicalis oblonga, obtusa vel truncata,
0,15-0,20 x 0,19—0,23 mm; caudae 0,04—0,10 mm
longae. Stylus circa 2,6 x 0,19—0,21 mm, ad apicem
versus biramosus, ramis 0,63—0,66 mm longis, trunca-
tis. Stylopodium 0,34—0,46 mm latum, 0,17x0,23 mm
altum, cellulae parietibus valde incrassatis. Nectarium
0,29-0,32 mm latum, 0,10—0, 1 1 mm altum, sub stylo-
podio. Ovarium 1 ,7-2,7 mm longum, tetracostatum,
ubi maturum complanatum, alis marginalibus angustis
sed sursum latioribus, glandibus superficiaribus, bicellu-
losis vel breviter biseriatis (usque ad tres cellulae altis).
Pappus nullus.
TYPE. — Transvaal, 2329 (Petersburg): Iron Crown
Mountain near Haenertsburg, grassy slopes, alt. 5 500'
(— DD). L. E. Codd 9440 , 24.1.1956 (PRE, holo.; K,
MO, iso.). Figure 1.
FIGURE 1. — Inezia speciosa Brusse, habit. L. E. Codd 9440, holo-
type.
Erect perennial herb, up to 450 mm high. Main stem
simple to two-branched at base, erect, 2-4 mm thick,
produced into scape, hirsute, sparsely leafy, subbasally
many-branched (without the main stem branching).
28
Bothalia 19,1 (1989)
Secondary branches subbasal, numerous, simple to sub-
simple, densely leafy, hirsute, ascending, terminating at
subequal height. Leaves up to 20 mm long, moderately
densely hirsute, the upper (younger) simple, the lower
(older) imparipinnatisect, up to trijugate, main segments
cuneate to linear, terminal segments linear-lanceolate;
margins involute; apices glabrous, sclerotic, often
brown. Axillary leaves common in axils of pinnatisect
leaves, simple, filiform to linear-lanceolate, one- or two-
paired, the second (younger) pair always shorter. Inflor-
escence simple. Scape solitary, produced from main
stem, hirsute, very sparsely leafy. Head solitary,
radiate, up to 25 mm diam. excluding rays, more than
100-flowered. Involucral bracts 3— 4-seriate, subequal,
narrowly lanceolate, rufous hirsute on outer surface,
glabrous on inner surface, 1 — 1,5 mm wide, 6—9 mm
long, the outer narrower than the inner. Receptacle nude.
Ray florets 25-30, female but sterile. Rays white,
much longer than bracts, simple to deeply trilobate, spar-
sely pubescent, with superficial bicellular glands (Figure
2); cells of the upper surface shortly (dumpy) botuliform,
radially lightly striate (fingerprinted); tube short, about 1
mm long. Androecium absent. Style 1,7— 1,8 x
0,16—0,23 mm, two-branched towards apex; branches
0,46—0,54 mm long, truncate; base only slightly
thickened. Nectary absent. Ovary not well developed,
with superficial bicellular glands. Ovule reduced. Pap-
pus absent. Disc florets bisexual, fertile, more than 100,
4,2— 5,7 mm long. Corolla yellow, 2, 5-3,0 mm long,
with superficial bicellular glands, narrowly funnel-
shaped to cylindrical, tetralobate; lobes 0,4-0,5 mm
long, spreading when mature; margins thickened, with
protruding elongated cells, with thickened and longitudi-
nally striate (fingerprinted) walls (Figure 4); cells of the
inner surface transversely striate (fingerprinted). Sta-
mens 4, attached to corolla tube above base but below
middle. Filament short, 0,44—0,55 mm long; filament
collar 0,29—0,33 x 0,10—0,11 mm, cells 6-8-seriate
in longitudinal section; anther thecae 0,96—1,11 mm
long; apical appendage oblong, obtuse to truncate,
0,15—0,20 x 0,19—0,23 mm; tails 0,04-0,10 mm
long. Style around 2,6 x 0,19—0,21 mm, two-branched
towards apex, branches 0,63 —0,66 mm long, truncate.
Stylopodium 0,34—0,46 mm wide, 0,17—0,23 mm
high; cells with thick walls. Nectary 0,29—0,32 mm
wide, 0,10—0,11 mm high, under stylopodium. Ovary
1,7 -2, 7 mm long, four-ribbed, flattened when mature,
marginal wings narrow but broadened above, superficial
glands, bicellular to shortly biseriate (up to three cells
high; Figure 3). Pappus absent.
In placing this new species in the monotypic Inezia
Phill., the circumscription of the genus has had to be
changed. Phillips (1932) originally separated Inezia from
Lidbeckia Berg, (a genus with two species) mainly on
the fertile ray florets, and the distribution — Lidbeckia
being a genus of the Cape floral area, whereas Inezia is
a genus of mountainous grassland in the eastern
Transvaal and Swaziland. However, this new species
breaks this distinction down, because it has ray florets
with ovules of similar reduction to Lidbeckia quinque-
loba (L.f.) Cass. In any case, the ray florets of the only
hitherto known species of Inezia, I. integrifolia, are not
perfectly fertile, the ovules being slightly degenerate,
but nevertheless present. I. speciosa has an even more
reduced ovule in the ray achenes. The type of Lidbeckia,
L. pectinata Berg. , has ray florets which are completely
sterile, with no style and the achene severely reduced,
with no ovule present. Lidbeckia quinqueloba (L.f.)
Cass., on the other hand, has the style developed, with
two unequal branches with rounded, rather than truncate
apices. A rudimentary ovule is also discernible in the
achene. All four species have the achene surfaces with
broad bicellular glands (Figure 2) which sometimes
become three-tiered (Figure 3). These are also present on
the rays, but are more sparse here. The degree of reduc-
tion of the ray floret no longer seems to hold well as a
distinguishing character between these two genera.
FIGURE 2. — Inezia speciosa Brusse, photomicrograph of the bicel-
lular glands. L. E. Codd 9440, holotype. Bar = 0, 10 mm.
FIGURE 3. — Inezia speciosa Brusse, photomicrograph of the three-
tiered biseriate glands. L. E. Codd 9440, holotype. Bar = 0,10
mm.
Bothalia 19,1 (1989)
29
'
FIGURE 4. — Inezia speciosa Brusse, photomicrograph of the elong-
ated cells on the margin of the lobes of the disc florets, showing
the longitudinal striations. L. E. Codd 9440 , holotype. Bar =
0,10 mm.
The stylopodium, however, proves to be a useful dis-
tinguishing character between these two genera. The sty-
lopodium of Inezia, consists of cells that are thick-
walled, whereas those of Lidbeckia are thin-walled. The
nectary, a doughnut-shaped mass of delicate cells, with
many stomatal openings on the surface, occurring just
below the stylopodium, and surrounding the core of the
stigma base, is conspicuously larger in Lidbeckia than
in Inezia disc florets. The filament collars are also more
robust in Lidbeckia, with 9—10 rows of cells across,
than Inezia, with 5—8 rows of cells across.
Otherwise the four species (two of Inezia and two of
Lidbeckia ) are fairly similar, particularly in the flat-
tened, narrowly winged achenes, the oblong apical
anther appendages which are obtuse to truncate, and the
presence of broad bicellular glands (Figure 2) on the
surfaces of both the disc and the ray achenes and corol-
las, as well as on the bracts.
These characters are like those of Osmitopsis (Bremer
1972), but this genus has a paleate receptacle, long-
tailed anthers, five corolla lobes and sometimes a lacer-
ate, scarious, sleeve-like pappus. The bicellular glands,
although similar in shape to those of Inezia and Lid-
beckia, appear to contain resin or terpenes to the exclu-
sion of everything else, and appear submacroscopically
as resin droplets. The glands of Inezia and Lidbeckia do
not have this appearance, and the contents appear
aqueous (Figure 2).
The cells on the inner surface of the disc corolla lobes
of all four species are transversely striate (fingerprinted).
The cells on the ray surface are more specialized. The
ray surface cells of /. integrifolia are rounded with thick
walls and cross striations (fingerprinting). The ray sur-
face cells of I. speciosa are obtusely acuminate, or botu-
liform, and faintly radially striate. Lidbeckia pectinata
ray surface cells are ellipsoid, and faintly radially striate.
The ray surface cells of L. quinqueloba are the most
spectacular of the four, with obtusely acuminate or botu-
liform shape, with radial striations, and with the cell
bases 5—8 crenate with the axils of the crenations thick-
ened. However these various structures do appear to be
generically significant.
At present this new species is known only from the
type collection, from the lower slopes of Iron Crown
Mountain near Haenertsburg, in the north-eastern
Transvaal.
REFERENCES
BREMER, K. 1972. The genus Osmitopsis (Compositae). Botaniska
Notiser 125:9—48.
PHILLIPS, E. P. 1932. Inezia, a new genus of Compositae from South
Africa. Kew Bulletin 1932: 297—298.
F. BRUSSE*
* Botanical Research Institute, Private Bag X101 , Pretoria 000 1 .
MS. received: 1988.01.31.
A NEW PHYMASPERMUM (ANTHEMIDEAE) SPECIES FROM DOLOMITE AREAS OF THE WOLKBERG
Phymaspermum argenteum Brusse, sp. nov.
Sufffutex usque ad 850 mm altus, ex caudice ligneo
bulbiformi 40-50 mm diametro perenni oriens. Caules
basi simplices vel versus apicem ramosi, 2—8 mm dia-
metro, praeter partes basales dense foliati, parce vel
dense tomentosi. Folia usque ad 25 mm longa, impari-
pinnatisecta vel simplicia, non- vel trijugata, segmentis
filiformibus vel linearibus, circa 1 mm latis, segmentis
inferioribus quam segmentis superioribus longioribus,
argenteo-hirsutis, apicibus glabris, scleroticis. Inflor-
escentia terminalis, corymbosa, capitulis numerosis,
pedunculis foliis parvis. Capitula discoidea, cylindrica,
4-7 x 2-3 mm, 10- 17-flora. Bracteae involucri
3—4- seriatae, interiores exterioribus longiores, lanceo-
latae, interiores 5-6 x 1 mm, concavae, marginibus
apicibusque scariosis. Receptaculum nudum. Flores
omnes fertiles, omnes hermaphroditi, 4,5 -6,0 mm
longi. Corolla lutea, 3-3,5 mm longa, prope basin an-
guste cylindrica, supra basin campanulata, quinquelo-
bata, lobis 0, 4-0,8 mm longis, glanduloso-pilosa, glan-
dibus biseriatis. Filamentum breve, 0,65—1,00 mm
longum; collum filamenti 0,24—0,37 mm longum, ad
basin 0,09—0,14 mm diametro; cellulae in sectione
longitudinali 7—10 seriatae; thecae 0,93—1,15 mm
longae; appendix apicalis lanceolata, 0,23—0,37 x
0,13-0,16 mm; caudae usque ad 0,17 mm longae (Fig-
ure 7). Stylus 2,4-2, 7 x 0,13-0,24 mm, ad apicem
30
Bothalia 19,1 (1989)
versus biramosus, ramis 0,5— 0,7 mm longis, truncatis.
Stylopodium 0,24—0,42 mm latum, cellulis non vel le-
viter incrassatis. Nectarium sub stylopodio, 0,27—0,42
mm latum, 0,14—0,20 mm altum. Ovarium 1,0— 2,5
mm longum, decacostatum, glandibus bicellulosis super-
ficiaribus numerosis (Figure 6); apex annulo incrassato,
laevis vel decadentatus.
TYPE. — Transvaal, 2429 (Zebediela): 18 km from the
main Pietersburg-Tzaneen road near Boyne to Wolkberg
Estates. One km from the turn-off to Frosch Farm to
Wolkberg Estates. On highest point on road. Erect to
reclining, 0,5—0,85 m tall, silvery subshrub, with yel-
low flower heads, growing as isolated plants in grassland
or open woodland, near low rock outcrops. On rolling
dolomite hill at NW base of prominent NE-SW running
dolomite ridge. Crushed leaves without strong odour.
Soil a brown loam. Not seen on dolomite ridge, but
fairly common on rolling hill below (-BB). F. Brusse
5567, 30.05.1988 (PRE, holo.; B, BM, BR, C, E,
GRA, K, LD, M, MEL, MO, NBG, P, S, UPS, US,
WAG, iso.). Figure 5.
base, sparsely to densely tomentose. Leaves up to 25
mm long, imparipinnatisect to simple, non- to trijugate,
segments filiform or linear, about 1 mm wide, lower
segments longer than upper segments, silvery-hirsute,
tips glabrous, sclerotic. Inflorescence terminal, corym-
bose, heads numerous, peduncles small-leafy. Heads
discoid, cylindrical, 4—7 x 2—3 mm, 10-17-flowered.
Involucral bracts 3-4-seriate, inner ones longer than
outer ones, lanceolate, inner bracts 5-6x1 mm, con-
cave, margins and apices scarious. Receptacle nude.
Flowers all fertile, all hermaphrodite, 4,5— 6,0 mm
long. Corolla yellow, 3,0— 3,5 mm long, narrowly
cylindrical below, campanulate above, five-lobed, lobes
0,4— 0,8 mm long, glandular pilose, glands biseriate.
Filament short, 0,65—1,00 mm long; filament collar
0,24-0,37 x 0,09— 0,14 mm below; cells 7— 10 seriate
in longitudinal section; thecae 0,93 - 1 ,15 mm long; api-
cal appendage lanceolate, 0,23—0,37 x 0,13-0,16
mm; tails up to 0,17 mm long (Figure 7). Style 2,4— 2,7
x 0,13-0,24 mm, two-branched towards apex,
branches 0,5— 0,7 mm long, truncate. Stylopodium
0,24—0,42 mm wide, cells not to only slightly thick-
ened. Nectary below stylopodium, 0,27—0,42 mm
wide, 0,14—0,20 mm high. Ovary 1,0-2, 5 mm long,
ten-ribbed, superficial bicellular glands numerous
(between ribs; Figure 6); apex with thickened rim,
smooth to ten-toothed.
FIGURE 6. — Phymaspermum argenteum Brusse, photomicrograph
showing the bicellular glands on the achene surface. D. R. J.
van Vuuren 1533 , paratype. Bar = 0, 10 mm.
FIGURE 5. — Phymaspermum argenteum Brusse, habit. F. Brusse
5567 , holotype.
Subshrub to 850 mm high, arising from a woody
perennial bulb-shaped caudex, 40 - 50 mm diam., just
below soil level. Stems simple at base to branched
above, 2—8 mm diam., densely leafy except towards
This new species of Phymaspermum superficially
resembles Phymaspermum acerosum (DC.) Kallersjo,
except for the dense silvery hairiness of the leaves. The
leaves of P. acerosum are normally almost hairless
when mature. The hairs, mainly on the undersurface, are
not visible to the unaided eye and are longer than those
of P. argenteum. The inflorescences of the two species
are quite similar, but the heads contain more flowers in
P. argenteum (10-17) than in P. acerosum (4-9; Hil-
liard 1977).
Bothalia 19,1 (1989)
31
FIGURE 7. — Phymaspermum argenteum Brusse, photomicro-
graph showing the anther tails. D. R. J. van Vuuren 1533 ,
paratype. Bar = 0,10 mm.
The anther tails of Phymaspermum argenteum are
longer (Figure 7) than those of P. acerosum, which are
quite short, and the anther bases could be described as
being merely acute. However, this would be misleading
in this case as the cellular structure at the base seems
quite similar, and the cells of the tails of P. argenteum,
are more elongated than those of P. acerosum.
Phymaspermum acerosum is one of the three problem
species in this genus, in that the achenes lack all traces of
superficial bicellular glands (Kallersjo 1985), but P.
argenteum is typical, with abundant superficial bicellu-
lar glands, between the blunt ribs on the achene (Figure
6).
P. argenteum is not a very conspicuous plant and
does not grow much taller than 0,85 m. P. acerosum, on
the other hand, can grow up to two metres (2 m) tall, and
is a large and conspicuous plant of higher altitudes.
Phymaspermum acerosum grows in acidic soils, and
has not been recorded from alkaline soils. P. argenteum,
however, grows in alkaline soils in dolomite areas of the
eastern Transvaal, and so far has only been recorded
from the dolomitic regions of the Wolkberg.
TRANSVAAL. — 2429 (Zebediela): erect semiwoody herb from
stout rootstock. Viscous. Heads yellow. Dolomite formation, Wolk-
berg Plateaux 14 m. SE of Boyne (-BB). D. R. J. van Vuuren 1533,
3.6. 1962 (PRE, K). Wolkberg Estates. Wooded grassland on mountain
slope. Gentle. E aspect. Gritty sandy soil. Herb forming isolated
stands. Rare. Alt. 1 600 m(— BB). S. P. Fourie2759 (ex TPA Nature
Cons. Herbarium 633988), 1982.05.26 (PRE).
REFERENCES
HILLIARD, O. M. 1977. Compositae in Natal, pp. 659. University of
Natal Press, Pietermaritzburg.
KALLERSJO, M. 1985. Fruit structure and generic delimitation of
Athanasia (Asteraceae- Anthemideae) and related South African
genera. Nordic Journal of Botany 5: 527—542.
F. BRUSSE*
* Botanical Research Institute, Private Bag X 101, Pretoria 0001.
MS. received: 1988.01.31.
MELIACEAE
TURRAEA PULCHELLA REDISCOVERED
The first and previously only collections of Turraea
pulchella (Harms) Pennington were made by Alice M.
Pegler in the Kentani District, Transkei, between April
1900 and October 1913. Originally described by Harms
(1917) as the sole member of the genus Nurmonia, it was
transferred to Turraea L. by Pennington & Styles
(1975). As more than 70 years have passed without re-
collections, the species had been considered to be either
extremely rare or extinct.
On September 17, 1987, a few plants of T. pulchella
were found on the Matabetule Plateau ± 20 km north-
west of Durban during a plant collecting trip organized
by staff of the Natal Herbarium. During several subse-
quent visits to the area, T. pulchella was found to be
locally abundant, particularly in heavily grazed grass-
land burnt in July/ August. The plants had prominent,
perennial rootstocks 80-200 mm long, ± prostrate to
spreading annual shoots and pure white flowers with a
singular spicy fragrance.
The vegetation of the Matabetule Plateau is a mosaic
of Aristida junciformis Trin. & Rupr. subsp. juncifor-
mis grassland and bush clumps associated with termita-
ria. Prominent among the trees are Syzygium cordatum
Hochst., Spirostachys africana Sond., Combretum
molle R. Br. ex G. Don and Faurea saligna Harv.
Although many plants of T. pulchella grow in open
grassland, they also tend to be associated with the
isolated low-growing woody shrublets scattered between
bush clumps — particularly geoxylic suffrutex forms of
Faurea saligna and Sclerocarya birrea (A. Rich.)
Hochst. subsp. caffra (Sond.) Kokwaro.
The soils on the plateau are very sandy, light grey to
reddish and derived from Natal Group Sandstone (for-
merly known as Table Mountain Sandstone). Fortunately
the Matabetule Plateau (which is in KwaZulu) is ear-
marked as a nature reserve and will form part of the
Durban Metropolitan Open Space System (M.O.S.S.).
32
Bothalia 19,1 (1989)
In its subherbaceous suffrutescent habit T. pulchella is
very different from most other members of the genus. A
rather similar habit is shown by T. streyi F. White &
B. T. Styles, described in 1986, another rare species
known from only two localities in Natal. Subsequent to
the rediscovery of T. pulchella , several plants of T.
streyi were relocated at the type locality near St.
Michael’s-on-Sea. Detailed studies on these two interest-
ing species can now be undertaken to elucidate aspects of
their taxonomy, biology and ecology.
VOUCHER SPECIMENS
NATAL — 2930 (Pietermaritzburg): Matabetule Plateau. ± 20 km
NW of Durban (-DB). Van Wxk 814 1.8238 (PRU): Williams 36. 84
(NH).
REFERENCES
HARMS, H. 1917. Uber die asiatische Meliaceen-Gattung Munronia
Wight und eine verwandte siidafrikanische Gattung. Berichte der
Deutschen Botanisehen Gesellschaft 35: 74-82.
PENNINGTON. T. D. & STYLES. B. T. 1975. A generic monograph
of the Meliaceae. Blumea 22: 419-540.
WHITE, F. 1986. The taxonomy, chorology and reproductive biology
of southern African Meliaceae and Ptaeroxylaceae. Bothalia 16:
143-168.
R. WILLIAMS*
* Natal Herbarium, Botanical Research Unit, Botanic Gardens Road.
Durban 4001.
MS. received: 1988.08.11.
FABACEAE
THE IDENTITY OF LOTONONIS ELONGATA (CROTALARIEAE)
Several specimens of a species superficially similar to
Lotononis prostrata (L.) Benth. and L. azurea (Eckl.
& Zeyh.) Benth. have accumulated in southern African
herbaria since 1953. The herbarium collections and an in
situ study have shown that the species is quite distinct
from all other species of the section Telina (E. Mey.)
Benth. It was written up to be described as a new species
under the name L. repens (Van Wyk unpublished
manuscript).
A recent examination of the Thunberg Herbarium in
Uppsala has revealed that ‘L. repens’ is identical to Ono-
nis elongata Thunb., a species treated by both Harvey
(1862) and Diimmer (1913) as synonymous with L. pros-
trata (L.) Benth. Unlike Harvey, Diimmer apparently
did not see the type specimen and presumably followed
Harvey’s interpretation of the species. Bentham (1843)
did not study the Thunberg collection and did not cite
Ononis elongata in his revision of Lotononis. Dietrich
(1847) somewhat hesitantly transferred most of Thun-
berg’s Ononis species to the genus Lotononis — amongst
others also O. elongata. These and several other new
combinations by Dietrich have apparently been over-
looked by subsequent workers.
The type specimen of Ononis elongata is a small
piece of flowering material only, so that the unusual
pods, distinctive habit and other diagnostic characters
were not known before.
Lotononis elongata (Thunb.) D. Dietr., Synopsis
plantarum 4: 960 ( 1 847).
Ononis elongata Thunb.. Prodr omits plantarum capensiarn 2: 129
(1800); Thunb.: 587 (1823): DC.: 167 (1825); Harv. 2: 53 (1862), as
synonym of L prostrata: Diimmer: 296 (1913), as synonym of L.
prostrata Type: South Africa, Cape Province, in Cap. b. Spei'. Thun-
berg s.n. sub THUNB-UPS 16607 (UPS!, lecto., here designated).
Prostrate perennial herb up to 1 ,2 m wide and no more
than 0,1 m high. Branches prostrate and creeping,
spreading from a central rootstock; twigs densely pilose.
Leaves digitately trifoliolate, very variable in size and
shape; petiole (2—) 4-8 (—11) mm long, sparsely
pubescent; leaflets oblanceolate, obovate or elliptic,
(3—) 7—16 (—20) x (1,5—) 2-5 (-8) mm, abaxially
sparsely pubescent, adaxially glabrous. Stipules con-
sistently present, single at each node, lanceolate to
broadly ovate, (1 — ) 2—8 (—11) mm long. Inflor-
escences leaf-opposed at each node, slender, long-
pedunculate, 25—75 mm long, invariably single-flow-
ered; bracts small, up to 2,5 mm long; bracteoles absent.
Flowers 10-14 mm long, yellow; pedicel 1-3 mm
long. Calyx 7-9 mm long, with the lateral lobes on
either side fused higher up in pairs, sparsely pubescent.
Standard large, broadly ovate to orbicular, 11 — 16 mm
long, with a line of hairs dorsally along the middle,
adaxially yellow, abaxially pale brown. Wing petals
dimorphic, the one oblanceolate with an obliquely trun-
cate apex, the other oblong-obovate with a rounded
apex, the former positioned higher than the latter at
anthesis; both longer than the keel; sculpturing similar in
both, upper basal and upper left central, in 5— 6 rows of
inter- and intracostal lunae. Keel petals semicircular,
obtuse, auriculate and pocketed near the base. Anthers
dimorphic. Pistil 9- 12 mm long; ovary linear, 6-8 mm
long, the upper part without ovules and similar to the
style, the lower fertile part very short, 2-3 mm long.
Fruit ovate, terete, very short, 8—14 mm long (exclud-
ing the style), 3—6 mm in diameter; apex gradually
tapering to the style; pubescent. Seed suborbicular in
side view, up to 2 mm long, distinctly tuberculate, pale
brown or with some irregular black marks; hilar area
markedly swollen and black in colour, obscuring the
hilar valve (Figure 8). Chromosome number. 2n = 28!
Voucher specimen: Van W\k 2573 (BOL, C. GRA, JRAU. K. M,
MO. NBG. NH, PRE, SAAS.' STE).
Lotononis elongata is similar to L. azurea and L.
prostrata but differs from these and related species in
the shape of the flowers, the dimorphic wing petals, the
sterile upper part of the ovary and in the short, ovate and
distinctly pointed fruit. It differs from L. azurea also in
the yellow colour of the flowers, the shape of the leaflets
and the more hairy twigs and leaves. From L. prostrata it
also differs in the more spreading habit, which can be up
to 1 ,2 m wide (± 0,3 m in L. prostrata).
Bothalia 19,1 (1989)
33
FIGURE 8. — -Lot ononis elongata. A1 , A2, flowering branches, showing the prostrate habit and slender peduncles: B 1 , B2. leaves in abaxial
view; Cl. C2. calyx opened out, upper lobes to the left, showing the fusion of the lateral lobes; Dl, D2, standard petals (note the
difference in size); El. E2, wing petals from the same flower, showing the dimorphic shape; F. keel petal; Gl, G2. pistils, showing the
sterile upper parts of the ovaries; HI, H2. mature fmit (note the size, shape and also the pointed, tapering apices, which are diagnostic for
the species); I, androecium; J 1 , J2, seeds in side view, showing tuberculate surface; J3, seed in hilar view, showing the raised area around
the hilar valve; K1 . K2, bracts; LI , L2, L3, stipules, showing variation in size and shape. All from Van Wxk 2573 except A1 & B2 from
Vlok 1762 and C2. Dl , G2, 1 & L2 from Vlok 1718. Scales in mm.
34
Bothalia 19,1 (1989)
FIGURE 9. — The known geographical distribution of Lotononis elongata.
The species has a relatively wide distribution in the
southern Cape (Figure 9). It seems to have escaped the
notice of collectors for nearly 200 years — recent collec-
tions date back to 1953. Thunberg must have collected
the type specimen on one of his two journeys to the
eastern Cape between 1772 and 1774. It is known that he
crossed the Outeniqua Mountains on several occasions
during this period (Gunn & Codd 1981). Recent collec-
tions by Mr J. H. J. Vlok (Saasveld Forestry Research
Centre, George) have considerably expanded the known
distribution range.
The peculiar pistil, in which the upper sterile part of
the ovary closely resembles the style, has not been
observed in any other species of the section Telina. This
development approaches the usual condition in the genus
Crotalaria (where the style is geniculate and hairy), but
in L. elongata the hairy part of the pistil is undoubtedly
part of the ovary and not the style. The presence of
macrocyclic pyrrolizidine alkaloids in L. elongata (Van
Wyk & Verdoom in prep.) also suggests an affinity with
Crotalaria. The single stipules, calyx structure, tubercu-
late seed surface and chromosome number, however,
leave little doubt about the correct generic position in
Lotononis.
CAPE. — 3322 (Oudtshoom): Swartberg Mountains, mid-northern
slopes of Blesberg, next to forestry track (-BC), 15.12.1986, VI ok
1762 (JRAU, K, NBG, PRE); northern foothills of Outeniqua Moun-
tains, about 3 km west ofHerold at Camferskloof (-CD), 22. 1 1.1986,
Vlok 1718 (BOL, C, GRA, JRAU, K, M, MO, NBG, NH, PRE,
SAAS, STE); top of Outeniqua Pass (-CD), 23.1. 1987, B-E. van Wyk
2573 (PRE, BOL, C, GRA, JRAU, K, M, MO, NBG, NH, PRE,
SAAS, STE); Zebra, top of Outeniqua Pass (-CD), 19.10.1953,
Lewis 3734 (SAM); Zebra, George District (-CD), 19. 10. 1953, Bar-
ker 8195 (MO, NBG). 3323 (Willowmore): about 8 km from Union-
dale on road to Avontuur (-CA), 20.12.1967, Grobbelaar 1401
(PRE); 23,2 km east by south of Uniondale (-CB/-CD), 16.11 1958
Acocks 19986 (K, M).
ACKNOWLEDGEMENTS
I am indebted to Mr J. H. J. Vlok (Saasveld Forestry
Research Centre, George) for valuable collections of L.
elongata and to Dr and Mrs Charles McDonald (Taren-
taalbos, George) who kindly collected ripe fruits and
seeds for chromosome counting. The taxonomic study of
Lotononis is a registered Ph. D. project at the University
of Cape Town. Financial support from the Rand Afri-
kaans University enabled me to study the Thunberg col-
lection.
REFERENCES
BENTHAM, G. 1843. Enumeration of Leguminosae, indigenous to
southern Asia, and central and southern Africa. Hooker’s, London
Journal of Botany 2: 504 -613.
DE CANDOLLE, A. P. 1825. Prodromus systematis naturalis regni
vegetabilis 2: 166- 168. Treuttel & Wurtz, Paris.
DIETRICH, D. N. F. 1847. Synopsis plantarum 4: 959—961. Bern-
hard Voigt, Weimar.
DUMMER, R. A. 1913. A synopsis of the species of Lotononis Eckl.
& Zeyh. , and Pleiospora Harv. Transactions of the Royal Society
of South Africa 3: 275 — 335.
GUNN, M. & CODD, L. E. 1981. Botanical exploration of southern
Africa: 347-350. Balkema, Cape Town.
HARVEY, W. H. 1862. Leguminosae. In W. H. Harvey & O. W.
Sonder, Flora capensis 2: 47-66. Hodges, Smith, Dublin.
THUNBERG, C. P. 1800. Prodromus plantarum capensium 2:
129- 130. Edman, Uppsala.
THUNBERG, C. P. 1823. Flora capensis: 584-589. Schultes,
Stuttgardt.
B-E. VAN WYK*
* Department of Botany, Rand Afrikaans University, P.O. Box 524,
Johannesburg 2000.
MS. received: Sept. 1988.
Bothalia 19,1 (1989)
35
FUSCIDEACEAE
A NEW SPECIES OF FUSCIDEA (LICHENES) FROM THE CAPE FOLD MOUNTAINS
Fuscidea hottentotta Brusse, sp. nov.
Thallus crustosus, saxicola, violaceo-brunneus, ad 80
mm diametro, 100- 300 pm crassus, rimoso- vel dissi-
tiareolatus. Areolae 0,2— 2,0 mm diametro. Hypothal-
lus ater, thallum marginans, margine 0,5— 2,0 mm lato.
Cortex superior circa 20 pm crassus. Stratum gonidiale
20—100 pm crassum; algae Protococcoideae, 5—16 pm
diametro. Medulla alba, 50—200 pm crassa. Apothecia
nigella, adnata, lecideina, ad 1 ,2 mm diametro, margini-
bus leviter brunneis vel atris, discis atris, planis vel con-
cavis. Excipulum pallide brunneum, 35-45 pm cras-
sum, anticipate paraplectenchymatum, cellulis 4-6,5
/xm latis, 4—15,5 pm longis. Hypothecium hyalinum,
65—95 pm crassum. Hymenium hyalinum, 65—75 pm
altum, solum ascis J+ caeruleis; epihymenium brun-
neum. Asci clavati, tholis J+ caeruleis. Ascosporae
octonae, hyalinae sed mox fuscae, curvae vel reni-
formes, simplices vel interdum uniseptatae, 11,5-17,5
x 5— 8 pm. Pycnidia hyalina, ampulliformia, circa 80
pm lata et circa 100 pm profunda. Pycnidiosporae * hya-
linae, rectae, anguste ellipsoideae, acrogenae, 3-5 x
1 — 1,4 pm. Thallus acidum ramalinolicum continens.
TYPE. — 3322 (Oudtshoom): 40 km from the Olifants
River bridge near Oudtshoom to Mossel Bay, Robinson
Pass. SW slopes of Ruitersberg. On almost vertical SW
Table Mountain Sandstone rock face, on steep SW
slope. Alt. 790 m (-CC). F. Brusse 5312, 1988.03.12
(PRE, holo.; B, BM, LD, UPS, iso.). Figure 10.
FIGURE 10. — Fuscidea hottentotta Brusse. Habit. F. Brusse 5312,
holotype. Scale in mm.
Thallus crustose, saxicolous, violet-brown, to 80 mm
across, 100- 300 pm thick, rimose to scattered areolate.
Areoles 0, 2-2,0 mm across. Hypothallus black, lining
thallus; lining 0,5— 2,0 mm broad. Upper cortex about
20 gm thick. Algal layer 20— 100 pm thick; algae Pro-
tococcoid, 5—16 pm diam. Medulla white, 50-200
* The term pycnidiospore is used here, because it implies any type of
spore arising from within a pycnidium, and does not necessarily imply
asexuality. As these spores may actually be spermatia (non-motile
gametes), the term ‘conidium' is avoided, because of its definite
asexual connotations.
FIGURE 11. — Fuscidea hottentotta Brusse. Ascospores, ascus and
paraphyses. Stippling indicates the amyloid reaction in Lugol’s
iodine solution. Heavy lines delineating the paraphyses, indicate
brown walls. F. Brusse 5312, holotype. Bar = 10 /am.
pm thick. Apothecia blackish, adnate, lecideine, to 1,2
mm across, margins light brown to black, disc black,
plane to concave. Exciple pale brown, anticlinally para-
plectenchymatous, cells 4-6,5 /u,m wide, 4-15,5 pm
long. Hypothecium hyaline, 65—95 pm thick. Hyme-
nium hyaline, 65—75 pm high, only asci J+ blue; epi-
hymenium brown. Asci clavate, eight-spored, tholus J+
blue (Figure 11). Ascospores 8, hyaline but soon
fuscous, curved or reniform, simple but sometimes uni-
septate, 11,5—17,5 x 5-8 pm. Pycnidia hyaline,
ampulliform, about 80 pm wide by about 100 /u.m deep.
Pycnidio spores* narrowly ellipsoid, hyaline, straight,
acrogenous, 3-5 x 1 — 1,4 pm. Chemistry: ramalinolic
acid (major) and sekikaic acid (minor to trace).
This new species of Fuscidea, is fairly closely related
to Fuscidea cyathoides (Ach.) V. Wirth & V6zda, but
has slightly larger ascospores and significantly longer
pycnidiospores. The ascospores are 8,5-14,0 x
4, 0-6, 5 pm (James, Poelt & Wirth 1981; Oberhollenzer
& Wirth 1984), as opposed to 11,5-17,5 x 5—8 /umfor
F. hottentotta. The pycnidiospores of F. cyathoides are
2,8— 3,5 x 1 — 1,5 pm (Coppins in litt.), whereas those
of F. hottentotta are longer, being 3-5 x 1-1,4 pm.
However, the most conspicuous difference between
these two species is the chemistry, with F. cyathoides
containing fumarprotocetraric and protocetraric acids
36
Bothalia 19,1 (1989)
(James, Poelt & Wirth 1981; Oberhollenzer & Wirth,
1984), and F. hottentotta containing ramalinolic acid,
with a minor amount of sekikaic acid. This was deter-
mined by thin-layer chromatographic comparisons of the
hydrolysis products with those of divaricatic and homo-
sekikaic acids, and the hydrolysis data presented by
Culberson (1972). A negative homofluorescein test
(Asahina & Shibata 1954) and microcrystal tests con-
firmed this determination.
Thus far this new species has verifiable specimen
records from Robinson’s Pass, at the western end of the
Outeniqua Mountains and from Caledon, and is therefore
widespread in the southern Cape Province.
CAPE PROVINCE. — 3419 (Caledon): Caledon, Quarzitfelsen ober-
halb d. Bades, c. 300 m. J. Brunnthaler s.n., 28.10.1909 (W 460).
Cited by Zahlbruckner (1932) as Lecidea (Biatora) rivulosa Ach. (=
Fuscidea cyathoides (Ach.) V. Wirth & Vezda).
SPECIMENS OF OTHER SPECIES EXAMINED
Fuscidea cyathoides (Ach.) V. Wirth & V6zda
CZECHOSLOVAKIA. — 4820: Na kfemenci na PleSivci. V. Los
s.n. 1921 . (V. Kufak, Lichenes Bohemiae 574', PRE 890).
ACKNOWLEDGEMENTS
The author is grateful to the director of the Naturhisto-
risches Museum Wien (W) for the loan of the Brunn-
thaler specimen. Thanks are also extended to Dr B. J.
Coppins and Dr V. Wirth for reviewing this paper.
REFERENCES
ASAHINA, Y. & SHIBATA, S. 1954. Chemistry of lichen sub-
stances. Japan Society for the Promotion of Science, Tokyo.
CULBERSON, C. F. 1972. Improved conditions and new data for the
identification of lichen products by a standardized thin-layer chro-
matographic method. Journal of Chromatography 72: 113-125.
JAMES, P., POELT, J. & WIRTH, V. 1981. Fuscidea. In J. Poelt &
A. Vezda, Bestimmungsschlussel europaischer Flechten, Ergan-
zungsheft II: 150- 159. Cramer, Vaduz.
OBERHOLLENZER, H. & WIRTH, V. 1984. Beitrage zur Revision
der Flechtengattung Fuscidea. Beihefte zur Nova Hedwigia 79:
537-595.
ZAHLBRUCKNER, A. 1932. Lichenes in Africa lecti. Annales de
Cryptogamie Exotique 5,3—4: 198-275.
F. BRUSSE
* MS. received: 1988.07.29.
A NEW SPECIES OF M ARON EA (LICHENES) FROM THE DRAKENSBERG
Maronea afroalpina Brusse, sp. nov.
Thallus crustosus, basalticola, albidus, ad 20 mm
diametro, 0, 1 — 1 ,0 mm crassus, rimoso-areolatus. Areo-
lae 0,1 — 1,8 mm diametro. Prothallus non visus. Cor-
tex superior 15—25 /xm crassus. Stratum gonidiale
30—70 pm crassum; algae Protococcoideae, 5 — 11 pm
diametro. Medulla alba, 60- 900 pm crassa. Apothecia
nigella, sessilia, ad 1,5 mm diametro, lecideina, margi-
nibus pallide brunneis vel atrobrunneis, discis atris
pruina alba tectis. Excipulum infeme 40-50 pm cras-
sum, in lateribus 50—70 /im crassum, interne hyalinum
vel stramineum, exteme brunneum, radiatim paraplec-
tenchymatum, cellulis 3—7 pm diametro, interdum
elongatis et usque ad 14 mm longis. Hypothecium stra-
mineum, 50-100 pm crassum, paraplectenchymatum,
cellulis 3-6 pm diametro. Hymenium fertile 70-95
mm altum, hyalinum; epihymenium brunneum. Para-
physes simplices sed apices versus ramosae, septatae,
1,3 — 1,7 pm crassae, capitatae, capitibus 2,8— 5,2 mim
crassis, brunneis. Asci clavati vel acuminate clavati,
tholis J+ caeruleis. Ascosporae numerosae (± 100 vel
ultra), simplices, hyalinae, ellipsoideae, parvae, 5-9 x
3, 2-5, 2 mm. Pycnidia non visa. Thallus acidumdiva-
ricaticum solum continens.
TYPE. — 2828 (Bethlehem): 31 km S of Phuthadi-
tjhaba (Witsieshoek), summit of Western Buttress
(Mont-aux-Sources), on vertical S face of basalt boulder
outcrop on gentle S slope, alt. 3 080 m (— DB). F.
Brusse 5553, 1988.04.05 (PRE, holo.; BM, LD, iso.).
Figure 12.
Thallus crustose, basalticolous, whitish, to 20 mm
diam., 0,1 -1,0 mm thick, rimose-areolate. Areoles
0,1 -1,8 mm diam. Prothallus not seen. Upper cortex
15—25 pm thick. Algal layer 30-70 pm thick; algae
Protococcoid, 5—11 pm diam. Medulla white, 60—900
pm thick. Apothecia blackish, sessile, to 1,5 mm
diam., lecideine; margins light to dark brown, discs
black with white pruina. Exciple 40—50 pm thick
below, 50-70 mm thick on sides, hyaline to stramineous
within, brown towards exterior, radiately paraplecten-
chymatous, cells 3—7 pm diam., sometimes becoming
14 pm long. Hypothecium stramineous, 50—100 pm
thick, paraplectenchymatous, cells 3—6 pm diam.
Hymenium 70—95 /urn high wen fertile, hyaline; epihy-
menium brown. Paraphyses simple but branched
towards apex (Figure 13), septate, 1,3— 1,7 pm thick,
capitate, heads 2,8— 5,2 pm thick, brown. Asci clavate
FIGURE 12. — Maronea afroalpina Brusse, habit. F. Brusse 5553,
holotype. Scale in mm.
Bothalia 19,1 (1989)
37
FIGURE 13. — Maronea afroalpina Brusse, ascus and paraphyses.
The heavy lines at the tips of the paraphyses indicate brown walls.
Stippling indicates the reaction in Lugol’s iodine solution. F.
Brusse 5553 , holotype. Bar = 10 pm.
or acuminate-clavate, tholus J+ blue (Figure 13). Asco-
spores numerous (± 100 or more), simple, hyaline,
ellipsoid, small, 5—9 x 3, 2-5, 2 ^tm. Pycnidia not
seen. Chemistry, divaricatic acid only.
This is a unique new species of Maronea , because the
apothecia are lecideine and the thallus is saxicolous,
whereas all other species of Maronea known up to now
have lecanorine apothecia and are corticolous. The leca-
norine condition is characterized by a very reduced
proper exciple and a well developed thalline exciple, not
always in a protruded state. Oberhollenzer & Wirth
(1984) treated several types of exciples in a single octo-
sporous genus, Fuscidea V. Wirth & V£zda. The spe-
cies with sunken apothecia and reduced exciples could
be considered to have lecanorine apothecia, e.g. Fusci-
dea atlantica (Magn.) James & Poelt, originally de-
scribed as a Lecanora. In a similar vein, Hertel (1984)
has treated several lichens with Lecanora- type ascus
apices, and curved acrogenous pycnidiospores with leci-
deine exciples, as species of Lecanora rather than of
Lecidea, as would have been done in the past. The crea-
tion of a new genus for this lichen, therefore, seemed
unwarranted.
Under the old system of classification of lichens, this
species would key out at Sarcogyne Fw. (Magnusson
1935; Poelt 1969; Zahlbruckner 1926), but this genus
often has a poorly developed thallus (mostly cryptothal-
line), and the exciple is of a different structure and is
dark brown to carbonized in colour. The paraphyses in
Sarcogyne are ecapitate and strongly gelled, unlike the
loose capitate paraphyses of Maronea afroalpina, and
M. constans (Nyl.) Hepp, the type of Maronea
(Hafellner 1984).
The major difference, the one which places Sarcogyne
and Maronea in two different families, is the ascus apex,
which is amply illustrated in Figure 2 and by Hafellner
(1984) for Maronea, and by Brusse (1987, 1988) for the
Acarosporaceae, of which Sarcogyne is a typical
member.
It is as well to state here, that several species of Maro-
nea [section Pseudomaronea (Miill. Arg.) Magn.],
treated by Magnusson (1934), such as Lecanora crassi-
labra Mull. Arg. , have Lecanora- type ascus apices, and
are not true Maronea species.
Maronea afroalpina Brusse is presently known only
from high altitudes in the Drakensberg, at Mont-aux-
Sources.
REFERENCES
BRUSSE, F. 1987. A new yellow Acarospora (Lichenes) from the
Waterberg, South West Africa/Namibia. Bothalia 17: 179-181.
BRUSSE, F. 1988. Lithoglypha, a new lichen genus from Clarens
Sandstone. Bothalia 18: 89-90.
HAFELLNER, J. 1984. Studien in Richtung einer natiirlicheren
Gliederung der Sammelfamilien Lecanoraceae und Lecideaceae.
Beihefte zur Nova Hedwigia 79: 24 1 - 37 1 .
HERTEL, H. 1984. Uber saxicole, lecideoide Flechten der Subantark-
tis. Beihefte zur Nova Hedwigia 79: 399-499.
MAGNUSSON, A. H. 1934. Die Flechtengattung Maronea Mass.
Acta Horti Gotoburgensis 9: 4 1 —66.
MAGNUSSON, A. H. 1935. On the species of Biatorella and Sarco-
gyne in America. Annales de Cryptogamie Exotique 7:
115-146.
OBERHOLLENZER, H. & WIRTH, V 1984. Beitrage zur Revision
der Flechtengattung Fuscidea. Beihefte zur Nova Hedwigia 79:
537-595.
POELT, J. 1969. Bestimmungsschliissel europaischer Flechten.
Cramer, Lehre.
ZAHLBRUCKNER, A. 1926. Spezieller Teil. In A. Engler & K.
Prantl, Die natiirlichen Pflanzenfamilien , edn. 2, vol. 8: 61—270.
Engelmann, Leipzig.
F. BRUSSE
MS. received: 1988.07.29.
38
Bothalia 19,1 (1989)
COMBRETACEAE
A NEW SPECIES OF COMBRETUM FROM NATAL
Combretum mkuzense Carr & Retief, sp. nov., a
C. kirkii Laws, frutice semi-scandenti ramulis termi-
nalibus rigidis nec sinuosis, inflorescentiis puberulis dif-
fert.
TYPE. — Natal, 2732 (Ubombo): Mkuze Game
Reserve headquarters (— CA), Carr 187 (PRE, holo.;
K).
Large shrub up to 5 m tall, widely spreading (to 12 m)
with many near-horizontal lateral branches with apices
sometimes twining; bark mainly smooth, pale buff-
coloured with longitudinal reticulation and occasionally
with dark grey bands, flaking; young branchlets light
green, glabrous to sparsely puberulous and lepidote.
Leaves opposite, exstipulate, petiolate; lamina elliptic
to oblong-elliptic, (30-)45(-80) x (15-)20(-30)
mm, base rounded, apex obtuse or occasionally retuse,
slightly discolorous, upper surface deep green, glabrous
except for scales, sometimes with a few cilia at the base,
veins slightly immersed, lower surface paler green, gla-
brous except for scales and a few trichomes on the mid-
rib, hairy pockets sometimes present, 5-7 pairs of lat-
eral veins raised; petioles 4—9 mm long, lepidote,
puberulous. Scales 45-85 pm in diam., 8-celled with a
number of tangential walls. Inflorescence a spike, up to
25 mm long, singly or in pairs at the apices of previous
year’s growth and in axils on current extensions;
peduncle and rhachis light green, sparsely puberulous,
densely lepidote. Flowers sessile, 4-merous; bracteoles
± 0,5 mm long, lepidote; lower receptacle 2—2,3 mm
long including restriction at junction with upper recep-
tacle, brownish green, sparsely puberulous with dense
yellowish stalked scales; upper receptacle cupuliform,
surrounding the disc, distal portion infundibuliform,
overall length ± 3 mm, width at sepal apices ± 3 mm,
green, sparsely puberulous with whitish scales. Sepals
triangular, ± 0,8 mm long, apices fringed with short
whitish hairs. Petals 4, pale green, spathulate, gla-
brous, ± 1,5 mm long. Stamens 8; filaments light
green, 6-6,5 mm long; anthers light yellow, 1 mm long.
FIGURE 14. — Combretum mkuzense
Carr & Retief, holotype in PRE.
Bothalia 19,1 (1989)
39
Style light green, ±3,5 mm long with stigma slightly
expanded and darkened. Disc square in outline with long
silvery hairs. Fruit 4-winged, up to 50 x 50 mm, outline
subcircular with a wide shallow basal notch and a small
apical notch, apical peg up to 1 mm long, lepidote, stipe
up to 20 mm long, densely lepidote; wings brown-tinged
limegreen when ripening, cinnamon when ripe. Seed
ellipsoidal, up to 16 x 9 mm, dark purplish brown.
Cotyledons 2, up to 33 x 45 mm, transversely elliptic,
arising above soil level; petioles ± 3 mm long. Figure
14.
NATAL. — 2632 (Bella Vista): 15 km ENE of Makane’s Drift
(-CD), Stephen 717 (PRE). 2732 (Ubombo): 3 miles W of Sihang-
wane store (-AA), Moll 5370 (PRE); 1 mile E of Pongola pont
(— AB), Strey & Moll 3778 (PRE); Mkuze Game Reserve (-CA),
White 10388 (FHO, PRE); eastern side of farm ‘Shotton 13810’
(-CD), Ward 8793 (PRE).
Combretum mkuzense, which is evidently rare, occurs
in the northernmost part of Natal. It is found in mixed
woodland in association with other combretaceous spe-
cies, Acacia, Sclerocarya, Strychnos, Dialium, Newto-
nia and Albizia. It grows in deep sand at altitudes of up
to 100 m and within 60 km of the coast.
C. mkuzense normally flowers in September but a
second flowering late in March has also been recorded.
The flowers are sweetly scented and appear to be bee-
pollinated.
C. mkuzense is placed in the subgenus Combretum,
section Macrostigmatea. According to Exell (1978) this
section comprises three species, namely C. schumannii,
C. kirkii and C. gillettianum. The section can be divided
into two subsections on the basis of the disc, which is
glabrous with only a very short free margin in C. schu-
mannii and with a pilose margin free for ± 1 mm in C.
kirkii and C. gillettianum. C. mkuzense has a disc with
a pilose margin and is therefore thought to be more
closely related to C. kirkii and C. gillettianum and to
C. schumannii (Table 1).
C. mkuzense is most closely allied to C. kirkii. There
are, however, several differences between the two spe-
cies. The seeds of C. kirkii are as large as 25 x 9 mm
while the seeds of C. mkuzense seen did not exceed 16
x 9 mm. Unlike the new species which has a peduncle
and rachis sparsely puberulous and densely lepidote, the
inflorescence of C. kirkii is glabrous except for the
scales. Habit differences clearly separate these two spec-
ies. C. kirkii is a liane with slender, sinuous, flexible,
readily damaged apices many of which die back in
winter. C. mkuzense is a scrambling shrub with
numerous characteristic elongated lateral branches
taking off in opposite pairs and at right angles. These
branches are fairly straight and maintain appreciable
rigidity right up to the apices but sometimes the apices
may twine. It has been established, using thin film chro-
matography, that for each species of Combretum there
is a characteristic profile of compounds in the leaf mate-
rial (Carr & Rogers 1987). An examination of the pro-
files of C. mkuzense and C. kirkii shows similarities
but also a significant difference.
Fruits of C. zeyheri, a tree which occurs in the same
area as the new species, are similar to those of C. mku-
zense but the habits of the two species differ so widely
that they should not be easily confused. C. zeyheri is a
small to medium-sized tree while C. mkuzense is a
scrambling shrub. The scales of the new species (Figure
15) agree well with those of other representatives of the
section Macrostigmatea. They differ markedly from
those of C. xanthothyrsum (sect. Chionanthoideae)
(Figure 16) a species which has sometimes been con-
fused with our species.
TABLE 1. — A comparison of Combretum mkuzense, C. kirkii and C. gillettianum. Based partly on Exell (1978)
40
Bothalia 19,1 (1989)
FIGURE 15. — Scales on the leaf surface of Combretum mkuzense Carr
& Relief, X 360. Carr 187. Scanning electron micrograph taken
at the Royal Botanic Gardens, Kew.
FIGURE 16. — A scale on the leaf surface of Combretum xanthothyr-
sum Engl. & Diels, x 600. Proctor 2759. Scanning electron
micrograph taken at the Royal Botanic Gardens, Kew.
REFERENCES
CARR, J. D. & ROGERS, B. C. 1987. Chemosystematic studies of
the genus Combretum (Combretaceae), part 1. South African
Journal of Botany 53: 173—176.
EXELL, A. W. 1978. Combretaceae in Flora Zambesiaca 4:
100-183.
J.D. CARR and E. RETIEF
MS. received: 1987.12.28.
Bothalia 19, 1:41^3(1989)
Leaf anatomy of the South African Danthonieae (Poaceae). XVIII.
Centropodia mossamedensis
R. P. ELLIS*
Keywords: Centropodia mossamedensis, C4, Danthonieae, Kranz, leaf anatomy, Poaceae
ABSTRACT
The leaf blade anatomy of Centropodia mossamedensis (Rendle) T. A. Cope [= Asthenatherum mossamedense
(Rendle) Conert] is described and illustrated. This description is based on freshly fixed material and confirms that this
species has Kranz anatomy with the C4 photosynthetic pathway. The anatomy differs little from that of C. glauca and both
undoubtedly belong to the same genus which is justifiably separated from the other danthonoid genera.
UITTREKSEL
Die blaaranatomie van Centropodia mossamedensis (Rendle) T. A. Cope [= Asthenatherum mossamedense (Rendle)
Conert] word beskryf en ge'illustreer. Hierdie beskrywing is gebaseer op vars gefikseerde materiaal en bevestig dat hierdie
spesie die Kranz-tipe anatomie en C4-fotosintese besit. Die blaaranatomie wyk weinig af van die van C. glauca en albei
spesies behoort ongetwyfeld aan dieselfde genus. Hierdie studie bevestig dat Centropodia van die ander genera in die
Danthonieae geskei behoort te word.
INTRODUCTION
In a previous paper (Ellis 1984) in this series the ana-
tomy of Centropodia mossamedensis (Rendle) T. A.
Cope [= Asthenatherum mossamedense (Rendle)
Conert] (Cope 1983) was briefly described. This descrip-
tion was based on herbarium material and the anatomical
preparations were not of a very high quality. Subsequent
to the above study, fresh material of C. mossamedensis
was collected and fixed in the field, yielding good qual-
ity leaf blade transverse sections. The results are de-
scribed, illustrated and compared with the leaf anatomy
of C. glauca (Nees) T. A. Cope [= Asthenatherum glau-
cum (Nees) Nevski],
MATERIALS AND METHODS
Plants of C. mossamedensis were collected in South
West Africa/Namibia. Herbarium voucher specimens
were prepared for verification by the National Herbarium
(PRE) where they are now housed.
Leaf blade segments were removed and immediately
fixed in FA A. Leaf blade transverse sections and abaxial
epidermal scrapes were prepared following the methods
outlined in a previous paper in this series (Ellis 1988).
The standardized terminology of Ellis (1976, 1979)
was used for the anatomical descriptions together with
the following abbreviations:
vb/s — vascular bundle/s
1’ vb/s — first order vascular bundle/s
2'vb/s — second order vascular bundle/s
3’ vb/s — third order vascular bundle/s
ibs — inner bundle sheath; mestome sheath
obs — outer bundle sheath; parenchyma sheath
* Botanical Research Institute, Department of Agriculture and Water
Supply , Private Bag X 1 0 1 , Pretoria 000 1 .
, MS. received: 1988.06.09.
Specimens examined:
Centropodia mossamedensis
SWA/NAMIBIA. — 2014 (Welwitschia): Damaraland, 60 km W of
Khorixas on road to Skeleton Coast (-AB). Ellis 4750. 2116 (Oka-
handja): Okahandja Dist. ; 26 km W on road to Swakopmund, Otji-
tundu River crossing (-DD), Ellis 4725, 4726.
ANATOMICAL DESCRIPTION OF CENTROPODIA
MOSSAMEDENSIS
Leaf in transverse section
Leaf outline : expanded and flat lamina (Figure 1A &
E). Ribs and furrows : very slight adaxial ribs (Figure IB
& F); slight furrows between all vbs; ribs rounded.
Abaxial ribs and furrows more pronounced than adaxial
ones (Figure IB & F); furrows between all vbs and ribs
rounded. Median vascular bundle : structurally indisting-
uishable from lateral l’vbs. Vascular bundle arrange-
ment: 9 or 13 l’vbs in leaf section: 3, 4 or 5 3 ’vbs
between consecutive l’vbs except laterally where fewer
3’ vbs are present (Figure 1A & E); 2’ vbs absent; all vbs
centrally located in blade. Vascular bundle structure:
3’vbs slightly elliptical with well developed xylem and
phloem tissue and an ibs (Figure IB, C & F); l’vbs
elliptical (Figure IB, C & F); phloem adjoins the ibs;
metaxylem vessels narrow, with a diameter slightly less
than that of the obs cells; diameter greater than that of the
ibs cells. Vascular bundle sheaths: double; slightly ellip-
tical to almost rounded; both sheaths entire around all
vbs (Figure 1C); no extensions although a few Kranz
cells may be located outside the outer sheath (Figure
1C); parenchyma sheath cells very numerous (15—26),
regular in size and shape, fan-shaped with straight radial
walls and inflated outer tangential walls; specialized,
large, centripetally situated chloroplasts conspicuous; ibs
complete around V and 3’vbs; cells with slight secon-
dary thickening. Sclerenchyma: small adaxial girders as-
sociated with all l’vbs and strands with the 3’vbs; taper
toward the bundles; similar abaxial girders and strands
42
Bothalia 19,1 (1989)
FIGURE 1. — Leaf anatomy of Centropodia mossamedensis. A-D, Ellis 4726: A, outline of flat, expanded blade, X 100; B, transectional
anatomy, x 250; C, anatomical detail showing Kranz anatomy with centripetal specialized parenchyma sheath chloroplasts, x 400; D,
abaxial epidermis with costal prickles and intercostal stomata, interference contrast, x 250. E-G, Ellis 4750: E, blade outline, x 100; F,
Kranz transectional anatomy, x 250; G, abaxial epidermis with few costal prickles, X 250. H, Ellis 4725, abaxial epidermis with well
developed prickles and intercostal microhairs, X 250.
associated with all vbs except that strands sometimes
also associated even with l’vbs (Figure IB & C); fibres
thick- walled (Figure IF) or thin- walled (Figure IB & C)
but never lignified. No sclerenchyma between bundles.
Small sclerenchyma cap in margin. Mesophyll : radiate
chlorenchyma (Figure IB, C & F); single layer of tabular
cells surround bundles completely (Figure IB & C) or
with small interruptions due to girders (Figure IF); late-
ral cell count 2 or 3. No colourless cells associated with
the bulliform cells. Adaxial epidermis : fan-shaped bulli-
form cell groups with central cell shield-shaped; occupy
less than half the leaf thickness; epidermal cells with
slightly thickened outer walls; macrohairs not present;
small costal hooks present (Figure IB) or absent (Figure
IF) ; no papillae. Abaxial epidermis : small bulliform-like
cells at bases of furrows; macrohairs absent; costal
prickles present (Figure IB) or absent (Figure IF); no
papillae.
Abaxial epidermis in surface view
Intercostal long cells: elongated with side walls al-
most parallel (Figure ID) to inflated, fusiform (Figure
IG) ; walls not sinuous; cells adjoin one another or sepa-
Bothalia 19,1 (1989)
43
rated by short cells or stomata. Stomata : low dome-
shaped (Figure ID & G); not in regular files and occur in
most long cell files; 1 or 2 interstomatal cells between
successive stomata in a file. Intercostal short cells:
absent or very rare (Figure ID) or irregular (Figure 1G);
no cork or silica cells but just very short epidermal cells.
Papillae: absent. Prickles: costal, either very common
(Figure ID & H) or rare (Figure 1G); barbs either well
developed or virtually absent. Hooks: absent. Micro-
hairs: present on all specimens but rare (Figure 1H); both
cells elongated but distal cell not tapering to a pointed
apex; two cells about equal in length. Macrohairs: ab-
sent. Silica bodies: variable, irregular dumbbell-shaped
(Figure 1G), short and narrow (Figure 1H) or horizon-
tally elongated rectangular (Figure ID); occur through-
out costal zones; sometimes associated with cork cells
but often not.
DISCUSSION AND CONCLUSIONS
The transectional leaf anatomy compares very closely
with that of C. glauca (Ellis 1984). Both are undoub-
tedly Kranz with radiate mesophyll, and no chloren-
chyma cells are more than one cell distant from a Kranz
cell. This indicates the presence of the C4 photosynthetic
pathway which is confirmed by a 12c/13c ratio of — 12,6
°/00 (De Winter & Hardy 8021). The outer bundle sheath
has a regular outline and is Kranz with centripetally lo-
cated specialized chloroplasts. This structure is typical of
that characteristic of the NAD-me subtype of the C4 pho-
tosynthetic pathway but this has yet to be confirmed as
Centropodia has not yet been biochemically typed
(Hattersley 1987).
Anatomical differences between C. mossamedensis
and C. glauca are only minor, particularly the leaf in
transverse section. Vessel element diameter is proportio-
nally greater in C. mossamedensis where they are
slightly wider than the inner bundle sheath cells but they
are, nevertheless, still relatively narrow. The bulliform
cells occupy less than half the leaf thickness in C. mossa-
medensis but in C. glauca they are equal to at least half
the leaf thickness. In transection no elongated prickle
hairs are evident as in many C. glauca specimens.
Superficially the abaxial epidermis differs consider-
ably from that of C. glauca. No interlocking prickles
resembling macrohairs are present and the unique macro-
hairs with corrugated cell walls, as in C. glauca var.
lasiophyllum, were not observed.
These epidermal differences are visually very striking
but it must be remembered that C. glauca exhibits conti-
nuous anatomical variation from those specimens with
conspicuous interlocking prickle hairs to specimens wi-
thout this hair type (Ellis 1984). This variation pattern is
associated with an ecological dine from the extremely
arid Namib Desert eastward to the Kalahari. The anat-
omy of C. mossamedensis appears to be a northward
expression of this cline along a moisture gradient and C.
mossamedensis may merely represent a continuation of
this reduction trend evident in C. glauca.
The two species are distinct morphologically (Conert
1962) and also appear to occupy different niches. C.
glauca is a species of the loose red sands of the Kalahari
dunes whereas C. mossamedensis is confined to gravelly
or coarse waterborne sands in dry watercourses. Their
separation at species level is, therefore, not questioned
by this study even though these two species do not ex-
hibit significant leaf anatomical differences.
ACKNOWLEDGEMENTS
H. Ebertsohn is thanked for technical assistance,
A. Romanowski for the photography and M. van der
Merwe for typing the manuscript.
REFERENCES
CONERT, H. J. 1962. Uber die Gramineen-Gattung Asthenatherum
Nevski. Senckenbergiana Biologica 43: 239-266.
COPE, T. A. 1983. Centropodia. an earlier name for Asthenatherum
(Gramineae). Kew Bulletin 37: 657-658.
ELLIS, R. P. 1976. A procedure for standardizing comparative leaf
anatomy in the Poaceae. I. The leaf blade as viewed in transverse
section. Bothalia 12: 65-109.
ELLIS, R. P. 1979. A procedure for standardizing comparative leaf
anatomy in the Poaceae II. The epidermis as seen in surface
view. Bothalia 12: 641-672.
ELLIS, R. P. 1984. Leaf anatomy of the South African Danthonieae
(Poaceae). I X. Asthenatherum glaucum. Bothalia 15: 153—159.
ELLIS, R. P. 1988. Leaf anatomy of the South African Danthonieae
(Poaceae). XVI. The genus Urochlaena. Bothalia 18: 101-104.
HATTERSLEY, P. W. 1987. Variations in photosynthetic pathway. In
T. R. Soderstrom et a!.. Grass systematics and evolution. Smith-
sonian Institution Press, Washington DC.
Bothalia 19, 1:45-52(1989)
Kranz distinctive cells in the culm of Arundinella (Arundinelleae
Panicoideae; Poaceae)
EV ANGELINA SANCHEZ*, MIRTA O. ARRIAGA* and ROGER P. ELLIS**
Keywords: anatomy, Arundinella, C4, culm, distinctive cells, double bundle sheath, NADP-me
ABSTRACT
The transectional anatomy of photosynthetic flowering culms of Arundinella berteroniana (Schult.) Hitchc. & Chase
and A. hispida (Willd.) Kuntze from South America and A. nepalensis Trin. from Africa is described and illustrated. The
vascular bundles are arranged in three distinct rings, the outermost being external to a continuous sclerenchymatous band.
Each of these peripheral bundles is surrounded by two bundle sheaths, a complete mestome sheath and an incomplete,
outer, parenchymatous Kranz sheath, the cells of which contain large, specialized chloroplasts. Kranz bundle sheath
extensions are also present. The chlorenchyma tissue is also located in this narrow peripheral zone and is interrupted by the
vascular bundles and their associated sclerenchyma. Dispersed throughout the chlorenchyma are small groups of Kranz
distinctive cells, identical in structure to the outer bundle sheath cells. No chlorenchyma cell is, therefore, more than two
cells distant from a Kranz cell. The structure of the chlorenchyma and bundle sheaths indicates that the C4 photosynthetic
pathway is operative in these culms.
This study clearly demonstrates the presence of the peculiar distinctive cells in the culms as well as in the leaves of
Arundinella. Also of interest is the presence of an inner bundle sheath in the vascular bundles of the culm whereas the
bundles of the leaves possess only a single sheath. It has already been shown that Arundinella is a NADP-me C4 type and the
anatomical predictor of a single Kranz sheath for NADP-me species, therefore, either does not hold in the culms of this
genus or the culms are not NADP-me. This is only the second reported breakdown of this association between MS anatomy
and the NADP-me biochemical C4 type.
UITTREKSEL
Die anatomie van dwarssnee van fotosintetiese bloeiwysehalms van Arundinella berteroniana (Schult.) Hitchc. &
Chase en A. hispida (Willd.) Kuntze vanaf Suid-Amerika en A. nepalensis Trin. van Afrika word beskryf en geillustreer.
Die vaatbondels is in drie duidelike ringe gerangskik, met die buitenste vaatbondels aan die buitekant van ‘n aaneenlopende
sklerenchiemband. Hierdie periferale vaatbondels word elk omring deur twee vaatbondelskedes, ’n volledige
mestoomskede en ’n onvolledige, buitenste parenchiematiese Kranz-skede waarvan die selle groot, gespesialiseerde chloro-
plaste bevat. Kranz-verlengings van die bondelskede is ook teenwoordig. Chlorenchiemweefsel is ook teenwoordig in
hierdie smal periferale sone en word onderbreek deur die vaatbondels en hul meegaande sklerenchiem. Klein groepies
Kranz-kenmerkende selle wat struktureel identies is aan die buitenste vaatbondelskedeselle is deur die chlorenchiem
versprei. Geen chlorenchiemsel is dus meer as twee selle van ’n Kranz-sel verwyder nie. Die struktuur van die chlorenchiem
en die vaatbondelskede dui aan dat die C4- fotosintetiese weg in die halms gevolg word.
Hierdie studie toon duidelik die teenwoordigheid van die unieke kenmerkende selle in beide die halms en blare van
Arundinella. Ook opvallend was die teenwoordigheid van ’n binneste vaatbondelskede in die vaatbondels van die halm,
terwyl die vaatbondels van die blare slegs ’n enkele skede besit. Daar is reeds getoon dat Arundinella ’n NADP-me C4- tipe
is, en die anatomiese aanwyser van ’n enkele Kranz-skede vir NADP-me-spesie is dus of nie van toepassing op die halms
van hierdie genus nie of die halms is nie NADP-me nie. Hierdie is slegs die tweede bevestiging van ’n NADP-me
biochemiese C4- tipe sonder MS anatomie.
INTRODUCTION
Some species of Arundinella Radii are characterized
by the presence of Kranz distinctive cells in the meso-
phyll of the leaf blades. Arundinella is a C4 genus which
possesses the Kranz syndrome (Brown 1977) and these
distinctive cells are very similar to those of the Kranz
mestome sheath which surrounds the vascular bundles
(Brown 1975). The distinctive cells have thicker walls
than those of the radially arranged chlorenchyma cells
between which they are embedded (Carolin et al. 1973)
and these walls also stain heavily. They contain abun-
dant specialized chloroplasts which store starch (Brown
1975; Renvoize 1982a). In transection they occur either
* CONICET, Museo Argentine) de Ciencias Naturales ‘Bernardino
Rivadavia’, Av. A. Gallardo 470, 1405 Buenos Aires, Argentina.
** Botanical Research Institute, Department of Agriculture and Water
Supply , Private Bag X 1 0 1 , Pretoria 000 1 .
MS. received: 1988.06.22.
singly between the vascular bundles or are found in
groups of two to six cells without accompanying vascu-
lar tissue.
Some authors are of the opinion that the distinctive
cell files connect with the parenchyma sheath cells
(Carolin et al. 1973; Ellis 1977). However, in parader-
mal view it is evident that they are not continuous with
the vascular tissue but are contiguous isolated Kranz cell
strands that lie parallel to the vascular bundles and are
not in contact with them (Crookston & Moss 1973).
They are presumably functionally linked to the vascular
bundles at intervals by cross veins (Crookston & Moss
1973; Crookston 1980; Renvoize 1982a). Where the
cross veins traverse strands of distinctive cells some of
these Kranz cells become appressed to the cross vein, so
forming a functional link with the vascular tissue.
These cells were first reported by Vickery (1935) in
the leaf blade transection of A. nepalensis Trin. Tateoka
46
Bothalia 19,1 (1989)
TABLE 1. — Historical review of the study of Kranz distinctive cells in grass leaf blades
Bothalia 19,1(1989)
47
demonstrated that these cells are isolated photosynthetic
carbon reduction (PCR, Kranz) strands embedded in the
primary carbon assimilation (PCA) chlorenchyma tissue;
and they exhibit NADP-me activity (Reger & Yates
1979). Ultrastructurally they are also seen to be similar
to the Kranz mestome sheath cells. Both have large agra-
nal chloroplasts containing numerous starch grains
(Crookston & Moss 1973; Crookston 1980) and a sub-
erized lamella is located in the cell walls (Hattersley &
Browning 1981). The chloroplasts of the chlorenchyma
cells, on the other hand, are free of starch and have well
developed grana and the cell walls lack a suberized
lamella. The distinctive cells are, therefore, undoubtedly
Kranz cells and will be designated as such.
From Table 1 it can also be seen that distinctive cells
have only been reliably reported in four genera belong-
ing to four small tribes of the Panicoideae: Arundinella
of the Arundinelleae (Tateoka 1956a, 1958); Garnotia of
the Gamotieae (Tateoka 1956b, 1958); Arthropogon of
the Arthropogoneae (Tateoka 1963) and Anthephora of
the Anthephoreae (Johnson & Brown 1973). Watson et
al. (1986) record ‘circular cells’ in nine genera of the
Panicoideae. The reports of distinctive cells in Trichop-
teryx and Loudetia of the Arundinelleae (Brown 1977)
appear to be misleading and probably refer to the very
reduced vascular bundles surrounded by only three or
four Kranz sheath cells which are known from these
genera which also lack cross veins (Renvoize 1982a).
Many authors have confirmed the occurrence of distinc-
tive cells in Arundinella and Garnotia (Table 1) and
Arthropogon xerachne (Sanchez & Arriaga 1988) but
verification of their reported presence in Anthephora is
required. We have examined leaves of 25 specimens of
four species of Anthephora, all of which have very small
minor vascular bundles consisting of only three or four
bundle sheath cells surrounding a minute vascular
strand. Consequently we query the reported presence of
distinctive cells in this genus. We have also examined
leaves of Tristachya lejostachya and Loudetia flammida
(Sanchez & Arriaga 1988), L. pedicellata and L. simplex
(Ellis 1977) without detecting the presence of distinctive
cells. In all these cases a few xylem vessels were de-
tected in association with the Kranz cells but these are
not considered to be distinctive cells.
Nevertheless, as presently known, distinctive cells are
characteristic of and unique to these four small tribes of
the Panicoideae and may indicate phylogenetic
relationships between them. Johnson & Brown (1973)
consider the possession of distinctive cells to be suffi-
cient grounds for considering these four tribes as consti-
tuting one tribe or even a supertribe. Garnotia and Arun-
dinella, in particular, are very closely related (Renvoize
1982b) and appear to constitute a distinct and related
group of genera sharing this interesting anatomical fea-
ture, as well as spikelet characteristics. Distinctive cells
are not a characteristic of the tribe Arundinelleae but are
only a feature of some species of Arundinella (Renvoize
1982a).
The culms (aerial stems) of grasses display consider-
able anatomical variation but, in contrast to the leaf
blade, have been poorly documented (Sabnis 1921; Can-
field 1933; De Wet 1960; Metcalfe 1960; Auquier &
Somers 1967). Some of these studies include members
of the Arundinelleae. De Wet (1960) describes the peri-
pheral vascular bundles of the culm of Arundinella as
being surrounded by a parenchymatous bundle sheath
composed of small cells. Auquier & Somers (1967) con-
sider the anatomical structure of the culm of Arundinella
as belonging to the ‘panicoid type’ with the peripheral
bundles surrounded by a well developed parenchymatous
sheath. None of these authors refer to the presence of
Kranz anatomy in the cortical zone of the culm. Sanchez
(1979, 1981a, 1981b, 1983a, 1983b, 1984) is the first
worker to report the presence and development of Kranz
anatomy in flowering and stoloniferous culms. Kranz
anatomy is only developed in the upper exposed parts of
flowering culms and not the basal parts which are
covered by the leaf sheath (Sanchez 1981a); it is there-
fore essential to examine comparative material.
The objective of this study is to determine whether
Arundinella exhibits Kranz structure in the flowering
culm. If this is so then it will also be of interest to see
whether distinctive Kranz cells are also present. This
paper describes the structure and arrangement of these
cells in the culms of three species of Arundinella : A.
berteroniana (Schult.) Hitchc. & Chase and A. hispida
(Willd.) Kuntze from Argentina and A. nepalensis from
South Africa. A. hispida from the New World and
A. nepalensis from the Old World appear to be closely
related and Phipps (1967) included them both in the
Nepalenses series which he considered to be central to
the genus.
MATERIALS AND METHODS
Transverse and longitudinal sections of flowering
culms were made from segments taken from the centre of
the first intemode below the inflorescence. Both herba-
rium and field collected material fixed in FAA was used.
Sections were either free-hand or the material was desili-
cified, embedded in wax and sectioned on a rotary
microtome. These sections were stained with Alcian
Blue and Safranin (Cutler 1978) or Fast Green and Saffa-
nin (Johansen 1940). Uncleared sections were soaked in
5% NaOH for 5-10 minutes to restore turgidity and
were then used to observe chloroplast position in the
Kranz cells.
Material examined
A. berteroniana
BA 27/2263; BA 18993; Giusti 1214 (BA); Parodi 1781 (BA); Ver-
voorst & Cuezzo 7731 (CTES).
A. hispida
BA 11258; BA 16098; Schinini & Tressens 24545 (BA); Schinini et
al. 17348 (CTES); Quarin 409 (CTES); Schulz 3469 (CTES); Royo
238 (CTES).
A. nepalensis
BAA 19752; Ellis 479 (PRE 61722)*, 1218 (PRE 61723)*, 1368
(PRE 61724)*, 1436 (PRE 61725)*, 1481 (PRE 61726)*, 1617 (PRE
61727)*, 2116 (PRE 61728)*, 3358 (PRE 622413)*, 4977.
ANATOMICAL DESCRIPTION OF THE CULM
The general shape of the sections is circular (Figures
1A; 2A, C & E; 3A; 4A) with a smooth or slightly undu-
lating outline. The diameter of the transections was
* only leaf blade material examined.
48
Bothalia 19,1 (1989)
FIGURE 1 . — Schematic diagrams of the culm anatomy of Arundinella.
A, transection of the first intemode of a flowering culm; B, tran-
sectional detail showing arrangement of vascular bundles (v) and
the location of the sclerenchymatous ring (s); C, detail of the
peripheral zone showing distinctive cells (d) in the chlorenchyma
(c); D, paradermal longitudinal view of Kranz distinctive cells (d)
with a transverse connection of xylem (x).
found to be ± 2 mm in A . berteroniana and A . hispida
and ± 1 ,5 mm in A. nepalensis.
The epidermis is simple. Stomata were observed adja-
cent to the chlorenchyma zones and the subsidiary cells
are at the same level as the epidermal cells (Figures 2D;
4C). No prickle hairs or hooks were observed.
A discontinuous ring of chlorenchyma is present be-
low the epidermis. This ring consists of 1-6 layers of
rachymorph cells none of which are more than two cells
distant from a Kranz cell. This tissue is interrupted at
regular intervals by the sclerenchyma girders of the peri-
pheral vascular bundles (Figures IB; 2B, D & F; 3C;
4B). This peripheral zone is narrow, occupying V10— ‘/12
of the total diameter of the culm transection.
This peripheral chlorenchymatous zone is bounded
internally by a continuous ring of sclerenchyma consist-
ing of 1 — 3( — 4) layers of lignified, fibrous cells (Figures
IB; 2B, D & F; 3C; 4B). This unicylindrical sclerenchy-
matous ring encloses the parenchymatous pith, the centre
of which may be hollow (Figures 2A; 4A) or not (Figures
2C; 3A).
The vascular bundles are arranged in 3( — 6) distinct
circles or rings and are alternately spaced (Figures 1A;
2A, C & E; 3A; 4A), although rarely 4 or 5 circles of
bundles are present. The bundles can be divided into two
types: peripheral bundles and non- peripheral bundles.
The peripheral vascular bundles are external to the
sclerenchymatous ring and occur in two distinct size
classes. The larger first or second order bundles with
metaxylem vessels are partially attached to the scleren-
chyma ring and are linked to the epidermis by scleren-
chyma girders (Figures 1C; 2B, D & F; 3C; 4B). These
bundles are surrounded by two bundle sheaths, a com-
plete mestome sheath and an incomplete Kranz paren-
chyma sheath (Figures 1C; 2B & D; 4C). Bundle sheath
extensions of the latter sheath may extend along the outer
surface of the sclerenchymatous ring for a distance of
from 2-3 cells (Figures 1C; 4C). Some of the larger
bundles may exhibit a partial or complete periphloematic
sheath (Caro 1961). The smaller third order peripheral
bundles without metaxylem vessels do not have scleren-
chyma girders attaching them to the epidermis (Figures
1C; 2F; 4C). These bundles are surrounded by only a
single incomplete Kranz parenchyma sheath, interrupted
where it adjoins the sclerenchymatous ring. These smal-
ler bundles may also possess bundle sheath extensions.
Distinctive Kranz cells are present in the peripheral
zone (Figures 1C; 2B; 4B & C). These cells are similar
in structure to the Kranz parenchyma sheath cells of all
the vascular bundles of this zone. They have thicker
walls and larger, predominately centrifugally located
chloroplasts than do the chlorenchyma cells and are
found singly or in groups of 1 — 3( — 4) without associated
xylem or phloem cells.
In paradermal view the Kranz distinctive cells form
long rows, 1 or 2 cells wide, that lie parallel to the
vascular bundles but are not accompanied by vascular
tissue (Figure ID). They are connected by lateral cross-
veins which traverse from one vascular bundle to an-
other. These interconnections consist only of xylem ele-
ments and they are not accompanied by bundle sheath
cells. No phloem cells were seen. Interconnections are
relatively common in the chlorenchymatous zone in the
culms of Arundinella.
A second and third (seldom a fourth or fifth) circle of
collateral vascular bundles is situated on the inner side of
Bothalia 19,1 (1989)
49
FIGURE 2. — Culm anatomy of Arundinella hispida as seen in transverse section. A-B, Schinini & Tressens 24545: A, culm outline with
hollow pith (p), X 50; B, peripheral zone with vascular bundles, chlorenchyma and distinctive cells (d) external to the continuous
sclerencyma ring (s), X 400. C-D, Schinini etal. 17348: C, culm outline showing solid pith (p), x 50; D, detail of peripheral Kranz
tissue with distinctive cells in the chlorenchyma external to the sclerenchyma ring (s), X 400. E— F, BA 11258: E, culm outline,
X 50; F, detail of peripheral zone with external vascular bundles, the sclerenchyma ring (s) and interal vascular bundles in the pith,
x 250.
the sclerenchymatous ring. The second circle is partially
embedded in these fibres but the other circles are located
in the parenchymatous ground tissue of the pith. These
inner two circles of bundles consist only of larger, first
order vascular bundles surrounded by a single mestome
sheath (Figures IB; 2B, D & F; 3B, C;4B).
DISCUSSION AND CONCLUSIONS
The presence of these rare and specialized Kranz dis-
tinctive cells has previously been confirmed in only two
genera, Arundinella and Garnotia. However, there has
been much confusion in the literature regarding the ter-
minology for these cells (Table 1) and, although Watson
et al. (1986) record circular cells in nine genera, it is not
clear whether these are all homologous with the particu-
lar cells described here. This situation is confusing and
unsatisfactory and it is proposed that the term Kranz
distinctive cells should in future be employed only for
isolated groups of, or single, Kranz cells in the meso-
phyll which are not associated with contiguous vascular
tissue. The term distinctive cells enjoys historical prece-
dent (Tateoka 1956b) and Kranz distinctive cells also
gives an indication of their function. Furthermore this
designation is explicit even when translated into other
50
Bothalia 19,1 (1989)
FIGURE 3. — Transectional culm anatomy of Arundinella bertero-
niana, Giusti 1214. A, outline of culm with solid pith (p), X 100;
B, detail of peripheral zone with internal vascular bundles, scle-
renchymatous ring (s) and outermost Kranz zone, x 250; C,
outer, peripheral zone showing darkly stained, thickened lignified
tissue of the mestome sheaths, sclerenchyma girders and the scle-
renchyma ring, x 250.
FIGURE 4. — Culm anatomy of Arundinella nepalensis, Ellis 4977. A,
hollow pith (p) with two rings of vascular bundles in the outermost
pith layers, x 100; B, peripheral zone showing inner vascular
bundles, sclerenchyma ring and chlorenchyma tissue (c) inter-
rupted by outer ring of Kranz vascular bundles, x 250; C, detail
of chlorenchyma, distinctive cells (d), Kranz outer bundle sheath
and sheath extensions, inner mestome sheath and sclerenchyma
ring, x 400.
FIGURE 5. — Leaf blade anatomy of Arundinella in transverse section. A-B, A. nepalensis: A, distribution of distinctive cells in the
chlorenchyma between all vascular bundles, Ellis 1481, x 100; B, anatomical detail of distinctive cells (d) and single, Kranz vascular
bundle sheath (m), Ellis 1617, x 400. C, A. berteroniana with distinctive cell groups dispersed in the chlorenchyma tissue, Davidse
32217, x 400.
Bothalia 19,1 (1989)
51
languages. Standardization on this most commonly used
terminology should ensure the elimination of any future
misunderstanding.
This study has clearly shown that the unique Kranz
distinctive cells of the leaf blades of Arundinella (Figure
5A, B & C) also occur in the photosynthetic culms. In
culm transections they are seen to be rounded cells with
thickened walls which contain specialized chloroplasts.
They are distinctly larger than the chlorenchyma cells in
which they are embedded. These distinctive cells occur
as isolated groups or strands in the chlorenchyma com-
prising one to three contiguous Kranz cells without ac-
companying vascular tissue. In paradermal view it is
clear that they do not form part of the vascular tissue
and, therefore cannot be considered as degenerate inter-
calary vascular bundles (Brown 1977). Instead they are
seen to be long, isolated Kranz cell columns not physi-
cally connected to the vascular bundles but presumably
functionally linked at regular intervals by vascular
strands. This structure is virtually identical to that
described for the distinctive cells of the leaf blades of
several Arundinella species. There can be no doubt that
these cells which are reported here in the culms of Arun-
dinella for the first time, represent homologous struc-
tures in leaf blades and photosynthetic culms. The occur-
rence of Kranz anatomy in both culms and leaves is
noteworthy, because grass species with Kranz leaf ana-
tomy do not necessarily exhibit Kranz structure in the
culm as well (Sanchez unpublished).
The fact that the peripheral first order vascular bundles
are surrounded by two bundle sheaths, a complete mes-
tome sheath and an interrupted Kranz parenchyma
sheath, is of considerable interest because this configura-
tion (Figure 5) differs from the condition in the leaf blade
as reported in the literature. Many authors record a single
bundle sheath in the leaf of Arundinella species, which is
referred to as the XyMS condition by Hattersley & Wat-
son (1976) or as the MS type by Brown (1977). Exam-
ples are Vickery (1935), Brown (1958, 1977), Metcalfe
(I960), Jacques-Felix (1962), Crookston & Moss
(1973), Hattersley & Watson (1975, 1976), Ellis (1977)
and Renvoize (1982a). Other workers have reported dou-
ble bundle sheaths in the leaf blades of Arundinella.
Tateoka (1956a) illustrates this PS condition for A. hirta,
Conert (1957) for A. decempedalis, Tateoka (1958) for
A. leptochloa and A. villosa and Li & Phipps (1973) for
A. bengalensis. Some of these latter workers (Tateoka
1958; Li & Phipps 1973) studied A. nepalensis and A.
berteroniana, which were also examined in the present
study, and found them to have only a single bundle
sheath. Eight specimens of A. nepalensis and one of A.
berteroniana examined in this study were all observed to
have only a single Kranz sheath in the leaf blade. It was,
therefore, most unexpected to observe a definite, inner
fibrous sheath in the first order bundles of the culm of all
three species studied. This may reflect a general condi-
tion present in all NADP-me grasses or may be an excep-
tional condition limited to Arundinella and other taxa
with distinctive cells. This interesting observation re-
quires further study as it represents a rare exception to
the XyMS character for predicting the NADP-me bio-
chemical C4 type (Hattersley 1987). This anatomical pre-
dictor for the NADP-me type may, therefore, apply to
leaf blades only or leaves and culms of a given grass may
have different photosynthetic pathways.
ACKNOWLEDGEMENTS
The authors are grateful to Aurelio Schinini for co-
operation with collection of fresh material, A. Botha for
technical assistance, A. Romanowski for the photo-
graphy, G. Condy for the line drawings and M. van der
Merwe for typing the manuscript.
REFERENCES
AUQUIER, P. & SOMERS, Y. 1967. Recherches histotaxonomiques
sur le chaume des Poaceae. Bulletin de la Societe Royale Botani-
que de Belgique 100: 95— 140.
BROWN, W. V. 1958. Leaf anatomy in grass systematics. Botanical
Gazette 119: 170-178.
BROWN, W. V. 1975. Variations in anatomy, associations, and ori-
gins of Kranz tissue. American Journal of Botany 62: 395^t02.
BROWN, W. V. 1977. The Kranz syndrome and its subtypes in grass
systematics. Memoirs of the Torrey Botanical Club 23: 1 —97.
CANFIELD, R. H. 1933. Stem structure of grasses on the Jomadas
Experimental Range. Botanical Gazette 95: 636—648.
CARO, J. A. 1961 . Las especies de Aristida (Gramineae) del centra de
la Republica Argentina. Kurtziana 1: 123—206.
CAROLIN, R. C., JACOBS, S. W. L. & VESK, M. 1973. The struc-
ture of the cells of the mesophyll and parenchymatous bundle
sheath of the Gramineae. Botanical Journal of the Linnean Society
66: 259-275.
CLAYTON, W. D. & RENVOIZE, S. A. 1986. Genera graminum:
grasses of the world. Kew Bulletin Additional Series 13: 1—389.
CLIFFORD, H. T. & WATSON, L. 1977. Identifying grasses: data,
methods and illustrations. University of Queensland Press, St
Lucia.
CONERT, H. J. 1957. Beitrage zur Monographic der Arundinelleae.
Botanische Jahrbiicher 77: 226—354.
CROOKSTON, R. K. 1980. The structure and function of C4 vascular
tissue — some unanswered questions. Berichte der Deutschen
Botanischen Gesellschaft 93: 7 1 -78.
CROOKSTON, R. K. & MOSS, D. N. 1973. A variation of C4 leaf
anatomy in Arundinella hirta (Gramineae). Plant Physiology 52:
397-402.
CUTLER, D. 1978. Applied plant anatomy . Academic Press, London.
DE WET, J. M. J. 1960. Culm anatomy in relation to taxonomy.
Bothalia 7:311-316.
ELLIS, R. P. 1977. Distribution of the Kranz syndrome in the southern
African Eragrostoideae and Panicoideae according to bundle
sheath anatomy and cytology. Agroplantae 9: 73- 1 10.
GOULD, F. W. 1972. A systematic treatment of Garnotia (Grami-
neae). Kew Bulletin 27: 515—562.
HATTERSLEY, P. W. 1987. Variations in photosynthetic pathway. In
T.R. Soderstrom et al., Grass systematics and evolution. Smith-
sonian Institution Press, Washington D.C.
HATTERSLEY, P. W. & BROWNING, A. J. 1981. Occurrence of
the suberized lamella in leaves of grasses of different photo-
synthetic types. 1. In parenchymatous bundle sheaths and PCR
(Kranz) sheaths. Protoplasma 109: 371-401.
HATTERSLEY, P. W. & WATSON, L. 1975. Anatomical parameters
for predicting photosynthetic pathways of grass leaves: the ‘maxi-
mum lateral cell count’ and the 'maximum cells distant count’.
Phytomorphology 25: 325—333.
HATTERSLEY, P. W. & WATSON, L. 1976. C4 grasses: an anatomi-
cal criterion for distinguishing between NADP-malic enzyme
species and PCK or NAD-malic enzyme species. Australian Jour-
nal of Botany 24: 297 — 308.
HATTERSLEY, P. W., WATSON, L. & OSMOND, C. B. 1977. In
situ immunofluorescent labelling of Ribulose- 1,5-biphosphate
52
carboxylase in leaves of C3 and C4 plants. Australian Journal of
Plant Physiology 4: 523-539.
JACQUES-FELIX H. 1962. Les graminees d’Afrique Tropicale. I.
Generalites, classification, description des genres. Institut de
Recherches Agronomiques Tropicales et des Cultures Vivieres,
Paris.
JOHANSEN, D. A. 1940. Plant microtechnique. MacGraw Hill, New
York.
JOHNSON, M. C. 1964. An electron microscope study of the photo-
synthetic apparatus in plants with special reference to the Grami-
neae. Ph.D. thesis. University of Texas, Austin.
JOHNSON, M. C. & BROWN, W. V. 1973. Grass leaf ultrastructural
variations. American Journal of Botany 60: 727—735.
LI, Y-H.C. & PHIPPS, J. B. 1973. Studies in the Arundinelleae. XV.
Taximetrics of leaf anatomy. Canadian Journal of Botany 51:
657-680.
METCALFE, C. R. 1960. Anatomy of the Monocotyledons. 1. Grami-
neae. Clarendon Press, Oxford.
PHIPPS, J. B. 1967. Studies in the Arundinelleae (Gramineae). V. The
series of the genus Arundinella . Canadian Journal of Botany 45:
1 047-1 057.
REGER, B. J. & YATES, I. E. 1979. Distribution of photosynthetic
enzymes between mesophyll specialized chlorenchyma and
bundle sheath cells of Arundinella hirta. Plant Physiology 63:
209-212.
RENVOIZE, S. A. 1982a. A survey of leaf-blade anatomy in grasses.
II. Arundinelleae. Kew Bulletin 37: 489-495.
RENVOIZE, S. A. 1982b. A survey of leaf-blade anatomy in grasses.
III. Gamotieae. Kew Bulletin 37: 497—500.
SABNIS, T. S. 1921. The physiological anatomy of the plants of the
Indian desert. Journal of Indian Botany 2: 217—227.
SANCHEZ, E. 1979. Estructura Kranz en tallos de Gramineae (Era-
grosteae). Kurtziana 12-13: 113-118.
Bothalia 19,1 (1989)
SANCHEZ, E. 1981a. Desarrollo de la estructura Kranz en tallos de
Gramineae. Lilloa 35: 37-40.
SANCHEZ, E. 1981b. Variacion de la estructura Kranz en el tallo de
Diandrochloa glomerata. Lilloa 35: 41-46.
SANCHEZ, E. 1983a. Estudios anatomicos en Blepharidachne Hackel
(Poaceae, Eragrostoideae, Eragrosteae). Revista de Museo Argen-
tina de Ciencias Naturales, Botanica 6: 73-87.
SANCHEZ, E. 1983b. Dasyochloa Willdenow ex Rydberg (Poaceae),
genero monotipico de Norteamerica. Lilloa 36: 130-138.
SANCHEZ, E. 1984. Estudios anatomicos en el genero Munroa
(Poaceae, Chloridoideae, Eragrostideae). Darwiniana 25: 43-57.
SANCHEZ, E. & ARRIAGA, M. 1988. El sindrome de Kranz en
Arundinelleae (Panicoideae, Poaceae) de la Flora Sudamericana.
Parodiana 6.
TATEOKA, T. 1956a. Re-examination of anatomical characteristics of
the leaves of Eragrostoideae and Panicoideae (Poaceae). Journal
of Japanese Botany 31:210-218.
TATEOKA, T. 1956b. Notes on some grasses. 1. Botanical Magazine,
Tokyo 69: 311 — 315.
TATEOKA, T. 1958. Notes on some grasses. VIII. On leaf structure
of Arundinella and Garnotia. Botanical Gazette 120: 101 — 109.
TATEOKA, T. 1963. Notes on some grasses. XV. Affinities and
species relationships of Arthropogon and relatives, with reference
to leaf structure. Botanical Magazine, Tokyo 76: 286-291.
TURPE, A. M. 1970. Sobre la anatomia foliar de Jansenella griffi-
thiana (Poaceae, Arundinelleae). Senckenbergiana Biologica 51:
277-285.
VICKERY, J. W. 1935. Leaf anatomy and vegetative characters of the
indigenous grasses of New South Wales. Proceedings of the
Linnean Society of New South Wales 60: 340—373.
WATSON, L., DALLWITZ, M. J. & JOHNSTON, C. R. 1986. Grass
genera of the world: 728 detailed descriptions from an automated
database. Australian Journal of Botany 34: 223-230.
Bothalia 19,1:53-67(1989)
The vegetation ecology of the Eastern Transvaal Escarpment in the
Sabie area. 1. Physical environment
G. B. DEALL*, J. C. SCHEEPERS* and C. J. SCHUTZ**
Keywords: climate, Eastern Transvaal Escarpment, environment, geology, physiography, soil
ABSTRACT
The physiography, geology, soil and climate of a broad transect of the Eastern Transvaal Escarpment in the Sabie area
are described. The transect extends from Hazyview (530 m elevation) to Mount Anderson (2 280 m elevation). The
description provides a contextual framework for the subsequent vegetation classification.
UITTREKSEL
Die fisiografie, geologie, grand en klimaat van ’n bree lynopname van die Oos-Transvaalse platorand in die Sabie-
gebied word beskryf. Die lynopname strek vanaf Hazyview (530 m bo seespieel) tot Mount Anderson (2 280 m bo
seespieel). Die beskrywing verskaf ’n koordinerende raamwerk vir die daaropvolgende plantegroeiklassifikasie.
INTRODUCTION
The Eastern Transvaal Escarpment, representing
Acocks’s (1975) Veld Types 8 (North-Eastern Mountain
Sourveld) and 9 (Lowveld Sour Bushveld), is an area
subject to land-use conflict. The need to produce food
and timber comes into increasing conflict with the need
to protect mountain catchments, to conserve natural
ecosystems, and to preserve scenic landscapes. Rational
land-use planning is required to resolve such conflict
(Ferrarefa/. 1988).
The relevance of plant-ecological studies to land-use
planning and management is well known (cf. Pentz
1938, 1945; Bayer 1970; Walker 1976; Edwards 1967;
Muller 1983).
In the past, Acocks’s (1975) veld types have provided
useful information at the scale of broad landscape types.
However, to meet the present needs for regional and
subregional planning, attention must be focused on sec-
tions of these landscapes (Van der Meulen & Scheepers
1978). A more detailed classification of vegetation into
ecologically interpretable vegetation units is therefore
necessary. A description of the physical environment is
fundamental to this task.
STUDY AREA
This study is confined to a broad transect located in
the Sabie area of the Eastern Transvaal Escarpment. The
transect is situated between the Olifants River in the
north and the Crocodile River in the south. Spanning
both the Pilgrims Rest and the Nelspruit Districts, it is
bounded by latitudes 25°00' and 25°10' south and longi-
tudes 30°30' and 31°10' east (Figure 1). It is 65 km long
and 20 km broad, covering approximately 1 300 km2.
* Botanical Research Institute, Private Bag X 10 1 , Pretoria 0001,
RSA.
** D. R. de Wet Forestry Research Centre, Private Bag X520, Sabie
1260, RSA.
MS. received: 1988.08.10.
The village of Sabie ( 1 050 m) is centrally situated in the
transect, whilst Hazyview (530 m) and Mount Anderson
(2 280 m) are situated at the eastern and western extrem-
ities respectively. Its orientation is perpendicular to that
of the Escarpment, which has a roughly north-south
orientation in this area.
PHYSIOGRAPHY
Following the rationale of Scheepers (1978) and with
certain modifications, the study area may be divided into
the following broad physiographic belts (Figure 2):
1, the Subalpine Belt consisting of the rolling plains,
level terraces and prominent peaks of the mountain sum-
mits (above 1 900 m);
2, the steeply sloping Montane Belt of the mountain
slopes to the west of the Escarpment Plateau (±
1 200-1 900 m);
3, the steeply sloping, much dissected Submontane
Belt of the escarpment slopes to the east of the Escarp-
ment Crest, and including the Escarpment Plateau (±
900-1 400 m);
4, the gently sloping, undulating Upland Belt of the
foothills (± 500— 900 m).
A finer subdivision of these physiographic belts leads
to the identification of ten physiographic zones (Figure
2). The main criteria for physiographic zonation are
geomorphology and altitude.
Since geomorphology does not always vary con-
sistently with altitude, the altitudinal limits of the phy-
siographic zones are arbitrary.
(a) Summit (above 1 900 m). It can be divided into
Summit Plateau (± 1 900—1 950 m). Summit Slopes (±
1 950—2 100 m), and Summit Peaks (more than 2 100
m).
(b) Mountain Slopes (± 1 200-1 900 m). This zone
is found between the Escarpment and Summit. Alti-
tudinal subdivision facilitates the recognition of Lower
(± 1 200-1 500 m) and Upper (± 1 500-1 900 m)
METRES
54
Bothalia 19,1 (1989)
FIGURE 1. — Location of the study area (inset), showing a hypothetical profile (above) and the detailed topography (below). Based on S.A.
1:250 000 Topographical Sheet 2530 Barberton.
KILOMETRES
FIGURE 2. — Hypothetical profile of the study area showing physiographic and climatic zonation in relation to geology.
Slopes. Generally, this is a zone of steeply east-sloping
hills intersected by valleys and rivulets.
(c) Escarpment Plateau (±1 200—1 400 m). This is
a flat, gently west-sloping zone forming a shelf or pla-
teau immediately west of, and including, the main
Escarpment Crest. The Escarpment Plateau may be
divided into the Plateau Interior (± 1 200- 1 300 m) and
the Plateau Crest ( ± 1 300-1 400 m).
(d) Escarpment Upper Slopes (± 1 100—1 300 m).
This is a steeply sloping zone immediately below, and to
Bothalia 19,1 (1989)
55
the east of the main Escarpment Crest. The slopes are
intersected by numerous deeply-incised gorges, kloofs
and valleys.
(e) Escarpment Lower Slopes (± 900—1 100 m).
This zone is due east of the Escarpment Upper Slopes.
Valleys are wider and less deeply incised.
(f) Upper Foothills (± 700—900 m). This and the
following zone represent a transition from the Escarp-
ment Slopes to the Lowveld. The Upper Foothills are
characterized by broken undulating terrain.
(g) Lower Foothills (± 500—700 m). This zone lies
between the Upper Foothills and the Lowveld. Valleys
are wide and flat-bottomed, and slopes are gentle.
In Kayser’s (1983) morphogenetic classification of
south-eastern Africa our Foothills and Escarpment
Slopes would be described as comprising ‘peneplains,
clusters of residual peneplains, regions with mountains
and knobs’. His concept of ‘peneplains on the lower
escarpment level with moderate degree of dissection’
finds equivalence in our Lower Eoothills, whereas
‘peneplains on the upper escarpment level with high
degree of dissection’ corresponds to our Escarpment
Slopes and Upper Foothills. The Escarpment Plateau
matches ‘marginal zones on the inland plateau level with
strong relief (Kayser 1983). The Plateau Crest Zone is
identified as a particularly high benchland scarp, steep
and divided into sections (presumably co-incident with
the Black Reef Quartzite Formation). The transition
from Escarpment Plateau to Mountain Slopes is marked
by a medium-height single benchland scarp (presumably
co-incident with the Rooihoogte Formation). The Moun-
tain Slopes and Summit would be described as ‘high,
closed benchland highland’ (Kayser 1983).
The main drainage systems eastward are the Sabie and
Mac-Mac Rivers (Figure 1). The Sabie River has its
source in the Mount Anderson range and flows east-
north-east to the Lowveld. The Mac-Mac River drains
the dolomite country southwards from Driekop near
Graskop until it swings abruptly east at Mac-Mac Falls
and then continues its course to the Lowveld.
GEOLOGY
A major geological survey resulted in the publication
of a map of the area south of Sabie in 1960 (Visser &
Verwoerd 1960). Since then surveys and lithostratigra-
phic studies have been undertaken on the Transvaal
Supergroup (Zietsman 1964; Button 1973a), the Wolk-
berg Group (Button 1973b), the Malmani Subgroup
(Button 1973c), the Timeball Hill Formation (Eriksson
1973) and the Archaean granite basement (Robb 1978;
Lageat & Robb 1984). A great deal of confusion sur-
rounding the lithostratigraphy of the area was cleared up
by the recent handbook published by the Geological Sur-
vey (SACS 1980).
The whole Escarpment region forms the eastern rim of
the Bushveld Igneous Complex of the central Transvaal.
The stratigraphy and field nomenclature are shown in
Table 1 . The varying relief of the Escarpment is deter-
mined largely by the underlying geological structure.
Essentially, four different geological systems are repre-
sented. They are traversed by a network of diabase in-
trusions. Contacts between the systems are roughly
parallel and in a north-south direction.
Nelspruit Granite
Nelspruit Granite, an intrusive of Swazian age, is the
oldest rock formation (Table 1). It comprises the biotite-
bearing gneissose granites and migmatites forming the
undulating terrain of the Escarpment Slopes and Foot-
hills (Figure 2).
Overlying the granite, and reaching from the Escarp-
ment Plateau to the Summit, are the stratified rocks of
Vaalian age, namely the Transvaal Sequence, repre-
sented mainly by the Wolkberg, Chuniespoort and Preto-
ria Groups (Table 1 ; Figure 2).
The Wolkberg Group
In the study area, the Wolkberg Group is represented
mainly by the Sekororo, Selati Shale and Black Reef
Quartzite Formations (Table 1). The Black Reef
Quartzite Formation, with its extreme resistance to
weathering, is largely responsible for the formation of
the Escarpment Plateau with its numerous associated
waterfalls, viz. Sabie, Mac-Mac, Lisbon and Berlin
Falls. Accelerated weathering along joints results in the
characteristic ‘pillar and passage’ topography with its
weird weathering forms (‘gendarmes’). The Black Reef
Quartzite Formation rarely exceeds 10 m in thickness
south of Sabie, but reaches nearly 100 m near Marieps-
kop in the north. It has a westerly dip of 5°. Shale is
always present, particularly near the top, close to the
contact with the overlying dolomite (SACS 1980) (Table
1). North-south flowing streams often follow the course
of this shale, thus forming a natural division between the
Black Reef Quartzite and the dolomites.
The Chuniespoort Group
Overlying the Black Reef Quartzite Formation and
rising into the Mountains from the Escarpment Plateau,
are the chemical sediments of the Chuniespoort Group,
including dolomite, shale, limestone and chert (Table 1;
Figure 2). The dolomite is mostly fine-grained and blue-
grey in colour with a black weathered surface (owing to
manganese oxides) and a wrinkled texture resembling
the skin of an elephant, thus deriving the name ‘Oli-
fantsklip’.
The incorporation of carbonaceous shale and quartzite
with the overlying dolomite produces the dark dolomite
of the Oaktree Formation. The Monte Christo, Lyttelton,
Eccles and Frisco Formations are characterized on the
basis of chert content (Table 1).
Dolomite rock is not often exposed within the high
rainfall zone owing to the solubility of the limestone.
Exceptions are Spitskop and the cliff line of the Eccles
Formation, more resistant to weathering because of the
high-chert content. The solubility of the limestone has
resulted in the formation of caves, usually more visible
within the Eccles Formation.
The Pretoria Group
The dolomites mark the transition to the overlying
Pretoria Group, which consists predominantly of
quartzite and shale in the Rooihoogte, Timeball Hill and
TABLE 1. — Chronostratigraphic and lithostratigraphic divisions in the Sabie area of the Eastern Transvaal Escarpment (adapted from SACS 1980)
56
Bothalia 19,1 (1989)
Bothalia 19,1 (1989)
57
Boshoek Formations, and of prominent volcanic ele-
ments in the Hekpoort Andesite Formation (SACS 1980)
(Table 1; Figure 2). The Rooihoogte Formation consists
of a conglomerate/breccia base with quartzite above,
forming a clearly visible cliff line from Ceylon State
Forest southwards. Waterfalls such as Bridal Veil, Lone
Creek and Horseshoe originate on the brow of this cliff
line. The conglomerate is distinguished as the Bevet’s
Conglomerate Member.
The Timeball Hill Formation comprises a shale zone
up to 1 600 m thick. Two bands of quartzite, the Klap-
perkop Quartzite Member, run through the middle of the
zone, giving rise to broken cliff faces with waterfalls.
The lower band averages 10 m in thickness whereas the
upper band reaches up to 60 m. Its resistance to weather-
ing has resulted in the formation of the Summit Plateau
(e.g. at Hartebeestvlakte).
The shales and mudstones of the Timeball Hill Forma-
tion are carbonaceous, thinly bedded, highly jointed,
fissile and dark in colour. They weather to pale colours.
In the Sabie area there are anomalous dips due to slump-
ing into solution cavities in the dolomite beneath, and
further tilting of the strata sometimes occurs in the vicin-
ity of diabase dykes.
The Boshoek Formation forms a small, prominent
cliff above the Timeball Hill Shale. It consists mostly of
quartzite, conglomerate, shale and pyroclastics.
In the Hekpoort Andesite Formation flows of basaltic
andesite are the most prominent rock types. The Forma-
tion reaches a thickness of 200 m.
Transvaal Diabase
Transvaal Diabase, an intrusive of Mokolian age,
criss-crosses the entire area in the form of dykes and
interbedded sills (Table 1). It is rare in the Wolkberg
Group, but most common in the Pretoria Group, particu-
larly in the form of sills. Other basic rocks also make up
these sills, those in the upper Chuniespoort Group and
Pretoria Group being composed of pyroxenitic material.
Sills occur most frequently between the two bands of
Klapperkop Quartzite and within the Boshoek Forma-
tion. Dykes run mostly in a north-north-easterly direc-
tion.
SOILS
Von Christen (1959, 1964) studied 40 soil profiles in
the Escarpment area in an attempt to gain insight into
nutrient cycling and podzolization processes under pine
stands. Subsequently a land type survey was conducted
by Schoeman et al. (1980), which included the samples
of 10 profiles.
The description which follows is based largely on the
study of 439 profiles covering the widest possible range
in soil and site conditions (Schutz in prep.). Soil proper-
ties are strongly influenced by parent material and geo-
logical substrate (Table 2).
Nelspruit Granite. The soils derived from the granites
are deep, well drained, apedal, red, ferrallitic, and
highly leached. Well decomposed granite saprolite is
quite frequently encountered within the soil profile
below a depth of 800 mm. Quartz grit, often in the form
of a broken stoneline, is commonly found in the upper B
horizon. Coarse sand is prevalent, mainly towards the
surface, whereas the subsoil, or decomposed saprolite is
siltier. Although the clay content of the B horizons of
these soils is usually high (sandy clay loam to sandy
clay), the water-holding capacity of the topsoil is low. In
bottom-land positions and river terrace areas, hydromor-
phic soils tend to develop. These soils can be extremely
wet during summer.
Organic carbon content in the topsoil of granitic soils
is far lower than that of other soils in the area (2,3 %).
This is probably owing to accelerated decomposition of
organic substances occasioned by high temperatures
(Franzle 1984). Some pockets of organic rich or humic
topsoils (Oakleaf Form) do occur in narrow drainage
lines or concave footslopes. Otherwise topsoils are rela-
tively pale in colour (usually Hutton soils).
Selati Shale. Soils derived from Selati shale are not
extensive, occurring in a narrow band along the Escarp-
ment Crest, where the Black Reef Quartzite Formation
has been eroded. Parent materials are shale and
quartzite.
These are the shallowest of all the soils in the area
(740 mm) and the bedrock is usually overlain by a layer
of hardpan ferricrete. In summer, the water table is
generally high. In winter, however, slow but efficient
drainage is facilitated both by the sandy texture of the
soil and by the slight westerly dip of the strata (Figure 2).
These soils can thus be particularly dry in winter. Such
fluctuations of the water table together with eluviation
are conducive to the formation of bleached E horizons.
Soils derived from Selati shale are usually sandy clay
loams with a coarse sandgrade in the A horizon. Stone-
lines are common when the parent material is shale. On
the shallow soils of the Escarpment Crest the dominant
soil form is the Glenrosa. Profiles of the Hutton, Oakleaf
and Griffin Forms develop on the deeper soils.
Black Reef Quartzite. Soils on the Black Reef
Quartzite Formation extend further south than Selati
soils, but are also not extensive. South of Sabie all sandy
soils along the Escarpment Crest are derived from Black
Reef material. North of Sabie they occur in a band
parallel with Selati soils, but generally further west down
the dipslope.
Soils on Black Reef Quartzite are mostly very shallow
over solid bedrock. Under grassland, however, fairly
deep soils have developed. Of all soils in the area they
have the deepest A horizons (450 mm).
Soils on Black Reef Quartzite are sandy loams, with
the highest percentage sand (78 %) of all the soils. They
also have the darkest surface soils, with lowest values
and chromas. Organic, peaty topsoils are common, reac-
hing a maximum of 9,3 % organic carbon.
The dominant soil form is the Hutton, but Mispah,
Clovelly, Inanda and Oakleaf Forms are also common.
Less common soil forms include Champagne (under
marshy conditions) and Houwhoek and La Motte (under
highly leached conditions).
Oaktree. Towards the Plateau Interior, the Oaktree
soils begin to appear. They are distinguished from Black
Reef sands by an increase in clay content of the subsoil.
58
Bothalia 19,1 (1989)
E c
£ £
5 ii
Q 5
a
CO «N
•- II
JS c
u ^
o o O vo cn
« T) n m (N © CD O O
■a K Tf* Tt <N VO <N Ov VO
T3 CO CO i— <
IOOOONO OOTj-ON
o
s ^
3 m
x a
O o O CO <N
rf o o o a\ rn |h
co >n vo O) o o 'C
<N (J
O O O OO On O
tJ" t «-•
& i
10 oo co <n © © O
m" t h « oo n’ «
X Q
£ =
CO *0 CN_ CO
>0 ’t M 'O
O CD © co r-
^ ro" <N — < O
CO On <n r*^ CD O O »n O
rsiTj-i— ivooor^TfO-— i £
•5 4= -C T3
« « J2 o
’£ £
cd w
E? x
. MU •— I CU *— « »— < t— i
t-^EOjc^ojcooo
o o 3 J= o •*= „ -C .C J=
^.•c-0'-|r^'c— • o)
'a I E
E, £ ~
-2 X eu U U
X + o
i4 ^
E E
Bothalia 19,1 (1989)
59
A horizons are shallow but the solum is moderately
deep (about 1 090 mm). The topsoils have the highest
percentage of fine sand of all the soils (together with
Timeball Hill Formation). They are generally clay loams
except in cases where B horizons are overlain with a
colluvial, dark sandy A horizon derived from Black Reef
Quartzite. B horizons are classified as clay. Some of the
wetter B horizons tend to have massive structure. Stone
content of the B horizon is low (11 %).
Surface horizons are usually dark when associated
with Black Reef Quartzite soils. Yellow-brown apedals
over red apedals abound. The B horizon has the highest
colour value of all the soils.
Dolomite. Soils derived from dolomite are deep, and
well drained. Only on ridge tops and steep slopes does
the unweathered rock come close to the surface. The
presence of many chert stones and manganese concre-
tions suggests that these soils are transported.
Of all soils in the area, dolomite soils have the highest
pH and lowest exchange acidity, probably owing to their
limestone origin. Although P levels are low, the sum of
basic cations is high, making them the most fertile of
soils in the area. Often, there is such colluvial mixing of
unconsolidated rock with the red soil matrix that classifi-
cation of the soil becomes difficult. Otherwise, Hutton
and Griffin Forms are usually dominant.
Timeball Hill. Because they are developed in the dis-
sected terrain and steep slopes of the Mountain and Sum-
mit Zones, the soils derived from the Timeball Hill For-
mation tend to be shallow, except on terraces and valley
bottoms. The A horizons are shallower than those of
other soils in the area. Pockets of silty clay loam
400- 500 mm deep are nevertheless prevalent on the
slopes, and root penetration is further facilitated by
cracks in the shale. Where the shale strata dip (in asso-
ciation with diabase intrusions), weathering is usually
deeper than where strata are horizontal.
Soils are well drained clay loams with a fine sand-
grade in the topsoil. Subsoils are high in clay content,
but can be quite sandy in areas below the two Klapper-
kop Quartzite cliff lines. The high subsurface stone con-
tent (47 %) is typical of these soils. The stones may be
loose, or compacted into stonelines, particularly where
there is shale colluvium over diabase.
Transvaal Diabase. Soils derived from Transvaal
Diabase are not easily recognizable as they are always
buried below colluvial topsoil from the adjacent country
rock, from which they are separated by a stoneline at 300
to 500 mm depth.
Diabase soils therefore exist only as subsoils and their
position as an important subdivision of soils is question-
able.
CLIMATE
The Eastern Transvaal Escarpment lies in a seasonally
arid, subtropical region, with hot wet summers and cool
dry winters. It forms a transition area between the
warmer Lowveld to the east and the climatically more
extreme Highveld plateau to the west. Summer to winter
shifts in the position of the high pressure system of the
southern hemisphere play a major role in the constitution
of weather (Fabricius 1988).
A climatic gradient facilitates the recognition of two
major climatic belts based on altitudinal distribution of
mist. These belts are especially meaningful for identify-
ing gross climatic variability. Following Scheepers
(1978), there is the Mistbelt of the high, cooler altitudes
as opposed to the Low Country of the low, warmer alti-
tudes (Figure 2). The boundary between these two belts
is irregular and poorly defined in the study area, but
appears to fluctuate between 900 and 1 100 m elevation.
This zone of fluctuation may constitute a third climatic
belt, namely Transitional Mistbelt. Its range conforms to
Scheepers’s (1978) arbitrary lower limit of about 1 050
m elevation. A fourth climatic belt, derived from the
subdivision of Mistbelt into ‘moist’ and ‘dry’ sectors
(Humid and Subhumid Mistbelt respectively. Figure 2),
may also be justified (Deall 1985). The Subhumid Mist-
belt (represented by the Summit Zone), is a distinctly
cooler and drier sector of the Mistbelt (Figures 3 & 4). It
is frequently misty up to about 1 800 m elevation and
completely clear on the Summit above. The boundary
between Humid and Subhumid Mistbelt is therefore
assumed to co-incide with the transition from Montane to
Subalpine Belt (Figure 2).
Fabricius’s (1984, 1988) hygric classification of
south-eastern Africa, based on a humidity index (the
ratio of precipitation to potential evapotranspiration —
Papadakis 1966), concurs fairly well with the climatic
JASONDJFMAMJJ asondj fmamj jasondj fmamj
FIGURE 3.— Annual march of extreme daily maximum (edx) and minimum (edn) temperatures and mean monthly maximum (mmx) and
minimum (mmn) temperatures for three stations in the Sabie area of the Eastern Transvaal Escarpment. Compiled from Weather Bureau.
<; A e thp Snil anrl Irrioation Research Institute, Private Bag X79. Pretoria 000 1
60
Bothalia 19,1 (1989)
A SUBHUMID MISTBELT (Summit) B HUMID MISTBELT (Escarpment Plateau) C LOW COUNTRY (Lower Foothills)
FIGURE 4. — Annual march of mean monthly rainfall (histogram) and mean rainy days per month (graph) for three stations in the Sabie area of
the Eastern Transvaal Escarpment. Compiled from Weather Bureau, S.A. (1965b) and from climatological reports of the Soil and
Irrigation Research Institute, Private Bag X79, Pretoria 0001 .
belt concept outlined above. Low Country and Mistbelt
correspond to ‘subhumid’ and ‘humid’ types respect-
ively. Transitional and Humid Mistbelts together corre-
spond to ‘humid’ and ‘perhumid’ types. The Subhumid
Mistbelt peculiar to the Summit is inferred on Fabricius’s
(1984) map to be ‘perhumid’. Insolation, temperature,
rainfall, and humidity records (Tables 3, 4, 7 & 8) for
Long Tom on the Summit (25°08' south and 30°37' east),
however, suggest a ‘subhumid’ rather than a ‘perhumid’
climatic type. Moreover, the humidity index for Long
Tom is calculated as 0,698, a ‘subhumid’ category in
Fabricius’ (1984, 1988) classification.
According to Schulze & McGee (1978), light (insola-
tion), temperature, and moisture (precipitation) are the
most significant climatic factors in vegetation develop-
ment. These operate together to produce ‘homogeneous
environments in which certain plant communities attain
importance’ .
INSOLATION
Insolation may be measured directly in terms of the
quantity of incoming solar radiation, and indirectly by
means of sunshine duration. Maps of incoming solar
radiation patterns based on solar radiation measurements
in southern Africa have been compiled by Schulze &
McGee (1978). In addition, the Weather Bureau (1950)
has compiled maps showing broad, countrywide patterns
of sunshine duration and cloud cover based on a number
of stations in South Africa.
Schulze & McGee’s (1978) maps show that incoming
radiation is subject to seasonal variation. For instance,
radiant flux densities for the study area in summer are
apparently 80 x 105 Jm- 2 day-1 higher than in winter.
The magnitude of seasonal variation is supposedly tem-
pered by the interposing effect of cloud cover which,
during summer in the study area, reduces the average
duration of bright sunshine by 20—30 % (Weather
Bureau 1950).
There is also geographic variation of sunshine dura-
tion within the study area. For instance, the average
annual duration of bright sunshine in the Mistbelt is less
than 60 % of the possible sunshine, whereas it is
60— 70 % in the Low Country. Similarly, the Mistbelt
experiences less ‘bright’ days (days with 90—100 % of
possible sunshine) than the Low Country (Weather
Bureau 1950).
This trend of increasing cloudiness is broken in places
on the Summit (Subhumid Mistbelt) however, where,
owing to rain-shadow effects, the duration of daily
sunshine may be greater than in the Low Country. Thus
throughout the year, Long Tom (2 118 m) in the Sub-
humid Mistbelt experiences more sunshine than Hazy-
view (530 m) in the Low Country (Table 3).
Besides seasonal and geographic variability, there is
also physiographic variability of insolation owing to
slope and aspect. Daily incoming radiant flux densities
on sloping terrain as a function of slope, aspect, and
season have been presented for cloudless days in South
TABLE 3. — Annual march of mean daily sunshine (hours) for two stations in the Sabie area of the Eastern Transvaal Escarpment*
* Based on climatological reports of the Soil and Irrigation Research Institute, Private Bag X79, Pretoria 0001.
Bothalia 19,1 (1989)
61
TABLE 4. — Annua] march of temperature extremes, ranges and means (°C) for six stations in the Sabie area of the Eastern
Transvaal Escarpment*
Emmett
610
1931-1937
Hazyview
530
1971-1983
* Compiled and adapted from Weather Bureau. S.A. (1954a) and from climatological reports of the Soil and Irrigation Research In-
stitute, Private Bag X79, Pretoria 0001 . _
HMB = Humid Mistbelt: SMB = Subhumid Mistbelt; TMB = Transitional Mistbelt: LC Low Country.
62
Bothalia 19,1 (1989)
Africa for the latitudinal range 20°S to 35°S by Schulze
(1975). The following general trends for the study area
(at 25°S) are derived from these data.
In midsummer, steep slopes receive less radiation than
gentle slopes, regardless of aspect. Steep slopes with a
south-facing aspect receive greater radiation than those
with a north-facing aspect. Incoming radiation on gentle
slopes, however, is apparently unaffected by aspect.
At the equinoxes and in midwinter, north-facing
aspects receive greater radiation on steep slopes than
they do on gentle slopes. The converse applies to south-
facing aspects. Finally, regardless of steepness, slopes
experience decreasing radiation as they tend more
towards south-facing aspects.
TEMPERATURE
Although temperature alone may not be a significant
factor in determining major regional vegetation forma-
tions, it does play a part in determining floristic varia-
tions on a meso- and micro-scale (Schulze & McGee
1978). Such variations result from differential effects of
temperature on plant growth rates, seed germination,
seedling survival and flowering phenology.
Temperature data for several stations in the study area
are given in Tables 4 and 5 and in Figure 3. These data
will be discussed comparatively with respect to the dif-
ferent climatic belts.
TABLE 5. — Occurrence of severe frost (less than 0°C in Steven-
son screen) at two stations in the Sabic area of the Eastern
Transvaal Escarpment*
* Compiled from Weather Bureau, S.A. (1954a).
Subhumid Mistbelt
Long Tom station on the Summit Plateau is considered
to be representative of the Subhumid Mistbelt. Mean
monthly temperatures range from 8,9°C in June to 16,6°C
in January (Table 4E). Average summer and winter tem-
peratures are differentiated by 7,7°C. Extreme daily
maxima are greatest in early summer (November:
29, 1°C), and extreme daily minima are least in winter
(June: -4,0°C) (Table 4A, B; Figure 3A). Both the mean
and the extreme daily ranges are greatest in September
(11,3°C and 29,5°C respectively) (Table 4C, D).
Although data are lacking, frost is assumed to be a signi-
ficant factor in the climate of the Subhumid Mistbelt.
Humid Mistbelt
Data for the stations Sabie and Tweefontein on the
Escarpment Plateau are considered to be representative
of the Humid Mistbelt. Mean monthly temperatures in
the Humid Mistbelt range from 12,1°C in the coolest
month (July) to 21,1°C in the warmest month (January)
(Table 4E). Average summer and winter temperatures
are differentiated by 9,0°C. Extreme daily maxima are
greatest in late spring (October: 37,5°C), and extreme
daily minima are least in winter (July: — 4,2°C) (Table
4A, B; Figure 3B). Although the mean daily range is
least in February (9,9°C) and greatest in July (17,3°C)
(Table 4D), the extreme daily range can reach up to
37,5°C in early spring (August to September).
Severe frost (less than 0°C in Stevenson screen) is
prevalent in the winter months (June to July), but has
been known to occur as early as May and as late as
September (Table 5). Its average duration, however, is
only 31 days. The prevalence of extremely high day
temperatures in the absence of rain in late spring could
be inhibitory to many species, especially those on ex-
posed, north-facing (xeric) sites. Likewise, frosty condi-
tions would tend to exclude species which inhabit the
lower frost-free climatic belts.
Transitional Mistbelt
Temperature conditions at Bergvliet (Escarpment
Lower Slopes) are considered representative of the
Transitional Mistbelt. Mean monthly temperatures range
from 14,4°C in the coolest month (July) to 21 ,5°C in the
warmest month (January) (Table 4E). Seasonal differen-
tiation of mean monthly temperature is only 7,1°C.
As in the Humid Mistbelt, extreme daily maxima are
greatest in late spring (October: 38,3°C), and extreme
daily minima are least in winter (July: 0,0°C) (Table 4A,
B). Extreme daily maxima, and especially extreme daily
minima, are maintained at higher levels than in the
Humid Mistbelt. The nett effect is a narrowing of the
extreme daily range, its magnitude being greatest in
October (33,0°C) and smallest in March (23,9°C) (Table
4C). The greatest mean daily range, however, occurs in
August (13,0°C) and the smallest in February (9,2°C)
(Table 4D).
Severe frost (less than 0°C in Stevenson screen) has
never been recorded in 15 years of observation (Table 5).
Isolated pockets of frost may occur, however, in areas of
poor air drainage such as valley bottoms.
As in the Humid Mistbelt, the desiccating conditions
of late spring are probably inhibiting to many plants.
This is enhanced in the Transitional Mistbelt where the
dampening effect of mist is less prevalent. Conversely,
frost is no longer a factor limiting to plants in the Transi-
tional Mistbelt.
Low Country
Temperature characteristics for this climatic belt are
exemplified by the stations Emmett and Hazyview
(Lower Foothills). Mean monthly temperatures range
from 15,6°C in winter (July) to 24,6°C in summer
(January) (Table 4E). As in the Humid Mistbelt, sea-
sonal temperatures differ by 9,0°C. The greatest extreme
daily maximum for Emmett (40,4°C) occurred in Sep-
tember, a month earlier than in the Humid and Transi-
tional Mistbelts and two months earlier than in the
Subhumid Mistbelt. Conversely, the lowest extreme
daily minimum for Emmett ( 1 ,8°C) occurred in August, a
Bothalia 19,1 (1989)
63
month and two months later respectively (Figure 3;
Table 4A, B). Thus, the Low Country apparently warms
up sooner and cools down later than the Mistbelt as a
whole. It therefore has a longer warm-hot season than
the latter (Figure 3).
As in the Humid Mistbelt, the greatest extreme daily
range occurs in September (36,6°C). The smallest
extreme daily range, as in the Transitional Mistbelt,
occurs in March (20,8°C) (Table 4C). These extreme
ranges co-incide largely with the mean daily range for
Emmett, which is greatest in September (13,4°C) and
smallest in January (10,5°C) (Table 4D). This suggests
that temperatures in the Low Country are seasonally
more consistent than in the Humid and Transitional Mist-
belts. Frost (less than 0°C in Stevenson screen) is not an
important factor in the Low Country although it may
occur in isolated pockets, as in the Transitional Mistbelt.
As for the other climatic belts, the Low Country
experiences hot dry conditions in late spring prior to the
rainy season, and with the absence of mist, conditions
become quite arid. Compared to the Mistbelt, the long
warm-hot season of the Low Country means an extended
growing season for plants, without the inhibiting effect
of frost. Furthermore, the greater predictability of
temperature regimes in the Low Country probably mode-
rates demands on the adaptive capability of plants.
The Mistbelt as a whole is consistently cooler than the
Low Country (Figure 3). Mean monthly temperatures
decrease with increasing altitude. Long Tom being
approximately 7,5°C and Sabie 3,5°C cooler than Hazy-
view throughout the year (Table 4E). Likewise, but with
the exception of anomalies caused by local geomorpho-
logy, the temperature range generally narrows with in-
creasing altitude. For example, the annual average of the
mean daily range is 13,6°C at Hazyview (530 m) and
only 9,8°C at Long Tom (2 118 m) (Table 4D).
Wind
Although wind data for stations in the area are lack-
ing, wind conditions at nearby stations are considered
fairly representative and are therefore presented in
Table 6.
TABLE 6. — Seasonal variation in wind direction for three
stations in the vicinity of the study area*
* Compiled from Weather Bureau, S.A. (1960).
South-easterly to easterly to north-easterly winds pre-
dominate during summer (January). These winds blow
from the Indian Ocean and are often associated with
anticyclonic systems, but can also be associated with
cyclonic systems. Their persistence is often the har-
binger of the steady rains, drizzle and mist so typical of
Escarpment weather in summer.
Winds which blow from the southerly to south-west-
erly sectors during early summer, especially in the
afternoon, are often associated with thunderstorms. Dur-
ing winter, these same winds are associated with cold
fronts which are sometimes attended by mist and drizzle.
Violent berg wind conditions, such as the 115 kph
westerly wind gust recorded in Pietersburg one Septem-
ber, may occasionally occur (Weather Bureau 1960).
These winds become heated by compression as they drop
over the Escarpment from the Highveld Plateau and are
known to cause considerable physical damage to timber
plantations. Indigenous vegetation does not apparently
suffer much physical damage, but the low humidity of
such winds in the presence of high spring temperatures is
sure to have adverse physiological effects. Furthermore,
such winds may seriously increase the risk of fire in the
dry months of early spring.
Precipitation
Soil moisture is derived from precipitation mainly in
the form of rainfall, mist (and fog), dew, hail and snow.
(a) Rainfall. The study area falls within the summer
rainfall zone where the bulk (85 %) of annual rainfall
occurs between November and March. During this
period, rain is precipitated mainly in the form of
afternoon thunderstorms and instability showers caused
by convection in, and convergence of, tropical air
masses (Weather Bureau 1965a). The convergence is
often occasioned by a particularly pronounced trough of
low pressure over Botswana-Namibia (Fabricius 1988).
Light orographic rainfall (drizzle) without thunder, and
associated with advection, is also prevalent in the sum-
mer months, especially on the windward sides of the
Mountain and Escarpment Slopes. The small proportion
of winter rainfall is derived solely from orographic preci-
pitation, since conditions favouring thunderstorms and
instability showers do not persist in the winter months
(Weather Bureau 1965a).
On average, the Escarpment region experiences a
maximum of over 140 days with measurable rainfall per
annum, including 60 to 80 thunderstorms occurring early
in the rainy season (Weather Bureau 1965a). Prolonged
periods of rain, usually in the form of drizzle are fairly
common. For instance, periods with seven consecutive
rainy days are encountered on about two occasions per
year, and periods with four consecutive rainy days may
be encountered on as many as 10 occasions per year
(Weather Bureau 1965a).
Rainfall along the eastern Escarpment is generally
reliable. In 58 years of recording, 78 % of the annual
falls lie within about 20 % of the normal rainfall. A
further 10 % of annual falls may be regarded as ‘wet’
years (120-140 % of normal), and the remaining 12 %
as ‘dry’ years (60- 80 % of normal) (Weather Bureau
1965a).
The regional climate is of the monsoon type in which
three seasons can be recognized:
1, the rainy season of summer and late summer (i.e.
November to March); 2, the cool dry season of ‘autumn’
to early spring (i.e. April to August); 3, the warm dry
64
Bothalia 19,1 (1989)
season of spring and early summer (i.e. September to
October).
The strongly seasonal nature of the rainfall at three
stations in the study area is illustrated in Figure 4. (Cau-
tion should be exercised in comparing A and C because
of the latter’s relatively short period of recording). Maxi-
mum rainfall occurs during the warmer months, from
November to March. In the Low Country and Humid
Mistbelt, November to March are ‘superhumid’ months
in which the monthly rainfall consistently exceeds 100
mm (Walter 1971). In the Subhumid Mistbelt, however,
the superhumid period is limited largely to January and
February. Also, there is a general increase in rainfall
with altitude; the mean monthly January rainfall in the
Humid Mistbelt being approximately 52 mm more than
in the Low Country. This trend is interrupted on the
Summit, however, where rainfall is considerably less
than in the Low Country (Figure 4A), and where the
humid period is considerably shorter.
The average number of rainy days in both the Mistbelt
and Low Country shows much the same type of annual
' ariation as the rainfall amount (Figure 4B, C). Rainfall
is nevertheless consistently heavier in the Humid Mist-
belt than in the Low Country. For example, for the same
number of rainy days in December, Tweefontein in the
Humid Mistbelt experiences 45 mm more rainfall than
Emmett in the Low Country (Figure 4B, C). The heavier
rainfall in the Humid Mistbelt may be attributed to the
greater influence of orographic rainfall in this climatic
belt.
Rainfall data for specific stations within and around
the study area are given in Table 7. Stations are grouped
according to the physiographic zones they occupy, and
this facilitates the computation of a ‘composite’ mean or
index for each zone (except the Escarpment Upper
Slopes). This value broadly represents the mean annual
rainfall of each zone. Zonal variability can be explained
largely in terms of the prevailing physiography. Thus it
is clear from Table 7 that rainfall increases steadily from
the Lower Foothills (904 mm) up to a first maximum on
the Plateau Crest (1 607 mm). This increase is due to the
fact that elevations of land force air currents to rise and
cool, and hence to precipitate moisture. Furthermore,
even in the absence of general atmospheric movements,
mountain regions are known to have local ascending
currents that would enhance the precipitation process
(Killick 1963). From the Plateau Crest, rainfall
decreases into the Plateau Interior (1 200 mm). The
effect of the Plateau Crest in creating a local rain-shadow
may be responsible for this decrease. From the Plateau
Interior, rainfall again increases steadily up to a second
maximum on the Upper Mountain Slopes (1 853 mm).
Above this zone, rain-shadow effects and falling tempe-
ratures probably cause significant reductions in the
vapour content of the rising air masses, and rainfall con-
sequently decreases to 774 mm on the Summit (Table 7).
TABLE 7. — Mean annual and absolute maximum and minimum rainfall for 22 stations in the Sabie area of the Eastern Transvaal
Escarpment*
* Compiled from miscellaneous records of the Weather Bureau, Pretoria; from Weather Bureau, S. A. (1954b); and from climatological
reports of the Soil and Irrigation Research Institute, Private Bag X79, Pretoria 0001.
**Rainfall figures adjusted to the nearest millimetre,
t Corrected altitude.
Bothalia 19,1 (1989)
65
(b) Mist* Precipitation from mist (and fog) supple-
ments rainfall precipitation quite significantly. Prelimi-
nary results from ‘fog-catchers’ in the Eastern Transvaal
have yielded precipitation figures that exceed those of
standard rain gauges by between 105 and 280 % (Fabri-
cius 1969). Where mist or fog is carried through the
crown of trees, fine droplets of water collect on foliage
and branches. These droplets coalesce and eventually
become large enough to drip to the ground, thus contri-
buting directly to soil moisture. This phenomenon,
known as ‘fog drip’, is most significant when mist or fog
is moving through tall vegetation. Even when mist or fog
is stagnant or when it moves through short vegetation, it
contributes indirectly to soil moisture by reducing evapo-
transpiration. Conversely, stagnant fogs may prevent the
precipitation of dew by their inhibiting effect on
radiation.
In the Mistbelt, mist is experienced extensively. It
also occurs occasionally in the Low Country, on wind-
ward, mesic slopes during very wet periods in summer.
Cold-air drainage on calm winter nights results in the
formation of stagnant valley fogs. In the Low Country,
these fogs do not persist beyond early morning, being
soon dissipated by the heat of the sun.
(c) Dew. Radiation of heat at night causes surface
temperatures to fall and, under relatively humid condi-
tions, this results in precipitation of moisture in the form
of dew. River valleys are especially subject to dew pre-
cipitation.
Dew contributes to soil moisture by reducing the rate
and duration of evapotranspiration. Such reduction is
accomplished when, with the evaporation of dew, rapid
temperature-rises are checked and humidities raised.
Dew precipitation may thus serve to ease, or even posi-
tively counteract, drought conditions.
(d) Hail. Precipitation by hail does not contribute very
significantly to soil moisture and neither does it appear to
cause much damage to indigenous vegetation. This is
mainly because it occurs so infrequently, being prevalent
during only four or five spring thunderstorms annually
(Weather Bureau 1965a).
(e) Snow. Snowfall results from subpolar air masses
advancing from the south of the study area (Fabricius
1988). Apart from increasing soil moisture during
winter, snow acts as an insulating blanket protecting
plants from excessively cold temperatures and prevent-
ing the ground beneath from freezing (Killick 1963).
Owing to the dryness of the winter, snowfalls in the
study area are not particularly frequent. Between 1969
and 1983, only four instances of snow were recorded,
usually in early spring (August- September) (Mrs D.
Livingstone pers. comm.). The snow is confined to the
Summit and Mountain Slopes, where slopes as low as
1 600 m elevation may be affected. Snow in early spring
may therefore be particularly beneficial to high-altitude
plants whose soil moisture is largely depleted after the
prolonged dry conditions of winter.
* In this text, the term ‘mist’ is used to indicate suspended moisture
droplets condensed by the cooling of saturated air masses rising against
sloping ground. In contradistinction, the term ‘fog’ is used to denote
such suspended moisture droplets condensed from saturated air cooled
at night by radiation and temperature inversion, being restricted mainly
to low-lying and level terrain.
Humidity
Data for Hazy view in the Low Country and Long Tom
in the Subhumid Mistbelt are presented in Table 8. As
expected, humidity levels are correlated with precipita-
tion, temperature, and wind; the highest mean values
being recorded in April- May in the Low Country and in
February -March in the Subhumid Mistbelt. These
periods are associated with decreasing autumn tempera-
tures (Figure 3A, C) near the close of the rainy season,
when fairly high levels of soil and atmospheric moisture
still prevail (Figure 4A, C). The lowest mean humidity
values were recorded in June-July in both the Low
Country and the Subhumid Mistbelt. This is a period of
low rainfall and low temperatures (Figures 3 & 4). In
addition, prevailing winds tend to be dry south-westerly
(Table 6).
The period July -August in the Subhumid Mistbelt
may be subject to large fluctuations in humidity. For
example, the extreme minimum humidity may be as low
as 2%, whereas the extreme maximum may be as high as
96% (Table 8). The low minima may arise on occasions
when hot, dry south-westerly ‘bergwinds’ blow. Condi-
tions in the Low Country are not as dry, the lowest
extreme minimum humidity being 1 1% in September.
Table 8 shows the situation at the extremes of the
study area only. Humidities for the intermediate Humid
and Transitional Mistbelts are obviously expected to be
higher than those for either the Subhumid Mistbelt or the
Low Country.
CONCLUSION
The foregoing treatment has been necessary for estab-
lishing a basic environmental context for vegetation
classification (cf. Deall et al. 1989). Environmental
attributes are largely responsible for determining plant-
species distributions. Discussion of the environment,
therefore, draws attention to the potential determinants
of vegetation composition, vegetation structure, and
vegetation distribution. The Eastern Transvaal Escarp-
ment is a region with a great diversity of natural habitats.
It is this diversity which elicits the vegetation response
reflected in the numerous syntaxa described elsewhere
(cf. Deall etal. 1989).
ACKNOWLEDGEMENTS
Mr J. Schoeman provided valuable guidance on
geology and soils. Mrs J. Schaap drew the figures, Ms
A. P. Backer helped proofread the text, Ms H. M. Mool-
man translated the abstract, and Mrs M. van der Merwe
typed the text.
REFERENCES
ACOCKS, J. P. H. 1975. Veld types of South Africa. 2nd edn.
Memoirs of the Botanical Survey of South Africa No. 40: 1-128.
BAYER, A. W. 1970. Plant ecology in the service of man in southern
Africa. South African Journal of Science 66: 71— 77.
BUTTON, A. 1973a. A regional study of the stratigraphy and develop-
ment of the Transvaal basin in the eastern and north-eastern
Transvaal. Ph.D. thesis. University of the Witwatersrand.
BUTTON, A. 1973b. The depositional history of the Wolkberg proto-
basin, Transvaal. Transactions of the Geological Society of South
Africa 76: 19—25.
66
Bothalia 19,1 (1989)
TABLE 8. — Annual march of mean and extreme maximum and minimum humidity for two stations in the Sabie area of the
Eastern Transvaal Escarpment*
*Compiled from climatological reports of the Soil and Irrigation Research Institute, Private Bag X79, Pretoria 0001.
BUTTON, A. 1973c. The stratigraphic history of the Malmani dolo-
mite in the eastern and north-eastern Transvaal. Transactions of
the Geological Society of South Africa 76: 229-248.
DEALL, G. B. 1985. A plant-ecological study of the Eastern
Transvaal Escarpment in the Sabie area. M.Sc. thesis. University
of Pretoria.
DEALL. G. B.. THERON, G. K. & WESTFALL. R. H. 1989. The
vegetation ecology of the Eastern Transvaal Escarpment in the
Sabie area. 2. Floristic classification. Bothalia 19: 69-89.
EDWARDS, D. 1967. A plant ecology survey of the Tugela Basin.
Memoirs of the Botanical Survey of South Africa No. 36: 1 - 285 .
ERIKSSON, K. A. 1973. The Timeball Hill Formation — a fossil delta.
Journal of Sedimentary Petrology 43: 1046-1053.
FABRICIUS, A. F. 1969, cited in Schutte, J. M. 1971. Die onttrek-
king van water uit die newel laag en lae wolke by Mariepskop.
S.A. Department of Water Affairs, Division Hydrological
Research, Technical Note 20: 1—21.
FABRICIUS. A. F. 1984. Klimageographie. Afrika-Kartenwerk S5,
Blatt 5. Deutsche Forschungsgemeinschaft, Bonn-Bad Godes-
berg.
FABRICIUS, A. F. 1988. Klimageographie-Sudafrika (Mozambique,
Swaziland, Republik Siidafrika). Afrika-Kartenwerk S5, Beiheft
S5. Deutsche Forschungsgemeinschaft, Bonn-Bad Godesberg.
FERRAR, A. A., ISAACS, G. & STACEY, J. R. (eds). 1988.
Environmental conservation features of the Transvaal Escarp-
ment: an information base for regional planning, pp. 1 13. Occa-
sional Report No. 31. FRD, CSIR, Pretoria.
FRANZLE, O. 1984. Bodenkunde-Sudafrika (Mozambique, Swazi-
land, Republik Siidafrika). Afrika-Kartenwerk S4, Beiheft S4.
Deutsche Forschungsgemeinschaft, Bonn-Bad Godesberg.
KAYSER, K. 1983. Geomorphologie. Afrika-Kartenwerk S2, Blatt 2.
Deutsche Forschungsgemeinschaft, Bonn-Bad Godesberg.
KILLICK, D. J. B. 1963. An account of the plant ecology of the
Cathedral Peak Area of the Natal Drakensberg. Memoirs of the
Botancial Survey of South Africa No. 34: 1-178.
LAGEAT, Y. & ROBB, L. J. 1984. The relationships between struc-
tural landforms, erosion surfaces and geology of the Archaean
Granite basement in the Barberton region, eastern Transvaal.
Transactions of the Geological Socien of South Africa 87:
141-159.
MULLER, T. 1983. A case for a vegetation survey in a developing
country based on Zimbabwe. Bothalia 14: 721—723.
PAPADAKIS, J. 1966. Climates of the world and their agricultural
potentialities. Buenos Aires.
PENTZ, J . A. 1938. The value of botanical survey and the mapping of
vegetation as applied to farming systems in South Africa.
Memoirs of the Botanical Suney of South Africa No. 19: 1-15.
PENTZ, J. A. 1945. An agro-ecological suney of Natal. Department
of Agriculture and Forestry Bulletin No. 250, Govt. Printer,
Pretoria.
ROBB, L. J. 1978. A general geologic description of the Archaean
Granite terrain between Nelspruit and Bushbuckridge, eastern
Transvaal. Transactions of the Geological Socien of South Africa
81:331-338.
SACS 1980. South African Committee for Stratigraphy. Stratigraphy
of South Africa. Part 1 (Comp. L. E. Kent). Lithostratigraphy of
the Republic of South Africa, South West Africa/Namibia and the
Republics of Bophuthatswana, Transkei and Venda. Handbook of
the Geological Suney of South Africa 8: 1-690.
SCHEEPERS, J. C. 1978. Vegetation of Westfalia Estate on the
North-Eastern Transvaal Escarpment. Memoirs of the Botanical
Suney of South Africa No. 42: 1 — 230.
SCHOEMAN. J. L., TURNER, D. P. & FITZPATRICK, R. W.
1980. Land type map 2530, Barberton. Government Printer, Pre-
toria.
SCHULZE, R. E. 1975. Incoming radiation on sloping terrain: a gen-
eral model for use in southern Africa. Agrochemophvsica 7:
55-61.
SCHULZE, R. E. & McGEE, O. S. 1978. Climatic indices and classi-
fication in relation to the biogeography of southern Africa. In
M. J. A. Werger, Biogeography and ecology of southern Africa:
19-52. Junk, The Hague.
SCHUTZ, C. J. in prep. Site relationships for Pinus patula in the
Eastern Transvaal Escarpment area. Ph.D. thesis, University of
Natal, Pietermaritzburg.
VAN DER MEULEN, F. & SCHEEPERS, J. C. 1978. On vegetation
studies and land evaluation in South Africa. In R. Tiixen,
Assoziationskomplexe und ihre praktische Anwendung. Berichte
des Internationalen Symposien des Internationalen Vereinigung
furVegetationskunde, Rinteln, 1977: 481—489. Cramer, Lehre.
VISSER, H. N. & VERWOERD, W. J. 1960. The geology of the
country north of Nelspruit. An explanation of Sheet 22. pp. 128.
Geological Survey of South Africa.
VON CHRISTEN, H. C. 1959. The forest soils of the Transvaal mist-
belt. pp. 74. Report No. 1096/59, Division Chemical Services,
Department of Agriculture, Pretoria. Unpublished.
VON CHRISTEN, H. C. 1964. Some observations on the forest soils
of South Africa. Forestry in South Africa 5: 1—21.
WALKER, B. H. 1976. An approach to the monitoring of changes in
the composition and utilization of woodland and savanna vegeta-
tion. South African Journal of Wildlife Research 6: 1—32.
Bothalia 19,1 (1989)
67
WALTER, H. 1971. Ecology of tropical and subtropical vegetation.
Oliver & Boyd, Edinburgh.
WEATHER BUREAU, S.A. 1950. Sunshine and cloudiness in South
Africa. Climate of South Africa. Publication WB 14 South African
Weather Bureau.
WEATHER BUREAU. S.A. 1954a. Climate statistics. Climate of
South Africa. Publication WB i9 South African Weather Bureau.
WEATHER BUREAU, S.A. 1954b. Rainfall statistics. Climate of
South Africa. Publication WB 20 South African Weather Bureau.
WEATHER BUREAU, S.A. 1960. Surface winds. Climate of South
Africa. Publication WB 26 South African Weather Bureau.
WEATHER BUREAU, S.A. 1965a. General survey. Climate of South
Africa. Publication WB 28 South African Weather Bureau.
WEATHER BUREAU, S.A. 1965b. Average monthly rainfall. Cli-
mate of South Africa. Publication WB 29 South African Weather
Bureau.
Z1ETSMAN, A. L. 1964. The geology of the Sabie-Pilgrim' s Rest
goldfield. M.Sc. thesis. University of the Orange Free State.
Bothalia 19,1:69-89(1989)
The vegetation ecology of the Eastern Transvaal Escarpment in the
Sabie area. 2. Floristic classification
G. B. DEALL*, G. K. THERON** and R. H. WESTFALL*
Keywords: Eastern Transvaal Escarpment, floristic classification, phytosociology, PHYTOTAB
ABSTRACT
The indigenous vegetation of the Eastern Transvaal Escarpment in the Sabie area is classified with the aid of the
PHYTOTAB program package. Four ecological-formation classes (efc) based on floristics, physiognomy and climate
correspond to four data subsets. Plant communities in each efc are defined by means of 46 differential species-groups
distributed amongst forest, thicket, woodland, shrubland and grassland structural types. Environmental correlation is
facilitated by means of 21 habitat types.
UITTREKSEL
Die inheemse plantegroei van die Oos-Transvaalse platorand in die Sabiegebied is met behulp van die PHYTOTAB-
programpakket geklassifiseer. Vier ekologiese formasieklasse (efc) wat op floristiek, fisionomie en klimaat gebaseer is,
kom met vier datasubstelle ooreen. Plantgemeenskappe in elke efc word gedefinieer deur middel van 46 differensiele
spesiegroepe wat versprei is tussen woud-, ruigte-, bosveld-, struikveld- en grasveldstrukturele tipes. Omgewingskorrelasie
word vergemaklik deur middel van 21 habitattipes.
INTRODUCTION
To facilitate land-use planning and management of the
Eastern Transvaal Escarpment on a regional and sub-
regional basis, the scale of Acocks’s (1975) veld types
needs to be enlarged (Van der Meulen & Scheepers
1978). A more detailed classification of Veld Types 8
(North-Eastern Mountain Sourveld) and 9 (Lowveld
Sour Bushveld) is therefore required. Floristic classifica-
tions express relationships between plant communities
and their environment and are thus potentially suitable
planning tools. This paper describes a detailed floristic
classification of Veld Types 8 and 9 in the Sabie area,
within the environmental context established by Deall et
al. (1989). The classification is purely descriptive and
predictive, implying no explanation of causality.
METHODS
The study area comprises a broad transect whose lo-
cality and physical environment are described compre-
hensively by Deall et al. (1989). Within the transect 251
quadrats (measuring 10 x 20 m each) were distributed
subjectively amongst 50 different physiognomic/land
type stratification units based on 1: 250 000 Land Type
Series 2530 Barberton. The quadrats were distributed as
follows: 46 in forest, 64 in thicket, 76 in woodland, nine
in shrubland and 56 in grassland [physiognomy based on
Edwards’s (1983) formation classes]. In each quadrat,
all plant species were listed together with their Domin-
Krajina cover-abundance values and growth forms
(Deall 1985). The total canopy cover and estimated
height range of each stratum of vegetation was also
* Botanical Research Institute, Private Bag X 101 , Pretoria 0001 .
** Department of Botany, University of Pretoria, Pretoria 0002.
MS. received: 1988.10.11.
recorded for the purpose of physiognomic-structural
classification (Edwards 1983). Environmental parame-
ters recorded for each quadrat include the following:
(i) Climatic Belt, based on altitudinal distribution of
mist (cf. Deall etal. 1989).
(ii) Physiographic Zone, based on altitude and geo-
morphology (cf. Deall etal. 1989).
(iii) Geomorphology, based on position in local land-
scape. Eight classes were recognized: knolls, terraces,
upper slopes, midslopes, foot slopes, dry kloofs, moist
stream banks, marshes.
(iv) Aspect, expressed as 'mesoclinal’ (67°-247°) and
‘xeroclinal’ (248°-66°).
(v) Exposure in terms of the degree of exposure to sun
and wind, expressed as sheltered, partly sheltered or
exposed.
(vi) Lithology (field nomenclature), based on the stra-
tigraphic classification for South Africa (SACS 1980), as
adapted by Deall etal. (1989).
(vii) Rock cover based on the degree of limitation on
mechanical utilization (cf. Van der Meulen 1979).
By means of selected programs in PHYTOTAB
(Westfall et al. 1982), a provisional phytosociological
classification was made on the basis of the entire data set
(251 releves and 1 043 species, including unidentified
specimen numbers). Four data subsets were then identi-
fied on the basis of floristic and environmental disconti-
nuity (Deall 1985). Species in each subset were then
reclassified within the confines of their subset distribu-
tion (Coetzee 1983). Thus, both the number of syntaxa
(represented by releve-groups) and the species defining
them (represented by differential species-groups), were
increased relative to the provisional classification. For
the sake of brevity, poorly-defined syntaxa were re-
moved, leaving only those that are clearly defined by
70
Bothalia 19,1 (1989)
I £
O o
_ u,
■o £
E "n
•1 =3 'Z
o « M
O « c
(/) Cl. d
« O 43
£ C
1) r*
S .« C a
*j 05 " o
■ £ g I55”
"saS“
J5> o o £
CU X
O O o ^
E
-a •§ o,
<V M
L"ga3
c S-^
5 o< j=
3
— 'S
u 5
i2 A
CD ^
g. C
5 6
o« £
1) *N
II
C/3 3
•X §■
=3 •*
H U!
o a> - c o.
<u o ‘s c 2, _
. E ■§ § *>
§" ”c3 3 S w
&
C/3 o W ID
s
•S I
. o c
0 3 1)
5^0^,
SS —
« 'i § «>
•3 1 1 8.
c 3 "* O <■
' aM.S® '
^£.s'
-♦— • O * — \ cd
2 ail
<3 ^3
E «?
r? S
Bothalia 19,1 (1989)
71
FIGURE 1. — Protected valleys in
Lower Mountain grasslands
support the Hypoestes triflo-
ra-Dovyalis lucida Tall/Short
Forest (c.3).
differential species-groups. Both non-differential species
and species-groups defining more than a single syntaxon
were also removed*. Syntaxa were reduced to single
columns by transforming the cover-abundance values of
differential species to constancy values thereby creating
synoptic tables (Deall 1985).
RESULTS
The four data subsets correspond to four ecological-
formation classes (efc) based on floristics, physiognomy
and climate (especially mist). Each efc is represented by
a synoptic table (Tables 2, 4, 6 & 8). The syntaxa in each
efc are informally ranked as communities and are
grouped according to habitat type (Tables 1 , 3, 5 & 7).
Communities are named according to the recommen-
dations for a standardized South African syntaxonomic
nomenclature system proposed by the Botanical
Research Institute, Pretoria (Scheepers et al. MS). Each
name comprises a differential species followed by a
dominant species and a physiognomic-structural term
(Edwards 1983).
Further information on species mentioned in the text
but not included in synoptic tables is documented in an
annotated checklist (Deall & Backer 1989).
In instances where plants could not be identified be-
yond the generic level, the epithet sp. is appended (e.g.
Combretum sp.). It should be noted that such a name
may not necessarily refer to a consistent species concept,
but may include various species within the genus.
Differential species are not listed in the community
descriptions which follow but are referred to in the syn-
optic tables instead (Tables 2, 4, 6 & 8).
1 FOREST AND MESIC THICKET OF THE MISTBELT AND
LOW COUNTRY
Mesic Thicket (Table 1) refers to a denser, more lush
type of thicket which normally occurs in the Mistbelt.
* Full phytosociological tables are nevertheless available on request
from the Vegetation Ecology Division, Botanical Research Institute,
Private Bag X 1 0 1 , Pretoria 000 1 .
Where it occurs in the Low Country, it is usually in
mesic situations, i.e. riparian or south-facing. Total
vegetation cover is usually more than 75 %. Associated
communities are represented throughout the altitudinal
range but are notably absent on the Summit Peak and on
the Escarpment and Summit Plateaux.
1 . 1 Humid Mistbelt forest associated with Timeball
Hill Shale and Klapperkop Quartzite
c.l Buddleja salviifolia—Leucosidea sericea Tall
(riparian) Forest
Confined to steep moist kloofs on Summit and Upper
Mountain Slopes, notably at Mount Anderson and Blyf-
staanhoogte. Differential and dominant species are indi-
cated in Table 2 (Group A). Non-differential dominants
include Diospyros whyteana.
c.2 Ochna arborea var. arborea — Combretum kraussii
High Forest
Covering extensive areas (e.g. 'Maritzbos') on
mesoclinal Mountain Slopes. Differential and dominant
species are indicated in Table 2 (Group B). Non-differ-
ential dominants include Cussonia spicata, Syzygium
gerrardii, Oxyanthus speciosus subsp. gerrardii, Cassi-
pourea gerrardii, Sclerochiton harveyanus and Olea
capensis subsp. macrocarpa.
1.2 Humid Mistbelt forest associated with Dolo-
mite,Lower Mountains
c.3 Hypoestes triflora— Dovyalis lucida Tall/Short
Forest
Occurring on footslopes and stream banks, Vertroost-
ing Nature Reserve (Figure 1). Differential and dominant
species are indicated in Table 2 (Group C). Non-differ-
ential dominants include Xymalos monospora and Scle-
rochiton harveyanus.
c.4 Streptocarpus cyaneus— Dovyalis lucida Short
Forest
72
Bothalia 19,1 (1989)
FIGURE 2. — Ekebergia pterophylla
— Psychotria zombamontana
Tall/Short Forest associated
with Black Reef Quartzite out-
crops, Plateau Crest (c.5). Aloe
arborescens is noteworthy on
massive boulders protruding
through the canopy.
Confined to steep mesoclinal cliffs, notably at Sabie.
Differential species are indicated in Table 2 (Group D).
Dominant species include Kiggelaria africana, Diospy-
ros why team and Polystichum luctuosum.
1.3 Humid Mistbelt forests associated with Black
Reef Quartzite outcrops
c.5 Ekebergia pterophylla— Psychotria zombamontam
Tall/Short Forest
Scattered on rocky knolls of the Plateau Crest, notably
in Mac-Mac Nature Reserve (Figure 2). Differential and
dominant species are indicated in Table 2 (Group E).
Non-differential dominants include Clivia caulescens
and Syzygium gerrardii.
1.4 Humid Mistbelt forests associated with Nel-
spruit Granite
c.6 Clerodendrum myricoides— Syzygium gerrardii
Tall Forest
Occurring on steep mesoclinal slopes and kloofs of
Escarpment Upper Slopes (Figure 3). Differential
species are indicated in Table 2 (Group F). Dominant
species include Combretum kraussii, Oxyanthus specio-
sus subsp. gerrardii, Psychotria cape ns is and Tricalysia
cape ns is.
1.5 Transitional Misbelt thickets associated with
Nelspruit Granite
c.7 Clematis brachiata— Acacia ataxacantha Short
Thicket
Covering sheltered mesoclinal slopes of Upper Foot-
hills, especially Bergvliet Forest Reserve. Differential
species are indicated in Table 2 (Group G). Dominant
species include Brachylaem discolor subsp. transvml-
ensis, Protorhus longifolia, Maytenus mossambicensis
subsp. mossambicensis, Canthium gueinzii and Rhoicis-
sus tomentosa.
c.8 Tetradenia ‘complex’ -Acacia ataxacantha Short
Thicket
Occurring patchily on partly sheltered xeroclinal sites
of the Escarpment Lower Slopes (Figure 4). Differential
species are indicated in Table 2 (Group H). Dominant
species include Canthium inerme and Smilax kraussiana.
FIGURE 3. — Clerodendrum myri-
coides-Syzygium gerrardii Tall
Forest (c.6) showing Tricalysia
capensis dominating the under-
storey.
Bothalia 19,1 (1989)
73
FIGURE 4. — Woodland variant of
the Tetradenia ‘complex’ -Aca-
cia ataxacantha Short Thicket
on partly sheltered sites of the
Escarpment Lower Slopes (c.8).
1 .6 Low Country thickets associated with Nelspruit
Granite and Transvaal Diabase, Upper Foot-
hills
c.9 Pittosporum viridiflorum— Acacia ataxacantha
Short Thicket
Scattered on diverse xeroclinal slopes. Differential
species are indicated in Table 2 (Group I). Dominant
FIGURE 5. — Pavetta sp .—Celtis africana Tall Forest on sheltered
banks of the Sabie river, Lower Foothills (c. 1 2).
species include Syzygium cor datum , Parinari curatelli-
folia and Diospyros whyteana.
c.10 Pycnostachys urticifolia— Acacia ataxacantha
Short Thicket
Occurring intermittently on mesoclinal upper slopes.
Differential species are indicated in Table 2 (Group J).
Dominant species include Catha edulis, Diospyros why-
teana, Diospyros lycioides subsp. sericea, Smilax kraus-
siana and Euclea ‘complex’ .
1.7 Low Country forests associated with Nelspruit
Granite, Lower Foothills
c. 1 1 Schoenoplectus corymbosus— Syzygium cor datum
Tall (riparian) Forest
Confined to sheltered stream banks. Differential spe-
cies are indicated in Table 2 (Group K). Dominant spec-
ies include Anthocleista grandiflora, Syzygium corda-
tum, Diospyros whyteana, Oplismenus hirtellus and Dal-
bergia armata.
c. 12 Pavetta sp. —Celtis africana Tall Forest
Situated on mesoclinal midslopes and stream banks
(Figure 5). Differential species are indicated in Table 2
(Group L). Dominant species include Monanthotaxis
caffra, Dalbergia armata, Rhoicissus tomentosa. May te-
rns undata, Oplismenus hirtellus and Dietes iridioides.
2 WOODLAND AND XERIC THICKET OF THE LOW
COUNTRY
Xeric Thicket (Table 3) refers to the ‘scrubby’ type of
thicket normally associated with the Low Country. Asso-
ciated communities are represented mainly on xeroclinal
upper slopes and midslopes underlain by Nelspruit Gran-
ite or Transvaal Diabase.
2.1 Partly sheltered woodlands and thickets of
rocky sites, Lower Foothills
c.13 Monanthotaxis caffra— Rhus pentheri Short
Thicket
74
Bothalia 19,1 (1989)
TABLE 2. — Floristic classification of Forest and Mesic Thicket of the Mistbelt and Low Country
* Dominant species. Digits 1 -5 in matrix denote constancy values. For explanation of ‘complex see Deall & Backer (1989). Growth
forms: TR = Tree; SH = Shrub; LN = Lianoid; EP = Epiphyte; FB = Forb or herb; CY = Sedge: GR = Grass; PT = Fern.
Bothalia 19,1 (1989)
75
=:
5
A
76
Notable around Sandford, with a vegetation cover of
more than 80 %. Differential species are indicated in
Table 4 (Group A). Dominant species include Zanthoxy-
lum davyi. Acacia ataxacantha, Catha edulis and Mayte-
nus undata.
c.14 Combretum collinum subsp. gazense—Panicum
maximum Short Closed Woodland
Widespread, with a vegetation cover of less than
80 %. Differential species are indicated in Table 4
(Group B). Dominant species include Combretum colli-
num subsp. suluense. Heteropyxis natalensis, Panicum
‘complex’ and Aloe barbertoniae .
2.2 Partly sheltered woodlands and thickets of
rocky sites, Upper Foothills
c.15 Hyparrhenia gazensis—Bauhinia galpinii Short
Thicket
Situated around Evert and Emmett with a vegetation
cover of more than 75 %. Differential species are indi-
cated in Table 4 (Group C). Dominant species include
Dombeya rotundifolia subsp. rotundifolia, Acacia ataxa-
cantha, Maytenus heterophylla and Protasparagus sp.
FIGURE 6. — Dicoma zeyheri—Parituiri curatellifolia Short Closed
Woodland on xeroclinal granite slope. Upper Foothills (c. 16).
Loudetia simplex dominates the field layer.
Bothalia 19,1 (1989)
c.16 Dicoma zeyheri—Parinari curatellifolia Short
Thicket/Closed Woodland
Notable at Zeederberg with a vegetation cover of more
than 75 % (Figure 6). Differential species are indicated
in Table 4 (Group D). Dominant species include Ptero-
carpus angolensis, Catha edulis, Faurea speciosa, Dios-
pyros whyteana, Bauhinia galpinii, Smilax kraussiana
and Flemingia grahamiana.
c.17 Rhynchosia sordida—Parinari curatellifolia
Short Closed Woodland
Widespread, with a vegetation cover of more than
75 %. Differential species are indicated in Table 4
(Group E). Dominant species include Faurea saligna,
Themeda triandra, Flemingia grahamiana and Cymbo-
pogon ‘complex’.
2.3 Exposed woodlands and shrublands with more
than 50 % vegetation cover (disturbed) and low
rock cover
c.18 Mucuna coriacea subsp. irritans—Antidesma
venosum Low Thicket/Short Closed Woodland
Occurring on heavily utilized sites. Upper Foothills.
Differential species are indicated in Table 4 (Group F).
Dominant species include Parinari curatellifolia, Bauhi-
nia galpinii, Flemingia grahamiana and Hyperthelia dis-
soluta.
c.19 Diheteropogon amplectens— Parinari curatelli-
folia Low Open Woodland
Associated with old-land disturbance, mainly on Up-
per Foothills. Differential species are indicated in Table
4 (Group G). Dominant species include Trema orienta-
ls, Smilax kraussiana, Flemingia grahamiana and Schi-
zachyrium sanguineum.
c.20 Andropogon schirensis— Parinari curatellifolia
Short Open Woodland
Situated on disturbed sites of Escarpment Lower
Slopes, notably Bergvliet Forest Reserve. Differential
species are indicated in Table 4 (Group H). Dominant
species include Loudetia simplex and Schizachyrium
sanguineum.
c.21 Vernonia centaureoides— Hyperthelia dissoluta
Low Sparse Woodland/Tall Sparse Shrubland
Occurring on Upper Foothills in disturbed areas. Dif-
ferential species are indicated in Table 4 (Group I).
Dominant species include Parinari curatellifolia.
2.4 Exposed woodlands and shrublands with less
than 50 % vegetation cover and high rock cover
(usually granite sheet rock)
c.22 Ceratotheca triloba- Bequaertiodendron maga-
lismontanum Low Open Woodland
Occurring mainly on Upper Foothills and Escarpment
Lower Slopes. Differential species are indicated in Table
4 (Group J). Dominant species include Combretum molle
and Loudetia simplex.
Bothalia 19,1 (1989)
TABLE 4. — Floristic classification of Woodland and Xeric Thicket of the Low Country
Habitat type
Community number
Total releves per community
2.1
2.2
16
2
2.3
2.4
A. Differential species of Community 13
Monanthotaxis caffra (SH)
Mimusops zeyheri (TR)
Ansellia gigantea (EP)
Eulophia streptopetala (FB)
Scadoxus multifloras subsp. multiflorus (FB)
Plectranthus sp. (SH)
B. Differential species of Community 14
Combretum collinum subsp. gazense (TR)
Terminalia sericea (TR)
Senecio venosus (FB)
Vernonia natalensis (FB)
Lannea discolor (TR)
Chaetacanthus burchellii (FB)
Diospyros mespiliformis (TR)
Strychnos madagascariensis (TR)
Phyllanthus reticulatus (SH)
Maytenus mossambicensis subsp. mossambicensis (TR)
Differential species of Community 15
Hyparrhenia gazensis (GR)
Neonotonia wight ii (LN)
Triumfetta pilosa var. pilosa (FB)
Euclea natalensis (TR)
Ocimum urticifolium (SH)
Erianthemum dregei (EP)
Sphenostylis marginata subsp. marginata (LN)
D. Differential species of Community 16
Dicoma zeyheri (FB)
Thunbergia atriplicifolia (LN)
Aster sp. (FB)
Passiflora edulis (LN)
Tridactyle tricuspis (EP)
Trimeria grandifolia (TR)
lpomoea crassipes (FB)
Differential species of Community 17
Rhynchosia sordida (FB)
Senecio serratuloides var. serratuloides (FB)
F. Differential species of Community 18
Mucuna coriacea subsp. irritans (LN)
Triumfetta pilosa var. tomentosa (FB)
G. Differential species of Community 19
Diheteropogon amplectens (GR)
Acanthospermum australe (FB)
Cassia quarrei (FB)
Hemizygia transvaalensis (FB)
Cony z a sumatrensis (FB)
H. Differential species of Community 20
Andropogon schirensis (GR)
Gladiolus exiguus (FB)
Diospyros galpinii (FB)
Eriosema gunniae (FB)
I. Differential species of Community 21
Vernonia centaureoides (FB)
Indigofera oxalidea (FB)
J. Differential species of Community 22
Ceratotheca triloba (FB)
Crassula natalensis (FB)
Pellaea calomelanos (PT)
Helichrysum kraussii (FB)
Protorhus longifolia (TR)
Brachiaria serrata var. serrata (GR)
Cephalanthus natalensis (LN)
. Differential species of Community 23
*Aloe petricola (FB)
* Myrothamnus flabellifolia (FB)
* Coleochloa setifera (CY)
* Dominant species. Digits 1 -5 in matrix denote constancy values.
Epiphyte; FB =Forb or herb; CY = Sedge; GR = Grass; PT =Fern.
Growth forms; TR — Tree; SH — Shrub; LN — Lianoid; EP
78
Bothalia 19,1 (1989)
FIGURE 7. — Aloe petricola—Coleo-
chloa setifera Short Sparse
Shrubland on granite ‘dwala'
(c.23).
c.23 Aloe petricola—Coleochloa setifera Short Sparse
Shrubland
Widespread on Foothills and Escarpment Lower
Slopes (Figure 7). Differential and dominant species are
indicated in Table 4 (Group K). Non-differential domi-
nants include Cheilanthes viridis var. viridis.
3 WOODLAND AND SHRUBLAND OF THE MISTBELT
Associated communities are evenly distributed be-
tween Humid and Subhumid Mistbelts (Table 5).
3.1 Humid Mistbelt communities on partly shel-
tered midslopes with low rock cover
c.24 Galopina aspera—Faurea speciosa Low Open
Woodland/Low Thicket
Notable in Spitskop Forest Reserve on xeroclinal sites
underlain by Black Reef Quartzite, Escarpment Upper
Slopes (Figure 8). Differential species are indicated in
Table 6 (Group A). Dominant species include Faurea
speciosa, Rhynchosia komatiensis, Acalypha wilmsii.
Eulalia villosa, Smilax kraussiana and Flemingia gra-
hamiana.
c.25 Artemisia afra—Bowkeria cymosa Low Thicket/
Low Open Woodland
Occurring in Vertroosting Nature Reserve on mesocli-
nal sites underlain by dolomite, Lower Mountain Slopes.
Differential species are indicated in Table 6 (Group B).
Dominant species include Cussonia spicata, Bowkeria
cymosa, Rhus transvaalensis, Indigofera swaziensis,
Rhoicissus tridentata and Pteridium aquilinum.
3.2 Humid Mistbelt communities on exposed Black
Reef Quartzite outcrops, Plateau Crest and
Escarpment Upper Slopes
c.26 Tecomaria capensis subsp. capensis— Bequaer-
tiodendron magalismontanum Low Closed
Woodland
Situated on xeroclinal upper slopes at Sabie. Differen-
tial and dominant species are indicated in Table 6 (Group
C). Non-differential dominants include Psychotria
capensis, Helichrysum kraussii and Cyperus leptocla-
dus.
FIGURE 8. — Galopina aspera—
Faurea speciosa Low Open
Woodland of Escarpment Upper
Slopes (c.24). The community
is evidently maintained by fire.
TABLL 5. Woodland and Shrubland of the Mistbelt: an ecological basis for the recognition of five habitat types (3.1 -3.5) and 1 1 communities (c.24-c.34), confer Table 6
Bothalia 19,1 (1989)
79
c Ha ,u
S « s £ a
5 2 is ” £ u
K ts o g t .s
c 73 i 35 ft£ £
«> c js a. 5 "3
>. ~ "■ ^ .3 § 3
5
i"cEo
£5h
.. o =3
5(5 j; eo
£ ft-°
5 tr u
H O £ «
™ £ i§ 73 „
E 5 H j= <n
co m
8 >,E3i
0£.HI
Q- p , >
X q 3
W-S=a
80
Bothalia 19,1 (1989)
TABLE 6. — Floristic classification of Woodland and Shrubland of the Mistbelt
* Dominant species. Digits 1 -5 in matrix denote constancy values. For explanation of 'complex’ see Deall & Backer ( 1 989). Growth
forms: TR=Tree; SH=Shrub; LN=Lianoid; EP=Epiphyte; FB = Forb or herb; CY=Sedge; GR = Grass; PT = Fern.
Bothaha 19,1 (1989)
81
TABLE 6. — Floristic classification of Woodland and Shrubland of the Mistbelt (continued)
* Dominant species. Digits 1 -5 in matrix denote constancy values. For explanation of ‘complex’ see Deall& Backer (1989). Growth
forms: TR =Tree; SH = Shrub; LN=Lianoid; EP = Epiphyte; FB = Forb or herb; CY = Sedge; GR = Grass; PT=Fern.
c.27 Diospyros galpinii-Bequaertiodendron magalis-
montanum Tall Open Shrubland/Low Open
Woodland
Notable at Sabie on xeroclinal upper slopes. Differen-
tial and dominant species are indicated in Table 6 (Group
D). Non-differential dominants include Loudetia sim-
plex, Panicum natalense, and Rhynchosia monophylla.
c.28 Selago atherstonei—Syzygium cordatum Low
Open Woodland
Occurring in Spitskop Forest Reserve on xeroclinal
terraces and slopes (Figure 9). Differential and dominant
species are indicated in Table 6 (Group E). Non-differ-
ential dominants include Bequaertiodendron tnaga-
lismontanum and Loudetia simplex.
c.29 Clutia monticola— Loudetia simplex Low Open
Woodland
Widespread on xeroclinal upper slopes. Differential
species are indicated in Table 6 (Group F). Dominant
species include Helichrysum mimetes and Rhus tumuli-
cola.
3.3 Subhumid Mistbelt communities of exposed
rocky upper slopes with less than 75 % vegeta-
tion cover
c.30 Trachypogon spicatus—Rhus tumulicola Low
Open Woodland/Tall Open Shrubland
Confined to Klapperkop Quartzite outcrops on meso-
clinal Summit Slopes, Hartebeestvlakte (Figure 10). Dif-
ferential and dominant species are indicated in Table 6
(Group G). Non-differential dominants include Protea
caffra.
c.31 Festuca costata var. costata—Cliffortia nitidula
subsp. pilosa Short Closed/Open Shrubland
Occurring in association with Hekpoort Andesite out-
crops on xeroclinal Summit Peaks, notably Mount
Anderson. Differential and dominant species are indi-
cated in Table 6 (Group H). Non-differential dominants
include Myrsine africana, Vaccinium exul, Helichrysum
splendidum and Cliffortia nitidula subsp. pilosa.
Bothalia 19,1 (1989)
FIGURE 9. — Selago atherstonei—
Syzygium cordatum Low Open
Woodland of Black Reef
Quartzite outcrops, Plateau
Crest (c.28). Aloe petricola is
visible bottom right and centre
left.
i % +* ’
FIGURE 10. — Trachypogon spica-
tus—Rhus tumulicola Low Open
Woodland associated with Klap-
perkop Quartzite outcrops.
Summit Slopes (c.30). The con-
spicuous tree is Proiea cajfra.
is
FIGURE 11. — Gunnera perpensa —
Nidorella auriculata Short Open
(marshy) Shrubland on deep hy-
dromorphic soils overlying
Timeball Hill Shale (c.32).
Eucomis aulumnalis subsp. cla-
vata and Gunnera perpensa are
conspicuous.
Bothalia 19,1 ( 1989)
83
FIGURE 12. — Tall Closed (marshy)
Grassland variant of Commu-
nity 32. Andropogon appendi-
culatus is totally dominant.
3.4 Subhumid Mistbelt communities of marshes
and stream banks with low rock cover and more
than 75% vegetation cover, Summit Plateau
c.32 Gunnera perpensa—Nidorella auriculata Short
Open (marshy) Shrubland/Tall Closed (marshy)
Grassland
Confined to exposed, level terraces with deep hydro-
morphic alluvium overlying Timeball Hill Shale, Harte-
beestvlakte (Figures 11 & 12). Differential and dominant
species are indicated in Table 6 (Group I). Non-differen-
tial dominants include Rabdosiella calycina, Nerine an-
gustifolia and Andropogon appendiculatus .
c.33 Cyathea dregei— Hypericum revolutum Low
Open Woodland
Situated at Hartebeestvlakte on partly sheltered, meso-
clinal stream banks with fairly shallow soils overlying
Timeball Hill Shale or Klapperkop Quartzite (Figure
13). Differential and dominant species are indicated in
Table 6 (Group J). Non-differential dominants include
Cliffortia nitidula subsp. pilosa.
3.5 Subhumid Mistbelt communities of steep rocky
kloofs in Timeball Hill Shale, Summit Slopes
c.34 Aloe arborescens—Rhus pyroides var. gracilis
Low Bushland
Occurring in sheltered xeroclinal sites below Mount
Anderson (Figure 14). Differential and dominant species
are indicated in Table 6 (Group K). Non-differential
dominants include Hypericum revolutum.
4 GRASSLAND OF THE MISTBELT
Associated communities are all fire-maintained, oc-
curring on exposed sites with low rock cover. They are
represented over all geological substrates. Table 7.
4.1 Humid Mistbelt grasslands of Escarpment
Slopes overlying Nelspruit Granite (more than
75 % vegetation cover)
c.35 Gladiolus densiflorus—Loudetia simplex Short
Closed Grassland
Notable at Hebron as relic patches on mesoclinal foot
slopes with low rock cover. Differential and dominant
species are indicated in Table 8 (Group A). Non-differ-
ential dominants include Monocymbium ceresiiforme
and Athanasia acerosa.
c.36 Cliffortia repens— Loudetia simplex Short Open
(grassy) Shrubland
Occurring as relic patches on xeroclinal upper slopes
with moderate rock cover, Frankfort Forest Reserve.
Differential and dominant species are indicated in Table
8 (Group B). Non-differential dominants include Era-
grostis capensis, Pearsonia sessilifolia ‘complex’ and
Hemizygia subvelutina.
4.2 Humid Mistbelt grasslands of Escarpment Pla-
teau (less than 75 % vegetation cover)
c.37 Tephrosia elongata— Monocymbium ceresiiforme
Low Closed Grassland
Situated in Mac-Mac Nature Reserve on xeroclinal
terraces overlying Black Reef Quartzite with moderate
rock cover. Differential species are indicated in Table 8
(Group C). Dominant species include Bulbostylis schoe-
noides and Loudetia simplex.
c.38 Helichrysum cephaloideum— Monocymbium cere-
siiforme Low Closed Grassland
Confined to Mac-Mac Nature Reserve on xeroclinal
terraces overlying Black Reef Quartzite with low rock
cover (Figure 15). Differential species are indicated in
Table 8 (Group D). Dominant species include Rendlia
altera, Loudetia simplex and Becium obovatum.
c.39 Wahlenbergia huttonii—Eragrostis racemosa
Low Closed Grassland
Situated around Sabie vicinity on xeroclinal terraces
overlying Oaktree Dolomite with low rock cover. Differ-
ential species are indicated in Table 8 (Group E). Domi-
nant species include Loudetia simplex, Themeda trian-
dra, Bulbostylis schoenoides and Helichrysum pilosel-
lum.
84
Bothalia 19,1 (1989)
FIGURE 13. — Cyathea dregei—
Hypericum revolutum Low
Open Woodland in drainage
lines of Summit Plateau (c.33).
FIGURE 14. — Aloe arborescens-
Rhus pyroides var. gracilis Low
Bushland of steep rocky kloofs
in Timeball Hill Formation,
Summit Slopes (c.34).
FIGURE 15. — Helichrysum cepha-
loideum —Monocymbium cere-
siiforme Low Closed Grassland
of Escarpment Plateau (c.38).
Pleridium aquilinum is domi-
nant in patches.
TABLE 7.— Grassland of the Mist belt: an ecological basis for the recognition of five habitat types (4.1 -4.5) and 12 communities (c.35-c.46), confer Table 8
Bothalia 19,1 (1989)
85
1 °-
E -2
1/5 =8 3
■s gi
-o 5
2 3
w £
II?
3^2 O
CO fi- w
E 2
— i_ <u
• a S
a. o
^ o
Q
a o
E
Cl >
s ^
o °
w
SD
CC O
- o £
C* N
»
£ O
■3 i
a) 5 ^
O-XJ >
= I8
$
86
Bothalia 19,1 (1989)
TABLE 8.- — Floristic classification of Grassland of the Mistbelt
* Dominant species. Digits 1-5 in matrix denote constancy values. For explanation of 'complex’ see Deall& Backer (1989). Growth
forms: TR=Tree; SH = Shrub; LN = Lianoid; EP=Epiphyte; FB = Forb or herb; CY=Sedge; GR = Grass; PT=Fern.
Bothalia 19,1 (1989)
87
TABLE 8. — Floristic classification of Grassland of the Mistbelt (continued)
H.
I.
*
J.
K.
L.
Habitat type
Community number
Total releves per community
35
4
Differential species of Community 42
Lippia javanica (SH)
Flemingia grahamiana (FB)
Rhus discolor (SH)
Aristea woodii (FB)
Rhus dentata (TR)
Indigo f era hilaris (FB)
Cyphia elata ‘complex’ (FB)
Crassula natalensis (FB)
Heteromorpha pubescens (SH)
Acalypha caperonioides (FB)
Leonotis ‘complex’ (FB)
Helichrysum mixtum (FB)
Differential species of Community 43
Indigofera sp. 1 (FB)
Protea caffra (TR) 2
Dicoma anomala subsp. cirsioides (FB)
Rhynchosia anguiosa (FB)
Acalypha angustata vat. glabra (FB)
Cvperus obtusiflorus var. flavissimus (CY)
Faurea speciosa (TR) 2
Differential species of Community 44
Helichrysum wilmsii (FB)
Drosera burkeana (SH)
Helichrysum glomeratum (FB)
Helichrysum spiralepis (FB)
Helichrysum subluteum (FB)
Scleria dieterlenii (CY)
Monsonia transvaalensis (FB)
Clerodendrum triphyllum var. triphyllum (SH)
Differential species of Community 45
Eriosema kraussianum (FB)
Protea gaguedi (TR)
Gladiolus longicollis var. platypetalus (FB)
Gnidia caffra (FB)
Oxalis obliquifolia (FB)
Sporobolus centrifugus (GR)
Vernonia thodei (FB)
Differential species of Community 46
Erica atherstonei (SH)
Hypoxis galpinii (FB)
Alepidea sp. (FB)
Hebenstretia comosa (FB)
Peucedanum sp. (FB)
Psammotropha myriantha (FB)
4.1
4.2
39
5
4.3
4.4
4.5
46
3
1 4
2
2
3
9
2
1
2
2
1
2
1
1
2
1 3
2
1 2
2
1 2
2 2
2
3
1
4
4
5
4
4
4
* Dominant species. Digits 1-5 in matrix denote constancy values. For explanation of ‘complex’see Deall& Backer (1989). Growth
forms: TR=Tree; SH = Shrub; LN = Lianoid; EP= Epiphyte; FB = Forb or herb; CY = Sedge; GR = Grass; PT = Fern.
c.40 Digitaria monodactyla—Loudetia simplex Low
Closed Grassland
Notable in Sabie vicinity on xeroclinal terraces overly-
ing Black Reef Quartzite with moderate rock cover. Dif-
ferential and dominant species are indicated in Table 8
(Group F). Non-differential dominants include Eragros-
tis capensis.
c.41 Asclepias crassinervis—Andropogon schirensis
Short Closed Grassland
Situated on mesoclinal terraces overlying Black Reef
Quartzite with low rock cover, notably in Spitskop For-
est Reserve. Differential species are indicated in Table 8
(Group G). Dominant species include Themeda triandra.
4.3 Humid Mistbelt grasslands of Lower Mountain
Slopes
c.42 Lippia javanica— Loudetia simplex Short Closed
Grassland/Tall Open Shrubland
Occurring on xeroclinal slopes overlying Upper Dolo-
mite with low rock cover, Vertroosting Nature Reserve.
Differential species are indicated in Table 8 (Group H).
Dominant species include Themeda triandra and Hemi-
zygia transvaalensis.
c.43 Indigofera sp .—Monocymbium ceresiiforme Low
Closed Grassland/Sparse Woodland
Notable in Vertroosting Nature Reserve on xeroclinal
upper slopes overlying Bevet’s Conglomerate with low
Bothalia 19,1 (1989)
FIGURE 16. — Helichrysum wilmsii
—Andropogon schirensis Low
Closed Grassland of Summit
Plateau (c.44). Sopubia cana
var. cana is the dominant forb.
rock cover. Differential and dominant species are indi-
cated in Table 8 (Group I). Non-differential dominants
include Themeda triandra, Loudetia simplex and Erio-
sema ellipticifolium.
4.4 Subhumid Mistbelt grasslands overlying Time-
ball Hill Shale with low rock cover and mesocli-
nal aspect
c.44 Helichrysum wilmsii— Andropogon schirensis
Low Closed Grassland
Situated on terraces and slopes of the Summit Plateau,
Hartebeestvlakte (Figure 16). Differential species are in-
dicated in Table 8 (Group J). Dominant species include
Festuca costata var. costata , Themeda triandra , Allote-
ropsis semialata subsp. eckloniana , Loudetia densispica
and Sopubia cana var. cana.
c.45 Eriosema kraussianum—Rendlia altera Low
Closed Grassland
Occurring on eroded terraces and slopes of the Sum-
mit Plateau and Summit Slopes, Hartebeestvlakte (Fig-
ure 17). Differential species are indicated in Table 8
(Group K). Dominant species include Alloteropsis se-
mialata subsp. eckloniana, Andropogon schirensis, Fes-
tuca costata var. costata, Monocymbium ceresiiforme
and Loudetia densispica.
4.5 Subhumid Mistbelt grasslands overlying Hek-
poort Andesite with high rock cover and xero-
clinal aspect
c.46 Erica atherstonei—Harpochloa falx Low Closed
Grassland/Open (grassy) Shrubland
Confined to upper slopes of the Summit Peak, at
Mount Anderson. Differential and dominant species are
indicated in Table 8 (Group L). Non-differential domi-
nants include Rendlia altera, Festuca costata var. cos-
tata, Protea parvula and Erica cerinthoides var. cerin-
thoides.
CONCLUSION
At the finest level of detail, the classification of vege-
tation in the Sabie area has elicited 46 floristic entities
FIGURE 17. — Eriosema kraussia-
num— Rendlia altera Low
Closed Grassland of eroded ter-
races and slopes, Summit
(c.45).
Bothalia 19,1 (1989)
89
(communities), defined on the basis of differential spe-
cies. Environmental correlation enhances the descriptive
and predictive value of the classification. As a planning
and management tool, however, it is obviously incom-
plete. Aspects such as causality and community function
must yet be investigated in association with relevant key
questions. The floristic classification presented here
should provide a basis for such investigation.
ACKNOWLEDGEMENTS
Dr J. C. Scheepers provided overall supervision of the
project. The Directorate of Forestry, Mondi Forests, lo-
cal farmers and local foresters kindly permitted sampling
of vegetation on their properties. Dr P. J. Weisser and
Messrs P. Kgwedi and P. Maphangane assisted with
field work. Mrs B. C. de Wet and Messrs P. Gonsalves
and N. P. Barker were especially helpful with computer
operations. Staff of the National Herbarium verified
most of the plant specimens. Mr K. P. Kirkman and Ms
A. P. Backer provided technical input and Mrs A. J.
Romanowski handled photographic processing. Ms H.
M. Moolman helped proofread the text and Mrs E. du
Plessis translated the abstract. Mrs M. van der Merwe
typed the text.
REFERENCES
ACOCKS, J. P. H. 1975. Veld types of South Africa. No. 2nd edn.
Memoirs of the Botanical Survey of South Africa No. 40: 1-128.
COETZEE, B. J. 1983. Phytosociology, vegetation structure and land-
scapes of the Central District, Kruger National Park, South
Africa. Cramer, Vaduz.
DEALL, G. B. 1985. A plant-ecological study of the Eastern
Transvaal Escarpment in the Sabie area. M.Sc. thesis, University
of Pretoria.
DEALL, G. B. & BACKER, A. P. 1989. The vegetation ecology of
the Eastern Transvaal Escarpment in the Sabie area. 3. Annotated
checklist. Bothalia 19: 91 — 110.
DEALL, G. B., SCHEEPERS, J. C. & SCHUTZ, C. J. 1989. The
vegetation ecology of the Eastern Transvaal Escarpment in the
Sabie area. 1. Physical environment. Bothalia 19: 53-67.
EDWARDS, D. 1983. A broad-scale structural classification of vege-
tation for practical purposes. Bothalia 14: 705-712.
SACS 1980. South African Committee for Stratigraphy. Stratigraphy
of South Africa. Part 1 (Comp. L. E. Kent). Lithostratigraphy of
the Republic of South Africa, South West Africa/Namibia and the
Republics of Bophuthatswana, Transkei and Venda. Handbook of
the Geological Survey of South Africa 8: 1 —690.
SCHEEPERS, J. C., BOUCHER, C. & WESTFALL, R. H. no date.
The standardization of South African syntaxonomic nomencla-
ture. Unpublished document prepared for the South African Syn-
taxonomic Nomenclature Committee. Botanical Research Insti-
tute, Pretoria.
VAN DER MEULEN, F. 1979. Plant sociology of the western
Transvaal Bushveld. A syntaxonomic and synecological study.
Cramer, Vaduz.
VAN DER MEULEN, F. & SCHEEPERS, J. C. 1978. On vegetation
studies and land evaluation in South Africa. In R. Tuxen,
Assoziationskomplexe (Sigmeten) und ihre praktische Anwen-
dung. Berichte der Internationalen Symposien der Internationalen
Vereinigung fur Vegetationskunde, Rinteln, 1977: 481-489.
Cramer, Lehre.
WESTFALL, R. H., DEDNAM, G., VAN ROOYEN, N. & THE-
RON, G. K. 1982. PHYTOTAB — a program package for Braun-
Blanquet tables. Vegetatio 49: 35-37.
Bothalia 19,1:91-110(1989)
The vegetation ecology of the Eastern Transvaal Escarpment in the
Sabie area. 3. Annotated checklist
G. B. DEALL* and A. P. BACKER*
Keywords: Checklist, Eastern Transvaal Escarpment, Sabie area
ABSTRACT
Over 1 000 plant taxa occurring in a 1 300 km2 belt transect extending from Hazyview (530 m elevation) to Mount
Anderson (2 280 m elevation) in the Sabie area of the Eastern Transvaal Escarpment are listed by genus, with species
arranged alphabetically within genera. Annotations include genus author, species author, growth form, habitat, collector
and specimen number. Floristic analysis shows the Asteraceae as representing the largest family, followed by the Fabaceae,
Poaceae, Rubiaceae and Liliaceae. Floristic affinities are briefly discussed.
UITTREKSEL
In ’n 1 300 km2 bree lynopname, wat vanaf Hazyview (530 m bo seespieel) tot Mount Anderson (2 280 m bo
seespieel) in die Sabie-area van Oos-Transvaalse platorand strek, kom meer as 1 000 planttaksons voor wat volgens genus
gelys is. Die spesies is alfabeties onder die genusse gerangskik. Aantekeninge sluit die genus-outeur, spesie-outeur,
groeivorm, habitat, versamelaar en eksemplaamommer in. Volgens die floristiese analise verteenwoordig die Asteraceae
die grootste familie gevolg deur Fabaceae, Poaceae, Rubiaceae en Liliaceae. Floristiese verwantskappe word ook kortliks
bespreek.
INTRODUCTION
This checklist contains 1 009 identifiable plant taxa
(excluding mosses and lichens) recorded in the 251
releves of the study area (cf. Deall et al. 1989a & b).
Most taxa are represented by voucher specimens housed
in either the National Herbarium, Pretoria; the D. R. de
Wet Forestry Research Centre Herbarium, Sabie; or in
the senior author’s field herbarium. Botanical Research
Institute, Pretoria. A number of plants seen in the field
are not included in the checklist because they were in too
poor a condition to be identified. Voucher specimen
numbers are italicized at the end of each species citation.
All pertain to G. B. Deall except where indicated. Some
taxa can, if necesary, be verified by more than the single
voucher specimen quoted. Additional numbers are avail-
able from the senior author’s collection register kept at
the National Herbarium, Pretoria.
Classification of the Pteridophyta is according to
Schelpe & Anthony (1986) and that of the Spermato-
phyta is according to Gibbs Russell et al. (1985, 1987).
Species are arranged alphabetically within genera, exotic
species being indicated by an asterisk. In instances
where plants could not be identified below the generic
level, the epithet sp. is appended (e.g. Combretum sp. ).
It should be noted that such a name may not consistently
refer to the same species concept, but may include va-
rious species within the genus. Also, species designated
as ‘complex’ are those whose identity in the field was not
clear. For example, Tetradenia ‘complex’ is named thus
because, although the first specimens collected were
identified as Tetradenia riparia, subsequent specimens,
which cursorily appeared to be T. riparia were identified
as T. brevispicata. This distinction could not easily be
* Botanical Research Institute, Private Bag X101, Pretoria 000 1 .
MS. received: 1988.10.17.
discerned in the field without prior knowledge, and thus
the term ‘complex’ is used to indicate that either species
could be involved. Annotative terminology is in accord-
ance with the categories used by Deall et al. (1989a &
b).
FLORISTIC ANALYSIS
A floristic analysis of the taxa contained in the check-
list is summarized in Table 1. The study revealed a total
of 1 009 identifiable species (including subspecies and
varieties) distributed amongst 501 genera and 133 fami-
lies. As with Scheepers’ (1978) study, the Asteraceae
constitute the largest family (37 genera and 128 species).
The Fabaceae (36 genera, 91 species) and Poaceae (49
genera, 90 species) are next, followed by the Rubiaceae
(26 genera, 48 species) and Liliaceae (21 genera, 46
species). The largest genera include Helichrysum (39
TABLE 1. — Floristic analysis of taxa in the Sabie area of the
Eastern Transvaal Escarpment
* including subspecies and varieties
92
Bothalia 19,1 (1989)
spp.), Senecio (21 spp.), Vernonia (12 spp.), Asplenium
(11 spp.), Cyperus (11 spp.), Rhynchosia (10 spp.), In-
digofera (9 spp.), Crassula (9 spp.), Protasparagus (9
spp.), and Hyparrhenia (9 spp.).
Of the species whose distribution is limited to the
Montane and Subalpine Belts, 29 % are present in the
Natal Drakensberg (Killick 1963); 28 % in the northern
Transvaal Drakensberg (Scheepers 1978); and 17 % in
the Chimanimani Mountains of Zimbabwe (Goodier &
Phipps 1961). This suggests that the Montane and Subal-
pine flora of the Sabie area has both temperate and tropi-
cal affinites, although the former is apparently stronger.
ACKNOWLEDGEMENTS
Knowledge of the flora was gained initially by Dr J.
C. Scheepers’ assistance in the field. The Directorate of
Forestry, Mondi Forests, local farmers and local forest-
ers kindly permitted vegetation on their properties to be
sampled. Dr P. J. Weisser, Messrs P. Kgwedi and P.
Maphangane assisted with field work. Staff of the Natio-
nal Herbarium verified most of the plant specimens. Mr
K. P. Kirkman helped with cataloguing, Miss H. M.
Moolman helped proofread the text and Mrs W. Jones
translated the abstract.
REFERENCES
DEALL, G. B., SCHEEPERS, J. C. & SCHUTZ, C. J. 1989. The
vegetation ecology of the Eastern Transvaal Escarpment in the
Sabie area. 1 . Physical environment. Bothalia 19: 53-67.
DEALL, G. B., THERON, G. K. & WESTFALL, R. H. 1989. The
vegetation ecology of the Eastern Transvaal Escarpment in the
Sabie area. 2. Floristic classification. Bothalia 19: 69-89.
GIBBS RUSSELL, G. E., REID, C., VAN ROOY, J. & SMOOK, L.
1985. List of species of southern African plants. Edn 2, Part 1.
Cryptogams, Gymnosperms, Monocotyledons. Memoirs of the
Botanical Survey of South Africa No. 51: 1 — 152.
GIBBS RUSSELL, G. E., WELMAN, W. G., RETIEF, E., 1MMEL-
MAN, K. L., GERMISHUIZEN, G., PIENAAR, B. J., VAN
WYK, M. & NICHOLAS, A. 1987. List of species of southern
African plants. Edn. 2, Part 2. Dicotyledons. Memoirs of the
Botanical Survey of South Africa No. 56: 1 — 270.
GOODIER, R. & PHIPPS, J. B. 1961. A revised checklist of the
vascular plants of the Chimanimani Mountains. Kirkia 1: 44 -66.
KILLICK, D. J. B. 1963. An account of the plant ecology of the
Cathedral Peak area of the Natal Drakensberg. Memoirs of the
Botanical Survey of South Africa No. 34: 1 — 178.
SCHEEPERS, J. C. 1978. Vegetation of Westfalia Estate on the
North-Eastern Transvaal Escarpment. Memoirs of the Botanical
Survey of South Africa No. 42: 1 - 230.
SCHELPE, E. A. C. L. E. & ANTHONY, N. C. 1986. Pteridophyta.
In O. A. Leistner, Flora of southern Africa. Botanical Research
Institute, Pretoria.
CHECKLIST
PTERIDOPHYTA
LYCOPODIACEAE
Lycopodium L.
gnidioides L. f, trailing, on rocks in elfin-like Mistbelt forests,
1194
SELAGINELLACEAE
Selaginella Beauv.
dregei (Presl) Hieron. , mat-forming xerophytic pioneer in rock
depressions of Escarpment Slopes and Foothills, 1618
kraussiana (Kunze) A. Br. ex Kuhn , prostrate moss-like species
in field layer of Tall Forest, Escarpment and Mountain
Slopes, 326
MARATTIACEAE
Marattia Swartz
fraxinea J.E. Sm. ex J.F. Gmel. var. salicifolia (Schrad.) C.
Chr. , large waterside fern in dense shade of Tall (riparian)
Forest, Mountain Slopes and Upper Foothills, 1043
OSMUNDACEAE
Todea Willd. ex Bernh.
barbara (L.) T. Moore, large tree-like fern of Tall (swamp)
Forest, Escarpment Lower Slopes, 1120
GLEICHENIACEAE
Dicranopteris Bernh.
linearis (Burm. f.) Underw. , infrequent fern in forests and mesic
thickets of Foothills, 571
SCHIZAEACEAE
Mohria Swartz
caffrorum (L.) Desv. , rosette fern associated with rocky wood-
lands and grasslands of Mistbelt, 1023
CYATHEACEAE
Cyathea J.E. Sm.
dregei Kunze , large tree fern in moist (often riparian) sites of
Mistbelt and Low Country, 109
HYMENOPHYLLACEAL
Trichomanes L.
pyxidiferum L. var. melanotrichum (Schlechtd.) Schelpe , small-
lobed transluscent epiphyte of Mistbelt forests, 101 7
DENNSTAEDTIACEAE
Blotiella Tryon
glabra (Bory) Tryon, large tree-like fern of Tall (swamp) Forest,
Escarpment Lower Slopes, 1126
Pteridium Gled. ex Scop.
aquilinum (L.) Kuhn, widespread fern of disturbed grasslands
and thickets, especially Mistbelt, 29
Hypolepis Bernh.
sparsisora (Schrad.) Kuhn, large bracken-like fern m sunny
openings of Mistbelt forests, 1123
ADIANTACEAE
Adiantum L.
capillus-veneris L. , infrequent waterside fern in forests of Mist-
belt and Low Country, 894
Pteris L.
catoptera Kunze, large fern in field layer of moist forests, Humid
Mistbelt, 1063
cretica L., large fern in Tall (riparian) Forest, Mountain Slopes,
15 72
Cheilanthes Swartz
concolor (Langsd. & Fisch.) R. & A. Tryon, apparently rare
streambank fern in riparian forests of Low Country,
1846
hirta Swartz, occasional fern in rocky, sheltered mesic thickets
of Low Country, 1717
pentagona Schelpe & N.C. Anthony , small localized fern ot rock
outcrops in Summit Peak grasslands and shrublands, 2495
quadripinnata (Forssk.) Kuhn, localized fern of Summit shrub-
lands and woodlands, 2293
viridis (Forssk.) Swartz
var. glauca (Sim) Schelpe & N.C. Anthony, fern of exposed
rocky sites in grasslands and thickets of Humid Mist-
belt, 1414
var. viridis, widespread rosette fern, common in Mistbelt and
Low Country, 2077
Pellaea Link
calomelanos (Swartz) Link, rosette fern, common on rock
outcrops in w'oodlands and shrublands of Mistbelt and
Low' Country, 87
pcctiniformis Bak., rosette fern associated with rock outcrops
in forests and woodlands of Humid Mistbelt, 1391
POLYPODIACEAF
Polypodium L.
Bothalia 19,1 (1989)
93
polypodioides (L.) Hitchc., subsp. ecklonii (Kunze) Schelpe ,
creeping epiphytic fern, common in Mistbelt forests,
704
Pleopeltis H.B.K. ex Willd.
macrocarpa (Bory ex Willd.) Kaulf., creeping epiphytic fern,
common in Mistbelt forests, 717
DAVALLIACEAE
Arthropteris/. Sm.
monocarpa (Cordem.) C. Chr., infrequent shade fern associated
with exposed (rocky) woodlands of Plateau Crest, 1400
ASPLENIACEAE
Asplenium L.
aethiopicum (Bunn, f.) Becherer, occasional fern in deep bushy
ravines, Summit Slopes, 2677
anisophyllum Kunze, robust epiphytic fern in forests of Humid
Mistbelt, 1065
boltonii Hook, ex Schelpe, small fern in High Forest of Moun-
tain Slopes, 21 72
erectum Bory ex Willd. var. erectum, ground fern in field layer
of high Mistbelt forests, 2138
inaequilaterale Willd., occasional fern in rocky field layer of
Mistbelt forests, 1130A
lobatum Pappe & Rawson, occasional fern in rocky field layer
of Mistbelt forests, 1046A
lunulatum Swartz , fern in dense shade of tall Mistbelt forests,
1130
rutifolium (Berg.) Kunze, common epiphytic fern in Mistbelt
forests, 335
sandersonii Hook., small epiphytic fern in moist forest of
Lower Mountain Slopes, 2230
splendens Kunze, common epiphytic or ground fern in Mistbelt
forests, 321
varians Wall, ex Hook. & Grev. subsp. fimbriatum (Kunze)
Schelpe, apparently confined to cool, moist cliff forests
of Lower Mountain Slopes, 1380
THELYPTERIDACEAE
Thelypteris Schmidel
bergiana (Schlechtd.) Ching, rosette fern, infrequent along shady
stream banks in forests of Mistbelt and Low Country,
895
gueinziana (Mett.) Schelpe, large fern in Tall (riparian) Forest
of Low Country, 1909
interrupta (Willd.) K. Iwats., large infrequent fern in Tall (ripa-
rian) Forest of Low Country, 1923
madagascariensis (Fee) Schelpe, rosette fern in tall Mistbelt
forests, especially Escarpment Slopes, 1122
sp., large fern in Tall Forest of Escarpment Slopes, includes
1045
ATHYRIACEAE
Athyrium Roth
scandicinum (Willd.) Presl, small infrequent fern in Tall (riparian)
Forest of Escarpment Lower Slopes, 1125
LOMARIOPSIDACEAE
Elaphoglossum Schott ex J. Sm.
acrostichoides (Hook. & Grev.) Schelpe, fern in shelter of large
boulders. Summit Peak shrublands, 2496
ASPIDIACEAE
Dryopteris Adam.
athamantica (Kunze) Kuntze, rosette fern in marshy grasslands,
especially Mistbelt, 1655
inaequalis (Schlechtd.) Kuntze, frequent rosette fern of Mistbelt
forests, 1001
Cyrtomium Presl
caryotideum (Wall, ex Hook. & Grev.) Presl. var. micropterum
(Kunze) C. Chr., infrequent fern in forests of Humid
Mistbelt, 1575
Polystichum Roth
luctuosum (Kunze) T. Moore, infrequent fern in forests of
Humid Mistbelt, especially Lower Mountain Slopes,
1381
macleae (Bak.) Diels, large leathery fern, infrequent in Tall (ri-
parian) Forest, Summit Slopes, 2260
Rumohra Raddi
adiantiformis (G. Forst.) Ching, robust fern apparently confined
to rocky outcrops in forest openings, especially Plateau
Crest, 1226
Tectaria Cav.
gemmifera (Fee) Alston, fairly infrequent fern in Tall Forest of
Escarpment Slopes, 1139
blechnaceae
Blechnum L.
giganteum (Kaulf.) Schlechtd., fairly infrequent fern in Tall
Forest of Escarpment Slopes, 1121
tabulate (Thunb.) Kuhn, large rosette fern widespread in Mist-
belt woodlands and grasslands, 1146
GYMNOSPERMAE
PODOCARPACEAE
Podocarpus L 'Herit.
falcatus (Thunb.) R. Br. ex Mirb., tall understorey and canopy
tree, sometimes dominant in High Forest of Mountain
Slopes, 2223
latifolius (Thunb.) R. Br. ex Mirb., dominant canopy tree in
elfin-like forests, Plateau Crest and Mountain Slopes,
2050
PINACEAE
*Pinus L.
spp., exotic timber tree encroaching on indigenous vegetation
wherever plantations exist
ANGIOSPERMAE — MONOCOTYLEDONAE
TYPHACEAE
Typha L.
capensis (P. Rohrb.) 19. E. Br., robust rhizomatous herb on
stream banks in Low Country, Scheepers 12
POACEAE
Ischaemum L.
fascicuiatum Brongn. , localized in marshy grasslands of Escarp-
ment Lower Slopes and Foothills, 1905
Elionurus Kunth ex Willd.
sp., small tufted perennial grass of Summit Plateau grasslands,
2741
Cleistachne Benth.
sorghoides Benth., occasional annual grass in xeric thickets of
Upper Foothills, 684
Eulalia Kunth.
villosa (Thunb.) Nees, widespread tufted perennial grass in
thickets, woodlands and grasslands of Mistbelt and Low
Country, 633
Bothriochloa Kuntze
bladhii (Retz.) S.T. Blake, widespread perennial grass in dis-
turbed grasslands and woodlands of Low Country, 598
Schizachyrium Nees
sanguineum (Retz.) Alston, widespread tufted grass in wood-
lands and grasslands of Mistbelt and Low Country, 603
Andropogon L.
appendiculatus Nees, tall perennial grass dominating hygric
sites with deep black soils, Summit Slopes and Plateau,
2580
chinensis (Nees) Men., small tufted perennial grass, infrequent
in grasslands and woodlands of Low Country, 1919
huillensis Rendle, large tufted perennial grass, infrequent in
disturbed grasslands, especially Mistbelt, 1300
schirensis A. Rich. , medium-size tufted perennial grass, abun-
dant (often dominant) in woodlands and grasslands of
Mistbelt, 1321B
Cymbopogon Spreng.
'complex’, tall tufted perennial grass in thickets, woodlands
and grasslands of Mistbelt and Low Country:
excavatus (Hochst.) Stapf ex Burn Davy, 1518
validus (Stapf) Stapf ex Bunt Davy, 1321C
Hyparrhenia Anderss. ex Fount.
anamesa Clayton, robust tufted perennial grass, infrequent in
moist grasslands, Humid Mistbelt, 1658
cymbaria (L.) Stapf, large tufted perennial grass in serai thickets
and woodlands. Humid Mistbelt, 1601
dregeana (Nees) Stapf ex Stent, large tufted perennial grass in
rocky open woodlands of Humid Mistbelt, 1977
94
Bothalia 19,1 (1989)
filipendula (Hochst.) Stapf
‘ complex’, tufted perennial grass, common in thickets, wood-
lands and grasslands of Mistbelt and Low Country:
var. filipendula, 1321F
var. pilosa (Hochst.) Stapf, 1624A
gazensis (Rendle) Stapf, tufted perennial grass, localized in fair-
ly rocky woodlands and thickets of Low' Country, 84c.
hirta (L.) Stapf, fairly frequent tufted perennial grass in w'ood-
lands and grasslands of Mistbelt and Low Country, 132 ID
newtonii (Hack.) Stapf var. macra Stapf, infrequent perennial
grass associated with seepage areas, Escarpment Slopes,
164
variabilis Stapf, tall perennial grass in woodlands of Low Coun-
try, 3
Hyperthelia Clayton
dissoluta (Nees ex Steud.) Clayton, large tufted grass, wide-
spread in Woodland and Xeric Thicket of the Low Coun-
try, 2018
Monocymbium Stapf
ceresiiforme (Nees) Stapf, small tufted perennial grass, abundant
in grasslands of Mistbelt, 1119
Trachypogon Nees
spicatus (L. f.) Kuntze , frequent tufted perennial grass scattered
in woodlands and grasslands of Mistbelt and low Coun-
try, 1513
Heteropogon Pers.
contortus (L.) Roem. & Schult., small tufted perennial grass,
widespread in Woodland and Xeric Thicket of the Low
Country, 730
Diheteropogon Stapf
amplectens (Nees) Clayton, large tufted perennial grass, infre-
quent in disturbed woodlands of Upper Foothills, 623
filifolius (Nees) Clayton, small tufted perennial grass, scattered
in Summit grasslands, 1694
Themeda Forssk.
triandra Forssk., widespread tufted perennial grass in thickets,
woodlands and grasslands of Mistbelt and Low Country,
745
Digitaria Haller
diagonalis (Nees) Stapf var. diagonalis, tufted perennial grass,
occasional in Mountain Slopes grasslands, 1533
maitlandii Stapf & C.E. Hubb., tufted perennial grass in deep-
soil grasslands of Plateau Interior, 1312
monodactyla (Nees) Stapf, small tufted perennial grass in rocky
retarded grasslands, especially Escarpment Plateau, 1276
tricholaenoides Stapf, rare rhizomatous perennial grass in
Plateau grasslands, Humid Mistbelt, 1244A
Alloteropsis C.B. Presl
semialata (R. Br.) Hitchc., small tufted perennial grass, common
in Escarpment Plateau and Summit grasslands, 1113
Brachiaria Griseb.
brizantha (A. Rich.) Stapf, infrequent grass in disturbed grass-
lands of Low Country, 721
serrata (Thunb.) Stapf, infrequent tufted perennial grass, some-
times associated with sheet-rock outcrops on Upper
Foothills, 1320A
subulifolia (Mez) Clayton, localized grass in dolomite-soil grass-
lands, Plateau Interior, 1320
Paspalum L.
scrobiculatum L., infrequent perennial grass of disturbed grass-
lands and woodlands, Mistbelt and Low Country, 6 78
*urvillei Steud., naturalized perennial grass, infrequent in dis-
turbed woodlands, Low' Country, 114
Oplismenus Beauv.
hirtellus (L.) Beauv., low straggling perennial grass, widespread
and dominant in field layer of Forest and Mesic Thicket
of Mistbelt and Low Country, 682
Panicum L.
‘complex’, common grass in field layer of Woodland and Xeric
Thicket of Low Country:
deustum Thunb., 1695 A
maximum Jacq., 847
ecklonii Nees, tufted perennial grass, widespread in Mistbelt
grasslands, 1096
natalense Hochst., widespread tufted perennial grass in Mistbelt
grasslands and woodlands, 1321H
Setaria Beauv.
megaphylla (Steud.) Dur. <6 Schinz, robust tufted perennial
grass, widespread in forests and mesic woodlands and
thickets of Mistbelt and Low Country, 602
sphacelata (Schumach.) M.B. Moss, medium-size tufted grass,
w idespread in Humid Mistbelt grasslands and woodlands
and in Low Country w oodlands and xeric thickets, 81 7
Rhynchelytrum Nees
‘complex’, small tufted grass, widespread in Woodland and
Xeric Thicket of Low Country and in Mistbelt w oodlands
and grasslands:
nerviglume (Franch.) Chiov., 1292B
repens (Willd.) C.E. Hubb., 1401
rhodesianum (Rendle) Stapf & C.E. Hubb., 1623
Melinis Beauv.
tenuinervis (Stapf) Stapf, rare tufted grass in closed (rocky)
woodlands. Low Country, 1704
Pennisetum Rich.
macrorum Trin. , tall perennial grass of riparian woodlands.
Summit Plateau, 2710
thunbergii Kunth, tufted rhizomatous perennial grass of marshy
grasslands. Summit Plateau, 2652
Ehrharta Thunb.
erecta Lam., sporadic straggly perennial grass in Mistbelt forests
and thickets, 243
Tristachya Nees
leucothrix Nees, tufted perennial grass scattered in woodlands
and grasslands of Mistbelt, 1321J
Trichopteryx Nees
dregeana Nees, short straggling perennial grass, usually associa-
ted with seepage, Mistbelt, 1108
Loudetia Hochst. ex Steud.
densispica (Rendle) C.E. Hubb. , tufted perennial grass confined
to woodlands and grasslands of Mistbelt, 1094
simplex (Nees) C.E. Hubb , tufted perennial grass, dominant in
Humid Mistbelt grasslands and woodlands and also
present in Low Country woodlands, shrublands and
thickets, 616
Helictotrichon Bess, ex Schult.
natalense (Stapf) Schweick ., locally abundant perennial grass
of marshy grasslands, Summit Plateau, 2659
Pentaschistis Stapf
sp., tufted perennial grass, infrequent in grasslands of Summit
Peaks, 2733
Phragmites Trin.
mauritianus Kunth, tall robust perennial reed in dense stands
along rivers in Low Country, 1965
Agrostis L.
eriantha Hack. var. eriantha, infrequent grass localized in shrub-
by grasslands. Summit Plateau, 2647
lachnantha Nees, loosely tufted perennial grass, infrequent in
grasslands of Escarpment Slopes, 1078
Aristida L.
congesta Roem. & Schult. subsp. barbicollis (Trin. & Rupr.) De
Winter, small tufted grass, infrequent in Low Country
woodlands, 176 7
junciformis Trin. <£ Rupr. subsp. junciformis, small tufted
grass, infrequent in rocky grasslands of Plateau Crest
and Summit, 1428
transvaalensis Henr., tufted grass in moist seepage of sheet-rock
formations. Transitional Mistbelt, 1621
Perotis Ait.
patens Gand., small grass, infrequent in Low Country wood-
lands, 1760
Sporobolus R. Br.
centrifugus (Trin.) Nees, densely tufted perennial grass, fairly
frequent on Summit Peaks and Slopes, 2371
‘complex’, tufted grass, widespread in Mistbelt grasslands and
in woodlands and shrublands of Low Country:
africanus (Poir.) Robyns & Tournay, 1541
pyramidalis (Beauv.), 555
sp., 972
stapfianus Gand, small tufted grass localized and infrequent on
sheet-rock formations in Mistbelt and Low Country,
1898
Eragrostis Beauv.
caesia Stapf, tufted perennial grass, localized in sandy grasslands
of Plateau Crest, 54
capensis (Thunb.) Trin. , widespread tufted perennial grass in
grasslands of Escarpment Slopes and Plateau, 1097
curvula (Schrad.) Nees, widespread tufted perennial grass,
except in forests and mesic thickets, 1294B
gummiflua Nees, infrequent grass of disturbed sheet-rock sites,
Humid Mistbelt, 1299A
Bothalia 19,1 (1989)
95
hierniana Rendle, infrequent grass on sheet-rock formations in
Mistbelt and Low Country, 1920
racemosa (Thunb.) Steud., widespread tufted perennial grass in
grasslands of Mistbelt, 1028
sclerantha Nees subsp. sclerantha, localized grass of lithoseral
grasslands, especially Mistbelt, 1424A
Microchloa R. Br.
caffra Nees, small tufted perennial grass, localized in lithoseral
grasslands of Mistbelt and Low Country, 1143
Rendlia Chiov.
altera (Rendle) Chiov., small tufted perennial grass, locally
abundant in grasslands of Summit and Plateau Crest,
1171
Harpochloa Kunth
falx (L. f.) Kuntze, tufted perennial grass in sandy grasslands of
Escarpment Slopes, Plateau Crest and Summit Peak,
1208
Ctenium Panz.
concinnum Nees, large tufted perennial grass, infrequent in
grasslands of Summit and Mountain Slopes, 1511
Pogonarthria Stapf
squarrosa (Roem. & Schult.) Pilg., tufted grass, rare in sandy
woodlands of Low Country, 1769
Bewsia Goossens
biflora (Hack.) Goossens, tufted perennial grass in grasslands of
Lower Mountain Slopes, 1312G
Styppeiochloa De Winter
gynoglossa (Goossens) De Winter, small tufted perennial grass,
confined to grasslands of Escarpment Slopes, 1102
Trichoneura N.J. Anderss.
grandiglumis (Nees) Ekman, small tufted grass, rare in sandy
woodlands of Low Country, 1 759
Koeleria Pers.
capensis (Steud.) Nees, densely tufted perennial grass in rocky
woodlands and grasslands of Mistbelt, 1079
Stiburus Stapf
alopecuroides (Hack.) Stapf, small tufted perennial grass,
localized in sandy grasslands of Escarpment and Summit
Plateau, 1109
Festuca L.
caprina Nees var. caprina, infrequent tufted perennial grass of
Summit Peak grasslands, 2509
costata Nees var. costata, densely tufted perennial grass, often
dominant in Summit grasslands, 2619
Bromus L.
speciosus Nees, scattered tufted perennial grass of Summit
grasslands, 2618
CYPERACEAE
Cyperus L.
albostriatus Schrad., common sedge in field layer of forests,
mesic thickets and rocky woodlands, especially Mistbelt,
1418
denudatus L. f, occasional sedge of marshy grasslands. Summit
Plateau, 25 75
immensus C.B. CL, robust sedge, infrequent in Tall (riparian)
Forest of Low Country, 1908
leptocladus Kunth, apparently localized sedge in rocky wood-
lands and grasslands, Escarpment Plateau, 1289
obtusiflorus Vahl
var. flavissimus Boeck. , localized sedge in grasslands of Lower
Mountain Slopes, 1478
var. sphaerocephalus (Vahl) Kuekenth., frequent sedge of
Summit grasslands, 2512
pseudoleptocladus Kukenth.. medium size sedge, widespread in
Forest and Mesic Thicket of Mistbelt and Low Country,
1056
schlechteri C.B. Cl., infrequent sedge of moist grasslands. Sum-
mit Plateau, 2687
semitrifidus Schrad. , small sedge, widespread in lithoseral grass-
lands of Escarpment Plateau and Summit, 1278
sexangularis Nees, infrequent sedge in field layer of Tall (ripa-
rian) Forest, Low Country, 1841
thorncroftii McClean, sedge of sheet-rock shrublands. Transi-
tional Mistbelt, 1620
Pycreus Beauv.
muricatus (Kukenth.) Napper, small sedge, localized in grass-
lands of Escarpment Slopes, 1077
Mariscus Gaertn.
solidus (Kunth) ined. subsp. solidus, giant sedge of marshy
grasslands. Transitional Mistbelt, 1969
Kyllinga Rottb.
alba Nees, small sedge of Escarpment Plateau grasslands, 161
odorata Vahl, small sedge, infrequent in Mistbelt grasslands,
Scheepers 1 9
Ficinia Schrad.
bergiana Kunth, small sedge, infrequent in grasslands of Lower
Mountain Slopes, 1501
sp., tussock sedge, Mistbelt and Low Country, includes 680,
2527
Scirpus L.
ficinioides Kunth, locally abundant sedge of marshy grasslands,
Summit and Escarpment Plateau, 1262A
Schoenoplectus Palla
corymbosus (Roth, ex Roem. & Schult.) J. Raynal, waterside
sedge, localized in Tall (riparian) Forest of Low Country,
1910
Bulbostylis Kunth
burchellii (Fical. & Hiem) C.B. Cl. , occasional sedge in rocky
woodlands of Plateau Crest, 1449
oritrephes (Ridley) C.B. G.
‘complex’, small tufted sedge common in Summit grasslands
and shrublands:
subsp. australis B.L. Burtt, 2389
subsp. oritrephes, 2326
schoenoides (Kunth) C.B. Cl. , small sedge, widespread in grass-
lands of Escarpment Plateau and Slopes, 1088
Rhynchospora Vahl
brownii Roem. c£ Schult., occasional sedge associated with seep-
ages in woodlands and grasslands of Transitional Mistbelt,
749
Tetraria Beauv.
natalensis (C.B. Cl.) Koyama, localized sedge of moist grass-
lands, Summit Slopes, 2394
cf. sp. nov. (De Winter & Codd 202), occasional perennial herb
of moist rocky woodlands and shrublands. Summit
Slopes, 2368, 2427
Coleochloa Gilly
setifera (Ridley) Gilly, large-tussocked sedge, locally abundant
on sheet-rock formations, Mistbelt and Low Country,
1882
Scleria Berg.
bulbifera Hochst. ex A. Rich., infrequent sedge of Escarpment
Plateau grasslands, 1335
dieterlenii Turrill, small perennial sedge of moist grasslands.
Summit Plateau and Slopes, 2563
melanomphala Kunth, occasional sedge of moist grasslands.
Escarpment Lower Slopes, 1944
Schoenoxiphium Nees
lehmannii (Nees) Steud., infrequent perennial sedge of Tall
(riparian) Forest, Lower Mountains, 1582
rufum Nees, tufted perennial sedge, infrequent in riparian
woodlands, Summit Plateau, 2444
schweickerdtii Merxm. & Podlech, rhizomatous perennial sedge
of rock crevices, Summit Peak, 2469
sp. nov., tufted sedge of riparian woodlands. Summit Plateau,
2435
Carex /..
austro-africana (Kukenth.) Raymond, waterside sedge of
marshy grasslands. Summit Plateau, 2578
cognata Kunth. var. drakensbergensis (C.B. Cl.) Kukenth.,
waterside sedge. Summit Plateau, 2443
spicato-paniculata C.B. CL, fairly widespread sedge in forests,
thickets and woodlands of Mistbelt and Low Country,
595
arecaceae
Phoenix L.
reclinata Jacq., small tree on stream banks of High (riparian)
Forest, Low' Country, 1838
ARACEAE
Zantedeschia Spreng.
sp., infrequent perennial herb of marshy grasslands, Summit
Plateau, (no voucher specimen)
Stylochiton Lepr.
natalense Schott, perennial tufted herb, localized in shaded
field layer of forests, thickets and woodlands, Low
Country, 702
XYRIDACEAE
Xyris L.
gerrardii N.E. Br. , infrequent grass-like herb of moist and marshy
grasslands. Summit Slopes and Plateau, 2656
96
Bothalia 19,1 (1989)
COMMELINACEAE
Commelina L.
africana L.
vai. africana, trailing herb, infrequent in Summit Plateau
grasslands, 2540
var. krebsiana (Kunth) C.B. CL , herb, localized in sandy grass-
lands of Plateau Crest, 1285
var. lancispatha C.B. CL, herb, infrequent in field layer of
Woodland and Xeric Thicket of Low Country, 1 724
eckloniana Kunth, locally abundant herb of marshy grasslands,
Plateau Crest, 1265
livingstonii C.B. CL , occasional herb of mesic thickets, Mistbelt
and Low Country, 720
sp., widespread herb, includes 1188, 1406
Aneilema R. Br.
aequinoctiale (Beauv.) Kunth, trailing herb in undergrowth of
Tall (riparian) Forest, Low Country, 1837
Cyanotis D. Don
lanata Benth., small succulent herb, rare on sheet-rock forma-
tions in Low Country, 1894
lapidosa PhilL, small herb, localized in rock crevices of Plateau
Crest woodlands, 1409
pachyrrhiza Oberm., small succulent herb, localized in rock
depressions of Plateau Crest woodlands, 1241
speciosa (L. f.) Hassk., occasional herb in grasslands of Humid
Mistbelt, 1149A
Floscopa Lour.
glomerata (Willd. ex Schult. <£ Schult. f.) Hassk., localized herb
in seepage areas of Tall Forest, Low Country, 1924
JUNCACEAE
J uncus L.
lomatophyllus Spreng., hydrophytic rosette herb of marshy
grasslands, Summit Plateau, 2657
LILIACEAE
Bulbine Willd.
sp., succulent geophyte, infrequent in lithoseral woodlands of
Low Country
Trachyandra Kunth
reflexipilosa (Kuntze) Oberm., widespread geophytic herb of
Summit Plateau grasslands and shrublands, 2531
saltii (Bak.) Oberm. , low perennial herb, localized in sandy grass-
lands of Escarpment Slopes and Plateau, 1090
Anthericum L.
angulicaule Bak. , rare herb in sandy grasslands of Escarpment
Crest, 1270
cooperi Bak., widespread geophytic herb in grasslands of
Plateau Interior and Summit, 1364
galpinii Bak. var. galpinii, localized herb, infrequent in lithoseral
grasslands of Escarpment Plateau, 1280
sp. cf. A. galpinii Bak. , infrequent herb of sheet-rock formations.
Low Country, 1860
Chlorophytum Ker-GawL
sp., infrequent herb in field layer of rocky woodlands, Plateau
Crest, 1191A
Eriospermum Jacq. ex Willd.
burchellii Bak. , geophytic herb localized in rocky shrublands of
Transitional Mistbelt and in grasslands of Escarpment
Slopes and Plateau, 1030
cooperi Bak. , widespread geophytic herb, infrequent in Mistbelt
grasslands and woodlands, 756
luteo-rubrum (Bak.), infrequent geophyte of retarded grassland.
Summit Plateau, 2349
sp. 1, infrequent grassland geophyte of Summit Plateau and
Slopes, 2541
sp. 2, infrequent geophytic herb in woodlands of Escarpment
Slopes, includes 1858
Kniphofia Moench
sp., perennial herb, infrequent in lithoseral grasslands of Escarp-
ment Slopes, 1160
splendida E.A. Bruce, perennial herb, localized in grasslands of
Lower Mountain Slopes, E.A. Bruce 315
Aloe L.
arborescens Mill, widespread shrubby species in rocky wood-
lands and forests, Mistbelt, Scheepers 426
barbertoniae Pole Evans, succulent perennial herb, widespread
in Woodland and Xeric Thicket of Low Country, 1998
longibracteata Pole Evans, succulent perennial herb, apparently
confined to xeric thickets and woodlands, Mistbelt, 1992
petricola Pole Evans, widespread succulent rosette herb, local-
ized on sheet-rock formations, Low Country and in
rocky woodlands, Humid Mistbelt, 1995
sp., widespread succulent herb, includes 1993, 1994
chortolirioides Berger var. woolliana (Pole Evans) Glen & Hardy,
small succulent herb on steep rock ledges, Summit grass-
lands, PNBG 29952
Agapanthus L 'Herit.
inapertus Beauv. subsp. parviflorus Leighton, perennial herb
often associated with riparian sites on Summit, 2627
Tulbaghia L.
sp., infrequent perennial herb of rocky shrublands, Mistbelt,
2744
Albuca L.
setosa Jacq., low bulbous herb, infrequent in Plateau Interior
grasslands, 11 76
Urginea Steinh.
capitata (Hook.) Bak., occasional bulbous herb of rocky shrub-
lands, Summit Plateau, 2457
Dipcadi Medik.
marlothii Engl., localized herb, infrequent in Escarpment
Plateau grasslands, 1279
Scilla L.
nervosa (Burch.) Jessop, bulbous rosette herb, localized in
disturbed lithoseral grasslands of Escarpment and
Summit Plateau, 1296
Eucomis L Herit.
autumnalis (Mill.) Chitt. subsp. clavata (Bak.) Reyneke, fairly
common bulbous herb of moist (often riparian). Summit
grasslands, 2440
Ornithogalum L.
saundersiae Bak. , geophyte in Low Country woodlands, 1 752
Ledebouria Roth.
cooperi (Hook, f.) Jessop, bulbous rosette herb, widespread in
grasslands of Plateau Interior and Summit, 1369
revoluta (L. f.) Jessop, bulbous geophyte, infrequent in Mistbelt
grasslands, 1554
sp., flattened succulent rosette herb, widespread in grasslands
of Humid Mistbelt, includes 1212
Dracaena Vand. ex L.
hookeriana K. Koch, subwoody shrub of shady field layer,
widespread in Forest and Mesic Thicket of Mistbelt and
Low Country, 907
Protasparagus Oberm.
africanus (Lam.) Oberm., small bush, widespread in woodlands
and thickets, especially Low Country, 516
falcatus (L.) Oberm. , robust spiny climber, common in forests
and thickets of Mountain and Escarpment Slopes and
Upper Foothills, 93
laricinus (Burch.) Oberm., localized shrubby species of rocky
woodiand, Lower Mountain Slopes, 1567
natalensis (Bak.) Oberm., spiny climber, infrequent in Tall
(riparian) Forest of Low Country, 1980F
plumosus (Bak.) Oberm., shrubby species in undergrowth of
forests, Escarpment Slopes and Plateau, 1062
racemosus (Willd.) Oberm., infrequent undershrub of Low
Country woodlands and thickets, 1 706A
rigidus (Jessop) Oberm., small bush, localized in Plateau Crest
woodlands, 1186
setaceus (Kunth) Oberm., small bush, common in forests of
Mountain Slopes, 1385
sp., common undershrub in forests, woodlands and thickets,
especially Transitional Mistbelt and Low Country,
includes 43
Myrsiphyllum Willd.
asparagoides (L.) Oberm., herbaceous twiner, rare in under-
growth of moist thickets, riparian woodlands, Summit
and Lower Mountain Slopes, 2450
ramosissimum (Bak.) Oberm., small shrub of riparian forest,
Mountain and Summit Slopes, 2136
Behnia Didr.
reticulata (Thunb.) Didr., perennial herbaceous twiner, common
in forests and thickets of Mountain and Escarpment
Slopes and Upper Foothills, 282
Smilax L.
kraussiana Meisn. , common climber, widespread in forests, thick-
ets and woodlands, Mistbelt and Low Country, 122
AMARYLLIDACEAE
Haemanthus L.
carneus Ker-GawL, occasional herb of rocky woodlands, Lower
Mountain Slopes, 1561
Bothalia 19,1 (1989)
97
sp., infrequent herb in grasslands of Lower Mountain Slopes,
1489
Scadoxus Raf.
multiflorus (Martyn) Raf. subsp. multiflorus, occasional wide-
spread herb of forests and thickets, 1 728
Boophane Herb.
disticha (L. f.) Herb., bulbous herb, apparently localized in
grasslands and woodlands of Escarpment Slopes
Clivia Lindl.
caulescens R.A. Dyer , perennial herb, localized in shady field
layer of Mist belt forests, Scheepers 407
Nerine Herb.
angustifolia Bak., large bulbous herb of marshy grasslands.
Summit Plateau, 2569
Brunsvigia Heist.
radulosa Herb., large bulbous herb, rare in grasslands of Lower
Mountain Slopes
Crinum L.
sp., bulbous herb, occasional in lithoseral woodlands, Low
Country, 974
Cyrtanthus L. f.
bicolor R.A. Dyer, herb, infrequent in lithoseral grasslands of
Escarpment Slopes, 1161
HYPOXIDACEAE
Hypoxis L.
‘complex’, robust geophytic herb, widespread in grasslands and
woodlands of Mistbelt and Low Country:
rigidula Bak., 1302
rooperi S. Moore, 66 7
costataSo^. , infrequent herb amongst rocks, Summit Peak shrub-
lands, 2489
filiformis Bak., delicate geophytic herb, widespread in Mistbelt
grasslands, 1093
galpinii Bak. , widespread geophytic herb of rocky grasslands
and woodlands, Mistbelt, 1112
gerrardii Bak., rare geophyte of Lower Mountain grasslands,
1502
multiceps Buchinger, robust geophyte, localized in dolomite-
soil grasslands of Plateau Interior, 131 7
VELLOZIACEAE
Xerophyta Juss.
retinervis Bak. , short woody perennial, common in early litho-
sere of Mistbelt and Low Country, 2003
DIOSCOREACEAE
Dioscorea L.
‘complex’, soft perennial twiner, frequent in forests and thickets
of Mistbelt and Low- Country:
cotinifolia Kunth, 655
sylvatica (Kunth) Eckl. var. sylvatica, 377
dregeana (Kunth) Dur. & Schinz var. dregeana, soft perennial
twiner, infrequent in closed rocky woodlands of Escarp-
ment Slopes, 1443 A
1RIDACEAE
Moraea Mill.
elliotii Bak. , small herb, localized in moist lithoseral grasslands
of Escarpment Slopes, 1118
muddii N.E. Br., medium-size herb, occasional in Mistbelt grass-
lands, 1426
sp., herb, scattered in woodlands and grasslands of Mistbelt,
includes 1628B
Dietes Salisb. ex Klatt
iridioides (L.) Sweet ex Klatt, gregarious herb, widespread but
localized (sometimes dominant) in Forest and Mesic
Thicket of Mistbelt and Low Country, 362
Aristea A it.
angolensis Bak. , infrequent herb of Summit Slope grasslands,
2310
woodii N.E. Br. , widespread herb in woodlands and grasslands
of Mistbelt and Low Country, 681
Schizostylis Backh. & Harv.
coccinea Backh. & Harv. , localized herb of marshy grasslands
and riparian woodlands. Summit Plateau, 2655
Hesperantha Ker-Gawl.
baurii Bak. , apparently rare bulbous herb of rocky Protea
woodland. Summit Slopes, 2599
Dierama K. Koch
sp. cf. galpinii N.E. Br., grass-like herb of Mistbelt grasslands.
includes 1282A
Crocosmia Planch.
aurea Planch., geophytic herb in light shade of Short (riparian)
Forest, Lower Mountain Slopes, 366
paniculata (Klatt) Goldbl., large geophytic herb of rocky shrub-
lands, Mistbelt, 2743
Gladiolus L.
densiflorus Bak. , large herb, scattered in grasslands and wood-
lands of Mistbelt and Low Country, 1 708
ecklonii Lehm., subsp. ecklonii, widespread, infrequent herb in
grasslands and woodlands of Mistbelt and Low Country,
1688
exiguus G.J. Lewis, widespread herb in woodlands and grass-
lands, Mistbelt, 1482
longicollis Bak. var. platypetalus (Bak.) Oberm. , widespread
herb of Summit Plateau grasslands, 2350
sp., widespread herb in grasslands and woodlands of Mistbelt
and Low Country, includes 782, 1318
varius F. Bol. var. micranthus (Bak.) Oberm. , widespread herb
of Summit grasslands, 2740
Watsonia Mill.
sp. cf. transvaalensis Bak., perennial herb of rocky shrublands,
Summit Slopes and Plateau, 2751
STRELITZIACEAE
Strelitzia Ait.
caudata R.A. Dyer, herbaceous tree, infrequent in Tall Forest
of Escarpment Slopes
ORCHIDACEAE
Stenoglottis Lindl.
fimbriata Lindl., small orchid on rocks and bark of elfin-like
forests. Plateau Crest, 1237
Holothrix L.C. Rich, ex Hook.
scopularia (Lindl.) Reichb. f, small ground orchid, apparently
rare in Protea woodland. Summit Slopes, 2429
Satyrium Swartz
cristatum Sond. var. longilabiatum A. V. Hall, infrequent geo-
phytic herb of rocky shrublands, Mistbelt. 2746
longicauda Lindl. var. longicauda, small geophytic herb, in-
frequent in grasslands of Summit Slopes, 2697
Browmleea Harv. ex Lindl.
coerulea Harv. ex Lindl., small herb, infrequent in closed wood-
lands of Low Country, 830
Disa Berg.
stachyoides Reichb. f, infrequent ground orchid of encroached
grasslands, Summit Plateau, 2550
Disperis Swartz
fanniniae Harv., small herb on rocks of elfin-like forests, Plateau
Crest, 1239
Polystachya Hook.
concreta (Jacq.) Garay & Sweet, epiphytic orchid, occasional
in mesic thickets of Transitional Mistbelt, 1622
ottoniana Reichb. f, small epiphytic orchid with row's of pseudo-
bulbs forming mats on trees in elfin-like forests, Plateau
Crest, 1201
sp., epiphytic orchid localized in mesic thickets, Transitional
Mistbelt, includes 926, 1049
Ansellia Lindl.
gigantea Reichb. f, large-tufted epiphytic orchid, apparently
localized in xeric thickets of Low Country, 803
Eulophia R. Br. ex Lindl.
horsfallii (Batem.) Summerh. , large ground orchid in Low Coun-
try riparian forests
streptopetala Lindl. , large ground orchid in light shade of forests,
woodlands and thickets, Mistbelt and Low Country,
1446
BulbophyUum Thouars
sandersonii Reichb. f, orchid on rocks and bark of elfin-like
forests, Plateau Crest, 1198
Tridactyle Schltr.
tricuspis (H. Bol) Schltr., widespread epiphyte in Forest and
Mesic Thicket of Mistbelt and Low Country, 658A
ANGIOSPERMAE — DICOTYLEDONAE
PIPERACEAE
Piper L.
capense L. f, soft shrub, sometimes dominant in Tall Forest of
Escarpment and Mountain Slopes, 2055
98
Bothalia 19,1 (1989)
Peperomia Ruiz. & Pav.
blanda (Jacq.) H.B.K. var. leptostachya (Hook. & Am) Duell,
small occasional herb on rocks in mesic thickets. Transi-
tional Mistbelt, 708
retusa (L. f.) A. Dietr. , small epiphytic herb (also on rocks) com-
mon in forests and mesic thickets, Mistbelt, 715
tetraphylla (G. Forst.) Hook. & Am, small epiphytic herb
(also on rocks) localized in forests of Humid Mistbelt,
1192
MYR1CACEAE
Myrica L.
pilulifera Rendle, small tree, widespread in rocky woodlands of
Mistbelt, 960
serrata Lam. . small tree of river banks. Low Country, 897
ULMACEAE
Celtis L.
africana Burnt f. , tall tree, widespread in forests, w oodlands
and thickets of Mistbelt and Low Country, 256
Trema Lour.
orientalis (L.) Blume , medium-size pioneer tree, w idespread in
thickets and woodlands of Mistbelt and Low Country,
794
MORACEAE
Ficus L.
ingens (Miq.) Miq. var. ingens, fairly small tree, localized on
rocks of Mistbelt and Low Country, 811
sur Forssk., fairly large tree, widespread in forests, woodlands
and thickets of Mistbelt and Low Country, 858
thonningii Blume, medium-size tree, widespread in forests,
woodlands and thickets of Mistbelt and Low Country,
872
URTICACEAE
Laportea Gaudich.
peduncularis ( Wedd .) Chew, shrubby perennial, infrequent in
rocky forested kloofs of Lower Foothills, 1 756
PROTEACEAE
Faurea Harv.
galpinii Phill. , rare understorey tree of Tall Forest, Upper Moun-
tain Slopes
saligna Harv., medium-size tree, often dominant in woodlands
of Low Country, 1 788
speciosa (Welw.) Welw., small tree, common in woodlands and
thickets of Mistbelt and Low Country, 644
Protea L.
caffra Meisn., small tree of rocky grasslands and woodlands,
Mistbelt, 1488
gaguedi Gmel., small tree, occasional in rocky grasslands of
Summit and Escarpment Slopes, 1145 A
parvula Beard, low prostrate shrublet, widespread in Summit
grasslands and shrublands, 2330
roupelliae Meisn. subsp. roupelliae, small tree of rocky wood-
lands, Summit Slopes, 2594
welwitschii EngL, small tree, common in localized open wood-
lands of Plateau Interior, 1666
LORANTHACEAE
Tapinanthus (Blume) Reichb.
rubromarginatus (Engl.) Danser, woody epiphytic parasite in
Protea woodland, Summit Slopes, 2610
Erianthemum V. Tieghem
dregei (EckL & Zeyh.) V. Tieghem, woody hemiparisitic epi-
phytic bush, fairly localized in rocky xeric thickets of
Low Country, 846
SANTALACEAE
Osyridicarpos A. DC.
schimperianus (Hochst. ex A. Rich.) A. DC., shrubby hemi-
parasitic climber, localized in thickets of Transitional
Mistbelt, 701
Thesium L.
costatum A. W. Hill, small perennial forb (hemiparasitic), infre-
quent in Mistbelt shrublands and grasslands, 1216
cytisoides A. W. Hill, infrequent forb in marshy grasslands,
Plateau Crest, 1268
goetzeanum Engl. , small perennial forb, infrequent in retarded
grasslands, Summit Plateau, 2321
sp., infrequent forb in Mistbelt grasslands, includes 1500, 255 7
OLACACEAE
Ximenia L.
caffra Sond. , small tree, occasional in Low Country woodlands,
1772
POLYGONACEAE
Rumex L.
bequaertii De Wild., occasional herb of marshy grasslands. Sum-
mit Plateau, 2574
sagittatus Thunb., shrubby perennial climber, localized in
rocky woodlands and thickets of Plateau Crest, 1411
sp., occasional waterside herb of riparian woodlands. Summit
Plateau, 2441
Polygonum L.
salicifolium Willd., small infrequent herb of stream banks,
Low'er Foothills, 1911
Oxygonum Burch.
dregeanum Meisn. var. strictum (C.H. Wr.) R.A. Grab., rare
semi-succulent bushy herb of Summit Plateau grasslands,
2553
AMARANTHACEAE
Cyathula Blume
cylindrica Moq., infrequent shrubby scrambler of Tall Forest,
especially Low Country, 1410
Pupalia Juss.
atropurpurea Moq., shrubby climber, infrequent in Low Coun-
try forests, 175 7
*Achyranthes L.
sicula (L.) All., infrequent forb, widespread throughout Mist-
belt and Low Country, 1599
PHYTOLACCACEAE
Phytolacca L.
octandra L., infrequent shrub in Tall Forest of Escarpment
Slopes, 1066
AIZOACEAE
Psammotropha Eckl. & Zeyh.
myriantha Sond., low tufted herb, localized in Summit Peak
grasslands, 2734
RANUNCULACEAE
Knowltonia Salisb.
transvaalensis Szyszyl. var. transvaalensis, rosette forb, wide-
spread in grasslands of Escarpment Slopes and Plateau,
744
Clematis L.
brachiata Thunb., infrequent liane, widespread in Forest and
Mesic Thicket of Mistbelt and Low Country, 1383
Ranunculus L.
multifidus Forssk. , small infrequent herb of riparian woodlands,
Summit Plateau, 2436
Thalictrum L.
rhynchocarpum Dill. & Rich., robust perennial herb of cool
moist shady undergrowrth, Mountain Slopes forests,
1353A
MENISPERMACEAE
Cocculus DC.
hirsutus (L.) Diels, slender twiner, infrequent in sheltered
thickets of Lower Foothills, 2027
Stephania Lour.
abyssinica Dill. & Rich., slender climber, infrequent in riparian
woodlands and forests of Mistbelt, 216
Cissampelos L.
torulosa E. Mey. ex Harv., soft twiner, abundant in forests and
thickets of Mistbelt and Low Country, 200
ANNONACEAE
Monanthotaxis BailL
caffra (Sond.) Verde., scrambling shrub, common in Forest and
Mesic Thicket of Mistbelt and Low Country, especially
below Escarpment Plateau, 707
Annona L.
senegalensis Pers.. low spreading tree, localized and w idespread
in Woodland and Xeric Thicket of Low Country, 738
trimeniaceae
Xymalos Baill.
monospora (Harv.) Badl. , medium-size tree, often dominating
the understorey of forests in Humid Mistbelt, 1002
Bothalia 19,1 (1989)
99
LAURACEAE
Cryptocarya R. Br.
liebertiana Engl., dominant canopy tree of Tall Forest, Moun-
tain Slopes, 2244
Ocotea AubL
kenyensis (Chiov.) Robyns, large tree, infrequent in forests of
Mistbelt and Low Country. 1388
BRASSICACEAE
Rorippa Scop.
*nasturtium-aquaticum (L.j Hayek , infrequent herb in shelter
of large boulders. Summit Peak shrublands, 2494
CAPPARACEAE
Capparis L.
brassii DC. .fairly rare scrambling shrub of Tall (riparian) Forest,
Low Country, 896
sepiaria L. var. subglabra (Oliv.) De Wolf, rare scrambler in Tall
(riparian) Forest, Lower Foothills, 1874
DROSERACEAE
Drosera L.
burkeana Planch., fairly common insectivorous herb of Summit
grasslands, 2533
sp., small insectivorous herb, infrequent in sandy grasslands of
Plateau Crest
CRASSULACEAE
Kalanchoe Adans.
rotundifolia (Haw.) Haw. , widespread succulent herb of Mistbelt
and Low Country lithosere, 779
Crassula L.
acinaciformis Schinz, giant subsucculent herb, infrequent in
woodlands and thickets of Low Country
alba Forssk.
‘complex’, succulent herb, common in lithoseral grasslands and
w'oodlands, especially Mistbelt:
var. alba, 1416 A
var. parvisepala (Schonl.) Toelken, R71
natalensis Schonl. , small succulent herb of rock depressions,
Mistbelt woodlands, 1521
pellucida L. subsp. brachypetala (Drege ex Harv.) Toelken,
small infrequent herb, gregarious in moist field layer of
forests, Escarpment Slopes, 1125
sarcocaulis Eckl & Zeyh. subsp. sarcocaulis, succulent shrublet,
widespread in Mistbelt rock formations, 1228
sp., occasional herb of moist grasslands, Escarpment Slopes,
includes 1115
swaziensis Schonl, small herb of rock formations in woodlands
and forests, Plateau Crest, 1232
vaginata Eckl. & Zeyh., succulent herb, infrequent in Mistbelt
grasslands, 1483
ESCALLONIACEAE
Choristylis Harv.
rhamnoides Harv., shrub of Mistbelt thickets and forests, 831
PITTOSPORACEAE
Pittosporum Banks ex Soland.
viridiflorum Sims, small tree, mainly localized in understorey
of thickets and woodlands. Transitional Mistbelt, 1793
myrothamnaceae
Myrothamnus Welw.
flabellifolia (Sond.) Welw. , xerophytic shrublet of sheet-rock
formations, especially Low Country, 1853
hamamelidaceae
Trichocladus Pers.
grandiflorus Oliv., large tree in canopy of Tall Forest, Plateau
Crest and Escarpment Upper Slopes, 1059
ROSACEAE
Rubus L.
pinnatus Willd.. widespread scrambler in forests and woodlands
of Mistbelt and Low Country, 2023
sp.. widespread scrambler in Forest and Mesic Thicket of Mist-
belt and Low Country, includes 480
Alchemilla L.
rehmannii Engl., infrequent herb of stream banks, Summit
Plateau, 2438
Agrimonia L.
*odorata Mill. , subwoody herb, infrequent in moist low thickets
of Mountain Slopes, 1596
Leucosidea Eckl. & Zeyh.
sericea Eckl. & Zeyh., small (usually riparian) tree of Summit
and Mountain Slopes. 56
Cliffortia L.
nitidula (Engl.) R.E. & Th. Fries Jr subsp. piiosa Weim., com-
mon ericoid shrub of open rocky woodlands and forest
margins, Mistbelt, 1693
repens Schltr., ericoid shrub, infrequent in Mistbelt grasslands,
1082
serpyllifolia Cham. & Schlechtd. , infrequent shrub amongst
rocks. Summit Peak shrublands, 2502
Prunus L.
atricana (Hook.f.) Kalkm., large canopy tree, occasional (some-
times dominant) in Mistbelt forests, 966
chrysobalanaceae
Parinari Aubl.
capensis Harv. subsp. capensis, low shrublet often associated
with rock outcrops, w idespread in grasslands of Escarp-
ment Plateau, 1295
curatellifolia Planch, ex Benth., fairly large canopy tree, abun-
dant (often dominant) in thickets and woodlands of
Mistbelt and Low Country, 7
connaraceae
Cnestis Juss.
natalensis (Hochst.) Planch. & Sond., small understorey tree in
Mistbelt forests and mesic thickets, 908
FABACEAE
Albizia Durazz.
versicolor It'e/w. ex Oliv. , medium-size tree, infrequent in wood-
lands of Upper Foothills, 2039
Acacia Mill.
ataxacantha DC., abundant and vigorous scandent tree, domi-
nant in thickets of Mistbelt and Low Country, 650
caffra (Thunb.) Willd. , small tree, w idespread in Woodland and
Xeric Thicket of Low Country, 818
davyi N.E. Br., small tree, widespread in Woodland and Xeric
Thicket of Low Country, 76 7
*mearnsii De Wild., naturalized exotic tree, fairly widespread
sp., small tree scattered infrequently in woodlands and thickets
of Mistbelt and Low Country, includes 868
Dichrostachys (A. DC.) Wight & Am.
cinerea (L.) Wight & Am.
‘complex’, small tree, localized and widespread in Woodland
and Xeric Thicket of Low' Country:
subsp. africana Brenan & Brumm. var. africana, 628
subsp. nyassana (Taub.) Brenan, 723
Entada Adans.
spicata (E. Mey.) Druce, robust prickly liane, localized in Tall
Forest of Mistbelt and Low Country, 1008
Bauhinia L.
galpinii N.E. Br., abundant scrambling shrub, often dominant
in woodlands, thickets and forests, especially Low
Country, 2030
Piliostigma Hochst.
thonningii (Schumach.) Milne-Redh. , infrequent small tree in
Low Country woodlands, 471
Tylosema (Schweinf.) Torre & Hillc.
fassoglensis (Schweinf.) Torre & Hillc., widespread robust cree-
per, a persistent pioneer of lithoseral woodlands and
thickets; exclusively Low Country, 519
Cassia L.
*bicapsularis L., shrubby climber, infrequent in mesic thickets
of Upper Foothills, 1800
*floribunda Cav. , occasional shrub in mesic thickets of Low
Country, 257
mimosoides L. , small subwoody forb, infrequent in sandy grass-
lands of Plateau Crest, 1643
petersiana Bolle, widespread shrub in Woodland and Xeric
Thicket of Low Country, 738
plumosa (E. Mey.) Vogel var. erecta Schorn & Gordon-Gray,
small subwoody forb, infrequent in rocky grasslands of
Lower Mountain Slopes, 1552
quarrei (Ghesq.) Steyaert, subw'oody forb, localized in disturbed
open woodlands of Upper Foothills, 599
Peltophorum (Vogel) Benth.
100
africanum Sond., medium-size tree, widespread in thickets and
woodlands of Foothills, 803A
Calpurnia E. Mey.
aurea (Ait.) Benth. , small localized tree in understorey of forests,
Lower Mountain Slopes, 295
Lotononis (DC.) Eckl. & Zeyh.
pulchra Duemmer, infrequent shrublet in grasslands of Lower
Mountain Slopes, 1476
Pearsonia Duemmer
aristata (Schinz) Duemmer, subwoody shrublet, widespread
but infrequent in grasslands and woodlands of Mistbelt
and Low Country, 1518A
obovata (Schinz) Polhill, prostrate subwoody forb, infrequent
in Mistbelt grasslands, 1522
sessilifolia (Harv.) Duemmer
'complex’, subwoody bushy forb, widespread and common in
woodlands and grasslands of Mistbelt and Low Country:
subsp. marginata (Schinz) Polhill, 1452
subsp. sessilifolia, 1707
uniflora (Kensit) Polhill, subwoody forb, rare in xeric thickets
of Low Country, 821
Crotalaria L.
capensis Jacq ., small shrub in Tall (riparian) Forest, Low Coun-
try, 888
recta Steud. ex A. Rich., small shrub, localized in disturbed
woodlands and thickets of Upper Foothills, 1624
Argyrolobium Eckl. £ Zeyh.
harveyanum Oliv. , slender perennial herb, infrequent in grass-
lands of Summit Slopes, 2373
speciosum Eckl. £ Zeyh., infrequent subwoody forb scattered
in woodlands, grasslands and thickets of Mistbelt and
Low Country, 709
transvaalenseSclnhz.infrequent subwoody forb in xeric thickets,
Low Country, 1840
Lotus L.
discolor E. Mey. subsp. discolor, fairly rare shrublet in moist
grasslands of Escarpment Lower Slopes, 1946
Indigofera L.
comosa N.E. Br. , infrequent shrublet of sheet-rock formations,
Low Country, 1883
hedyantha Eckl. £ Zeyh. , infrequent subwoody forb of retarded
grasslands. Summit Plateau, 2356
hilaris Eckl £ Zeyh., subwoody forb, infrequent in grasslands
of Mountain Slopes, 1535
oxalidea Welw. ex Bak., prostrate forb, infrequent in grasslands
and woodlands of Mistbelt and Low Country, 731
sanguinea N.E. Br. , subwoody forb, fairly frequent in grasslands
of Summit and Escarpment Plateau, 2073
sp., shrublet or prostrate forb of moist grasslands. Escarpment
Lower Slopes, includes 1945
sp. 1, subwoody shrublet localized in grasslands of Lower
Mountain Slopes, 1460
sp. 2, shrublet localized in grasslands of Mountain Slopes, 1475
swaziensis H. Bol, small shrub, widespread and common in
woodlands, grasslands and xeric thickets of Mistbelt and
Low Country, 665
tristoides N.E. Br., small shrub, infrequent in grasslands and
disturbed (sometimes riparian) woodlands of Mistbelt,
1939
Otholobium C.H. Stirton
polystictum (Benth. ex Harv.) C.H. Stirton, infrequent shrub
of wooded rocky knolls, Summit, 2311
Tephrosia Pers.
‘complex’, large forb, mainly in Low Country thickets and
woodlands:
polystachya E. Mey. var. latifolia Harv., 765A
shiluwanensis Schinz, 765
elongata E. Mey. var. elongata, lax forb, infrequent in grasslands,
woodlands and thickets of Mistbelt and Low Country,
1153
macropoda (E. Mey.) Harv., lax forb, infrequent in grasslands
of Escarpment Slopes, 1117
semiglabra Sond., forb, infrequent in rocky shrublands of
Lower Mountain Slopes, 1553
Aeschynomene L.
nyassana Taub., subwoody forb, common in rocky woodlands
and grasslands of Humid Mistbelt, 1182
rehmannii Schinz, subwoody forb, infrequent and localized in
sandy grasslands of Plateau Crest, 1246
var. leptobotrya (Harms ex Bak. f.) J.B. Gillett, subwoody
forb, localized in rocky grasslands, shrublands and
Bothalia 19,1 (1989)
woodlands of Escarpment Upper Slopes, Plateau Crest
and Summit, 1420
Smithia Ait.
erubescens (E. Mey.) Bak. f, shrublet in grasslands of Transi-
tional Mistbelt, 1950
Stylosanthes Swartz
fruticosa (Retz.) Alston, infrequent subwoody forb of dis-
turbed woodlands, Upper Foothills, 604
Zornia J.F. Gmel.
milneana Mohlenbr. , rare ground creeper of disturbed grasslands.
Plateau Crest, 1298
Desmodium Desv.
dregeanum Benth., small shrub, infrequent in disturbed grass-
lands of Upper Foothills, 737
gangeticum (L.) DC., infrequent lax forb of woodlands and
thickets, Low Country, 793
repandum (Vahl) DC., small undershrub, widespread and com-
mon in Forest and Mesic Thicket of Mistbelt and Low
Country, 1002A
setigerum (E. Mey.) Benth. ex Harv., ground creeper, infre-
quent in disturbed grasslands and woodlands of Mistbelt,
1288
Pseudarthria Wight £ Am.
hooked Wight & Am. var. hookeri, robust shrubby forb, wide-
spread and common in Woodland and Xeric Thicket of
Low Country and in Mistbelt grasslands and woodlands,
610
Dalbergia L. f.
armata E. Mey., robust spiny liane, common and widespread in
forests and mesic thickets, especially below Escarpment
Plateau, 21
Pterocarpus Jacq.
angolensis DC., fairly large canopy tree, mainly confined to
Woodland and Xeric Thicket of Low Country, 248
rotundifolius (Sond.) Druce subsp. rotundifolius, fairly large
canopy tree, confined to partially sheltered woodlands
and thickets on shallow soils of Lower Foothills, 2040
Abrus A dans.
laevigatus E. Mey. , slender subwoody twiner, common in forests,
thickets and woodlands below Escarpment Plateau, 713
Dumasia DC.
villosa DC. var. villosa, soft twiner in shrub layer of Tall Forest,
Mountain Slopes, 1573
Neonotonia Lackey
wightii (Am.) Lackey, localized climber of xeric thickets, Low
Country, 802
Erythrina L.
latissima E. Mey., small tree, infrequent in woodlands of Plateau
Crest and Escarpment Slopes, 1437
lysistemon Hutch., fairly large tree, widespread in forests,
thickets and woodlands of Mistbelt and Low Country,
874
Mucuna Adans.
coriacea Bak. subsp. irritans (Burtt Davy) Verde. , soft perennial
twiner, fairly common in woodlands and thickets of
Foothills, 1626
Rhynchosia Lour.
angulosa Schinz, localized forb in grasslands of Lower Moun-
tain Slopes, 1485
caribaea (Jacq.) DC., slender twiner, widespread in forests,
thickets and woodlands of Mistbelt and Low Country,
696
hirta (Andr.) Meikle & Verde., subwoody soft twiner, localized
in forests and mesic thickets of Foothils, 42
komatiensis Harms, small shrub, widespread and common but
localized mainly in Woodland and Xeric Thicket of Low
Country, 620
monophylla Schltr., herbaceous creeper, common in broken
lithoseral grasslands and woodlands of Mistbelt, 1413A
sordida (E. Mey.) Schinz, large subwoody forb, infrequent and
localized in disturbed woodlands of Upper Foothills,
1829
thorncroftii (Bak. f.) Burtt Davy, infrequent small shrub of
riparian forests, Escarpment Lower Slopes, 988
totta (Thunb.) DC., small delicate twiner, occasional in grass-
lands and woodlands of Mistbelt and Low Country,
1327
villosa (Meisn.) Druce , large forb, infrequent in moist grass-
lands of Summit and Plateau Crest, 1261
woodii Schinz, occasional prostrate subshrub of Summit grass-
lands, shrublands and woodlands, 2519
Bothalia 19,1 (1989)
101
Eriosema (DC.) G. Don
angustifolium Bunt Davy, widespread forb of Humid Mistbelt
grasslands, 1244
burkei Benth., infrequent forb of dolomite-soil grasslands,
Plateau Interior, 1345
cordatum E. Mey., infrequent forb of Mistbelt grasslands, 1362
ellipticifolium Schinz, widespread forb of Mistbelt grasslands,
1116
gunniae C.H. Stinon, occasional forb of Mistbelt grasslands,
1544
kiaussianum Meisn., common forb of retarded grasslands. Sum-
mit Plateau and Slopes, 2345
nutans Schinz, localized forb of Plateau Interior grasslands,
1657
psoraleoides (Lam.) G.Don, robust shrubby forb, infrequent in
Low Country woodlands, 1768
Flemingia Roxb. ex Ait. f.
gTahamiana Wight & Am., robust shrubby forb, infrequent in
Low Country woodlands, 2034
Vigna Savi
oblongifolia A. Rich. var. oblongifolia, soft slender twiner, rare
in sheltered w oodlands of Lower Foothills, 1780
nervosa Markoetter, delicate twiner, infrequent in grasslands of
Humid Mistbelt, 1481
Sphenostylis E. Mey.
angustifolia Sond, lax herbaceous twiner, infrequent and local-
ized in Escarpment Plateau grasslands, 1334
marginata E. Mey. subsp. marginata, vigorous climber, infre-
quent in thickets of Upper Foothills, 692
GERANIACEAE
Geranium L.
ornithopodon Eckl. & Zeyh., trailing herb, common in moist
(often riparian) grasslands and woodlands, Summit,
2433
Monsonia L.
attenuata Han. , rare forb of moist grasslands, Escarpment
Lower Slopes, 1688
transvaalensis Knuth, widespread forb of Summit grasslands,
2552
OXALIDACEAE
Oxalis L.
depressa Eckl. & Zeyh., very small delicate herb, infrequent in
Mistbelt grasslands and open woodlands, 1507
obliquifolia Steud. ex Rich., small herb, widespread in Summit
grasslands, w'oodlands and shrublands, 2516
LINACEAE
Linum L.
thunbergii Eckl & Zeyh., small localized herb of disturbed
lithoseral grasslands, Plateau Crest, 1284
RUTACEAE
Zanthoxylum L.
capense (Thunb.) Harv., small tree, localized in shrub layer of
forests, especially Escarpment Slopes, 334
davyi (Verdoorn) Waterm., widespread canopy tree of wood-
lands, forests and thickets of Mistbelt and Low Country,
694
thorncroftii (Verdoorn) Waterm., small tree, localized in shrub
layer of elfin-like forests. Plateau Crest
Vepris Comm, ex A. Juss.
undulata (Thunb.) Verdoorn & C.A. Sm., infrequent under-
storey tree of High Forest, Upper Mountain Slopes,
2254
Toddalia Juss.
asiatica (L.) Lam., robust prickly liane, commonly localized in
forest's and mesic thickets below Escarpment Plateau,
2025
Clausena Burm. f.
anisata (Willd.) Hook. f. ex Benth. , small understorey tree, wide-
spread in forests of Mistbelt and Low Country, 875
Citrus L.
*spp., exotic tree of cultivation, occasionally encroaching on
mesic thickets of Transitional Mistbelt
SIMAROUBACEAE
Kirkia Oliv.
acuminata Oliv. , medium size deciduous tree, infrequent in
moist woodlands and forests of Low Country, 1 718
PTAEROXYLACEAE
Ptaeroxylon Eckl. & Zeyh.
obliquum (Thunb.) Radik., infrequent understorey tree of
High Forest, Mountain Slopes, 2246
MELIACEAE
Ekebergia Sparrm.
capensis Sparrm. , small tree, localized and rare in understorey
of Tall (riparian) Forest, Low Country, 892
pterophylla (C. DC.) Hofmeyr, small tree, localized in elfin-like
forests and rocky woodlands of Plateau Crest, 1689
Trichilia P. Br.
emetica Vahl, fairly large canopy tree, infrequent in Low Coun-
try forests, 1 753
MALPHIGIACEAE
Sphedamnocarpus Planch, ex Benth. & Hook. f.
galphimiifolius (Juss.) Szyszyl.
subsp. galphimiifolius, slender subwoody climber, infrequent
in forests of Escarpment Lower Slopes, 1959
subsp. rehmannii Launert, robust climber, infrequent in xeric
thickets of Low Country, 819
pruriens (Juss.) Szyszyl.
var. lanceolatus Launen, subwoody climber, infrequent in
developing thickets of Escarpment Upper Slopes, 1674
var. pruriens, infrequent subwoody climber of xeric thickets,
Low Country, 500
POLYGALACEAE
Polygala L.
hottentotta Presl, small infrequent herb, widespread in wood-
lands and grasslands of Mistbelt and Low' Country, 625
sp., occasional forb of Summit shrublands and woodlands, in-
cludes 2640. 2648
uncinata E. Mey. ex Meisn., small infrequent herb, mainly local-
ized on sheet-rock formations supporting woodland,
Low Country, 978
virgata Thunb., small infrequent shrub of Upper Foothill
thickets, 943
Muraltia Juss.
flanaganii H. Bol., infrequent ericoid shrublet of rocky shrub-
lands, Summit Peak, 2507
sp., prostrate ericoid shrublet, infrequent in low shrublands of
exposed Summit Slopes, 2410
euphorbiaceae
Andrachne L.
ovalis (Sond.) Mull. Arg., infrequent shrub in riparian forests,
Upper Foothills, 1044
Securinega Comm, ex Juss.
virosa (Roxb. ex Willd.) Pax & K. Hoffrn., localized shrub in
woodlands and thickets, Low Country, 792
Phyllanthus L.
nummulariifolius Poir., infrequent shrub of xeric woodlands,
Lower Foothills, 1 774
reticulatus Poir. , untidy localized shrub of Low Country wood-
lands, especially on broken lithosere, 1698
Drypetes Vahl
gerrardii Hutch., small understorey tree, localized in riparian
forests of Mountain Slopes, 900
Antidesma L.
venosum E. Mey. ex. Tul., low spreading tree, common and
widespread in thickets and woodlands, especially Low
Country, 637
Bridelia Willd.
micrantha (Hochst.) Baill., fairly large tree, common in thickets
and woodlands. Often a pioneer on sheet-rock forma-
tions, mainly Low Country, 652
Adenocline Turcz.
acuta (Thunb.) Baill., soft undershrub, infrequent in kloof
forests of Upper Mountain Slopes, 2151
Acalypha L.
angustata Sond. vat. glabra Sond. , localized subwoody forb of
Lower Mountain Slope grasslands, 1458
caperonioides Baill. , widespread subwoody forb of Low'er Moun-
tain and Summit Plateau moist grasslands, 1534
petiolaris Hochst., localized subwoody forb of Foothill wood-
lands, 758
punctata Meisn., infrequent subwoody forb of Upper Foothill
woodlands and thickets, 622
wilmsii Pax ex Prain & Hutch., subwoody forb, common in
woodlands and grasslands of Humid Mistbelt, 1422
102
Tragia L.
okanyua Pax, slender twiner, infrequent in disturbed xeroclinal
woodlands of Upper Foothills, 6 72
rupestris Sond., slender twiner or creeper occurring in Plateau
Interior grasslands and Foothill thickets, 799
sp., infrequent twiner of Low Country woodlands, includes
1820
Ctenomeria Harv.
capensis (Thunb.) Harv. ex Sond., large twiner, infrequent in
mesic thickets of Escarpment Lower Slopes, 1401
Dalechampia L.
capensis Spreng. f, slender climber, localized in rocky mesic
thickets of Upper Foothills, 712A
Clutia L.
abyssinica Jaub. & Spach var. abyssinica, prostrate subwoody
herb, infrequent in disturbed woodlands of Upper
Foothills, 752
affinis Sond. , soft shrub of Cyathea riparian woodland, Summit
Plateau, 2120
hirsuta E. Mey. ex Sond., small shrub, infrequent in xeroclinal
thickets of Upper Foothills, 940
monticola S. Moore, frequent subwoody herb of woodlands
and grasslands, especially Humid Mist belt, 869
Euphorbia L.
epicyparissias E. Mey. ex Boiss. , undershrub in Leucosidea
riparian woodland. Summit Plateau. 2112
ingens E. Mey. ex Boiss., large succulent tree, usually associa-
ted with rocky xeric thickets and woodlands, Low-
Country
kraussiana Bernh ., small herbaceous perennial, rare in under-
growth of rocky riparian thickets, Escarpment Lower
Slopes, 932
striata Thunb. , localized forb of Plateau Crest grasslands, 1202
ANACARDIACEAE
Sclerocarya Hochst.
birrea (A. Rich.) Hochst. subsp. caffra (Sond.) Kokwaro, large
tree, widely scattered in Low- Country woodlands, 1844
Lannea A. Rich.
discolor (Sond.) Engl., small tree of lithoseral woodlands and
thickets. Low Country, 814
edulis (Sond.) Engl., low woody forb, widespread in Woodland
and Xeric Thicket of Low Country and in lithoseral
grasslands of Plateau Crest, 1421
Protorhus Engl.
longifolia (Bernh.) Engl., medium-size tree, common in forests
of Plateau Crest and Escarpment Lower Slopes and in
lithoseral woodlands and thickets, especially Upper
Foothills, 914
Rhus L.
chirindensis Bak. f. , occasional tree in understorey or canopy
of Escarpment Slopes forests, 939
dentata Thunb., widespread small tree occurring throughout
Mistbelt and Low Country, 675
discolor E. Mey. ex Sond. , small shrub in grasslands of Moun-
tain Slopes, precursor to open woodland, 1529
pentheri Zahlbr., small understorey tree of forests, woodlands
and thickets of Low Country, 757
pyroides Burch, var. pyroides, small bushy tree, widespread in
forests, woodlands and thickets of Mistbelt and Low
Country, 760
rehmanniana Engl., srqall tree, mainly localized in Plateau Crest
woodlands, 1436
transvaalensis Engl. , small tree, common and widespread in
woodlands and thickets, especially Woodland and Xeric
Thicket of Low Country, 675A
tumulicola S. Moore. , localized shrub of broken lithoseral shrub-
lands and woodlands, especially Summit and Plateau
Crest, 1185
AQUIFOLIACEAE
Ilex L.
mitis (L.) Radik., infrequent waterside tree, Mistbelt forests,
1354
CELASTRACEAE
Maytenus Molina
acuminata (L. f.) Loes. var. acuminata, frequent understorey
and canopy tree of Mountain Slopes forests, 2109
heterophylla (Eckl. & Zeyh.) N.K.B. Robson, small tree, com-
Bothalia 19,1 (1989)
mon in woodlands and thickets, especially Low Coun-
try, 601
mossambicensis (Klotzsch)Blakelock var. mossambicensis, small
tree, frequent in Forest and Mesic Thicket of Mistbelt
and Low Country, 269
peduncularis (Sond.) Loes, small tree, scattered infrequently in
Forest and Mesic Thicket of Mistbelt and Low Country,
900
undata (Thunb.) Blakelock, small tree of forests, woodlands
and thickets, especially Low Country, 837
Catha Forssk. ex Scop.
edulis (Vahl) Forssk. ex EndL, medium-size tree localized in
Low Country thickets, often on rocky sites, 644A
Pterocelastrus Meisn.
echinatus N.E. Br., occasional small tree in humid Mistbelt
forests and w'oodlands, 1067
Cassine L.
eucleiformis (Eckl. & Zeyh.) Kuntze, infrequent small tree of
Tall Forest, Mountain Slopes, 2284
papillosa (Hochst.) Kuntze, understorey tree of Tall Forest,
Mountain Slopes, 2279
tetragona (L. f.) Loes., infrequent understorey tree in High
Forest, Upper Mountain Slopes, 2169
Hippocratea L.
crenata (Klotzsch) K. Schum. & Loes., infrequent scandent
shrub of High Forest, Mountain Slopes, 2243
ICACINACEAE
Cassinopsis Sond.
ilicifolia (Hochst.) Kuntze , small understorey tree, widespread
in forests of Mistbelt and Low Country, 275
Apodytes E. Mey. ex Arn.
dimidiata E. Mey. ex Arn. subsp. dimidiata, medium-size tree,
w idespread and common in forests, thickets and wood-
lands, especially Forest and Mesic Thicket of Mistbelt
and Low Country, 666
Pyrenacantha Wight
grandiflora Baill., thick-branched climber, rare in rocky xeric
thickets of Upper Foothills, 1892
SAPINDACEAE
Allophyllus L.
‘complex’, small to large tree, infrequent but widespread in
Forest and Mesic Thicket of Mistbelt and Low Country:
cf. decipiens (Sond.) Radik., 2187
melanocarpus (Sond.) Radik., 1576
transvaalensis Rum Davy, 1129
Pappea Eckl. & Zeyh.
capensis Eckl. & Zeyh. , occasional small shrub in rocky wood-
lands, thickets and forests, Low Country, 1 705
MELIANTHACEAE
Bersama Fres.
transvaalensis Turrill, fairly large occasional tree of rocky forests
and woodlands, Mistbelt and Low Country, 1455
tysoniana Oliv., small tree, infrequent in forests of Mountain
and Escarpment Slopes, 1957
GREYIACEAE
Greyia Hook. & Harv.
radlkoferi Szyszyl., small tree of Mistbelt lithosere, occurring
as pioneer on sheet-rock formations and persisting in
forests, 1563
BALSAM IN ACE AE
Impatiens L.
hochstetteri Warb. subsp. hochstetteri, soft herb, localized in
moist field layer of riparian forests. Mountain and Escarp-
ment Slopes, 1013
RHAMNACEAE
Ziziphus Mill.
mucronata Willd. subsp. mucronata, small straggly tree, common
in woodlands and thickets, especially Low Country and
Transitional Mistbelt, 700
Berchemia Neck, ex DC.
zeyheri (Sond.) Grubov, small tree, infrequent in forests and
thickets of Lower Foothills, 1713
Scutia (Comm, ex DC.) Brongn.
Bothalia 19,1 (1989)
103
myrtina (Burm. f.) Kurz, robust scandcnt shrub, common in
High Forest of Mountain Slopes, 2195
Rhamnus L.
prinoides L’Herit., infrequent shrub in woodlands and forests.
Mist belt, 959
Phylica L.
paniculata Wittd. , infrequent shrub of broken lithoseral thickets.
Plateau Crest, 1227
V1TACEAE
Rhoicissus Planch.
revoilii Planch., robust woody liane of High Forest, Mountain
Slopes
rhomboidea (E. Mey. ex Harv.) Planch., robust woody liane,
frequent in forests and thickets of Mistbelt and Low
Country, 2048
tomentosa (Lam.) Wild & Drumm. , robust woody liane, frequent
in forests and thickets of Escarpment Slopes and Foot-
hills, 705
tridentata (L.f.) Wild & Drumm., small shrub, frequent in forests,
woodlands and thickets of Mistbelt and Low Country,
128
Cvphostemma (Planch.) Alston
anatomicum (C.A. Sm.) Wild & Drumm., occasional liane, scat-
tered in Mistbelt lithosere, 1392
simulans (C.A.Sm.) Wild & Drumm. ,rare twiner of xeric thickets,
Lower Foothills, 1878
woodii (Gilg& Brandt) Descoings, soft perennial forb, infrequent
in lithoseral shrublands and woodlands, mainly Low
Country, 810
TIL1ACEAE
Corchorus L.
confusus Wild, rare forb in xeric thickets. Upper Foothills, 741
sp., infrequent forb in grassy field layer of open woodlands,
Upper Foothills, 1903
Sparrmannia L. f.
ricinocarpa (Eckl. & Zeyh.) Kuntze, infrequent undershrub in
low thickets, Mountain Slopes, 1592
Grewia L.
monticola Sond. , small tree, localized in rocky woodlands of
Low Country, 1697
occidentalis L. , common and widespread shrubby tree of forests,
thickets and woodlands, Mistbelt and Low Country, 657
Triumfetta L.
pilosa Roth
var. effusa (E. Mey. ex Harv.) Wild, annual forb, scattered in
Low Country thickets and fairly localized in wood-
lands of Humid Mistbelt, 627
var. pilosa, annual forb, widespread in Low Country woodlands
and thickets, 733
var. tomentosa Szyszyl. ex Sprague & Hutch., annual forb,
localized in xeric thickets of Upper Foothills, 1803
rhomboidea Jacq., annual forb, widespread and infrequent in
woodlands and thickets of Mistbelt and Low Country,
685
welwitschii Mast. var. hirsuta (Sprague & Hutch.) Wild, peren-
nial forb, mainly localized in dolomite-soil grasslands of
Plateau Interior, 1323
MALVACEAE
Abutilon Mill.
sonneratianum (Cav.) Sweet, robust herb, infrequent in sheltered
woodlands of Low Country, 835
Sida L.
dregei Bum Davy, infrequent forb of xeric thickets, Lower Foot-
hills, 871
Pavonia Cav.
columella Cav. , large shrubby forb in moist disturbed field layer
of Low Country forests, 1925
Hibiscus L.
aethiopicus L. var. ovatus Harv., subwoody forb, localized in
Mistbelt grasslands, especially Plateau Interior, 1531
surattensis L., subwoody forb, infrequent in disturbed wood-
lands of Upper Foothills, 607
STERCULIACEAE
Dombeya Cav.
rotundifolia (Hochst.) Planch, var. rotundifolia, small tree, fre-
quent to abundant (often dominant) in Woodland and
Xeric Thicket of Low Country, 424
pulchra N.E.Br. , widespread occasional shrub of forests, thickets
and woodlands, Mistbelt and Low Country, 611
Hermannia L.
grandiflora N.E. Br., infrequent but widespread trailing forb of
disturbed woodlands, especially Low Country, 617
lancifolia Szyszyl., infrequent forb of Mistbelt grasslands, 1315
montana N.E. Br., localized forb of Mountain Slopes grasslands,
1490
Waltheria L.
indica L. , large shrubby forb of lithoseral woodlands, Low Coun-
try, 785
Sterculia L.
murex Hemsl., large canopy tree, widespread in woodlands and
thickets of Low Country, 2033
OCHNACEAE
Ochna L.
arborea Burch, ex. DC. var. arborea, fairly large understorey
tree, frequent (often dominant) in forests of Mountain
Slopes, 2163
gamostigmata Du Toit, small shrub, localized and common in
mesic thickets of Transitional Mistbelt and Low Coun-
try, 632
holstii Engl., medium-size understorey tree of forests and wood-
lands, Humid Mistbelt, 915
natalitia (Meisn.) Walp., small shrub, widespread in grasslands,
woodlands, thickets and forests of Mistbelt and Low
Country, 778
CLUSIACEAE
Hypericum L.
aethiopicum Thunb. subsp. sonderi (Bred.) N.K.B. Robson,
small subwoody forb, infrequent in Plateau Crest grass-
lands, 1642
lalandii Choisy, small perennial herb, localized in moist grass-
lands, Summit Plateau, 2565
revolutum Vahl, bushy shrub, locally abundant in riparian wood-
lands of forest margins. Summit, 2125
sp., shrub of riparian woodlands, Summit Plateau, 2131
FLACOURTIACEAE
Rawsonia Harv. & Sond.
lucida Harv. & Sond., understorey tree, infrequent in forests of
Escarpment and Mountain Slopes, 970
Kiggelaria L.
africana L. , fairly large canopy tree, frequent in forests of Escarp-
ment and Mountain Slopes, 989
Scolopia Schreb.
mundii (Eckl. & Zeyh.) Warb., infrequent shrub of kloof forests.
Mountain and Escarpment Slopes, 993
zeyheri (Nees) Harv., small to large tree, apparently localized
and infrequent in Low Country woodlands and thickets,
1720
Trimeria Harv.
grandifolia (Hochst.) Warb., understorey tree, common and
abundant in Forest and Mesic Thicket of Mistbelt and
Low Country, excluding Plateau Crest forests, 664
Flacourtia Comm, ex L ’Herit.
indica (Burm. f.) Merr., infrequent shrub in mesic woodlands
of Low Country, 813
Dovyalis E. Mey. ex Am.
lucida Sim, small tree or shrub, widespread in forests of Moun-
tain Slopes, 1360
zeyheri (Sond.) Warb., small tree or shrub, infrequent in Short
(cliff) Forest of Lower Mountain Slopes, 1386
PASSIFLORACEAE
Adenia Forssk.
digitata (Harv.) EngL, widespread and infrequent herbaceous
twiner in forests, thickets and woodlands, mainly Low
Country, 382
gummifera (Harv.) Harms var. gummifera, occasional climber in
Mistbelt and Low Country thickets, 1917
Passiflora L.
*edulis Sims, naturalized exotic climber, common in mesic
thickets of Transitional Mistbelt and Low Country, 654
BEGONIACEAE
Begonia L.
sp., occasional perennial herb in moist shady rock crevices,
Humid Mistbelt
104
Bothalia 19,1 (1989)
OLINIACEAE
Olinia Thunb.
emarginata Burtt Davy, fairly frequent understorey and canopy
tree of Tall (often riparian) Forest, Upper Mountain and
Summit Slopes, 2155
THYMELAEACEAE
Peddiea Harv.
africana Harv., small localized tree, common in Forest and
Mesic Thicket of Mist belt and Low Country, 719
Gnidia L.
caffra Meisn. , infrequent subwoody forb in grasslands, shrub-
lands and woodlands of Sub-Humid Mistbelt and Low
Country, 634
kraussiana Meisn. var. kraussiana, infrequent subwoody forb of
Sub-Humid Mistbelt and Low Country woodlands, 1815
microcephala Meisn. , infrequent subwoody forb of moist Escarp-
ment Plateau grasslands, 1653
nodiflora Meisn., infrequent subwoody forb of shallow rocky
sites, Summit Slopes shrublands, 2404
sp., subwoody forb of Mistbelt grasslands, includes 976, 1461
Passerina L.
montana Thoday, ericoid shrub, infrequent in rocky shrublands,
Summit Plateau
RHIZOPHORACEAE
Cassipourea Aubl
gerrardii (Schinz) Alston, frequent (sometimes dominant) under-
storey tree in forests of Mountain and Escarpment Upper
Slopes, 1060
COMBRETACEAE
Combretum Loefl.
apiculatum Sond. subsp. apiculatum, medium-size canopy tree,
occasional in woodlands of Low Country, 845
collinum Fresen.
subsp. gazense (Swynn. & Bak. f.) Okafor, medium-size canopy
tree, localized in woodlands of Lower Foothills
subsp. suluense (Engl & Diels) Okafor, localized canopy tree
of Low Country woodlands and thickets, 829A
kraussii Hochst., large canopy tree, common in Forest and
Mesic Thicket of Mistbelt and Low Country, 861
mode R. Br. ex G. Don, widespread small tree, especially com-
mon in Woodland and Xeric Thicket of Low Country,
631
sp., large waterside tree of Low Country forests, 887
zeyheri Sond., fairly small tree of Low Country woodlands and
thickets, 843
Quisqualis L.
parviflora Gerr. ex Harv., occasional scandent shrub of High
Forest, Upper Mountain Slopes, 2108
Terminalia L.
phanerophlebia Engl. & Diels, small infrequent tree of sheltered
xeric woodlands, Low Country, 862
sericea Burch, ex DC., medium-size canopy tree, scattered in
Low Country woodlands, 1758
MYRTACEAE
Psidium L.
*guajava L. , small spreading exotic tree, widespread and natural-
ized from cultivation
Eugenia L.
natalitia Sond., small understorey tree, widespread and fairly
common in forests and mesic thickets, especially Escarp-
ment Slopes, 884
Syzygium Gaertn.
cordatum Hochst., variable abundant tree, large in sheltered
forests, thickets and woodlands below Escarpment
Plateau; but stunted in rocky woodlands of Plateau
Crest, 883
gerrardii (Harv. ex Hook, f.) Burtt Davy, large canopy and
emergent tree, mainly localized in forests of Humid
Mistbelt, 909
guineense (Willd.) DC., small tree associated with S. cordatum
in rocky woodlands of Plateau Crest, 2721
Heteropyxis Harv.
natalensis Harv. , widespread medium-size tree, mainly localized
in forests, thickets and woodlands of Low Country, 608
MELASTOMATACEAE
Dissotis Benth.
phaeotricha (Hochst.) Hook. f. var. phaeotricha, large forb,
infrequent in Transitional Mistbelt grasslands, 1967
HALORAGACEAE
Gunnera L.
perpensa L., large perennial rhizomatous herb, abundant in
marshy grasslands of Summit Plateau, 2576
ARALIACEAE
Schefflera J.R. & G. Forst.
umbellifera (Sond.) Baill., localized canopy tree of Mistbelt
forests; Plateau Crest and Escarpment Slopes, 1009
Cussonia Thunb.
spicata Thunb., canopy and emergent tree, common and wide-
spread in forests, thickets and woodlands of Mistbelt
and Low Country, Scheepers 392
APIACEAE
Sanicula L.
elata Buch.-Ham., perennial herb of moist forest floor, Moun-
tain and Escarpment Slopes, 1011
Alepidea De la Roche
amatymbica Eckl. & Zeyh. var. amatymbica, perennial herb,
infrequent on Summit stream banks, 2739
basinuda Pott var. basinuda, perennial herb, rare in Plateau
Crest and Summit Plateau grasslands, 1203
gracilis Duemmer var. major Weim. , common perennial herb,
localized in grasslands and woodlands of Humid Mist-
belt, 1492
longifolia E. Mey. subsp. longifolia, perennial herb confined to
rocky woodlands, Summit, 2298
sp., perennial herb in grasslands of Summit Slopes and Peaks,
2407
Heteromorpha Cham. & Schlechtd.
pubescens Burtt Davy, widespread infrequent shrub of wood-
lands, Mistbelt and Low Country, 1808
transvaalensis Schltr. & Wolff, infrequent subwoody forb in
burnt woodlands, Escarpment Upper Slopes, 1645
trifolia ta (Wendl.) Eckl. & Zeyh., small infrequent tree of wood-
lands and forests, Humid Mistbelt, 991
Pimpinella L.
transvaalensis Wolff, infrequent perennial herb of rocky wood-
lands, Lower Mountain Slopes, 1514A
Sium L.
repandum Welw. ex Hiem, robust aquatic herb in riparian
woodlands, Summit Plateau, 2127
Annesorrhiza Cham. & Schlechtd.
flagellifolia Burtt Davy, perennial herb, infrequent in rocky
shrublands, Summit Plateau, 2465
Peucedanum L.
capense (Thunb.) Sond. var. capense, shrubby forb, infrequent
in disturbed thickets of Mountain Slopes, 1604
magalismontanum Sond., infrequent perennial herb, widespread
in grasslands and woodlands of Mistbelt and Low Coun-
try, 1087
sp., localized perennial herb of Summit Peak grasslands, 2408,
2472
ERICACEAE
Vaccinium L.
exul H. Bol., small shrub, frequent amongst rocks. Summit,
2306
Erica L.
atherstonei Diels ex Guth. & Bol, small shrub, common in
rocky shrublands, Summit Peaks and Slopes, 2613
caffrorum H. Bol. var. caffrorum, localized shrub of rocky
knolls. Summit Slopes and Peaks, 2304
cerinthoides L., localized shrublet of exposed stoney slopes,
Summit, Weisser 9037
drakensbergensis Guth. & Bol, localized shrub, scattered in
grasslands, shrublands and woodlands of Mistbelt, 1584
sp., small shrub of Low Open (grassy) shrubland, Summit Slopes,
2362
woodii H. Bol., infrequent shrub of Mountain Slopes and Sum-
mit grasslands, 1480
MYRSINACEAE
Maesa Forssk.
lanceolata Eorssk. var. rufescens (A. DC.) Taton, small tree,
common in understorey (or as pioneer) of Forest and
Mesic Thicket of Mistbelt and Low Country, 923
Bothalia 19,1 (1989)
105
Myrsine L.
africana L., small shrub, common in rocky shrublands, wood-
lands and forests of Mistbelt, 1193
Rapanea Aubl.
melanophloeos (L.) Mez , medium-size understorey tree of (main-
ly) Mistbelt forests, thickets and woodlands, 965
SAPOTACEAE
Bequaertiodendron De Wild.
magalismontanum (Sond.) Heine & J.H. Hemsl., large tree-like
shrub common in forests, thickets and woodlands below
Escarpment Plateau, and especially localized in rocky
woodlands of Plateau Crest, 2015
Mimusops L.
zeyheri Sond., fairly large tree, apparently confined to moist
rocky thickets and forests of Low Country, 1 712
EBENACEAE
Euclea Murray
‘complex’, widespread small understorey tree, mainly in wood-
lands and thickets of Transitional Mistbelt and Low
Country:
crispa (Thunb.) Guerke subsp. crispa, 918
divinorum Hiern, 28
schimperi (A. DC.) Dandy var. schimperi, 633
natalensis A. DC., infrequent small tree of Low Country
forests, thickets and woodlands, 833
Diospyros L.
galpinii (Hiern) De Winter, low shrublet of Mistbelt grasslands
and open woodlands, 1415
lycioides Desf. subsp. sericea (Bernh.) De Winter, widespread
small tree, abundant in less-shaded forests, thickets,
woodlands and grasslands of Mistbelt and Low Country,
624
mespiliformis Hochst. ex A. DC., large localized tree, infrequent
in low Country woodlands and thickets, often associated
with diabase outcrops and termitaria, 1980C
whyteana (Hiern) F. White, frequent shrub, widespread (often
dominant) in forests, thickets and w oodlands of Mistbelt
and Low Country, also as pioneer in broken lithoscre,
612
OLEACEAE
Schrebera Roxb.
alata (Hochst.) Welw., fairly large canopy tree, infrequent in
forest of Escarpment Slopes, 1068
Chionanthus L.
foveolata (E. Mey .) Steam
subsp. foveolata, fairly large tree, infrequent in forests of
Mistbelt and Low Country, 1848
subsp. major (Verdoorn) Steam, large tree in Tall Forest,
Mountain Slopes, 2283
Olea L.
capensis L. subsp. macrocarpa (C.H. Wr.) Verdoorn, fairly large
canopy tree, common (often dominant) in forests of
Mountain Slopes, 1357
europaea L. subsp. africana (Mill.) P.S. Green, small tree, local-
ized in understorey of Short (cliff) Forest, Lower
Mountains, 1393
Jasminum L.
angulare Vahl, fairly frequent shrub of High Forest, Lower
Mountain Slopes, 2201
sp., twiner of Tall Forest, Escarpment Slopes, includes 962A
streptopus E. Mey., localized twiner of forests and thickets,
Lower Mountain and Escarpment Slopes, 916
LOGANIACEAE
Strychnos L.
madagascariensis Poir. , small tree, mainly localized in wood-
lands of Lower Foothills, 1 750
spinosa Lam., small tree, fairly common in Low Country wood-
lands and thickets; occasionally in broken lithosere,
Mistbelt, 1 751
Anthocleista A fzel. ex R. Br.
grandiflora Gilg, tall canopy and emergent tree, common in
forests and mesic thickets of Transitional Mistbelt and
Low Country, Scheepers 797
Nuxia Comm, ex Lam.
congesta R. Br. ex Fresen., fairly small tree, infrequent in
rocky woodlands of Plateau Crest, 1071
Buddlcja /..
auriculata Benth., localized shrub of moist (often riparian)
forests, Mountain and Summit Slopes, 1373
salviifolia (L.) Lam., occasional shrub of rocky woodlands and
moist (often riparian) forests, Mountain and Summit
Slopes, 206 7
GENTIANACEAE
Sebaea Soland. ex R. Br.
leiostyla Gilg, infrequent perennial forb of Mountain and Sum-
mit grasslands, 1472
APOCYNACEAE
Carissa L.
bispinosa (L.) Desf. ex Brenan var. acuminata (E. Mey.) Codd,
widespread and common shrub in undergrowth of
Forest and Mesic Thicket of Mistbelt and Low Country,
92
Rauvolfia L.
caffra Sond., large canopy tree, infrequent in moist forests of
Escarpment Slopes, 211
PERIPLOCACEAE
Cryptolepis R. Br.
oblongifolia Schltr., subwoody half-tw ining forb, common and
widespread in Woodland and Xeric Thicket of Low
Country, and fairly frequent in lithoseral grasslands and
woodlands of Humid Mistbelt, 1032
Raphionacine Harv.
data N.E. Br., localized forb of dolomite-soil grasslands, Plateau
Interior, 1343
hirsuta (E. Mey.) R.A. Dyer ex Phill. , w idespread forb in grass-
lands of Humid Mistbelt, 1169
asclepiadaceae
Xysmalobium R. Br.
aceratoides (Schltr.) N.E. Br., prostrate perennial forb, wide-
spread on Summit, 2384
confusum Scott Elliott, robust perennnial forb, rare in grass-
lands of Lower Mountain Slopes, 1464
Asclepias L.
crassinervis N.E. Br., succulent herb, scattered in woodlands
and grasslands of Mistbelt and Low Country, 1351
dregeana Schltr. , rare herb of Mistbelt grasslands, 1352
Pentarrhinum E. Mey.
insipidum E. Mey. , vigorous climber, infrequent in xeric thickets
of Low Country, 863
Cynanchum L.
ellipticum (Harv.) R.A. Dyer, rare twiner of Low Country xeric
thickets, 1890
Sarcostemma R. Br.
viminale (L.) R. Br .leafless succulent twiner of sheet-rock shrub-
lands, Transitional Mistbelt, 1955
Secamone R. Br.
alpinii Schultes, vigorous slender liane, infrequent in forests
and thickets of Escarpment and Mountain Lower
Slopes, 91 7
gerrardii Harv. ex Benth. , slender climber, widespread and com-
mon in Forest and Mesic Thicket of Mistbelt and Low
Country, 697
parvifolia ( Olbv .) Bullock, localized twiner of Transitional Mist-
belt thickets, 931
Ceropegia L.
meyeri Decne., infrequent soft twiner of Plateau Crest wood-
lands, 1444
racemosa N.E. Br. subsp. setifera (Schltr.) Huber, a twiner of
Transitional Mistbelt mesic thickets, 995
sp., rare twiner of Low Country xeric thickets, 1875
woodii Schltr., infrequent soft twiner, widespread in forests of
Low Country and in rocky woodlands of Mistbelt, 1457
Tylophora R. Br.
anomala N.E. Br., widespread climber or creeper associated
with Low Country lithoseral woodlands and apparently
persisting in thickets and forests, 648
flanaganii Schltr., slender twining liane, occasional in Forests
of Mountain Slopes, 1569
Pcrgularia L.
daemia (Forssk.) Chiov. var. daemia, infrequent creeper ot
disturbed thickets, Upper Foothills, 1810
CONVOLVULACEAE
Cuscuta L.
sp., twining parasitic forb. localized in Woodland and Xeric
Thicket of Low Country
106
Ipomoea L.
bathycolpos Hallier f vai. bathycolpos, infrequent creeper of
grasslands and rocky woodlands, Escarpment Plateau,
1178
crassipes Hook., infrequent perennial forb, localized an wood-
lands and thickets of Upper Foothills, 640
sp., scrambler or twiner in Low Country woodlands, includes
1745 , 1904
BORAGINACEAE
Ehretia P. Br.
amoena Klotzsch, infrequent shrub of Low Country xeric
woodlands, 748A
VERBENACEAE
Lantana L.
*camara /.., fairly common naturalized shrub, especially in
Low Country w'oodlands, 949
mearnsii Moldenke var. latibracteolata Moldenke, small soft
shrub, infrequent in mesic thickets of Transitional Mist-
belt, 1072
Lippia L.
javanica (Bunn, f.) Spreng. , widespread small shrub (possibly a
forest prescursor), especially Mistbelt grasslands, 734
wilmsii H. Pearson , infrequent forb of disturbance, localized in
Upper Foothill woodlands and thickets, 742
Clerodendrum L.
glabrum E. Mey. var. glabrum, widespread small tree, infrequent
in woodlands and forests of Mistbelt and Low Country,
1442
myricoides (Hochst.) Vatke, canopy tree in forests of Escarp-
ment Slopes or understorey shrub in woodlands, thickets
and forests of Foothills, 1137
sp., infrequent shrub or forb in woodlands and thickets of
Foothills, includes 1723. 1819
suffruticosum Guerke var. suffruticosum, infrequent shrub
associated with sheet-rock formations, Transitional
Mistbelt, 1050
triphyllum (Harv.) H. Pearson var, triphyllum, subwoody peren-
nial forb, infrequent in Summit Plateau grasslands, 2537
LAMIACEAE
Acrotome Benth.
hispida Benth., infrequent forb, localized mainly in dolomite-
soil grasslands of Plateau Interior, 1307
Leonotis (Pers.) R. Br.
‘complex’, soft shrubby perennial, widespread in thickets,
woodlands and grasslands of Mistbelt and Low Country:
ocymifolia (Burm. f. ) Iwarsson
var. ocymifolia, 2305
var. raineriana (Visiani) Iwarsson, 739
sp., rare herb of Transitional Mistbelt woodlands, 1053
Stachys L.
grandifolia E. Mey. ex Benth., soft undershrub, infrequent in
moist shady thickets of Upper Foothills, 699
natalensis Hochst. var. galpinii (Briq.) Codd, infrequent forb of
rocky woodlands. Lower Mountain Slopes, 1498
nigricans Benth., slender forb, localized in grasslands of Escarp-
ment Plateau and Slopes, 1259
Tetradenia Benth.
‘complex’, small soft shrub, widespread as chasmophyte in early
lithosere and persisting in lithoseral woodlands and
thickets from Foothills to Plateau Crest:
brevispicata (N.E. Br.) Codd, 777
riparia (Hochst.) Codd, 397
Aeollanthus Mart, ex K. Spreng.
rehmannii Guerke, shrubby herb, localized in rocky woodlands
of Escarpment Slopes and Plateau Crest, 1408
Endostemon N.E. Br.
obtusifoliusYE". Mey. ex Benth.) N.E. Br., infrequent soft shrub
of mesic woodlands and thickets, Transitional Mistbelt,
929
Pycnostachys Hook.
reticulata (E. Mey.) Benth., tall shrubby forb of retarded grass-
lands, Summit Plateau, 2726
urticifolia Hook. , soft shrub, w idespread in lithoseral woodlands
and thickets of Upper Foothills and Escarpment Lower
Slopes, 614B
Plectranthus L 'Herit.
ciliatus E. Mey. ex Benth., infrequent shrub of High Forest,
Mountain Slopes, 2208
Bothalia 19,1 (1989)
fruticosus L’Herit., soft undershrub of rocky forests and wood-
lands, Plateau Crest, 1439
grandidentatus Guerke, shaded understorey shrub, infrequent in
lithoseral woodlands and forests of Plateau Crest and
Mountain Slopes, 1560
hadiensis (Forssk.) Schweinf. ex Spreng. var. hadiensis, localized
shrub (chasmophyte) of Plateau Crest open woodlands,
1425
laxiflorus Benth., infrequent soft undershrub of Transitional
Mistbelt forests, 1958
rubropunctatus Codd, soft undershrub, sometimes forming pure
stands in shrub layer of rocky forests and riparian wood-
lands of Escarpment and Summit Plateau, occasionally a
pioneer in Mistbelt sheet-rock formations, 1508
sp., w idespread shrub of forests, thickets, woodlands and grass-
lands, includes 834, 1379
spicatus E. Mey. ex Benth., succulent herb in soil pockets of
sheet-rock formations, Low Country, 1880
verticillatus (L. f.) Druce, usually a lax succulent herb covering
rocks in shade of Low Country forests, 1 714
Rabdosiella Codd
calycina (Benth.) Codd, subwoody perennial forb, widespread
in fire-protected shrublands and woodlands. Summit,
2318
Hoslundia Vahl
opposita Vahl, tall shrubby forb, rare in Low Country wood-
lands, 1869
Hemizygia (Benth.) Briq.
albiflora (N.E. Br.) Ashby, low bush of rocky shrublands and
grasslands, especially windy sites with shallow soil, Sum-
mit Slopes and Peaks, 2405
canescens (Guerke) Ashby, variable forb of lithoseral wood-
lands, Plateau Crest and Foothills, 1764
subvelutina (Guerke) Ashby, localized forb, common in grass-
lands of Escarpment Plateau and Slopes, 533
transvaalensis (Schltr.) Ashby, widespread forb of woodlands
and grasslands, Mistbelt and Low Country, 1290
Ocimum L.
urticifolium Roth., infrequent forb of xeric woodlands and
thickets, Low Country, 791
Becium Lindl.
obovatum (E. Mey. ex Benth.) N.E. Br. var. obovatum, local-
ized herb, common in grasslands of Escarpment Plateau
and Slopes, 1083
SOLANACEAE
Solanum L.
*mauritianum Scop., naturalized small tree, common as a weed
of timber plantations and often encroaching on indige-
nous woodlands and forests, especially Mistbelt
sp. cf. aculeastrum Dun., small shrub of sheet-rock formations
and disturbed forest openings, Low Country
terminale Forssk. subsp. terminale, subwoody climber of Tall
(kloof) Forest, Upper Mountain Slopes, 2153
SCROPHULAR1ACEAE (Part A)
Nemesia Vent.
rupicola ined., infrequent herb of riparian woodlands, Summit
Plateau, 2706
Diclis Benth.
reptans Benth. , small herb of riparian forests and woodlands.
Summit Plateau and Slopes, 2121
Halleria L.
lucida L., small tree, widespread from Low Country forests to
Mistbelt forests, thickets and woodlands, 1404B
Phygelius E. Mey. ex Benth.
aequalis Harv. ex Hiern, waterside shrub of riparian woodlands,
Summit Plateau, 2130
Bowkeria Harv.
cymosa Macowan, small tree, localized in rocky Mountain forests
and woodlands, 1513A
Sutera Roth
grandiflora (Galpin) Hiern, infrequent forb of Low Country
woodlands, 855
Zaluzianskya F. W. Schmidt
spathacea (Benth.) Walp. , infrequent forb of moist grasslands.
Summit Plateau, 2689
Ilysanthes Rafin.
wilmsii EngL ex Diels, semi-aquatic herb in shallow pools of
sheet-rock formations, Low Country, 1916
Bothalia 19,1 (1989)
107
SELAGINACEAE
Hebenstretia L.
comosa Hochst. , occasional herb in Summit grasslands, 2377
oatesii Rolfe, large herb, locally abundant in riparian woodlands,
Summit Plateau, 2437
Selago L.
atherstonei Rolfe, infrequent subwoody forb of lithoseral grass-
lands and woodlands, Mist belt, 536
elata Rolfe, rare shrubby forb in grasslands of Transitional Mist-
belt, 1964
hyssopifolia E. Mey., infrequent subwoody forb, localized in
lithoseral woodlands of Plateau Crest, 1424
lydenburgensis Rolfe, subwoody forb, infrequent in grasslands
of Summit Slopes, 2688
muddii Rolfe, subwoody forb, localized mainly in Plateau Crest
grasslands, 1098
sp., infrequent subwoody forb of Protea woodland, Summit
Slopes, 241 7
Tetraselago Junell
natalensis (Rolfe) Junell, subwoody forb, localized in lithoseral
grasslands and woodlands of Plateau Crest and Escarp-
ment Slopes, 1111
wilmsii (Rolfe) Hilliard & Burn, widespread subwoody forb of
shallow rocky soils, Summit grasslands, shrublands and
woodlands, 2301
SCROPHULARIACEAE (Part B)
Alectra Thunb.
sessiliflora (Vahl) Kuntze var. sessiliflora, apparently rare para-
sitic herb in grasslands of Mountain Slopes, 1486
Sopubia Buch.-Ham. ex D. Don
cana Harv. var. cana, frequent forb in grasslands of Summit and
Mountain Slopes, 1470
simplex (Hochst.) Hochst., infrequent forb of moist grasslands,
Summit Plateau, 2568
Buchnera L.
dura Benth., infrequent forb in grasslands of Lower Mountain
Slopes, 1491
longispicata Schinz, rare forb of Low Country woodlands, 1 763
Cycnium E. Mey. ex Benth.
racemosum Benth., subwoody forb, common on shallow rocky
soils, Summit grasslands, shrublands and woodlands,
2378
tubulosum (L. f.) Engl, small rare herb of disturbed grasslands.
Plateau Crest, 1293
Striga Lour.
bilabiata (Thunb.) Kuntze, small hemi-parasitic herb, rare in
grasslands of Escarpment Slopes, 1346
BIGNONIACEAE
Tecomaria Spach.
capensis (Thunb.) Spach., small scandent shrub or tree, infre-
quent in shrub layer and understorey of lithoseral wood-
lands, thickets and forests, Mistbelt and Low Country,
1404
J acaranda Juss.
*mimosifolia D. Don. , occasional exotic tree, naturalized in
mesic thickets of Low Country, 937
PEDALIACEAE
Ceratotheca Endl
triloba (Bemh.) Hook, f, tall shrubby forb, common on sheet-
rock formations and early lithoseral grasslands and
woodlands, Mistbelt and Low Country, 1282
gesneriaceae
Streptocarpus Lindl
cyaneus S. Moore, perennial herb, apparently localized in Short
(cliff) Forest, Lower Mountain Slopes, 1384
dunnii Hook, f, localized herb of rocky open shrublands and
woodlands, Mistbelt, 1423A
pentherianus Fritsch, epiphyte on trees and rocks of Tall (kloof)
Forest, Upper Mountain and Summit Slopes, 2160
polyanthus Hook, subsp. dracomontanus Hilliard, infrequent
herb of Escarpment Slopes forests, 1062B
ACANTHACEAE
Thunbergia Retz.
atriplicifolia E. Mey. ex Nees, infrequent localized twiner in
w oodlands of Upper Foothills, 643
neglecta Sond., infrequent twiner of mesic thickets, Upper
Foothills, 695
sp., herb in forests of Escarpment Lower Slopes, includes 986
Phaulopsis Willd.
imbricata (Forssk.) Sweet, small undershrub, common in forests,
thickets and woodlands of Transitional Mistbelt and
Low Country, 600
Dyschoriste Nees
depressa (L.) Nees, infrequent forb in shrubby grassland of
old-land successions, Upper Foothills, 725
Chaetacanthus Nees
burchellii Nees, shrubby forb in field layer of thickets and wood-
lands, mainly Low'er Foothills, 1737
Ruellia L.
sp., rare forb of closed woodlands, Lower Foothills, 1742
Crabbea Harv.
hirsuta Harv., low subwoody forb, scattered in Mistbelt grass-
lands and Low Country woodlands, 748
Barleria L.
gueinzii Sond., rare shrubby forb in rocky forests. Lower Foot-
hills, 1711
ovata E. Mey. ex Nees, widespread forb of Mistbelt grasslands
and woodlands, 1523
Sclerochiton Harv.
harveyanus Nees, undershrub, often dominant and abundant in
forests of Humid Mistbelt, 1062 A
Diclip tera Juss.
clinopodia Nees, undershrub, widespread in Forest and Mesic
Thicket of Mistbelt and Low Country, 686
Hypoestes Soland. ex R. Br.
aristata R. Br. , localized undershrub of closed w oodlands and
xeric thickets. Low Country, 740
triflora (Forssk.) Roem. & Schult., localized undershrub of Tall
(riparian) Forest, Mountain Slopes, 1570
Isoglossa Oerst.
eckloniana (Nees) Lindau, lax subwoody forb, rare in rocky
mesic thickets of Upper Foothills, 876
PLANTAGINACEAE
Plantago L.
*major L ., waterside herb of Tall (riparian) Forest, Low' Coun-
try, 901
RUBIACEAE
Kohautia Cham. & Schlechtd.
amatymbica Eckl. & Zeyh., small forb, infrequent in Mistbelt
grasslands, 1209
Conostomium Cuf.
natalense (Hochst.) Brem. var. glabrum Brem., infrequent forb,
scattered in rocky woodlands of Mistbelt and grasslands
of Low Country, 1516
Agathisanthemum Klotzsch
bojeri Klotzsch var. bojeri, perennial herb, infrequent in grass-
lands and woodlands of old-land successions, Low
Country, 726
Pentas Benth.
sp., rare forb of disturbed woodlands. Upper Foothills, 743
Breonadia Ridsdale
salicina (Vahl) Hepper& Wood, large waterside tree, conspicuous
in Tall (riparian) Forest of Low Country, Scheepers 606
Cephalanthus L.
natalensis Oliv. , woody liane and scrambler, occasional as chas-
mophyte in Mistbelt sheet-rock formations, persisting in
rocky woodlands of Plateau Crest, and frequent in forests
and mesic thickets of Mistbelt, 706
Burchellia R. Br.
bubalina (L. f.) Sims, occasional small tree of broken lithoseral
woodlands, Mistbelt, 1494
Coddia Verde.
rudis (E. Mey. ex Harv.) Verde. , occasional undershrub in xeric
thickets of Low Country, 1 783 A
Rothmannia Thunb.
capensis Thunb., canopy and understorcy tree of elfin-like
forests, Plateau Crest and Mountain Slopes, 1199
globosa (Hochst.) Keay , infrequent shrub or small tree in under-
storey of mesic thickets, Transitional Mistbelt, 886
Hyperacanthus E. Mey. ex Bridson
amoenus (Sims) Bridson, shrub or small tree, fairly frequent in
forests and thickets of Low Country, 885
Oxyanthus DC.
108
speciosus DC, subsp. gerrardii ( Sond .) Bridson, large under-
storey tree, frequent in forests and mesic thickets of
Escarpment and Mountain Slopes, 1014
Tricalysia A. Rich.
‘complex’, small understorey tree, common in Forest and Mesic
Thicket of Mistbelt and Low Country, occasional in
rocky woodlands of Plateau Crest:
capensis (Meisn.) Sim, 1136
lanceolata (Sond.) Burn Davy, 1397
Kraussia Harv.
floribunda Harv., localized undershrub of Low Country forests
and sheltered thickets, 1721
Pentanisia Harv.
angustifolia (Hochst.) Hochst., perennial herb, infrequent in
Mountain Slopes grasslands, 1473
prunelloides (Eckl. & Zeyh.) Walp., low perennial herb, wide-
spread and abundant in grasslands of Mistbelt, 6 76
Vangueria Juss.
infausta Burch, subsp. infausta, small tree of Mistbelt and Low
Country lithosere, persisting in woodlands and thickets,
790
Canthium Lam.
cibatum (Klotzsch) Kuntze, small understorey tree, localized in
Tall forests of Escarpment and Mountain Slopes, 1057
gilfillanu (N.E. Br.) O.B. Miller, shrub or small tree, rare in Low
Country woodlands, 829
gueinzii Sond., woody bane, common in forests and mesic
thickets below Escarpment Plateau, 889
inerme (L. f.) Kuntze, shrub or small tree, widespread in forests,
thickets and woodlands of Mistbelt and Low Country,
613
mundianum Cham. & Schlechtd., small tree, scattered in Foot-
hill woodlands and xeric thickets and in Plateau Crest
woodlands, 1407
pauciflorum ( Klotzsch ) Kuntze, understorey tree in High
Forest, Upper Mountain Slopes, 2148
sp., infrequent shrub of Low Country woodlands, 841
Psydrax Gaertn.
bvida (Hiern) Bridson, shrub or smab tree, locabzed in Short
(chff) Forest of Lower Mountain Slopes, 1371
locuples (K. Schum.) Bridson, rare shrub or smaU tree in under-
storey of Mistbelt forests, 1224
obovata (EckL & Zeyh.) Bridson subsp. obovata, fairly large
understorey tree, infrequent in tab forests of Mountain
and Escarpment Slopes, 1168
Pachystigma Hochst.
macrocalyx (Sond.) Robyns, infrequent shrub of rocky wood-
lands, Humid Mistbelt, 1190
sp., infrequent shrub of early lithoseral woodlands, Lower
Foothills, includes 1884
Fadogia Schweinf.
homblei De Wild, infrequent forb in broken bthoseral grass-
lands, Plateau Crest, 1419
tetraquetra Krause, widepsread forb, common in grasslands and
woodlands of Mistbelt, 1448
Pavetta L.
cooperi Harv. & Sond., shrub or smab tree, infrequent in Mist-
belt forests, 1956
galpinu Brem., occasional shrub in forests and mesic thickets of
Escarpment and Mountain Slopes, 997
gardenufoba A. Rich. var. gardenhfolia, infrequent shrub, chas-
mophyte in broken bthoseral woodlands of Transitional
Mistbelt, 1632
gardeniifoba A. Rich. var. subtomentosa K. Schum., infrequent
shrub, chasmophyte on sheet-rock formations, Low
Country, 806.
schumanniana F. Hoffm. ex K. Schum., fairly frequent shrub
or small tree, localized in partially sheltered woodlands
and xeric thickets of Lower Foothills, 784
sp., variable shrub of forests and thickets. Low Country, includes
1847
Psychotria L.
capensis (Eckl.) Vatke, smab tree, widespread and abundant in
Forest and Mesic Thicket of Mistbelt and Low Country
and in Humid Mistbelt woodlands, 971
zombamontana (Kuntze) Petit , smab understorey tree, frequent
in Mistbelt forests, 2053
Galopina Thunb.
aspcra (Eckl. & Zeyh.) Walp., infrequent forb, scattered in grass-
lands and woodlands of Humid Mistbelt, 1540
circaeoides Thunb., widespread perennial herb, common in un-
Bothalia 19,1 (1989)
dergrowth of forests, thickets and woodlands, Mistbelt
and Low Country, 691
Anthospermum L.
herbaceum L. f., herbaceous bushy perennial, scattered in Sum-
mit grasslands and woodlands, 2329
welwitschii Hiern, ericoid shrub, occasional in woodlands of
Plateau Crest and Escarpment Slopes, 1627
Richardia L.
*brasbiensis Gomes, naturabzed herb of old-land disturbance,
Upper Foothibs, 626
Spermacoce Gaertn.
natalensis Hochst. , rare forb, chasmophyte in broken bthoseral
woodlands of Plateau Crest, 1454
Rubia L.
cordifoba L. subsp. conotricha (Gand.) Verde., herbaceous
perennial creeper, infrequent in xeric thickets of Low
Country, 851
petiolaris DC. , herbaceous perennial cbmber, rare in Low Coun-
try woodlands, 1805
valerianaceae
Valeriana L.
capensis Thunb., locabzed perennial forb in association with
Andropogon appendiculatus vleis, Summit, 2582
DIPSACACEAE
Cephalaria Schrad.
pungens Szabo, fairly widespread herb, locabzed and common,
mainly in grasslands of Humid Mistbelt, 1471
Scabiosa L.
columbaria L., infrequent perennial herb of grasslands. Summit
and Escarpment Plateau, 747
CUCURBITACEAE
Momordica L.
balsamina L. , slender twiner, infrequent in sheltered xeric
thickets of Low Country, 865
boivinh Bail l, infrequent soft twiner, locabzed in rocky (dia-
base) woodlands of Low Country, 1697A
Cucumis L.
sp., occasional creeper in Low Country rocky thickets, 1368
zeyheri Sond., infrequent creeper in dolomite-sob grasslands,
Plateau Interior, 1368
Trochomeria Hook. f.
sagittata (Harv. ex Sond.) Cogn., infrequent slender twiner of
rank grasslands, Upper Foothbls, 1935
Coccinia Wight <& Am.
palmata (Sond.) Cogn., infrequent herbaceous climber, local-
ized in mesic thickets of Transitional Mistbelt, 892
CAMPANULACEAE
Wahlenbergia (Schrad. ex Roth)
huttonii (Sond.) Thulin, infrequent perennial herb of Mistbelt
grasslands, 1308
lycopodioides Schltr. & V. Brehm., infrequent herb of btho-
seral grasslands, Plateau Crest, 1636
undulata (L. f.) A. DC., subwoody herb, infrequent in btho-
seral grasslands of Humid Mistbelt, 1550
virgata Engl., subwoody herb, widespread in grasslands of
Plateau Crest and Summit, 1175
Craterocapsa Hilliard & Burn
tarsodes Hilliard & Burtt, widespread low perennial herb in
grasslands of Summit Slopes and Plateau, 2515
LOBELIACEAE
Cyphia Berg.
elata Harv.
‘complex’, occasional forb, scattered in Mistbelt grasslands and
woodlands:
var. elata Harv., 441
var. glabra Harv., 1416
stenopetala Diels, rare slender herb of retarded grasslands, Sum-
mit Plateau, 2532
Lobelia 1 .
erinus /.. . rare herb of Summit Plateau grasslands, 2558
flaccida ( Presl ) A. DC. subsp. flaccida, small herb of disturbed
grasslands. Summit Peak. 2720
Monopsis Salisb.
decipiens (Sond.) Thulin, small infrequent herb in moist grass-
lands of Escarpment Slopes and Plateau Crest, 1092
Bothalia 19,1 (1989)
109
ASTERACEAE
Vernonia Schreb.
adoensis Sch. Bip. ex Walp., small infrequent shrub, widely
scattered in Mistbelt grasslands and thickets, 942
amygdalina Del, large herb, infrequent in field layer of mesic
thickets, Upper Foothills, 703
centaureoides Klatt, infrequent forb in rank grasslands of
Upper Foothills, 1930
colorata (Willd.) Drake , small shrub, localized in rocky (dia-
base) woodlands of Lower Foothills, 1 700
hirsuta (DC.) Sch. Bip., subwoody forb, widespread in Mistbelt
grasslands and shrublands, 1477
natalensis Sch. Bip., widespread subwoody forb of grasslands
and woodlands, Mistbelt and Low Country, 1474
neocorymbosa Hilliard, fairly common subwoody forb, local-
ized in woodlands of Escarpment Slopes and Upper
Foothills, 1624B
oligocephala (DC.) Sch. Bip. ex Walp., shrubby forb, scattered
in woodlands and thickets of Mistbelt and Low Country,
815
poskeana Vatke & Hildebr. subsp. botswanica Pope, widespread
forb of lithoseral woodlands and grasslands, Mistbelt
and Low Country, 804
stipulacea Klatt, widespread robust shrub of forests, thickets
and woodlands, mainly Low Country, 1405
thodei Phill. , occasional forb of retarded grasslands. Summit
Plateau, 2353
wollastonii 5. Moore, occasional undershrub in forests of
Plateau Crest and Mountain Slopes, 1195
Ageratum L.
*conyzoides L., forb of disturbed woodlands. Low Country,
864
Stomatanthes R.M. King & H. Robinson
africanus (Oliv. & Hiern) R.M. King & H. Robinson, subwoody
forb of woodlands and grasslands. Escarpment Slopes,
1647
Mikania Willd.
natalensis DC., twining liane of forests and mesic thickets,
especially below Escarpment Plateau, 898
Aster L.
comptonii Lippert, infrequent subwoody forb of Mistbelt
grasslands and woodlands, 1594
harveyanus Kuntze, rare forb of Foothill woodlands, 768
lydenburgensis Lippert, rare forb of Mistbelt grasslands, 1338
sp., forb of disturbed woodlands, Foothills, includes 649, 750
Felicia Cass.
rosulata Yeo, rare forb in rocky shrublands of Summit Peak,
2503
Nidorella Cass.
auriculata DC., robust herbaceous bush in moist (often marshy)
grasslands and thickets, Mistbelt, 605
sp., infrequent forb of disturbed moist grasslands, Mistbelt,
includes 1249, 2322
Conyza Less.
*sumatrensis (Retz.) E.H. Walker, widespread introduced weed
of disturbance, Mistbelt and Low Country, 609
Brachylaena R. Br.
discolor DC. subsp. transvaalensis (Phill. & Schweick.) J. Paiva,
fairly large canopy and emergent tree in Forest and
Mesic Thicket of Mistbelt and Low Country, 679
Tarchonanthus L.
trilobus DC. var. galpinii (Hutch. & Phill) J. Paiva, localized
tree of rocky woodlands. Plateau Crest, 1433
Blumea DC.
alata (D. Don) DC., widespread herbaceous weed of disturbed
woodlands and thickets, Low Country, 606
Gnaphalium L.
sp., subwoody herb of retarded grasslands. Summit Plateau,
2327
Tenrhynea Hilliard & Burtt
phylicifolia (DC.) Hilliard & Burtt, w idespread shrubby forb of
woodlands and grasslands, Mistbelt and Low' Country,
766
Helichrysum Mill.
acutatum DC., common subwoody forb of Summit woodlands,
shrublands and grasslands, 235 7
adenocarpum DC., infrequent forb of broken lithoseral grass-
lands, Plateau Crest, 11 79
aureonitens Sch. Bip., small herb, common in grasslands of
Mistbelt, 1100
aureum (Houtt.) Merr. var. monocephalum (DC.) Hilliard, small
subwoody forb, infrequent in Mistbelt grasslands, 1242
cephaloideum DC., localized herb of Plateau Crest grasslands,
1205
chionosphaerum DC., infrequent forb in Low Open Shrubland
on exposed stoney slopes, Summit, Weisser 9036
chrysargyrum Moeser, widespread, infrequent bushy forb of
serai woodlands, Mistbelt and Low Country, 6 73
‘complex’, common subwoody forb, widespread in grasslands
and woodlands of Humid Mistbelt:
acutatum DC, 1099
thapsus (Kuntze) Moeser, 1520
cooperi Harv., localized forb of Plateau Crest woodlands, 1413
coriaceum Harv., subwoody herb, infrequent in Mistbelt grass-
lands, 1656
difficile Hilliard, large infrequent forb bordering marshy grass-
lands, Summit Plateau, 2737
edwardsii Wild, large forb, infrequent in Summit grasslands,
2482
glomeratum Klatt, infrequent forb of Summit Plateau grass-
lands, 2544
harveyanum Wild, infrequent forb. localized in dolomite-soil
grasslands of Plateau Interior, 1303
homilochrysum S. Moore, shrubby bush in sheltered ravines,
Summit Slopes, 2703
kraussii Sch. Bip., shrubby ericoid forb, localized in lithoseral
woodlands of Escarpment Slopes and Plateau Crest, 975
mariepscopicum Hilliard, infrequent forb of Summit grasslands,
2680
mimetes S. Moore, lax forb, widespread in lithoseral grasslands
and woodlands of Mistbelt, 1607
mixtum (Kuntze) Moeser, infrequent forb of woodlands and
grasslands, Mistbelt, 1525
nudifolium (L.) Less., widespread subwoody forb of thickets,
woodlands and grasslands, Mistbelt and Low Country,
618
odoratissimum (L.) Sweet, widespread bushy forb of open
woodlands and grasslands, especially Mistbelt, 1932
opacum Klatt, rare forb of Summit grasslands, 2692
pallidum DC., subwoody herb, infrequent in lithoseral wood-
lands of Humid Mistbelt, 1558
panduratum O. Hoffm. var. transvaalense Moeser, widespread
forb of mesic thickets, woodlands and grasslands,
especially Mistbelt, 712
pilosellum (L. f.) Less., subwoody herb, common in Mistbelt
grasslands, 1164
platypterum DC., robust perennial forb, occasional in Mistbelt
grasslands, 1585
reflexum N.E. Br., infrequent shrubby forb, localized in broken
lithoseral woodlands of Low'er Mountain Slopes, 1497
setosum Harv., rare forb of riparian woodlands, Summit
Plateau, 2126
sp., variable forb, widespread in Mistbelt grasslands and wood-
lands and in Low Country woodlands and thickets, in-
cludes 1157, 1253, 2299
sp. nov. 1 (Werdermann & Oberdieck 2179), infrequent sub-
woody herb, widely scattered in Mistbelt grasslands,
especially in broken lithosere of Lower Mountain Slopes,
1493
sp. nov. 2 (Scheepers s.n.), infrequent subwoody forb of dis-
turbed seepage thickets, Lower Mountain Slopes, 1590
sp. nov. 3 (Van der Schijff 4335A), localized forb, rare in
marshy grasslands of Escarpment Plateau, 1263
spiralepis Hilliard & Burtt, large forb, fairly common in grass-
lands of Summit Plateau, 2536
splendidum (Thunb.) Less., low dense bush invading moist
fire-protected sites on Summit, 2641
subluteum Burtt Davy, small forb, fairly common in grasslands
of Summit Slopes and Plateau, 2393
truncatum Burtt Davy, small tufted forb, infrequent in grass-
lands of Summit Plateau, 2562
umbraculigerum Less., lax soft subwoody forb, infrequent in
woodlands and grasslands of Humid Mistbelt, 1154
wilmsii Moeser, low shrubby forb, widespread in Mistbelt grass-
lands and woodlands, 1106
Stoebe L.
vulgaris Levyns, scattered ericoid shrub of Summit Plateau and
Slopes, 2390
Athrixia Ker-Gawl.
phylicoides DC., widespread shrubby forb, localized mainly in
110
Bothalia 19,1 (1989)
thickets and woodlands of Transitional Mistbclt and
Low Country, 645
Inula L.
glomerata Oliv. & Hiern, robust herb, localized mainly in more
exposed woodlands and xeric thickets of Low Country,
635
Geigeria Griessslich
burkei Harv. subsp. burkei var. elata Merxm., infrequent forb,
widely scattered in Low Country woodlands, 842
burkei Harv. subsp. burkei var. hirtella Merxm.. infrequent forb
of rank grasslands. Mountain Slopes, 1515
Anisopappus Hook. <& Am.
latifolius [S. Moore) B.L. Bum. trailing forb, infrequent in
Summit grasslands, 2554
Acanthospermum Schrank
*australe (Loefl.) Kuntze, lax forb of disturbed grasslands and
woodlands, especially from Plateau Crest to Upper
Foothills, 596
Bidens L.
kirkii (Oliv. & Hiern) Sherff, infrequent forb of lithoseral w ood-
lands, Plateau Crest, 1640
*pilosa L.. widespread forb, characteristic weed of disturbance,
Mistbelt and Low Country
Inulanthera Kallersjd
calva (Hutch.) Kallersjd, herbaceous bush, localized in grass-
lands and rocky woodlands of Plateau Crest and Lower
Mountain Slopes, 1976
Phymaspermum Less, emend. Kallersjd
acerosum (DC.) Kallersjd, herbaceous bush, localized in grass-
lands and rocky w'oodlands of Kscarpment and Summit
(Plateau and Slopes), 1111
Inezia Phill.
integrifolia (Klatt) Phill, perennial herb, widespread in grass-
lands and rocky woodlands of Mountain Slopes and
Summit, 1462
Schistostephium Less.
crataegifolium (DC.) Fenzl ex Harv. , subwoody forb, infrequent
in grasslands of Escarpment and Low er Mountain Slopes,
1530
heptalobum (DC.) Oliv. & Hiern , subwoody perennial forb,
widely scattered in undergrow'th of forests and mesic
thickets of Mistbelt, 928
rotundifolium (DC.) Fenzl ex Harv., infrequent shrub of Closed
Woodland on rocky knolls, Summit Slopes, 2294
Artemisia L.
afra Jacq. ex Willd., shrubby herb, localized in thickets and
woodlands of Mountain Slopes, 1559
Lopholaena DC.
disticha (N.E. Br.) S. Moore , subsucculent herb, infrequent in
grasslands of Escarpment Slopes, 1947
segmentata (Oliv.) S. Moore, perennial herb, infrequent in
grasslands of Escarpment Plateau and Lower Mountain
Slopes, 1532
Cineraria L.
sp., field layer herb of moist bushy ravines. Summit Slopes,
2679
Senecio L.
affinis DC., infrequent subwoody forb of Mistbelt grasslands,
1546
conrathii N.E. Br., infrequent subwoody forb of Mountain
Slope grasslands, 1616
coronatus (Thunb.) Harv., low forb, widely scattered in open
woodlands of Mistbelt and Low Country, 1511
deltoideus Less., infrequent climber in mesic thickets of Transi-
tional Mistbelt, 920
erubescens Ait. var. crepidifolius DC. , localized forb of Plateau
Crest grasslands, 1121
erubescens A it. var. dichotomus DC. , infrequent forb of Humid
Mistbelt grasslands, 1180
gerrardii Harv., large herb, localized in Plateau Crest grasslands,
1214
glaberrimus DC., widespread subwoody forb of grassslands and
woodlands, Mistbelt and Low Country, 1274
latifolius DC. (= S. sceleratus), localized forb, mainly in grass-
lands of Escarpment Plateau, 26
lydenburgensis Hutch. & Bum Davy, infrequent forb of Sum-
mit and Escarpment Plateau grasslands, 1314
niacrocephalus DC., rosette forb, infrequent in rocky grass-
lands of Escarpment Slopes, 1145
mikanioides Otto ex Harv. , slender liane, occasional in tall forests
of Escarpment Slopes, 1142
oxyriifolius DC. , small subsucculent herb, often associated wdth
rocks, w idespread in grasslands and woodlands of Mist-
belt and Low Country, 1038
panduriformis Hilliard, infrequent tall herb of disturbed
thickets, Escarpment .Slopes, 1250
polyodon DC. var. polyodon, infrequent forb in moist Mistbelt
grasslands, 1105
pterophorus DC. , subwoody forb, infrequent in serai grasslands
of Escarpment and Lower Mountain Slopes, 111
scitus Hutch, ex Burn Davy, occasional subsucculent forb of
Summit grasslands and shrublands, 2635
serratuloides DC. var. serratuloides, robust forb, localized in
disturbed woodlands of Upper Foothills, 1828
sp., forb of Summit grasslands, 2555
tamoides DC., succulent climber, in forests and mesic thickets
of Mountain and Escarpment Slopes, 2059
venosus Harv., subwoody forb, infrequently scattered in wood-
lands of Escarpment Slopes and Foothills, 1545
Euryops Cass.
pcdunculatus N.E. Br., subwoody herb, localized in lithoseral
grasslands and shrublands of Mistbelt, 1637
transvaalensis Klatt subsp. setilobus (N.E. Br.) B. Nord., small
subwoody herb, infrequent in Mistbelt grasslands, 1499
Haplocarpha Less.
scaposa Han., low perennial herb, widespread in woodlands,
shrublands and grasslands of Mistbelt and Low Country,
669
Berkheya Ehrh.
echinacea (Harv.) O. Hoffm. ex Burtt Davy subsp. echinacea,
infrequent forb, widespread in Low Country woodlands
and Mistbelt grasslands and shrublands, 769
insignis (Harv.) Thell., small forb, widespread in woodlands,
shrublands and grasslands of Mistbelt and Low Country,
641
latifolia Wood & Evans, infrequent forb of Plateau Crest open
woodlands, 1980
radula (Harv.) De Wild., large forb associated with wooded
knolls. Summit, 2302
setifera DC., infrequent forb of Low Country woodlands, 1823
sp., variable forb in undergrow'th of mesic thickets, Escarpment
Lower Slopes, includes 1075
Dicoma Cass.
anomala Sond. subsp. cirsioides (Harv.) Wild, perennial forb,
infrequent in Summit and Mountain Slopes grasslands,
1459
zeyheri Sond., perennial forb, widespread in woodlands and
grasslands of Mistbelt and Low Country, 636
Gerbera L.
ambigua (Cass.) Sch. Bip., widespread rosette forb of Plateau
Crest and Summit grasslands and woodlands, Weisser
9031
aurantiaca Sch. Bip., rosette forb, widespread in woodlands
and grasslands of Mistbelt and Low Country, 656
jamesonii H. Bol. ex Adlam, rosette forb, mainly localized in
Woodland and Xeric Thicket of Low Country, 84
piloselloides (L.) Cass, occasional rosette forb of Mistbelt
grasslands, 1333
viridifolia (DC.) Sch. Bip. subsp. viridifolia, small rosette forb,
infrequent in Low Country woodlands, 1 743
Tolpis Adans.
capensis (L.) Sch. Bip., rosette forb, localized in Plateau Crest
and Summit grasslands, 1210
Hypochoeris L.
*microcephala (Sch. Bip.) Cabrera var. albiflora (Kuntze)
Cabrera, rare rosette forb of dolomite-soil grasslands,
Plateau Interior, 1367
*radicata L., infrequent weed of disturbed rank grasslands,
Summit Plateau, 2642
Sonchus L.
integrifolius Han., small herb with milky latex, localized in
dolomite-soil grasslands of Plateau Interior. 1325
sp., infrequent herb on termitaria, grasslands of Lower Moun-
tain Slopes, 1543
wilmsii R E. Fries, herb with milky latex, infrequent in Mist-
belt grasslands, 1537
Bothalia 19,1: 111-123(1989)
A survey and analysis of traditional medicinal plants as used by the
Zulu, Xhosa and Sotho
ANNE HUTCHINGS*
Keywords: ailment, medicinal plants, Sotho, Xhosa, Zulu
ABSTRACT
A coded list of 794 traditional Zulu medicinal plants is presented with a key to the ailments concerned. Xhosa and
Sotho usage of these plants is incorporated. Medicinal usage in the pteridophytes, gymnosperms, monocotyledons and
dicotyledons is tabulated. Ailments are categorized and discussed with an analysis of the plant families involved in their
treatment. Patterns of usage between related plants are observed and some potentially effective or dangerous characteristic
family constituents are briefly outlined.
UITTREKSEL
’n Gekodeerde lys van 794 tradisionele medisinale plante van die Zoeloes word aangebied met 'n sleutel tot die
betrokke ongesteldhede. Die gebruik van hierdie plante deur Xhosas en Sotho’s word ook vermeld. Benutting van die
pteridofiete, gimnosperms, monokotiele en dikotiele is getabuleer. Ongesteldhede word gekategoriseer en bespreek met 'n
ontleding van die plantfamilies wat by die behandeling betrokke is. Benuttingspatrone tussen verwante plante is waarge-
neem en sekere potensieel doeltreffende of gevaarlike stowwe wat kenmerkend in families voorkom, word kortliks aange-
toon.
INTRODUCTION
The data on which this paper is based come mainly
from a literature survey conducted by the author towards
the compilation of a Pharmacapoeia of Zulu Folk Medi-
cine, which was initiated at the University of Zululand
by Dr A. H. Scott in 1986. Xhosa and Sotho usage of the
plants, which is closely allied, has been included. Data
from the author’s previous fieldwork in Transkei and
limited personal interviews with Zulu and Xhosa tradi-
tional healers is included and further data have been
abstracted from a list of the Botanical Research Insti-
tute’s holdings of medicinal plants. The main sources of
information on usage have been 1, A. T. Bryant, whose
work on Zulu medicine was originally published in the
Annals of the Natal Museum in 1909 but later destroyed
by fire and republished in 1966 (Bryant 1966); 2, J.
Gerstner, who published his work on Zulu plant names
and usage between 1938 and 1941 (Gerstner 1938, 1939,
1941); and 3, Maim Hulme, who included Zulu usage
and also illustrated her book (Hulme 1954). Watt &
Breyer-Brandwijk (1962) was also consulted and addi-
tional information on Sotho usage is from A. Jacot Guil-
larmod (Jacot Guillarmod 1971).
It would be of great interest to compare the number of
plant species used with the number of plant species
known to be available, but research in this field, embrac-
ing both ecological and trading aspects, is beyond the
scope of this paper. Comprehensive research into the
* Zulu Folk Medicine Research, Department of Biochemistry, Univer-
sity of Zululand, Private Bag X1001 , Kwadlangezwa 3886.
MS. received: 1988.07.15.
toxicity and active chemical principles in the plants used
is also beyond the scope of the paper.
DISCUSSION OF LIST OF MEDICINAL PLANTS
A total number of 794 plant species with specific
usage known has been recorded. These are grouped as
pteridopytes, gymnosperms, monocotyledons and dico-
tyledons. They are arranged in alphabetic order accord-
ing to family and within the family alphabetically
according to genus and species. All plant names used in
the literature surveyed have been recorded and the names
listed have been updated according to Gibbs Russell et
al. (1985, 1987). Authors’ names have been omitted
from the coded list to save space so that uses could be
tabulated and similarities in the usage of related species
could be easily seen.
Plants of the same genus which could not be identified
to species level have been counted as one species unless,
as in the case of the Plectranthus spp. illustrated by
Hulme (1954), it is quite evident that different species
are being referred to.
The species listed have been coded according to ail-
ment. These ailments have been very broadly catego-
rized and determined largely by symptom, as is tradi-
tional treatment. Overlaps occur, as in the case of
bladder and urethral disorders, which have been classed
by the author as a common renal ailment but could be
caused or affected by procreation-related ailments such
as sterility and venereal disease.
The presence of a known or suspected toxic element
recorded in the literature surveyed is indicated by an
asterisk in the coded list at the end of this article. The
toxic element may not always be known in the part of the
plant used.
112
Bothalia 19,1 (1989)
Key to ailments:
A — Sexual
B — Sterility
C — Venereal
D — Gynaecological
E — Newly bom infants
F — Gastro-intestinal
G — Renal
H — Tonic
I — Growths
J — Respiratory
K — Febrile
L — Headaches
M — Heart
N — Nervous
0 — Pain producing
P — Leprosy
Q — Dental
R — Sorcery
S — Snake-bite
T — Skin
U — Sprains, fractures
V — Eyes, ears and nose
W — Insecticidal
X — Charm
Y — Animal
ANALYSIS
In this analysis of ailments and methods of treatment,
families, of which three or more species are used for the
same ailment, are highlighted.
Families marked with an asterisk are those of which
one or more species used for the specific ailment have
been recorded as toxic.
Procreation-related ailments (A-E in key to ailments
and coded list of plants)
A. Sexual remedies include aphrodisiacs and a large
number of love charm emetics, which are normally taken
by men. Some medicines are administered as charms to
secure the fidelity of the beloved or to harm a rival in
cases of suspected infidelity or to protect the user against
the effects of such medicine.
Of the 109 species recorded as sexual remedies, three
or more occur in the following 12 families, accounting
for 79:
13 Orchidaceae
13 Fabaceae*
1 1 Liliaceae*
7 Rubiaceae
5 Euphorbiaceae*
4 Apiaceae
4 Acanthaceae
95 of the 109 species are used as love charm emetics.
B. Sterility remedies include cures for both women and
men. Barrenness in women is usually treated with a pur-
gative or an enema or medicine may be directly intro-
duced into the womb. Sterility and impotence in men is
treated with orally taken infusions, or powdered medi-
cines may be blown through the urethra (Bryant 1966).
Also included are medicines taken, usually orally, by a
couple desiring a particular gender in a child, or medi-
cines taken by both parents after a miscarriage.
Of the 45 species recorded as sterility remedies, three
or more occur in the following six families, accounting
for 25:
7 Orchidaceae 3 Liliaceae*
4 Iridaceae* 3 Amaryllidaceae
5 Fabaceae 3 Rubiaceae
C. Venereal diseases (syphilis and gonorrhoea) may be
treated with orally administered decoctions or, in cases
of discharge, medicine may be inserted directly into the
penis or vagina in liquid form or as a pill of pounded
leaves, and charred powdered leaves or a poultice may
be applied directly to venereal sores (Bryant 1966).
Of the 34 species recorded as venereal disease reme-
dies, three or more occur in the following five families,
accounting for 19:
6 Liliaceae* 3 Fabaceae*
4 Asteraceae 3 Solanaceae*
3 Ranunculaceae*
D. Gynaecological remedies include medicines taken
regularly during pregnancy to ensure a safe delivery and
a healthy child. These may be mixtures of various ingre-
dients, usually roots (Gerstner 1941) or may be made
from the roots of a single plant, soaked in water, which
is drunk daily by the expectant mother. Also included are
medicines administered during childbirth, or for painful
or delayed menstruation, to stimulate breast devel-
opment or the flow of milk, or to procure abortions.
Of the 78 species recorded, three or more occur in the
following six families, accounting for 37:
12 Liliaceae* 4 Ebenaceae*
10 Asteraceae* 4 Scrophulariaceae
4 Amaryllidaceae* 3 Euphorbiaceae*
Of the 78 species used for gynaecological purposes,
38 are recorded as taken regularly during pregnancy,
three or more of which occur in the following three fami-
lies, accounting for 16:
8 Liliaceae* 3 Amaryllidaceae*
5 Asteraceae*
E. Newly-born infants are sometimes given purification
purges at birth and later at weaning; these may be admin-
istered orally or as an enema. Medicines may be applied
to the navel or fontanel at birth and others may be ap-
plied to the mother’s breast at weaning. Sometimes
newly bom infants are held in the smoke of a burning
plant in a protective or purification ritual.
Of the 18 species recorded, seven occur in Liliaceae.
Stress-related ailments (L-O in key and coded list of
plants)
{Note — the term stress-related is used here to refer to
ailments that are caused by psychological stress, al-
though there may be other causes.)
L. Headaches are considered by traditional healers to be
caused by emotional or mental disturbance. They are
often treated by snuffed or inhaled medicines and occa-
sionally leaves are wrapped around the head.
Of the 44 species recorded, three or more come from
the following five families, acounting for 22:
6 Euphorbiaceae* 3 Fabaceae*
6 Asteraceae* 3 Asclepiadaceae*
4 Ranunculaceae*
M. Cardiac ailments include high blood pressure, chest
pain not associated with respiratory complaints and bad
dreams believed to be caused by heart problems (Bryant
a — Includes use as anthel-
mintic
c — Includes enema admin-
istration
d — Includes use against
diarrhoea
e — Includes use as an
emetic
i — Given to infants or
children
p — Taken regularly during
pregnancy
r — Rubbed into scarifica-
tions
s — Snuffed or inhaled
* — Toxic
4 Gentianaceae
4 Scrophulariaceae
4 Acanthaceae
4 Asteraceae*
3 Amaryllidaceae*
3 Caryophyllaceae
Bothalia 19,1 (1989)
113
1966). Treatment may be administered in the form of an
emetic or enema.
Of the 21 species recorded, three occur in the follow-
ing two families, accounting for six:
3 Liliaceae* 3 Fabaceae*
N. Nervous or psychological ailments include hysteria,
mental disturbance, nightmares, states of believed be-
witchment, states of impurification after the death of a
kraal member and states of trance which need to be in-
duced in the diviner to enable her to fulfil her function.
Emetics and snuffed or inhaled medicines are frequently
used for these purposes.
Of the 133 species recorded for nervous ailments,
three or more come from the following 12 families,
accounting for 8 1 :
16 Fabaceae*
12 Asteraceae*
8 Liliaceae*
8 Rubiaceae
7 Orchidaceae
8 Asclepiadaceae*
5 Iridaceae*
4 Euphorbiaceae*
4 Apiaceae
3 Hypoxidaceae*
3 Sapindaceae
3 Solanaceae*
Of the 318 species recorded as gastro-intestinal rem-
edies 46 are recorded as anthelmintics and 54 as diar-
rhoea and dysentery remedies. These species may also
be used for other gastro-intestinal purposes.
Of the 46 recorded anthelmintics, three or more occur
in the following four families, accounting for 22:
8 Asteraceae* 4 Myrsinaceae*
7 Liliaceae* 3 Aspidiaceae*
O. Pain producing ailments include pain in the joints or
back, rheumatism and also paralysis. Although these
may obviously not always have a psychological cause,
there is a traditional belief that the joints are the most
vulnerable area to the entrance of evil spirits (Ngubane
1977) and conversion disorders often do take the form of
pain in the joints and sometimes paralysis. Dried pow-
dered medicine is frequently rubbed into freshly cut
scarifications to treat these conditions and other forms of
administration such as enemas and emetics are also used.
Of the 46 species recorded, three or more come from
the following families, accounting for 13:
7 Liliaceae* 3 Meliaceae*
3 Fabaceae*
Miscellaneous ailments (F-K and Q-Y in key and
coded list of plants)
F. Gastro-intestinal ailments include:
1, stomach ache and constipation, which are treated
with enemas or orally administered purges;
2, intestinal worms which are treated with orally
administered anthelmintics or enemas;
3, diarrhoea and dysentery which are treated with an
orally administered medicine, frequently followed by an
enema of the same medicine;
4, nausea or the accumulation of bile is treated with an
emetic;
5, indigestion, for which small pieces of root, bark or
leaves may be chewed;
6, haemorrhoids and a condition which Bryant (1966)
and Ngubane (1977) refer to as gangrenous rectitis,
which is frequently treated with an enema, or a locally
applied lotion or powder and may also be treated with an
orally taken decoction.
Of the 318 species recorded for gastro-intestinal com-
plaints, three or more occur in the following 35 families,
accounting for 252:
Of the 54 recorded diarrhoea and dysentery remedies,
three or more occur in the following six families,
accounting for 30:
10 Fabaceae* 3 Sapindaceae*
7 Iridaceae* 3 Rubiaceae
4 Geraniaceae 3 Asteraceae
G. Renal ailments include kidney and urinary tract com-
plaints. Medicines may be orally administered, some-
times followed by an enema of the same medicine after
three days of treatment, or medicines may be rubbed into
incisions cut in the loins or inserted directly into the
urethra or a poultice may be applied externally.
Of the 44 species recorded for renal ailments, three or
more are found in the following five families, accounting
for 19:
5 Amaryllidaceae* 3 Apocynaceae*
5 Asteraceae* 3 Euphorbiaceae*
3 Liliaceae*
H. Debility ailments include general weakness, a low
resistance to infections, blood impurities and a lack of
appetite. They are treated with orally taken tonics,
emetics or enemas.
Of the 58 species recorded for debility, three or more
come from the following six families, accounting for 24:
8 Asteraceae 3 Apocynaceae*
4 Euphorbiaceae* 3 Periplocaceae
3 Fabaceae 3 Rubiaceae
I. Swellings or growths include swollen glands, scrofu-
lous and dropsical swellings and external or internal
swellings or lumps which may or may not be cancerous.
Medicines may be administered orally, sometimes fol-
lowed by a poultice made from the same ingredients, or
powdered medicines may be rubbed into incisions cut
around the swelling.
Of the 3 1 species used to treat swellings, three or more
come from the following two families, accounting for
eight:
5 Euphorbiaceae* 3 Asteraceae*
114
J. Respiratory ailments include chest pain from pleurisy
or bronchitis, coughs, sore throats and asthma or catarrh.
They are frequently treated with emetics, which perform
an expectorant function, or medicines may be chewed,
drunk, snuffed, inhaled or rubbed into incisions on the
chest.
Of the 144 species recorded for respiratory ailments,
three or more come from the following 14 families, ac-
counting for 92:
27 Asteraceae*
12 Fabaceae*
8 Euphorbiaceae*
6 Liliaceae*
5 Amaryllidaceae*
5 Apiaceae*
5 Lamiaceae
5 Verbenaceae
4 Rubiaceae
3 Phytolaccaceae*
3 Brassicaceae
3 Capparaceae*
3 Ebenaceae*
3 Celastraceae*
K. Febrile complaints include influenza, colds and
fevers, including malaria and rheumatic fever. They are
frequently treated with emetics. Snuffed or inhaled
medicines may also be administered and the patient may
be steamed or bathed to reduce the fever.
Of the 123 species involved, three or more come from
the following 13 families, accounting for 82:
21 Asteraceae*
12 Fabaceae
10 Lamiaceae
6 Liliaceae*
6 Euphorbiaceae*
5 Rubiaceae
4 Amaryllidaceae*
3 Iridaceae*
3 Rutaceae
3 Amaranthaceae
3 Apiaceae
3 Scrophulariaceae
3 Lobeliaceae
P. Leprosy is usually now referred by traditional healers
to the hospitals. Only two remedies have been recorded
and they are from Melianthaceae and Passifloraceae.
Q. Toothache and sore gums are treated with lotions or
powders rubbed on to the painful area.
Of the 30 species recorded for toothache, three or
more are from the following three families, accounting
for 12:
5 Solanaceae* 3 Euphorbiaceae*
4 Asteraceae*
R. Sorcery is believed to be the cause of many illnesses
and certain plants are believed to be used by sorcerers to
bring about evil. The same plants may be taken as an
antidote to the disease thus brought about, usually in the
form of an emetic.
Of the 16 species recorded as sorcerer’s medicines,
three each are from the following two families, account-
ing for six:
3 Euphorbiaceae* 3 Apiaceae*
S. Snake bite remedies may be taken or locally applied,
the same medicine often being administered at the same
time in both forms. Dried ground snakes’ heads are
sometimes an ingredient in the medicine.
Of the 43 species recorded, three or more come from
the following six families, accounting for 20:
4 Thymelaeaceae* 3 Fabaceae*
4 Asteraceae* 3 Euphorbiaceae*
3 Phytolaccaceae* 3 Apocynaceae*
T. Skin complaints include sores, wounds, bums and
rashes. These complaints may be treated by applied
lotions, poultices or washes. Washes may also be given
Bothalia 19,1 (1989)
to reduce the temperature in fevers. Also included are
hair restorers.
Of the 100 species recorded, three or more come from
the following 10 families, accounting for 64:
15 Asteraceae*
11 Fabaceae*
9 Solanaceae*
7 Euphorbiaceae*
4 Liliaceae*
4 Amaryllidaceae*
4 Thymelaeaceae
4 Acanthaceae
3 Verbenaceae
3 Lamiaceae
U. Fractures, sprains and bruises may be treated with
lotions, or powder may be rubbed into scarifications as
an anti-inflammant around the site of the damage.
Of the 40 species recorded, three or more come from
the following three families, accounting for 12:
6 Asteraceae* 3 Crassulaceae*
3 Euphorbiaceae*
V. Ear, eye and nose complaints are treated with lotions
or sap directly squeezed from the plants.
Of the 39 species recorded, three or more come from
the following four families, accounting for 19:
7 Liliaceae* 3 Crassulaceae*
6 Asteraceae 3 Rosaceae
W. Insecticides and piscicides include medicines applied
to rid the body of lice and fleas and those used to repel or
kill flies and those used to stun fish so that they may be
caught.
Of the 21 species recorded, three or more come from
the following two families, accounting for 12:
9 Fabaceae* 3 Asteraceae
X. Charms may be applied to placate evil spirits, for
protection against enemies and storms, for good luck or
to make the user liked. They may be bathed with, worn,
sprinkled or burnt in the home or fields or cultivated as
protective plants.
Of the 182 species recorded, three or more come from
the following 14 families, accounting for 126:
27 Liliaceae*
18 Orchidaceae
15 Asclepiadaceae*
1 1 Asteraceae*
9 Amaryllidaceae*
9 Crassulaceae*
8 Iridaceae*
7 Fabaceae*
7 Euphorbiaceae*
6 Apiaceae
3 Rhamnaceae
3 Scrophulariaceae
3 Rubiaceae
Y. Animals may be given medicines to prevent or cure
disease. Also included are medicines applied as theft
deterrents, either in the field or on eggs or drying animal
skins.
Of the 102 species used to treat or deter animals, three
or more come from the following 11 families, account-
ing for 53:
1 1 Asteraceae*
9 Fabaceae*
7 Liliaceae*
6 Asclepiadaceae*
5 Lamiaceae
3 Amaryllidaceae*
3 Dioscoreaceae*
3 Urticaceae
3 Geraniaceae
3 Vitaceae*
It may be observed (Table 1) that among the med-
icinally used plants recorded, a higher proportion of
monocotyledons are used as externally applied charms
and for procreation-related purposes and that there is a
slightly wider range of medicinal usage among the dico-
tyledons. Further analysis reveals that among the mono-
Bothalia 19,1 (1989)
115
cotyledons 43 % of the species are used for procreation-
related ailments, 37 % as externally applied charms and
26 % for possibly stress-related ailments whereas among
the dicotyledons 27 % are used for procreation-related
ailments, 26 % for possibly stress-related ailments and
19 % as externally applied charms. In both groups the
greatest proportion of plants used for other ailments are
used for gastro-intestinal purposes (32 % of monocotyle-
dons and 42 % of dicotyledons) and respiratory ailments
(11 % of monocotyledons and 20 % of dicotyledons).
This is reflected in Tables 2 & 3.
DISCUSSION
Perception of aetiology also determines treatment.
According to Bryant (1966) and Ngubane (1977), most
common ailments are believed to be caused by an excess
of bile or gall, which needs to be removed. Diseases
believed to be caused by evil spirits or pollution also
require catharsis. This explains the wide use of emetics,
enemas and purgatives. A total of 238 of the plants re-
corded for this study, i.e. 30 %, are used as emetics —
where use as an emetic has been recorded with no
specific ailment, the plant has been considered to be used
for gastro-intestinal purposes. Emetics are not normally
given to young children but enemas are and are consid-
ered by medical staff to be the cause of some of the
poisoning cases and liver damage seen in hospitals
(Savage & Hutchings 1987). Forms of administration are
not always recorded in the sources consulted and more
plants are probably used for enemas than appear on the
list. Species indicated as being used for children are
frequently administered in this way. Patterns of usage
between closely related species, such as the use of pteri-
dophytes as anthelmintics, Orchidaceae as love charm
emetics, Ranunculaceae species for headaches, Thy me -
laeaceae species for skin complaints and snake bite or
Lamiaceae for febrile conditions and various Asclepiada-
ceae for nervous complaints, are discernible in the list.
Such patterns of usage could obviously indicate that
related plants share chemical constituents, which would
in turn account for their possible effectiveness and/or
toxicity. Some characteristic family constituents with
some of their likely effects and potential dangers are
given below.
Widespread steroidal saponins, cardiac glycosides in
some of the Liliaceae and toxic alkaloids in the Amaryl-
lidaceae are among the potentially dangerous consituents
found in monocotyledonous plants, of which so many
are used in the procreation-related ailments. Steroidal
saponins may affect the sex hormones and are relatively
harmless when taken by mouth but found highly toxic if
they enter the blood stream (Trease & Evans 1983). This
may happen if there is any damage to the mucous lining
of the gastro-intestinal tract when enemas are adminis-
tered as, according to Ngubane (1977), the dosage is
controlled to enable the patient to retain the medicine for
a time. This custom would also make enemas made with
other toxic material more dangerous than emetics as ab-
sorption of material through the rectum is easier than
through the small intestine. Steroid or triterpenoid sapo-
nins (which are common in the dicotyledons) are often
found in the plants used as emetics and have the property
of foaming and also frequently irritate the mucosa. This
may account for their expectorant and decongestive
action when used for chest ailments. Anti-microbial, cy-
tostatic and anti-inflammatory activity have been demon-
strated in saponins (Lower 1985).
Cardiotonics, which can have a diuretic action by in-
creasing the renal bloodflow, may be found in mem-
bers of various other families apart from Liliaceae and
these include Apocynaceae and Asclepiadaceae, Rubia-
ceae and Solanaceae (see Oliver-Bever 1986). The di-
uretic action would be helpful in cases of gonorrhoea and
also in various kidney or heart diseases which cause
dropsical swellings. Various species of the above-men-
tioned families are used for venereal diseases, renal com-
plaints, dropsical swellings or heart complaints and may
be found effective.
The toxic Amaryllidaceae alkaloids produce gastro-
intestinal upset (Jaspherson-Schib 1970). Highly toxic
species are found in other alkaloid-containing families
such as Apocynaceae, Solanaceae and Euphorbiaceae.
These species also owe their effectiveness as purges and
their potentially dangerous properties to extreme gastric
irritation.
Antipyretic, protozoicidal and local anaesthetic pro-
perties are to be found in many of the West African
species of the alkaloid-rich Rubiaceae family (Oliver-
Bever 1986) and members of the family locally used for
febrile ailments would probably merit further investiga-
tion. Alkaloids have a marked action on the central ner-
vous system and can act as depressants (e.g. the sedative
reserpine from Rauvolfia vomitoria Afzel.) or stimulants
(e.g. the Strychnos alkaloids) which may account for the
use of various species from families such as Euphor-
biaceae and Solanaceae for nervous complaints.
Tannin has frequently been observed in the parts of the
plant used in the treatment of dysentery and diarrhoea or
for respiratory ailments and is a characteristic constituent
of many of the families thus used (e.g. Rosaceae,
Fabaceae, Geraniaceae). It is likely to be effective on
account of its protein-precipitating properties which, in
small doses, would form a protective, impermeable layer
and also tend to prevent the development of bacteria —
large doses would irritate the mucosa (Fliick 1976).
The presence of volatile oils with possible carminative
or antispasmodic activity is likely to account for the use
of the closely related Lamiaceae and Verbenaceae and
other aromatic families for coughs, colds, influenza and
digestive disorders.
The snake bite cures are interesting. Many are known
to be either toxic or else closely related to known toxic
species, notably from families where cardioactive toxins
(Melianthaceae, Loganiaceae, Apocynaceae and Ascle-
piadaceae) or alkaloids (Phytolaccaceae, Solanaceae and
Asteraceae) are known or else from families where cyto-
toxic activity has been observed (Euphorbiaceae and
Thymelaeaceae).
More fieldwork will undoubtedly reveal new usage of
plants as the practice of herbal medicine is still very
much alive. Patterns obviously exist in the usage of
plants, but the perception thereof is influenced by the
way in which the data are analysed and on the cultural
TABLE 1. — Medicinal usage in pteridophytes, gymnosperms, monocotyledons and dicotyledons
116 Bothalia 19.1 (1989)
: see key to list of medicinal plants.
Bothalia 19,1 (1989)
117
perceptions of the researcher. Nevertheless, plant usage
is patterned, and these patterns deserve further investiga-
tion.
ACKNOWLEDGEMENTS
The research for this study was partly undertaken
through Zulu Folk Medicine Research, which is headed
by Dr A. H. Scott and funded by the De Beers Chair-
man’s Fund Educational Trust, who are thanked for their
support. Mrs E. Brink (Grahamstown Botanical Unit) is
thanked for her help in updating the nomenclature and Dr
G. E. Gibbs Russell for making available the list of the
Botanical Research Institute holdings of medicinal
plants. Prof. S. E. Terblanche of the Department of Bio-
chemistry, University of Zululand and Dr L. M.
Gerritsma of the Department of Pharmaceutical Chemis-
try, University of Potchefstroom, are both thanked for
their helpful advice.
REFERENCES
BRYANT, A. T. 1966. Zulu medicine and medicine-men. Struik, Cape
Town (originally published in 1909 in the Annals of the Natal
Museum).
FLUCK, H. 1976. Medicinal plants. Rowson & Foulsham, London.
GERSTNER, J. 1938. A preliminary checklist of Zulu names of plants
with short notes. Bantu Studies 12: 215-236; 321-342.
GERSTNER, J. 1939. A preliminary checklist of Zulu names of plants
with short notes. Bantu Studies 13: 49-64; 131-149; 307-326.
GERSTNER, J. 1941. A preliminary checklist of Zulu names of plants
with short notes. Bantu Studies 15: 277-301; 369-383.
GIBBS RUSSELL. G. E., REID, C., VAN ROOY. J. & SMOOK. L.
1985. List of species of southern African plants. Edn 2, Part 1.
Memoirs of the Botanical Survey of South Africa No. 51.
GIBBS-RUSSELL, G. E., WELMAN, W. G., RETIEF, E..
IMMELMAN, K.. GERMISHU1ZEN, G., PIENAAR. B. J.,
VAN WYK. M. & NICHOLAS, A. 1987. List of species of
southern African plants. Edn 2, Part 2. Memoirs of the Botanical
Suney of South Africa No. 56.
HULME, M. M. 1954. Wild flowers of Natal. Shuter & Shooter.
Pietermaritzburg.
JACOT GUILLARMOD, A. 1971. Flora of Lesotho. Cramer, Lehre.
JASPERSEN-SCHIB, R. 1970. Toxische Amaryllidaceen. Plwrma-
ceutica Acta Helvetica 45: 424-433.
LOWER, E. S. 1985. Activity of the saponins. Drug and Cosmetic
Industry 135: 39-44.
NGUBANE. H. 1977. Body and mind in Zulu medicine. Academic
Press, London.
OLIVER-BEVER, B. 1986. Medicinal plants in tropical West Africa.
Cambridge University Press, Cambridge.
SAVAGE, A. & HUTCHINGS. A. 1987. Poisoned by herbs. British
Medical Journal 295: 1650-165 1 .
TREASE, G. E. & EVANS, W. C. 1983. Pharmacognosy. 12th edn
Bailliere Tindall, London.
WATT, J. M. & BREYER-BRANDWIJK, M. G. 1962. The medicinal
and poisonous plants of southern and eastern Africa. 2nd edn.
Livingstone, London.
LIST OF ZULU MEDICINAL PLANTS
Key to ai laents:
A - Sexual
B - Sterility
C - Venereal
D - Gynaecological
E - Nealy born infants
F - Gastro-intestinal
6 - Renal
H - Tonic
I - Sroaths
J - Respiratory
K - Febrile
L - Headaches
H - Heart
H - Nervous
0 - Pain producing
P - Leprosy
0 - Dental
R - Sorcery
S - Snake-bite
T - Skin
U - Sprains, fractures
V - Eyes, ears and nose
N - Insecticidal
1 - Chare
Y - Animal
a - Includes use as anthelaintic
c - Includes enesa administration
d - Includes use against diarrhoea
e - Includes use as an eeetic
i - 6iven to infants or children
p - Taken regularly during pregnancy
r - Rubbed into scarifications
s - Snuffed or inhaled
' - Toxic
A8CDEFGHIJKLHN0PQRSTUVMZV
118
Bothalia 19,1 (1989)
Bothalia 19,1 (1989)
119
120
Bothalia 19,1 (1989)
Vernonia oiigocephala
Vernonia sp.
BALAN I T ACEAE
Balanites laughann
BALANOPHORACEAE
Sarcophyte sanguines
BALSAHINACEAE
Iipatiens hochstetteri
BE50NI ACEAE
Fd I
Turbina oblongata
CRASSULACEAE
Cotyledon orbiculata*
Crassula alba
Crassula barbata
Crassula dependens
Crassula tulticava
Crassula orbicularis
Crassula sarcocaulis
Crassula sp.
Crassula vaginata
Kalanchoe crenata
T U V X
Fde
Fe
Fe
Fe
Fe
Ks Re
Bothalia 19,1 (1989)
122
Bothalia 19,1 (1989)
Bothalia 19,1 (1989)
123
T y
I
y
s
T
5 T
S
T
S 1 V
y
T
T
*
y
y
i
y
y
s
r
T
T
v
v
i
y
y
y
i
Bothalia 19,1: 125-132(1989)
Chromosome studies on African plants. 9. Chromosome numbers in
Ehrharta (Poaceae: Ehrharteae)
J. J. SPIES*, E. J. L. SAAYMAN*, S. P. VOGES** and G. DAVIDSE***
Keywords: aneuploidy, chromosome numbers, Ehrharta , meiosis, polyploidy, southern Africa, winter rainfall area
ABSTRACT
Cytogenetic studies of 53 specimens of 14 species of the genus Ehrharta Thunb. confirmed a basic chromosome
number of 12 for the genus. Chromosome numbers for 13 species are described for the first time. The highest ploidy level
yet observed in the genus (2n = lOx = 120) is reported for£. villosa var. villosa. B chromosomes were observed in several
specimens of four different species.
UITTREKSEL
Sitogenetiese studies van 53 eksemplare, verteenwoordigend van 14 spesies van die genus Ehrharta Thunb., bevestig
’n basiese chromosoomgetal van 12 vir die genus. Chromosoomgetalle van 13 spesies word vir die eerste keer beskryf. Die
hoogste poliplo'iede vlak wat nog in die genus beskryf is (2n = lOx = 120), is by E. villosa var. villosa gevind. B-
chromosome is by verskeie eksemplare van vier verskillende spesies waargeneem.
INTRODUCTION
The presentation of chromosome numbers in this
report continues with the format established in the first
publication in this series (Spies & Du Plessis 1986).
However, contrary to previous publications in this series
which all dealt with several different genera of the family
Poaceae, this paper covers the single genus, Ehrharta
Thunb. The results presented here are limited to the
species from the winter rainfall area of southern Africa.
The winter rainfall region of southern Africa, with its
Mediterranean climate, is located in the extreme south-
west of the continent. Though small in area, less than
4% of the total land surface of southern Africa, it is
floristically extremely important, being recognized as
one of the six Floral Kingdoms of the world (Goldblatt
1978). The flora of the Cape Floristic region, or Fynbos
Biome, is unique in many respects, being particularly
rich in species (Goldblatt 1978), with high levels of
endemism. Thus, in the 87 244 km2 of the Cape Region
(Fynbos and Succulent Karoo Biomes combined), there
are over 8 000 species of which 73% are endemic (Gibbs
Russell 1987). High species diversity characterizes this
fynbos vegetation (Gibbs Russell 1987) and this is also
the case with the grasses of this biome. Although rela-
tively inconspicuous in this vegetation, the fynbos
grasses are particularly interesting as many of them are
restricted to this region and are poorly known taxonomi-
cally.
Ehrharta is one of the grass genera with a high propor-
tion of its species endemic to the Fynbos Biome. All but
* Botanical Research Institute, Department of Agriculture and Water
Supply , Private Bag X 1 0 1 , Pretoria 000 1 .
** Current address: Biotechnological Research Centre, Private Bag
X293, Pretoria 0001.
*** Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri
63166-0299, USA.
MS. received: 1988.09.12.
three of the southern African species are endemic to the
Fynbos Biome (Gibbs Russell pers. comm.). The genus
belongs to the tribe Ehrharteae Nevski and, as currently
classified, is represented by 30 species (with 16 infra-
specific taxa) in southern Africa (Gibbs Russell & Ellis
1987). Gibbs Russell & Ellis (1987) subdivided the
South African representatives of the genus into seven
species groups, i.e. the Setacea, Capensis, Erecta, Caly-
cina, Villosa, Ramosa and Dura groups. The higher
classification of the tribe Ehrharteae, which has been
reviewed by Gibbs Russell & Ellis (1987), is still unre-
solved. Most recently Clayton & Renvoize (1986) in-
cluded the tribe in the subfamily Bambusoideae. How-
ever, both Soderstrom & Ellis (1987) and Kellogg &
Campbell (1987) excluded it from the Bambusoideae, to
which they considered the Ehrharteae to be a sister
group.
The aim of this study was to determine the chromo-
some numbers and the meiotic chromosome behaviour of
as many different species as possible. Future studies will
determine whether this information can contribute to the
delimitation of species in the genus, as well as to clarify-
ing the position of the Ehrharteae in the Poaceae.
MATERIALS AND METHODS
The material used during this study was collected and
fixed in the field. A list of the material used and the
localities are listed under results. Voucher herbarium
specimens are housed in PRE.
Inflorescences were fixed in Camoy’s fixative (1886).
After 24-48 hours of fixation, the fixative was replaced
by 70% ethanol. Anthers were squashed in aceto-car-
mine (Darlington & La Cour 1976). Slides were perma-
nently mounted by freezing them with liquid C02, fol-
lowed by dehydration in ethanol and mounting in
Euparal.
126
Bothalia 19,1 (1989)
RESULTS
Family Poaceae, Tribe Ehrharteae, Genus Ehrharta.
Capensis group
Capensis subgroup
Ehrharta barbinodis Nees ex Trin.: n = 12.
CAPE. — 2917 (Springbok): 1 km from Springbok to Hondeklip Bay
( — DD), Spies 3000.
E. capensis Thunb.: n = 12.
CAPE. — 3218 (Clanwilliam): Versveld Pass (-DC), .Spies 3168.
Longifolia subgroup
No results.
Erecta group
E. erecta Lam. var. erecta : n = 12.
CAPE. — 3325 (Port Elizabeth): King Neptune Beach (-DC), Spies
3478.
E. erecta Lam. var. natalensis Stapf: n = 12.
CAPE.— 3323 (Willowmore): Blouwkransrivier Pass (-DC),
Davidse 33656.
E. triandra Nees ex Trin.: n = 12 & 24.
CAPE. — 2917 (Springbok): 21 km from Springbok to Kleinsee
(-DA), Davidse 33240 (n = 24). 3018 (Kamiesberg): 6 km east of
Kamieskroon (-BD), Davidse 33320 (n = 12).
E. longiflora J. E. Sm.: n = 12 & 24.
CAPE. — 2918 (Gamoep): 21 km from Springbok to Kleinsee
(-CA), Spies 3343 (n = 12). 3018 (Kamiesberg): 52 km from Spring-
bok to Loeriesfontein (- AA), Spies 3375 (n = 12); Buffelsrivier Val-
ley between Pedroskloof and Bobbejaanshoek (- AA), Davidse 33308
(n = 12); 6 km east of Kamieskroon (— BD), Spies 2992 (n = 12).
3418 (Simonstown): Redhill, 4 km from Simonstown (-AB), Spies
3233 (n = 24).
Calycina group
E. brevifolia Schrad. var. brevifolia: n = 12.
CAPE. — 3017 (Hondeklipbaai): coastal dunes near Groenrivier
( - DC ) , Da vidse 33365 .
E. brevifolia Schrad. var. cuspidata Nees: n = 12 +
0-1B.
CAPE. — 2917 (Springbok): 36 km from Port Nolloth to Kleinsee
(-AC), Davidse 33273 ; Spies 3361, 3362. 3017 (Hondeklipbaai): 15
km east of Hondeklip Bay (-AB), Spies 3028; 42 km west of Garies
towards Groenriviersmond (-DC), Spies 3383.
E. calycina J. E. Sm.: n = 12, 12 + 0-2B, 24 & 24 +
2B.
CAPE. — 2917 (Springbok): 17 km from Okiep to Goodhouse
(-BD), Spies 2995 (n = 12); 1 km from Springbok to Hondeklip Bay
(-DD), Spies 3003 (n = 12 + 0-2B), 3005 (n = 24). 3017 (Honde-
klipbaai): 6 km east of Kamieskroon in Kamiesberg Pass (-BB),
Davidse 33321 (n = 24), 33326 (n = 12 + 0-2B); Spies 3380 (n =
12); 42 km from Garies to Groenrivier ( — DC), Spies 3393 (n = 12); 3
km south of Groenrivier on road to Kotzerus (-DD), Davidse 33371 (n
= 12 + 2B). 3018 (Kamiesberg): 16 km east of Kamieskroon (-AC),
Spies 3058 (n = 24), 3062 (n = 12); Rooiberg in Kamiesberg Range
(-AC), Davidse 33337 (n = 12), 33349 (n = 24 + 2B). 3118 (Van-
rhynsdorp): Gifberg Pass (—DC), Spies 3094 (12 + 2B), 3095 (n =
12); 24 km from Clanwilliam to Elizabethfontein (-DD), Davidse
33417 (n = 12). 3119 (Calvinia): Vanrhyns Pass (-AC), Spies 3110
(n = 12); top of Botterkloof Pass (-CD), Davidse 33401 (n = 12).
3318 (Cape Town): near Stellenbosch turnoff on road between Paarl
and Franschhoek (-DD), Spies 3198 (n = 12 + 2B). 3321 (Ladi-
smith): Seweweekspoort Pass (-AD), Spies 3272 (n = 12 + 0-2B).
3323 (Willowmore): 5 km from Avontuur to Uniondale (— CA),
Davidse 33687 (n = 24). 3419 (Caledon): 5 km from Hermanus to
Onrustrivier (-AC), Davidse 33828 (n = 12 + 0-2B). 3421
(Riversdale): 4 km from Kweekkraal to Droerivier ( — AA), Spies 3549
(n = 12 + 0-2B); 2 km from Vermaaklikheid to Puntjie (—AC), Spies
3562 (n = 12). 3424 (Humansdorp): Skuitbaai ( — AA), Davidse 33638
(n = 12).
E. delicatula (Nees) Stapf: n = 12.
SWA/NAMIBIA. — 2616 (Aus): near Aus (— CB), Spies 2927.
CAPE. — 2917 (Springbok): 57 km from Kleinsee to Springbok
(—DA), Davidse 33285.
E. melicoides Thunb. : n = 12.
CAPE. — 3018 (Kamiesberg): Rooiberg in the Kamiesberg Range
(-AC), Davidse 33346. 3119 (Calvinia): 25 km from Calvinia to
Clanwilliam (— CA), Spies 3122. 3420 (Bredasdorp): 6 km from
Malgas to Bredasdorp (- AD), Spies 3251.
E. pusilla Nees ex Trin.: n = 12.
CAPE. — 2918 (Gamoep): 53 km from Springbok to Gamoep
(— CC), Davidse 33302. 3018 (Kamiesberg): Buffelsrivier Valley be-
tween Pedroskloof and Bobbejaanshoek (-AA), Davidse 33310; 52
km from Springbok to Loeriesfontein (- AA), Spies 3374.
Villosa group
E. villosa Schult. f. var. villosa: n = 60.
CAPE. — 3424 (Humansdorp): 27 km from Humansdorp to Cape St.
Franc is ( - B B ) , Spies 3505 .
Ramosa group
E. ramosa (Thunb.) Thunb. subsp. ramosa: n = 12.
CAPE. — 3324 (Steytlerville): 71 km from Rocklands to Elandsrivier
( — DB), Spies 3490.
E. rehmannii Stapf subsp. rehmannii: n = 12 + 0-5B.
CAPE. — 3322 (Oudtshoom): Montagu Pass (-CD), Davidse
33708.
E. rehmannii Stapf subsp. subspicata Stapf: n = 36.
CAPE. — 3323 (Willowmore): 9 km from Coldstream to Humans-
dorp ( — DD), Spies 3515.
Dura group
E. dura Nees ex Trin.: n = 12 + 0— 4B.
CAPE. — 3322 (Oudtshoom): Robinson Pass (— CC), Davidse
33534.
DISCUSSION
According to Watson et al. (1986) the basic chromo-
some number of the genus Ehrharta is 12 and ploidy
levels vary from diploid to tetraploid. This study con-
firms the basic chromosome number but increases the
Bothalia 19,1 (1989)
127
range of polyploidy from diploid to decaploid. In order
to place the cytogenetic results in a taxonomic context,
the results are discussed according to the species groups
of Gibbs Russell & Ellis (1987).
Both species studied in the Capensis subgroup of the
Capensis group, i.e. E. barbinodis and E. capensis,
were diploid with no meiotic abnormalities (Figure 1A).
This is the first reported chromosome number for both
species.
brevifolia var. brevifolia and E. brevifolia var. cuspidata
(Figure 2A-C), as well as E. delicatula and E. meli-
coides, are the first reported numbers for the taxa.
A very high frequency of specimens from the Calycina
group were aneuploid (24%) with one or two additional
chromosomes, although most had two. With the excep-
tion of one E. brevifolia var. brevifolia specimen, these
additional chromosomes were restricted to E. calycina.
Love (1948), Tothill & Love (1964) and Spies & Voges
(1988) have previously reported one to six additional
chromosomes beyond the basic complement(s) in E. ca-
lycina. Tothill & Love (1964) referred to these additional
chromosomes as supernumerary chromosomes and Spies
& Voges (1988) called them B chromosomes. Both
groups of authors observed no size or behavioural differ-
ences between the additional and standard chromo-
somes. As in these three previous studies, we observed
meiotic irregularities, including late disjunction of biva-
lents during the first division (Figure 2L) and non-align-
ment of chromosomes on the metaphase plate (Figure
2J). As was also observed in these previous studies, the
extra chromosomes may sometimes pair, either as multi-
valents or as bivalents (Figure 2F). Additional abnorma-
lities observed during meiosis include the presence of
univalents in a diploid specimen Davidse 33371 (n = 12)
and up to six anaphase I laggards in Davidse 33321 (n =
24) (Figure 2L).
A very narrow spindle on one side of the cell was
observed in an E. delicatula specimen (Davidse 33285)
(Figure 3A & B). Specimens of both E. brevifolia and E.
delicatula had two chromosome pairs associated with the
nucleolus (Figure 2 A & B).
Only one specimen of the Villosa group was
successfully analysed. This specimen, Spies 3505 (E.
villosa var. villosa), provides the first chromosome
number report for a member of this group and it proved
to have the highest ploidy yet described in the Ehrhar-
teae, i.e. n = 60 (Figure 4A & B). Apart from a few
multivalents and univalents observed in this specimen,
meiosis in this decaploid was normal and no laggards
were observed, which is surprising in view of the high
ploidy level.
The described diploid and tetraploid levels in the
Erecta group (Parthasarathy 1939; De Wet & Anderson
1956; Tateoka 1965; Raven et al. 1965; Fernandes &
Queiros 1969; Davidse et al. 1986; Spies & Du Plessis
1986; Hoshino & Davidse 1988) are supported by this
study. Chromosome numbers are reported for the first
time for E. erecta var. natalensis (n = 12) and E. trian-
dra (n = 12 & 24) (Figure IB), as well as a new ploidy
level for E. longiflora (n = 12). Results reported up to
now indicate that 72 % of the sampled plants are diploid.
With the exception of E. erecta var. natalensis, of which
only one specimen has been studied, the four other taxa
of the group have both diploid and tetraploid specimens.
E. triandra and E. longiflora are closely related and have
many morphological similarities, with E. longiflora
being larger than E. triandra. However, since both spec-
ies have been shown to have diploid and tetraploid popu-
lations, this size difference cannot be attributed to differ-
ent ploidy levels.
The Calycina group was the most thoroughly studied
with counts from 45 plants, representing six of the seven
taxa. Both diploid (84%) and tetraploid (16%) specimens
were observed (Figure 2D-L) which corroborates pub-
lished results (Parthasarathy 1939; Love 1948; Tateoka
1957; De Wet 1960; Tothill & Love 1964; Spies & Du
Plessis 1986; Hoshinso & Davidse 1988; Spies & Voges
1988). The polymorphic E. calycina diploid forms in-
clude prostrate coastal plants with short, broad leaf
blades as well as the very common, widely distributed
plants with erect growth habit and flat leaf blades. The
tetraploid form of E. calycina seems to be restricted to
densely tufted plants with erect, narrow, often rolled leaf
blades. The chromosome numbers reported here for E.
FIGURE 1. — Meiotic chromosomes in:
A, the Capensis subgroup, E.
capensis, Spies 3168 , metaphase I
with 12 bivalents; B, the Erecta
group, E. triandra, Davidse 33320,
diakinesis with 12 bivalents of
which four are associated with the
nucleolus, x 1300.
128
Bothalia 19,1 (1989)
FIGURE 2. — Meiotic chromosomes in the Calycina group. A— C: E. brevifolia var. cuspidata , with 12 bivalents at diakinesis. A, Spies 3028',
B, Spies 3362, with B chromosome (indicated by an arrow); C, Davidse 33273. D-L: E. calycina, D, Spies 2995, with 12 bivalents at
diakinesis; E&F, Spies 3272, E, with 12 bivalents at metaphase I; F, with 13 bivalents at metaphase I; G, Davidse 33828 with 12
bivalents and 2 B chromosomes at diakinesis (indicated by arrows); H, Davidse 33371, with 12 bivalents and a B chromosome at
diakinesis (indicated by an arrow); I, Spies 3198, with 26 chromosomes during early anaphse I; J, Spies 3003, with 2 B chromosomes
during metaphase I (non-alignment of chromosomes indicated by arrows); K & L, Davidse 33321, with 24 bivalents at diakinesis and 3
lagging, late dividing bivalents at anaphse I (indicated by an arrow). D & J, x 730; rest, x 1200.
Bothalia 19,1 (1989)
129
FIGURE 3.— Meiotic chromosomes
in the Calycina group: A & B,
E. delicatula , Davidse 33285,
with 12 bivalents and a very dis-
tinct and narrow spindle, which
may be off-centre, B, X 1100.
Three specimens of the Ramosa group were counted
and the counts represent the first chromosome number
reports for these species. The E. ramosa subsp. ramosa
specimen was a normal diploid plant (Figure 5 A & B).
The E. rehmannii subsp. rehmannii specimen was a
diploid with 0-5 B chromosomes (Figure 5C — E).
During metaphase I the B chromosomes were indis-
tinguishable from the euchromosomes but at the end of
anaphase I they seemed to despiralize (Figure 5D). It
seems as if the division of these normal bivalents into
chromosomes and the division of the univalents into
chromatids is not synchronized. The E. rehmannii
subsp. subspicata specimen had the second highest
ploidy level found in the Ehrharteae, with n = 36. A low
frequency of multivalents (up to hexavalents) were ob-
served in this subspecies (Figure 5F & G). In addition,
FIGURE 4. — Meiotic chromosomes
in the Villosa group. A & B, E.
villosa var. villosa. Spies 3505,
A, 60 bivalents in metaphase;
B, 120 chromosomes during
anaphase I. Additional chromo-
some-like images in the photo-
graph were caused by crystaliza-
tion of stain which can be differ-
entiated from the chromosomes
under the microscope, x 930.
130
Bothalia 19,1 (1989)
FIGURE 5. — Meiotic chromosomes in the Ramosa group. A & B: E. ramosa subsp. ramosa , Spies 3490, with 12 bivalents. A, early
metaphase I; B, diakinesis. C— E: E. rehmannii subsp. rehmannii , Davidse 33708: C, metaphase I with 12 bivalents; D, anaphase I with 2
univalents dividing precociously into chromatids (indicated by arrows); E, anaphase I with unequal (12-14) distribution of chromo-
somes). F & G, E. rehmannii subsp. subspicata , Spies 3515, with 72 chromosomes which include multivalents (indicated by arrows). G,
x 745;A,B,C,E&F, X 1000;D, x 1200.
up to six univalents were observed during diakineses. No
laggards were observed in this specimen at anaphase I
and the rest of meiosis was normal.
The only representative of the Dura group studied,
was a diploid E. dura specimen with a B chromosome
present in approximately 90% of the cells studied (Figure
6A— C). The B chromosome appears as a round, lightly
stained body during diakinesis (Figure 6B). It is present
in all stages and during telophase I it is excluded from
the nucleus (Figure 6C).
In summary, the basic chromosome number of the
genus Ehrharta is 12 and ploidy levels range from di-
ploid to decaploid. Based on one earlier count for one
specimen of E. calycina with 2n=30, Spies & Voges
(1988) suggested that a basic chromosome number of six
is possible. However, the current, much more extensive
results, indicate that the 2n=30 plant is probably an
aneuploid with six additional chromosomes. It is also
interesting to note that ploidy levels exceeding tetraplo-
idy, were restricted to the eastern part of the distribution
area of the genus. Further cytogenetic studies on Ehr-
harta should include the production of artificial hybrids
between the different species and meiotic analyses of
these hybrids to clarify the genomic relationships in the
genus.
Bothalia 19,1 (1989)
131
ACKNOWLEDGEMENTS
Dr R. P. Ellis is thanked for his comments on the
manuscript and useful suggestions during this study. We
also thank Dr G. E. Gibbs Russell for identifying the
plant specimens, as well as for many useful suggestions
during the study. Dr Davidse’s work was made possible
by a research fellowship from the Department of Agri-
culture and Water Supply.
REFERENCES
CARNOY.J. B. 1886. La cytodierese de l'oeuf. Cellule 3: 1—92.
CLAYTON, W. D. & RENVOIZE, S.A. 1986. Genera graminum —
grasses of the world. Kew Bulletin Additional Series 13: 1—389.
DARLINGTON, C. D. & LA COUR, L. F. 1976. The handling of
chromosomes. Allen & Unwin, London.
DAVIDSE, G. HOSHINO, T. & SIMON, B. K. 1986. Chromosome
counts of Zimbabwean grasses (Poaceae) and an analysis of poly-
ploidy in the grass flora of Zimbabwe. South African Journal of
Botany 52: 521-528.
DE WET, J. M. J. 1960. Chromosome numbers and some morphologi-
cal attributes of various South African grasses. American Journal
of Botany 47: 44— 50.
DE WET, J. M. J. & ANDERSON, L. J. 1956. Chromosome numbers
in Transvaal grasses. Cytologia 21: 1-10.
FERNANDES, A. & QUEIROS, M. 1969. Contribution a la connais-
sance cytotaxonomique des Spermatophyta du Portugal. I.
Gramineae. Boletim da Sociedade Broteriana 43: 3— 140.
GIBBS RUSSELL, G. E. 1987. Preliminary floristic analysis of the
major biomes in southern Africa. Bothalia 17: 213—227.
GIBBS RUSSELL, G. E. & ELLIS, R. P. 1987. Species groups in the
genus Ehrharta (Poaceae) in southern Africa. Bothalia 17:
51-65.
GOLDBLATT, P. 1978. Analysis of the flora of southern Africa: its
characteristics, relationships and origins. Annals of the Missouri
Botanical Garden 65: 369—436.
HOSHINO, T. & DAVIDSE, G. 1988. Chromosome numbers of
grasses from southern Africa. I. Annals of the Missouri Botanical
Garden 75. In press.
KELLOGG, E. A. & CAMPBELL, C. S. 1987. Phylogenetic analysis
of the Gramineae: 310—322. In T. R. Soderstrom, K. W. Hilu,
C. S. Campbell & M. E. Barkworth, Grass systematics and evolu-
tion. Smithsonian Institution Press, Washington D C.
LOVE, R. M. 1948. Preliminary cytological studies of Ehrharta caly-
cina Smith. American Journal of Botany 35: 358—360.
PARTHASARATHY, N. 1939. Cytogenetical studies in Oryzeae and
Phalarideae. II. Cytological studies in Phalarideae. Annals of
Botany 3: 43—58.
132
Bothalia 19,1 (1989)
RAVEN, P. H., KYHOS, D. W. & HILL, A. J. 1965. Chromosome
numbers of spermatophytes, mostly Californian. Aliso 6:
105-113.
SODERSTROM, T. R. & ELLIS, R. P. 1987. The position of bamboo
genera and allies in a system of grass classification: 225-238. In
T. R. Soderstrom, K. W. Hilu, C. S. Campbell & M. E. Bark-
worth, Grass systematics and evolution. Smithsonian Institution
Press, Washington D.C.
SPIES, J. J. & DU PLESSIS, H. 1986. Chromosome studies on Afri-
can plants. 1 . Bothalia 16: 87-88.
SPIES, J. J. & VOGES, S. P. 1988. Chromosome studies on African
plants. 7. Bothalia 18: 1 14- 1 19.
TATEOKA, T. 1957. Notes on some grasses. VI. (9. Coelachne and
Sphaerocaryum. 10. Chromosomes and leaf structure of several
species). Botanical Magazine 827: 1 19- 125.
TATEOKA, T. 1965. Chromosome numbers of some East African
grasses. American Journal of Botany 52: 864—869.
TOTHILL, J. C. & LOVE, R. M. 1964. Supernumerary chromosomes
and variation on Ehrharta calycina Smith. Phyton 21:21-28.
WATSON, L., DALLWITZ, M. J. & JOHNSTON, L. R. 1986. Grass
genera of the world: 728 detailed descriptions from an automated
database. Australian Journal of Botany 34: 223 — 230.
Bothalia 19,1: 133-136(1989)
Miscellaneous notes
VARIOUS AUTHORS
TWO NOMENCLATURAL PROBLEMS INVOLVING ARTICLE 63
While preparing a course in plant nomenclature in
1982, I endeavoured to find examples of superfluous
names in South African taxonomic literature. Among
others, I found the following two cases but, in the first,
the rules, since the 1987 Berlin Congress, have changed
to make the second option in the example a case of later
synonymy rather than superfluity. However, it is such an
interesting case, that it deserves mention.
1 . Silene dewinteri or S. constantia?
In their protologue of S. constantia, Ecklon & Zeyher
included, in synonymy, S. crassifolia L. var. angustifo-
lia Bartl. (1832), based on Ecklon specimen/s from the
vicinity of ‘uppermost blockhouse’ at Devil’s Peak.
Bocquet in his ‘Observations’ on S. dewinteri stated that
because Ecklon & Zeyher cited S. crassifolia var. angus-
tifolia in synonymy under S. constantia the latter was
‘automatically typified by the type of the variety’.
According to Dr R. K. Brummitt of Kew, this could be
construed to constitute lectotypification of S. constantia
by the Ecklon ‘blockhouse’ specimen/s which, however,
Bocquet referred to 5. clandestina Jacq. However, it is
not certain whether this does constitute lectotypification
because, by definition (Article 7.5 of the International
code of botanical nomenclature, 1988), a lectotype can
only be a single specimen or illustration and Bocquet
clearly referred to specimens (plural). Unfortunately in
Bartling’s description of 5. crassifolia var. angustifolia
there is no indication of whether the varietal name is
based on one or more Ecklon specimens. I have corre-
sponded with Mr Daniel Masson of Geneva (G), who is
continuing with Bocquet’ s revision of the southern Afri-
can species of Silene, to establish whether one or two
specimens were involved and whether there were annota-
tions by Bocquet on the specimen/s. In his reply (letter of
1988.10.08) Masson mentions that he has two specimens
on loan from Stockholm which may be relevant. The
first is an Ecklon specimen bearing the names S. crassi-
folia var. angustifolia (det. Bartling, 1831) and 5. thun-
bergiana. No locality is given, but the label carries the
number 53 which may refer to ‘Stellenbosch,
Houwhoeksbergen’ , if it relates to the Ecklon & Zeyher
localities listed by Drege in Linnaea 19: 583-598
(1847). The second specimen, anonymous as to collec-
tor, bears three names viz. S.flexuosa (ined.), S. crassi-
folia var. angustifolia and S. constantia. The number 85
appears on the sheet and if it is an Ecklon & Zeyher
number (as listed by Drege in Linnaea 20: 258 (1847), it
refers to ‘eastern side of Table Mountain at Constantia’.
Masson identifies this specimen as 5. clandestina and
suggests that it is an Ecklon specimen. However, if the
number 85 is an Ecklon & Zeyher number, then the
locality is incorrect for the type of S. crassifolia var.
angustifolia. The same applies to the first specimen. On
the evidence available, it seems doubtful whether either
specimen is the Ecklon type of S. crassifolia var. angus-
tifolia from the ‘uppermost blockhouse’ at Devil’s Peak.
Neither specimen has been annotated by Bocquet -let
alone as lectotype of S. constantia.
Therefore, there are two aspects to the Silene dewin-
teri versus 5. constantia problem, one in which the lecto-
typification is regarded as having been effected and the
other not. The two aspects will be examined in turn.
1 . 1 Lectotypification effected
This situation concerns lectotypification of Silene con-
stantia irrespective of whether it is correct or not. Boc-
quet clearly stated that the description of 5. constantia,
but not the type, agreed with his concept of S. dewinteri.
In short, he admitted a discrepancy or conflict between
the designated lectotype of 5. constantia and its proto-
logue, specifically the description. What are the practical
implications of these facts? Two implications come to
mind: 1, if one ignores Bocquet’ s incorrect lectotypifica-
tion of S. constantia, then 5. dewinteri remains a correct
name and S. constantia remains a synonym of S. clan-
destina-, 2, if one rectifies the lectotypification, then S.
constantia becomes the correct name and 5. dewinteri a
later legitimate synonym in the context of Note 2 of
Article 63.1 (a superfluous name pre-the 1987 Berlin
Congress). There is no problem in designating a new
lectotype of S. constantia. Ecklon and Zeyher in describ-
ing S. constantia clearly used their own gathering Ecklon
& Zeyher s.n.: the specimen agrees with the description
and was collected near Constantia and Hottentots Hol-
land. However, such a step, if considered advisable, is
best left to someone working on the genus.
An obvious question arises. Why did Bocquet assert
that S. constantia was automatically typified by the type
of the cited synonym, 5. crassifolia var. angustifolia ? A
possible explanation is that Bocquet erroneously invoked
Article 7.11 (previously 7.9), which states that ‘a nomen
novum for an older name is typified by the type of the
older name’. The way in which Bocquet phrased the
typification, viz. ‘typified by the type of the variety
(older name*)’ seems to point to this. It is doubtful
whether Bocquet, in spite of possibly effecting lectotypi-
fication, deliberately intended lectotypifying 5. con-
stantia. Significantly, he did not use the word ‘lectotypi-
fication’.
* my parenthesis.
Bocquet in Bothalia 12: 309-311 (1977) described
Silene dewinteri as a new species with Bocquet 17774 as
holotype. He cited in synonymy ‘5. constantia Eckl. &
Zeyh., Enum. 32 (1834) quoad descr., typo et synony-
mia excludendis’ .
134
Bothalia 19,1 (1989)
1 .2 Lectotypification not effected
In this case one has the option of designating a lecto-
type for S. constantia, but this is not mandatory (Article
7.5 and Recommendation 7B). If a lectotype is desig-
nated, the Ecklon & Zeyher s.n. specimen mentioned
above would be a good choice. The result of this action,
as already indicated, would be that S. constantia be-
comes the correct name and S. dewinteri a later syno-
nym.
2. Antholyza caffra
In a note on Anapalina caffra (Ker ex Bak.) Lewis in
Journal of South African Botany 37: 235 (1971) Gold-
blatt states that Antholyza caffra Ker ex Bak. (1892) was
not a superfluous name, even though the earlier Anisan-
thus splendens Sweet (1831), a misidentification, was
cited in synonymy and he cites Stafleu’s interpretation of
the Seattle modification of Article 63 in support [see
Stafleu in Taxon 19: 41—42 (1970)]. The Seattle modifi-
cation relates to the explicit or implicit exclusion of the
type of the cited name. Stafleu interprets this as follows:
‘When it can be shown that the type of a cited name
cannot within reason have been included by the author
within the circumscription of his new taxon, his name
does not become automatically superfluous on account
of the mere citation of the older name’. Goldblatt writes:
‘As Baker probably named this plant Antholyza caffra
thinking that Ker’s name had priority over Anisanthus
splendens and because the latter cannot be included in
Baker’s circumscription of this species the question of
superfluity cannot really be raised, particularly if Staf-
leu’s interpretation of the modification of Article 63 is
followed’ .
Let us analyse Goldblatt’ s statement. Firstly, he states
that superfluity is not at issue because Baker believed
that Antholyza caffra Ker (1805) had priority over An-
isanthus splendens ( 1 83 1 ) . But Antholyza caffra Ker was
a nomen nudum and therefore not validly published. By
supplying a description Baker validated the name for the
first time and by citing Anisanthus splendens as a syno-
nym [‘the citation of the name itself (Article 63.2)]
without excluding its type either explicitly or implicitly
(there is no evidence of such exclusion) the name Antho-
lyza caffra becomes superfluous. Secondly, Goldblatt
asserts that because Anisanthus splendens, as figured and
described by Sweet, cannot be included in Baker’s cir-
cumscription of Antholyza caffra, the question of super-
fluity cannot be raised. The fact is that whatever taxo-
nomists of today think about the relationship between the
two species. Baker himself regarded Antholyza caffra as
conspecific with the earlier Anisanthus splendens. This
is not surprising if, as pointed out by Goldblatt, ‘the
plants are similar’ , though they are now known to belong
to different genera. Weresub & Hennebert [Taxon 12,6:
218—228 (1963)] would call this a case of facultative
superfluity involving a facultative synonym as opposed
to nomenclatural superfluity involving an obligate syno-
nym.
Clearly there seems to have been a misinterpretation
of the Seattle modification of Article 63, since the ques-
tion of explicit or implicit exclusion does not arise at all.
What is the implication of Antholyza caffra being super-
fluous? The name Antholyza caffra, being illegitimate,
the epithet caffra can only be used in Anapalina if the
combination Anapalina caffra is treated as new dating
from 1960 and attributed solely to Lewis.
I thank Dr R. K. Brummitt of the Royal Botanic Gar-
dens, Kew, for helpful comments on these two cases.
D. J. B. KILLICK
MS. received: 1988.02.15.
THE CARYOPSIS SURFACE OF PENTAMERIS AND PSEUDOPENTAMERIS ( ARUNDINOIDEAE, POACEAE) REVISITED
Barker (1986) reported on the structure of the surface
of the caryopses of five taxa in Pentameris Beauv. and
one in Pseudopentameris Conert. This study showed
there to be three types of surface sculpturing (colliculate,
rugose and reticulate) and three types of caryopsis shape
(cuneate, elliptic and globose-truncate). Free stylar hairs
were observed in all the examined taxa of Pentameris,
where they appear as a crown of short weak hairs (see
Barker 1986 for photomicrographs of these structures).
These structures were, however, absent from the cary-
opses of Pseudopentameris macrantha.
Subsequent to the work of Barker (1986), Clayton &
Renvoize (1986) have defined the fruit of Pentameris as
an achene, while that of Pentaschistis Stapf is considered
to be a caryopsis. This difference is recognized, and the
term caryopsis is used here in the broad sense, as advo-
cated by Sendulsky etal. (1987).
The caryopsis of Pseudopentameris brachyphylla was
predicted by Barker (1986) to have a narrowly elliptical
shape, reticulate surface sculpturing and no free stylar
hairs. The caryopsis of Pentameris longiglumis was ex-
pected to have an elliptic shape, colliculate surface fea-
tures and free stylar hairs.
This study was carried out to test these predictions,
and to augment the data on caryopsis structure in the
southern African Arundineae.
MATERIALS AND METHODS
Caryopses were obtained from herbarium specimens
housed in the National Herbarium, Pretoria (PRE). They
were gold-coated after being mounted on stubs using
two-sided sticky tape. Specimens were examined using
an ISI-SX-25 Scanning Electron Microscope. Photo-
graphs were taken using Tura 60 x 70 mm format black
and white 100 ASA film.
Specimens examined:
Pentameris longiglumis
CAPE. -33 18 (Cape Town): Table Mountain (-CD), Marloth
3078.
Pseudopentameris brachyphylla
CAPE. — 3419 (Hermanus): Die Mond se Kop (-AD), Barker 58.
Bothalia 19,1 (1989)
135
FIGURE 1. — A & B, Pentameris longiglumis, Marloth 3078: A, slightly malformed caryopsis, with colliculate sculpturing and free stylar
hairs, scale bar = 450 /ct; B, close-up of colliculate sculpturing, scale bar = 40 p. C & D, Pseudopentameris brachyphylla, Barker 58: C,
stylar end of caryopsis, note the reticulate sculpturing, linear-elliptical shape and absence of stylar hairs, scale bar = 420 p: D, high
magnification of reticulate sculpturing, scale bar = 155 p.
RESULTS AND DISCUSSION
The morphology of the caryopses of these two taxa
corroborate the predictions made in an earlier publication
(Barker 1986). Figure 1A shows the entire caryopsis of
Pentameris longiglumis. Although slightly malformed, it
is basically elliptical in shape, and the surface is of a
colliculate nature (Figure IB). This structure compares
closely with those of the previously examined Pentame-
ris obtusifolia, P. macrocalycina, and to a lesser extent,
P. thuarii, in that the former two species have elliptical
caryopses with colliculate sculpturing and free stylar
hairs (Barker 1986), while the latter species differs only
in having a globose-truncate caryopsis.
The caryopsis of Pseudopentameris brachyphylla is,
as predicted, narrowly elliptical and reticulately sculp-
tured, with no free stylar hairs. Figure 1C shows the
distal or stylar half of the grain. The remains of the style
and style base may be seen, but the small weak hairs,
which are obvious in Pentameris longiglumis, are lack-
ing. Figure ID shows the reticulate nature of the surface
of the caryopsis. The caryopsis of this species is visually
indistinguishable from that of the previously examined
Pseudopentameris macrantha.
Both species examined here are rare, and only a few
herbarium specimens were available. The small sample
size may be construed as inadequate, especially since the
caryopsis of the Pentameris longiglumis specimen is
slightly malformed. However, the results obtained con-
firm the predictions made previously (Barker 1986), and
a complete character set for all presently known taxa in
both genera is now available. These data are outlined in
the character by taxon matrix (Table 1) which is an up-
date of the table published earlier (Barker 1986).
Further work on Pentaschistis, Pentameris and Pseu-
dopentameris is in progress (Barker in prep., Linder
pers. comm, and Ellis pers. comm.), which will throw
further light on the taxonomy and phylogeny of these
related genera.
CONCLUSION
Species of Pentameris have elliptic or globose cary-
opses with colliculate sculpturing and free stylar hairs,
whereas the species of Pseudopentameris have narrowly
elliptic caryopses, reticulate surface features and no free
stylar hairs.
This study has provided additional information and
improved the understanding of the variation within and
between the genera Pentameris and Pseudopentameris.
The taxonomic importance of characters such as the pre-
sence or absence of free stylar hairs, has been reiterated.
136
Bothalia 19,1 (1989)
TABLE 1. — Caryopsis characters for all species in the genera Pentameris and Pseudopentameris (updated from Barker 1986)
ACKNOWLEDGEMENTS
I thank Dr R. P. Ellis, Dr G. E. Gibbs Russell, Mr
E. R. Robinson, Mrs L. Fish and two referees for their
comments on earlier manuscripts, Mrs A. Romanowski
for developing and printing the photographs, and Mrs
S. Perold for her help in the use of the SEM. Research
was carried out in part fulfilment for an M.Sc. disserta-
tion, University of the Witwatersrand.
REFERENCES
BARKER, N. P. 1986. The shape and ultrastructure of the caryopsis of
Pentameris and Pseudopentameris species (Arundinoideae, Poa-
ceae). Bothalia 16: 65-69.
BARKER, N. P. in prep. Leaf epidermal features in Pentameris
Beauv. and Pseudopentameris Conert (Arundinoideae, Poaceae).
Bothalia.
CLAYTON, W. D., & RENVOIZE, S.A. 1986. Genera graminum —
grasses of the World. Kew Bulletin Additional Series XIII. Her
Majesty’s Stationery Office, London.
SENDULSKY, T., FILGUEIRAS, T. S. & BURMAN, A. G. 1987.
Fruits, embryos, and seedlings. In The Systematics and evolution
of grasses. Smithsonian Institution Press, Washington D.C.
N. P. BARKER*
* Botanical Research Institute, Department of Agriculture and Water
Supply, Private BagXJOl, Pretoria 0001.
MS. received: 1988.06.22.
Bothalia 19,1: 137-139(1989)
OBITUARY
RUDOLF GEORG STREY (1907- 1988)
Rudolf Georg Strey (Figure 1) was bom in Templin in
the Uckermark region north of Berlin, Germany, on the
28th of April, 1907. His father, Wilhelm Georg, was a
pharmacist with a concession for the only pharmacy in a
town of some 7 000 inhabitants. Wilhelm Georg later
took over his brother’s pharmacy at the outbreak of the
First World War, in Stettin, and Strey attended school
there at the Friedrich-Wilhelm Gymnasium from 1914 to
1918. In 1916 Strey ’s father rented the Adler Pharmacy
in Angermunde in Neumark, halfway between Stettin
and Berlin, and Strey completed his schooling at the
Uckermarkische Real-Pro Gymnasium, in Angermunde,
from 1918 to 1923.
Strey was the second son by two years, and as the
tradition was at the time, the first son followed the
father’s profession and the second son was sent to study
agriculture. Strey attended a number of agricultural insti-
tutions between 1923 and 1928, in Genschmar im Oder-
bruch, at Gross Lubras in West Havelland, at Gross
Santersleben in the Magdeburger Borde, and finally at
Graeningen near Rathenow where his father had opened
the Grime Pharmacy after the war. In 1927—1928 he
went to an agricultural college, the Hohere Lehranstalt
fur Praktische Landwirte in Potsdam, and qualified with
the diploma ‘Staatlich gepriifter Landwirt’.
Owing to the depression and devaluation of the
currency in Germany after the war, and the Wall Street
disaster, Wilhelm Georg offered his son a passage to one
of the German colonies where his prospects would be
better. Strey sailed for Cape Town and arrived in South
West Africa in October 1929. He became apprenticed to
A. Stauch, a diamond merchant, to work on his farm
Dordabis, and it was here that he met his first wife on the
neighbouring farm Ibenstein. Shortly afterwards
Stauch’ s business went bankrupt and Strey attended a
course on karakul breeding at Neudamm Agricultural
College near Windhoek.
Strey’s father and uncle came out to SWA in 1930,
and Strey started on his own on the government farm
Buellspoort No. 172 District Rehoboth, near the Nau-
kluftberge. His father helped him buy the farm, but he
stayed only a short while before returning to Germany.
The farm proved profitable for dairy cattle until the big
drought of 1933, after which Strey switched to farming
with grey and black karakul. During this period he
became seriously ill with a combination of Malta fever,
amoebic dy sentry and malaria, and he returned to
Germany to the Tropen Institute Tubingen for treatment.
Strey developed an interest in botany because, as a
farmer, he wanted to know the plants that were either
palatable or poisonous to his stock. As there was little
literature available he made efforts to consult various
botanists. Dinter visited him at Buellspoort in December
1934 and he also met Prof. O. H. Volk and E. B.
Schoenfelder. In 1940 Strey was interned, first at
Baviaanspoort and Koffiefontein, and later he was sent
to the Andalusia Internment Camp at the present Jan
Kempdorp, near Warrenton, in the N Cape. A number of
visiting German botanists were also caught by the war,
and Prof. Volk, Dr Mueller-Stoll, Dr Rehm and Dr
Cloos started a University-type botany course in the
camp. With the help of other scientists, this course
started in 1940 and lasted until 1946. Students were
examined regularly and after the war the German univer-
sities acknowledged the course. Other lectures on
poisonous plants, ecology, pasture management and in-
digenous plants interested many of the farmers.
Strey built up a new Index of SWA plants, at the
suggestion of Prof. Volk, and he combined the names
from Dinter’ s index with Range’s Flora and the her-
barium lists of Schoenfelder and Rusch. He then started
a card index for all correct plant names with information
on literature, synonyms, types and voucher collections.
During this time the whole of Phillips’ Genera of South
African Flowering Plants was translated into German.
Strey’s index is housed at PRE, along with his indexes of
collectors, localities and botanical literature relating to
SWA.
On his 37th birthday, towards the end of the intern-
ment, Strey was presented with the following eulogy
which is framed and housed in the Natal Herbarium
museum:
Streyina aculeata spec. nov. unica.
(post annos 37 in captivitate se evolutans)
Discipulus scientiae amabilis; altus agilisque, perfectae, fere
patientiae, sed non rare distincte mucronatus.
FIGURE 1 .—Rudolf Georg Strey (1907- 1988).
138
Bothalia 19,1 (1989)
Initiator collection^ magnae cartarum; colonus, copias magnas ovium
cultivans, sed magno suo amore versans in regno variegato vegetabilio-
rum; omnes determinandi difficultates facillime superavit.
Habitat: in desertis extremis africae austro-occidentalis, in Porta Tau-
rina.
Coelesti dent ei mox libertatem, vitam longam luxurians et multos
successus.
Deterunt Andalusiae: ante diem quartam Calendas Majas anno Domini
1944.
Signed by O. Volk, H. Herre, A. Giess[?], Matz, Gunter, Kalun etc.
Loosely translated this means:
Streyina aculeata, a unique new species
(developing in captivity after 37 years)
A disciple of the amiable science [botany]; tall and agile, well built,
usually of perfect patience, but not rarely distinctly pointed.
Initiator of a large collection of cards; farmer, cultivating great quanti-
ties of eggs, but avidly studying the many-faceted plant kingdom; he
overcame all difficulties of identification most easily.
Habitat: in the extreme desert of South West Africa in Buellspoort.
May the inhabitants of heaven soon grant him freedom, a long luxu-
rious life and many successes.
Signed in Andalusia: 28.4. 1944.
After internment he was released to Stellenbosch
where he stayed for a few months with H. Herre, who
had resumed curatorship of the Stellenbosch University
Botanic Garden. Strey worked at the gardens and in the
herbarium of the University with Dr Rehm.
At the end of November 1946 he returned to
Buellspoort. Due to ill health, but also because of his
new and different interests, he gave up farming. He
started collecting from all parts of the country, sending
his plants for identification to Kew, Munich, Kirsten-
bosch (National Botanic Gardens) and Pretoria (Botani-
cal Research Institute). Bulbs, seeds and succulents were
sent to Herre in Stellenbosch. Buellspoort became a well
known type locality for plants and insects. Strey was a
keen insect collector, gathering the first blind beetles
from the Namib. He has a number of beetles and moths
named after him by Dr Koch and Prof. Janse of the
Transvaal Museum. He also collected fungi which are
mainly deposited in PREM.
He continued compiling the indexes of SWA collec-
tors, literature, place names and maps, but his chief in-
terest between 1947 and 1952 was archaeology. This
was principally because of his association with the re-
nowned Abbe H. Breuil, an expert in palaeolithic ar-
chaeology, who is perhaps best known for his studies in
cave art. Breuil occupied a chair at Witwatersrand Uni-
versity from 1944 to 1951, and he made three important
trips to SWA between 1947 and 1950. Strey guided and
helped organize these trips and together with Dr Scherz
and others they explored, recorded and mapped prehis-
toric sites in SWA and the northern Cape. A cave in the
Erongo Mountains is named after him. Strey also led an
expedition to the Okavango with Dr Gusinde from
America, in 1950, to study the Taung bushman tribe.
His collections of rock paintings, stone implements
and other museum pieces are housed in the Windhoek
Museum, the Swakopmund Museum and in the archaeo-
logical collections of the Witwatersrand University.
Strey met his second wife on one of the expeditions with
Breuil. She worked as an artist with the group reproduc-
ing various paintings, including many details of the
White Lady shelter in the Brandberg. An article by her
(Rita G. Strey 1962) provides a fascinating insight into
these excursions.
Strey sold the farm Buellspoort in 1950, and he trav-
elled and organized expeditions until going up to Pretoria
in 1953. He obtained employment working for a small
printing press, eventually buying his own press in Ger-
miston. On 17.2.1955 Strey was appointed as a techni-
cian to the Botanical Research Institute in Pretoria, to
work in the National Herbarium.
In 1961 he produced a manuscript on ‘Two hundred
years of Botanical Research in South West Africa’ which
was the culmination of his earlier research. This gives an
outline of the history of botany in SWA, divided up into
periods, with the main collectors listed and the herbaria
in which the collections are housed. The literature of
special interest to SWA botany is recorded, including
maps, ecology, physiology, medicinal and poisonous
plants. Strey also described a species of Cucumis with A.
Meeuse (Meeuse & Strey 1962). This is now a synonym
for Cucumella bryonifolia (Merxm.) C. Jeffrey.
He worked in Pretoria for seven years, living on a plot
in Silverton where he also bred chickens. In April 1962
he was transferred to Natal Herbarium in Durban where
he became curator and later officer-in charge of the
Botanical Research Unit.
His years in Natal marked the scientifically most ac-
tive phase of his career. He revitalized and re-organized
Natal Herbarium, building up a fine museum with early
memorabilia from Medley Wood, and own collections of
insects, wood, fruit and seeds. Strey developed the her-
barium garden, planting large numbers of rare Natal
plants. He realized that before Bews’s The Flora of Na-
tal and Zululand (1921) could be revised, a considerable
amount of collecting was necessary to fill in the gaps. He
collected from as wide an area as possible since many
species were known from only a few specimens, and his
collecting numbers which started at 4150 when he ar-
rived in Durban, reached 11300 by the time of his retire-
ment. Reference to Ross’s Flora of Natal (1972) bears
witness to the effective and untiring work of this
dedicated collector.
Strey also made a point of visiting type localities of
rare or once-collected Natal plants. His relocation of
Elephantorrhiza woodii, not collected since the Medley
Wood type from near Colenso, allowed J. H. Ross to
emend Phillips’s earlier description of the habit of the
species when Ross revised Elephantorrhiza in 1974.
This was possible because Strey succeeded in cultivating
the species at Natal Herbarium.
His earliest research at Natal Herbarium, concerned
the genus Dietes in 1965, and his notes, dissections of
petals and boxes of fruit and seeds are housed in PRE.
He then took considerable interest in palms, particularly
Hyphaene and Raphia. The latter resulted in his describ-
ing a new species with Mrs A. A. Mauve (Obermeyer &
Strey 1969).
Strey concentrated particularly on the rich flora of the
sandstone areas of southern Natal and Pondoland, with
Bothalia 19,1 (1989)
139
its many endemic species. He often collected with Mr
H. B. Nicholson of Skyline, St Michael’s-on-Sea, and
together they worked the region intensively. His publica-
tions arising from this activity are Strey & Leistner
(1968) and Strey (1972). Barringtonia racemosa drew
his attention and was written up for the Flowering Plants
of Africa (Strey 1973a).
Strey developed a great interest in the family Aralia-
ceae and his intensive research on Cussonia gave rise to
the following publications: Strey 1973b, 1975, 1981.
His original research in the Araliaceae also contributed
to, and was acknowledged in Burtt & Dickison 1975.
Plant species names which commemorate Strey are
Pseudosalacia streyi Codd (Celastraceae), Crassula
streyi Toelken (Crassulaceae), Sonderina streyi Merxm.
(Apiaceae); Piaranthus streyianus Nel (Asclepiadaceae),
nom. nud. and now a synonym for Orbea rangeana
(Dinter & Berger) Leach; Lithops streyi Schwantes (Me-
sembryanthemaceae), now a synonym for L. gracilideli-
neata Dinter [although this is not cited in the new Li-
thops book by Desmond T. Cole (1988).]; Lapeirousia
streyi Suesseng. (Iridaceae), now a synonym for L. cau-
data Schinz subsp. burchellii (Bak.) Marais & Goldblatt;
Turraea streyi F. White & B. T. Styles (Meliaceae);
Anthospermum streyi Puff (Rubiaceae); Indigofera
streyana Merxm. (Leguminosae), now /. hochstetteri
Bak. subsp. streyana (Merxm.) A. Schreib. and Indigo-
fera herrstreyi (my manuscript name).
Besides his published research, Strey built up very
comprehensive biographical notes on early Natal botani-
cal collectors, in particular M. J. Me Ken, J. Medley
Wood and Wilhelm Keit. Most of his notes, accumulated
during many hours of research in museums and libraries
in South Africa and Europe (particularly on one overseas
trip in 1978), are housed either at the Killie Campbell
Museum or the Natal Herbarium in Durban.
Strey retired at the end of April 1972, but was re-em-
ployed for a further three years, until the end of 1975,
before retiring permanently to his cottage at the Village
of Happiness, near Margate. He remained actively inter-
ested in Cussonia, growing a fairly representative collec-
tion of species around his home. Today his cottage can
still be easily singled out from a distance, as it is sur-
rounded by these distinctive plants. Rudolf Georg Strey
died peacefully on the 30th of June 1988 and was cre-
mated according to his wishes. He leaves two daughters
and a son by his first marriage, and one daughter by his
second marriage.
ACKNOWLEDGEMENTS
I would like to thank the following very much indeed
for all their help in providing certain details: Mr Rolf
Strey, Miss R. Williams, Mrs H. E. Noble, Mr H. B.
Nicholson and Dr O. A. Leistner, and Dr H. Glen for the
Latin translation.
REFERENCES AND PUBLICATIONS BY STREY
BURTT, B. L. & DICKISON, W.C. 1975. The morphology and
relationships of Seemannaralia (Araliaceae). Notes from the
Royal Botanic Garden, Edinburgh 33:449 —464.
MEEUSE, A. D. J. & STREY, R. G. 1962. Cucumis umbrosus A.
Meeuse & Strey. In A. D. J. Meeuse, The Cucurbitaceae of
southern Africa. Bothalia 8: 67.
OBERMEYER, A. A. & STREY, R. G. 1969. A new species of
Raphia from northern Zululand and southern Mozambique. Bo-
thalia 10: 29-37.
ROSS, J. H. 1974. The genus Elephantorrhiza. Bothalia 11:
241-251.
STREY, RITA, G. 1962. A tribute to M. l'Abbe H. Breuil. Lantern
11:48-57.
STREY, R. G. 1969. A note on the identity of Mesembryanthemum
brachyphyllum Welw. Bothalia 10: 55.
STREY, R. G. 1972. A new species of Eugenia from Pondoland and
southern Natal. Bothalia 10: 569—571.
STREY, R. G. 1973a. Barringtonia racemosa (L.) Roxb. Flowering
Plants of Africa 43: t.1706.
STREY, R. G. 1973b. Notes on the genus Cussonia in southern Af-
rica. Bothalia 11: 191-201.
STREY, R. G. 1975. A new species of Cussonia from the southern
Cape, Cussonia gamtoosensis. Bothalia 1 1: 515— 521 .
STREY, R. G. 1981. Observations on the morphology of the Aralia-
ceae in southern Africa. Journal of Dendrology 1 : 66—83.
STREY, R. G. & LEISTNER, O. A. 1968. The rediscovery of Rhyn-
chocalyx lawsonioides Oliv. Journal of South African Botany 34:
9— 13.
B. D. SCHRIRE
Bothalia 19,1: 141 - 143(1989)
Book Reviews
COMBRETACEAE IN SOUTHERN AFRICA by J. D. CARR. 1988.
Tree Society of southern Africa, P.O. Box 4116, Johannesburg 2000.
Pp. 236. Price R55,00 + GST.
This book covers the six genera which occur in South Africa, SWA-
/Namibia, Botswana, Swaziland and Zimbabwe. Unrest precluded the
inclusion of Angola, Zambia and Mozambique, but the author men-
tions for each species whether records exist from regions he was unable
to work in.
Carr's book is a practical ‘field identification guide based on certain
fruit characters, with species separated first of all by mature bark pat-
tern and then by reference to habit, habitat, average leaf size and
flowering period — in that order — followed by maximum and then
average dimensions of fruit’. The author’s descriptions of the plants are
mainly compiled from live material in the field but he used herbarium
material in order to account for variation within the species.
A key for the six genera is provided but, because four of the six are
represented by only one or two species in the region, the author ex-
cludes them from the specific guides for Combretum and Terminalia.
Carr describes each species in great detail. He provides synonyms,
common names, national tree numbers, as well as distribution maps
(with arrows indicating points on the map too small to shade). He also
provides geographical information in the form of regions and place
names. He then discusses the habit, habitat, bark, foliage, inflores-
cence, fruit and propagation and cultivation. Aside from the colour
plates by Elise Buitendag in the front of the book, the author provides a
full set of black and white photographs for each species. These photos
usually include the habit, inflorescences, fruit, the seedling and varia-
tion of bark patterns.
I tested the key using two people who had little knowledge of the
Combretaceae with specimens found in the garden. Five out of seven
species were identified quite easily and the other two required some
prompting.
Problems experienced were: (a) the key only provides the species
name and not the page on which the species description occurs. How-
ever, the author has arranged the genera and species in alphabetical
order, which helps to overcome this irritation; (b) the glossary is in-
complete; terms unfamiliar to non-taxonomists such as arcuate, reticu-
late and caducous, although found in the text, are not in the glossary;
(c) the use of the length of fruit for initial differentiation of species
caused some confusion. Carr divides the species with a 4-winged fruit
into three groups: 1, up to 25 mm long; 2, up to 40 mm long; 3, up to
77 mm long. The reader must realise that if the fruit of his specimen is
25 mm long, it does not necessarily fall into group 1. A fruit which is
60 mm long, however, can obviously only belong to group 3.
Apart from the minor irritations I have mentioned, this book is of
tremendous value not only to botanists who have to identify plants in
the field and horticulturists who need to know more about the propaga-
tion and cultivation of these indigenous plants, but also to laymen who
want to know more about our flora.
From an ecologist’s point of view this key is of immense value. The
author uses characters which are available at all times, with possible
exception of fruit, and his detailed habitat and distribution data are also
a great help. He covers the variation within a species admirably.
Denzil Carr has spent 12 or more years working on this book, travel-
ling extensively, using his own funds, to produce field notes on each
species in its natural habitat. He can, in my opinion, feel well satisfied
with his latest labour of love.
M. D. PANAGOS
SOUTHERN AFRICAN BOTANICAL LITERATURE 1600-1988
SABLIT compiled by A.S. KERKHAM. South African Library , P.O.
Box 496, Cape Town, 8000. Pp.256. 16 plates. ISBN 0 86968 083 8.
Price R35,00 + GST.
This book, Southern African botanical literature (SABLIT), began
as a revision of Flora africana, a work little known among South
African botanists, compiled in 1963 by W. Tyrrell-Glynn with the
assistance of M. R. Levyns in commemoration of the 50th anniversary
of Kirstenbosch. With the 75th anniversary celebrations taking place in
1988, the South African Library (SAL) offered to produce a revised
version. This involved more work than was first envisaged and several
important national bibliographies had to be consulted. The coverage of
Flora africana was 1600-1963 and included only South Africa. The
coverage of SABLIT has been extended to 1988 and includes southern
Africa ‘on a selective basis’ — unfortunately not defined, but obviously
including the RSA, Lesotho, Namibia, Swaziland, Zimbabwe and
Malawi, but excluding Mozambique and Botswana. The original Flora
africana was a selective bibliography based solely on the SAL collec-
tion. This is not the case with SABLIT: coverage has been widened and
major library collections have been consulted. The present biblio-
graphy has been brightened by the inclusion of 16 colour plates in-
tended to show the development of botanical illustration. An additional
plate, representing 'Oxalis speciosa' from Jacquin’s Oxalis (1784)
appears on the outer front cover.
In the foreword it is stated that the bibliography ‘is not intended to be
a masterpiece of bibliographical research, as detailed bibliographic
information of that nature is already available in F.A. Stafleu’s (&
Cowan’s!) multivolume Taxonomic literature.’ It should be pointed out
that Taxonomic literature ( 1976— 1988) covers the period 1753— 1940,
whereas SABLIT purports to cover the period 1600-1988 i.e. there is a
201-year difference in coverage between the two bibliographies. Fur-
thermore, Taxonomic literature includes only authors who have pub-
lished at least three generic names. This automatically excludes scores
of southern Africa authors, especially non-taxonomists. The point be-
ing made is that coverage of southern African literature in Taxonomic
literature is limited, because of the time period and generic name
criteria applied.
In the introduction the compiler, A S. Kerkham, describes the items
included in SABLIT. The main items included are literature references
dealing with botanical history, taxonomy and ecology, but excluding
horticulture, plant anatomy and physiology. Manuscripts, theses, jour-
nal articles, textbooks and juvenile books have been excluded. Certain
exceptions have been made where information in, say, a journal or
thesis is considered to be of a particular botanical interest.
The bibliography proper is preceded by an account of South African
botany (note: not southern African botany) with particular reference to
historical background and botanical illustration. In the foreword it is
said that this account is based on a paper delivered at Bibliophilia
Africana, the 4th South African Conference of Bibliophiles held in
September 1981. Unfortunately, the author of this paper is not
revealed, consequently the paper is not cited in the references used.
This historical account obviously derives much of its content from
previous works, such as Phillips (1930), Levyns (1970-1976), Dyer
(1977) and Gunn & Codd (1981) and is a concise, readable essay, but it
should not have been entitled ‘South African Botany’, since it is
restricted to works dealing with the flora and excludes vegetation. In
the discussion of Recent Publications a very notable omission is the
Flora of southern Africa (FSA) launched in 1963 to replace the now
outdated Flora capensis (1895- 1933). In the FSA series 24 volumes or
parts thereof have been published so far. Surely this definitive work
warrants inclusion rather than some of the somewhat light-weight
works included in the account. Another comment on Recent Publica-
tions. The compiler has been brave enough to single out a certain artist
as ‘currently South Africa’s leading botanical artist’. As editor of The
Flowering Plants of Africa for 15 years, I have had a lot to do with
botanical art and botanical artists but, in spite of this, I would not have
the temerity or presumption to choose South Africa's leading botanical
artist. The situation is that there is such a wealth of talent in botanical
art in the RSA at present, that it would be extremely difficult if not
impossible to designate any one artist as the best in the field.
The compiler also pronounces judgement on the scientific merit of
one of the publications, namely Auriol Batten’s Flowers of southern
Africa (1986). He describes it as ‘not necessarily a work of any great
scientific value’. Admittedly the text is written primarily for the lay-
man and is not a taxonomic treatise, but the paintings, as admitted by
Kerkham, are of a very high standard and represent extreme scientific
accuracy. Surely this contributes to its scientific value. To single out
this work as of little scientific value and not some of the very obvious
coffee table books which he cites is, to my mind, rather unfair.
The bibliography is divided into two parts, viz. monographs
(pp. 33-213) with 753 items and serials (pp. 214-224) with 64 items. I
shall consider monographs first. The very first item, 001 Acocks, John
142
Phillip Harrison, Veld types of South Africa (1953), probably the most
quoted botanical work in the RSA, contains one error and one omis-
sion. The error is in the spelling of Acocks’s third given name: it
should be Harison and not Harrison. The omission, surprisingly, is
mention of the accompanying veld type map, which is possibly consul-
ted more than the text. Item 002, the revised edition of the memoir,
fortunately, includes the map.
A considerable number of items have been omitted from the mono-
graph section. Some that immediately come to mind are: Bews, The
world’s grasses (1929), E.P. Phillips, An introduction to the study of
South African grasses (1931) and, more recently, Werger (ed.),
Biogeography and ecology of southern Africa, Vols 1 & 2 (1978),
Stirton (ed.), Plant invaders, beautiful but dangerous (1978), Farr,
Leussink & Stafleu's Index nominum genericorum plantarum, 3 vo-
lumes (1979), the extremely important work by White, The vegetation
of Africa + AETFAT/UNSO vegetation map of Africa ( 1983) and Reid
& Dyer, A review of the southern African species of Cyrtanthus
(1984).
In the introduction it was mentioned that in the bibliography, theses
and journal articles have been excluded unless of particular botanical
value. The assessment of botanical value has presumably been made by
the compiler. The result is that scores of theses and journal articles of
undeniable botanical value have been excluded — works of no less merit
than those included. Such theses include Irvine, The major veld types
of the northern Transvaal and their grazing management ( 1941 ), Van
der Schijff, Ekologiese studie van die flora van die Nasionale Kruger-
wildtuin ( 1958) and Grunow, Objective classification of plant commu-
nities: a synecological study in the sour-mixed bushveld of the
Transvaal ( 1965).
Regarding journal articles, four of Bews’s articles have been in-
cluded, but what of the remaining 25, all of a high standard? A signifi-
cant omission is E.P. Phillips, A contribution to the flora of the Leribe
Plateau and environs, with a discussion on the relationships of the
floras of Basutoland, the Kalahari and the south-eastern region [Annals
of the South African Museum 16: 1—379 (1917)]. Another is Henkel,
Ballenden & Bayer, An account of the plant ecology of the Dukuduku
Forest Reserve and adjoining areas of the Zululand coast belt [Annals
of the Natal Museum 8: 95—126 (1941)]. Finally, Edwards, A broad-
scale structural classification of vegetation for practical purposes
[Bothalia 14: 705-712 (1983)] has had considerable impact in local
ecological circles in recent times.
In a bibliography of this kind, one does not expect to find personal
assessments of botanical works by the compiler. For example, under
item 722 Wendland, Johann Christoph, Collectio plantarum ... earum,
(1805- 1819), was it necessary to say ‘Tolerable figures of many Cape
plants’ and under 600 Scott, Charles L. The genus Haworthia ( Lilia -
ceae): a taxonomic revision (1988), was it necessary to state, ‘A con-
troversial revision’? Out of 753 items, these two books seem to be the
only ones in the monograph section singled out for this sort of com-
ment.
Omissions have been referred to, but unnecessary inclusions also
deserve comment. Examples include items 303 and 304, International
Botanical Congress, Paris 1867 and Sydney 1981 respectively. The
Taws’ adopted at these congresses are international in character and
applicability and do not really form part of southern African botanical
literature. However, in the latter case (304) inclusion is probably justi-
fied because names of southern African plants are included in the
Nomina generica conservanda et rejicienda and Nomina familiarum
conservanda in the appendices. But why, if the codes are considered
part of southern African literature, are only two codes included, when
there have been many between 1867 and 1988?
There is little to say about the serial section except that several
serials have been omitted, for example Koedoe, Lammergeyer, Pro-
ceedings of the Grassland Society of southern Africa and the new
Euphorbia Journal ( 1983 — ).
Typographical errors do occur, for example Brabeium instead of
Brabejum on p.16, Reynold’s instead of Reynolds’s on p.27 and M.
Boulton instead of M. Bolton on p.47. Another thing noticed is that the
location of the items is not always complete: it is clear that the BRI
Library has not been fully consulted and is therefore not cited under
many of the items.
The book is of A5 size with a glossy soft cover and runs to 256
pages. It is published by the South African Library and costs R35 +
GST. The book will undoubtedly be of use to practising botanists for
reference purposes, as long as they do not expect to find everything
they are looking for. It is to be hoped that in future editions the input
from knowledgeable botanists will be greater. The purpose of the book
Bothalia 19,1 (1989)
is commendable, but unfortunately the product leaves much to be de-
sired.
D. J. B. KILLICK
THE BANKSIA ATLAS, Australian Flora and Fauna Series No. 8,
edited by ANNE TAYLOR and STEPHEN HOPPER. 1988. Austra-
lian Government Publishing Service, G.P.O. Box 84, Canberra ACT
2601, Australia. Pp.257.
The Banksia Atlas is a profoundly important pioneering work. Al-
though it has a line of forerunners it is unique in several significant
respects. It provides for the first time, at fine resolution, distribution
maps (with accompanying habitat and other data) of each species of a
diverse plant genus, in relatively undisturbed country, on a continental
basis.
The project was undertaken, and the Atlas produced, with the spe-
cific intent of evaluating the feasibility, and exploring the meth-
odology, of producing a plant distribution atlas at an Australian-wide
level based on a fully computerized data set derived from sight records
provided entirely by volunteer contributors. The three year project was
an eminent success and will hopefully provide the stimulus for a suc-
cession of similar programs not only in Australia but throughout the
world.
Banksia was an ideal choice for this atlas. With its 75 species, it is
diverse but yet manageable; it is taxonomically well studied and the
diagnostic characters of the species are clear; it is a distinctively Aus-
tralian native plant occurring in all the states and has strong aesthetic
appeal.
The published volume, though not particularly attractive such as
might catch the public eye, is an admirable production that is clearly
and systematically layed out and easy to use. The history, aims,
methods and results of the project are described in a 60-page introduc-
tory section, and this is followed by a double-page spread devoted to
each species.
For each species there is a distribution map with longitude/latitude
point locations based solely on sight records, together with an inset
map showing the occurrence as known from herbarium specimens. The
greatly expanded knowledge of distributions is clearly evident. Accom-
panying the maps are data on population sizes, plant form, times of
flowering and new shoot growth, response to fire, possible pollinators,
conservation status and a general discussion.
The data derives from 12 991 field record sheets received from 421
contributors from all walks of life and from across the land. The
number of sheets sent in by contributors ranged from one to over
1 000. Around half the individuals sent in 10 or fewer record sheets,
whilst just 15 % sent in more than 50. The record sheets were designed
for computerized storage and manipulation and the double-page
spreads were set directly from this data set.
The immediate spin-off from the project included new taxonomic
discoveries and the highlighting of taxonomic problem areas, as well as
increased knowledge of distributions, habitat, pollinators, conservation
status, and variation within populations. Longer term returns will no
doubt manifest themselves in diverse and possibly unexpected ways.
The most eloquent precursor to the Banksia Atlas is the Atlas of the
British flora, edited by Perring & Walters (1962). This work covered
all the indigenous and naturalized plants of Britain, totalling some
1 700 species. A 100 % sampling and plotting density on a 10 x 10 km
grid (3 500 cells) was achieved. Some 1 500 of the country’s botanists,
professional and amateur, were involved in the collection of field re-
cords over a period of five years. The atlas does not include habitat or
abundance data, but does include a series of loose overlays showing
such regional aspects as geology, rainfall and land-use.
The most obvious difference between the British and Australian
atlases lies in the meaning behind the distribution patterns. Those of
Britain can bear little resemblance to the original natural condition,
whereas those of Australia probably remain more or less true to the
situation before the arrival of European man. The British Isles include
an area but a tiny fraction of that of Australia, but support a human
population some four times larger. Agricultural and industrial man has
wrought havoc in Britain for several thousand years and in Australia for
only two hundred.
The latest crop of Nobel Prizes have gone to research on funda-
mental subatomic particles, on the precise atomic structure of photo-
synthesising molecules, and on certain drugs that are finely tuned to
home in on cancerous cells. Wonderful research, but it can wait. The
native animal populations and vegetation of the world cannot wait.
They are being eradicated at a catastrophic rate. There is mounting
Bothalia 19,1 (1989)
143
evidence that massive global extinctions have punctuated earth history
at roughly 30 million year intervals — caused possibly by outsized
meteorite impacts. There is rapidly mounting evidence also, that unless
man shifts his priorities radically, and literally overnight, he will be the
direct cause of a global extinction event more thorough than any pre-
viously felt.
The selection of Banksia for this seminal work is fortuitous also for
historical reasons. The genus was named after Sir Joseph Banks
(1743— 1820), who holds legitimate claim not only to the title of father
of Australian naturalists, but also to the nation itself. He accompanied
Captain Cook, as naturalist, on his first expedition to the Pacific and,
being a man of great independent wealth, paid for all his own equip-
ment as well as for an assistant (Solander, a student of Linnaeus) and
four artists. The expedition landed in Botany Bay (near todays’ Syd-
ney) in 1770, and here Banks had an unparalleled opportunity to ex-
plore the wholly unknown flora and fauna. It was largely through his
efforts in later years that the first colonies were established in Austra-
lia. His prestige back in England, where he held the Presidency of the
Royal Society for a record tenure of 41 years (1778—1820), was
without check.
What better beacon, naturalist’s torch, than Banksia to help light the
way towards a new consciousness of our precarious, fragile biosphere.
J. M. ANDERSON
.
** y'W ,
Bothalia 19,1: 145-150(1989)
Guide for authors to Bothalia
This guide is updated when necessary. The latest ver-
sion should therefore be consulted.
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 Botanical
Research Institute is devoted to the furtherance of
botanical science. The main fields covered are taxo-
nomy, ecology, anatomy and cytology. Two parts of the
journal and an index to contents, authors and subjects are
published annually.
EDITORIAL POLICY
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
reviews, are accepted. Manuscripts may be written in
either English or Afrikaans.
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 main-
tains an editorial board (see title page) to ensure that
international standards are upheld.
PRESENTATION OF MANUSCRIPT
Manuscripts should be typewritten on one side of good
quality A4-size paper, double-spaced throughout (in-
cluding abstract, tables, captions to figures, literature
references, etc.) and have a margin of at least 30 mm all
round. The original and three photocopies (preferably
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. Papers should
conform to the general style and layout of recent issues
of Bothalia (from volume 14 onwards). Material should
be presented in the following sequence: Title page with
title, name(s) of author(s), keywords, abstracts (in
English and Afrikaans) 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. The
sequence continues with Introduction and aims, Material
and methods, Results, Interpretation (Discussion),
Acknowledgements, Specimens examined (in revisions
and monographs). References, Index of names (recom-
mended 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
abstract are superfluous. All pages must be numbered
consecutively beginning with the title page to those with
references, tables and captions to figures.
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.
TITLE
The title should be as concise and as informative as
possible. In articles dealing with taxonomy or closely
related subjects the family of the taxon under discussion
(see also Names of taxa under General below) should be
mentioned in brackets but author citations should be
omitted from plant names.
KEYWORDS
Up to 10 keywords (or index terms) should be pro-
vided in English in alphabetical sequence. The following
points should be borne in mind when selecting key-
words:
1, Keywords should be unambiguous, internationally
acceptable words and not recently-coined little-known
words; 2, they should be in a noun form and verbs should
be avoided; 3, they should not consist of an adjective
alone; adjectives should be combined with nouns; 4, they
should not contain prepositions; 5, the singular form
should be used for processes and properties, e.g. evapo-
ration; 6, the plural form should be used for physical
objects, e.g. augers; 7, location (province and/or coun-
try); taxa (species, genus, family) and vegetation type
(community, veld type, biome) should be used as key-
words; 8, keywords should be selected hierarchically
where possible, e.g. both family and species should be
included; 9, they should include terms used in the title;
10, they should answer the following questions: 10.1,
what is the active concept in the document (activity,
operation or process); 10.2, what is the passive concept
or object of the active process (item on which the activ-
ity, operation or process takes place); 10.3, what is the
means of accomplishment or how is the active concept
achieved (technique, method, apparatus, operation or
process); 10.4, what is the environment in which the
active concept takes place (medium, location) and 10.5,
what are the independent (controlled) and dependent
variables? 11, questions 10.1 to 10.3 should preferably
also be answered in the title.
ABSTRACT
Abstracts of no more than 200 words should be pro-
vided in English and Afrikaans. Abstracts are of great
importance and should convey the essence of the article.
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. In articles dealing with taxonomy or
closely related subjects all taxa from the rank of genus
downwards should be accompanied by their author cita-
tions. Names of new taxa and new combinations should
not be underlined. If the article deals with too many taxa
only the important ones should be mentioned.
TABLE OF CONTENTS
A table of contents should be given for all articles
longer than about six typed pages, unless they follow the
strict format of a taxonomic revision.
146
Bothalia 19,1 (1989)
ACKNOWLEDGEMENTS
Acknowledgements should be kept to the minimum
compatible with the requirements of courtesy. Please
give all the initials of the person(s) your are thanking.
LITERATURE REFERENCES
In text
Literature references in the text should be cited as
follows: ‘Jones & Smith (1986) stated . . or \ . .
(Jones & Smith 1986)’ when giving a reference simply
as authority for a statement. When more than two
authors are involved use the name of the first author
followed by et al. When referring to more than one
literature reference, they should be arranged alpha-
betically according to author and separated by a semi-
colon, e.g. (Anon. 1981, 1984; Davis 1976; Nixon
1940). Titles of books and names of journals should
preferably not be mentioned in the text. If there is good
reason for doing so, they should be treated as described
in the paragraph In reference list below. Personal com-
munications are given only in the text, not in the list of
references. Please add the person’s full initials to iden-
tify the person more positively.
In taxonomic articles
The correct name (not underlined) is to be followed by
its author citation (underlined) and the full literature
reference, with the name of the publication written out in
full. 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 etal .: 14 (1966); Boris: 89 (1967);
Sims: t. 38 (1977); Sims: 67 (1980). Note that (1) refer-
ences are arranged in chronological sequence; (2) where
two or more references by the same author are listed in
succession, the author’s name is repeated with every
reference; (3) names of authors are written in the same
way (see Names of authors of plant names under
General), irrespective of whether the person in question
is cited as the author of a plant name or of a publication;
(4) the word ‘figure’ is written as ‘fig.’, and ‘t.’ is used
for both ‘plate’ and ‘tablet’.
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
synonymy.
In reference list
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. The references are arranged
alphabetically according 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. 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. Author names
are typed in capitals. Titles of journals and of books are
written out in full and are underlined as follows:
Transactions of the Linnean Society of London 5:
171—217, or Biology and ecology of weeds : 24. Titles
of books should be given as in Taxonomic literature, edn
2 by Stafleu & Cowan and names of journals as in World
list of scientific periodicals, edn 4. If the same author is
mentioned more than once, the name is written out in full
and not replaced by a line.
Examples of references:
Collective book or Flora
BROWN, N. E. 1909. Asclepiadaceae. In W. T. Thiselton-Dyer,
Flora capensis 6,2: 518-1036. Reeve, London.
BROWN, N. E. 1915. Asclepiadaceae. In W. T. Thiselton-Dyer,
Flora of tropical Africa 5,2: 500-600. Reeve, London.
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,
London.
Journal
DAVIS, G. 1988. Description of a proteoid-restioid stand in Mesic
Mountain Fynbos of the south-western Cape and some aspects of
its ecology. Bothalia 18:279-287.
STEBBINS, G. L. Jr 1952. Aridity as a stimulus to plant evolution.
American Naturalist 86:35—44.
SMOOK, L. & GIBBS RUSSELL, G. E. 1985. Poaceae. Memoirs of
the Botanical Survey of South Africa No. 51:45— 70.
In press, in preparation
TAYLOR, H. C. in press. A reconnaissance of the vegetation ofRooi-
berg State Forest. Department of Forestry, Technical Bulletin.
VOGEL, J. C. 1982. The age of the Kuiseb river silt terrace at Homeb.
Palaeoecology of Africa 1 5 . In press .
WEISSER, P. J., GARLAND, J. F. & DREWS, B. K. in prep. Dune
advancement 1937-1977 and preliminary vegetation succession
chronology at Mlalazi Nature Reserve, Natal, South Africa.
Bothalia.
Thesis
KRUGER, F. J. 1974. The physiography and plant communities of the
Jakkalsrivier Catchment. M. Sc. (Forestry) thesis, University of
Stellenbosch.
Miscellaneous paper, report, unpublished article, tech-
nical note, congress proceedings
ANON, no date. Eetbare plante van die Wolkberg. Botanical Research
Unit, Grahamstown. Unpublished.
BAWDEN, M. G. & CARROL, D. M. 1968. The land resources of
Lesotho. Land Resources Study No. 3, Land Resources Division,
Directorate of Overseas Surveys, Tolworth.
BOUCHER, C. 1981. Contributions of the Botanical Research Insti-
tute. 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.
TABLES
Each table should be presented on a separate sheet and
be assigned an Arabic numeral, i.e. the first table men-
Bothalia 19,1 (1989)
147
tioned in the text is marked ‘Table 1’. In the captions of
tables the word ‘table’ is written in capital letters. See
recent numbers of Bothalia for the format required.
Avoid vertical lines, if at all possible. Tables can often
be reduced in width by interchanging primary horizontal
and vertical heads.
FIGURES
General
Figures should be planned to fit, after reduction, into a
width of either 80, 118 or 165 mm, with a maximum
vertical length of 240 mm. Allow space for the caption in
the case of figures that will occupy a whole page. Letter-
ing and numbering on all figures should be done in letra-
set, 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. If the
lettering or wording on a figure is to be done by the
printer this information must be typed or neatly printed
on a photocopy of the figure or on an overlay attached to
the original. If several illustrations are treated as com-
ponents of a single composite figure they should be
designated by capital letters. Note that the word ‘figure’
should be written out in full, both in the text and the
captions. In the text the figure reference is then written
as in the following example: ‘The stamens (Figure 4A,
B, C) are . . .’.In captions, ‘figure’ is written in capital
letters. Magnification of figures should be given for the
size as submitted. It is recommended, however, that
scale bars or lines be used on figures. In figures accom-
panying taxonomic papers, voucher specimens should be
given in the relevant caption. Figures are numbered con-
secutively 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.
Authors should indicate in pencil in the text where they
would like the figures to appear. 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’. Authors wishing to
use illustrations already published must obtain written
permission before submitting the manuscript and inform
the editor of this fact. Captions for figures should be
collected together and typed on a separate sheet headed
Captions for figures.
Black and white drawings
Line drawings, including graphs and diagrams, should
be in jet-black Indian ink, preferably on bristol board or
tracing film. Lines should be bold enough to stand reduc-
tion. It is recommended that drawings should be twice
the size of the final reproduction.
Photographs
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. Photo-
graph mosaics should be submitted complete, the com-
ponent photographs mounted neatly on a white card base
leaving a narrow gap of uniform width between each
print. Note that grouping photograhps of markedly
divergent contrast results in poor reproductions.
Dot maps
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 dia-
meter). Blank maps are available from the editor.
Names of taxa
As a rule authors should use the names as listed by
Gibbs Russell et al. in Memoirs of the Botanical Survey
of South Africa Nos 48, 5 1 and 56. Names of genera and
infrageneric taxa are usually underlined with the author
citation (where relevant) not underlined. Exceptions in-
clude names of new taxa in the abstracts, 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 being not underlined and syno-
nyms underlined. Names above generic level are not
underlined. 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 abbre-
viated to the initial thereafter, except where intervening
references to other genera with the same initial could
cause confusion.
Names of authors of plant names
These should agree with the list compiled by the BRI
(TN TAX 2/1) which has also been implemented by
Gibbs Russell et al. in Memoirs of the Botanical Survey
of South Africa Nos 48, 51 and 56. Modem 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 recom-
mendations of the Code.
Names of authors of publications
These are written out in full except in the synonymy in
taxonomic articles where they are treated like names of
authors of plant names.
Names of plant collectors
These are underlined whenever they are linked to the
number of a specimen. The collection number is also
underlined, e.g. Acocks 14407. Surnames beginning
with ‘De’, ‘Du’ or ‘Van’ begin with a capital letter
unless preceded by an initial.
Measurements
Use only units of the International System of Units
(SI). Cm should not be used, only mm and/or m. The use
of ‘ ± ’ is preferred to c. or ca.
Numerals
Numbers ‘one’ to ‘nine’ are spelled out in normal text,
and from 10 onwards they are written in Arabic nume-
rals. In descriptions of plants, numerals are used
throughout. Write 2, 0-4,5 (not 2-4,5). When counting
members write 2 or 3 (not 2—3).
148
Bothalia 19,1 (1989)
Abbreviations
Abbreviations should be used sparingly but con-
sistently. No full stops are placed after abbreviations
ending with the last letter of the full word (e.g. edition =
edn; editor =ed.), after units of measure, after compass
directions and after herbarium designations.
KEYS TO TAXA
It is recommended that (apart from multi-access keys)
indented keys be used with couplets numbered la- lb,
2a— 2b, etc. (without full stops thereafter). Keys consist-
ing of a single couplet have no numbering. Manuscripts
of keys should be presented as in the example below.
Herbarium voucher specimens
Wherever possible authors should refer to one or more
voucher specimen(s) in a registered herbarium.
SPECIES TREATMENT IN TAXONOMIC PAPERS
General presentation
The procedure to be followed is illustrated in the
example (under Description and example of species
treatment, below), which should be referred to, because
not all steps are described in full detail. The correct name
(see also Names of taxa, under General), with its litera-
ture citations is followed by the synonymy (if any), the
description and the discussion, which should consist of
paragraphs commencing, where possible, with italicised
leader words such as flowering time, diagnostic charac-
ters, distribution and habitat.
Numbering
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, the first line of the paragraph to be indented. Infra-
specific taxa are marked with small letters, e.g. lb.,
12c., etc.
Literature references within synonymy
(See above under Literature references, paragraph 2.)
Citation of specimens
Type specimen in synopsis
The following should be given (if available): country
(if not in RSA), province, locality as given by original
collector, modem equivalent of collecting locality in
square brackets (if relevant), date of collection
(optional), collector’s name and collecting number (both
underlined). 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 speci-
men in question. The herbaria in which the relevant
type(s) are housed are indicated by means of the abbre-
viations given in the latest edition of Index Herbariorum.
The holotype (holo.) and its location are mentioned first,
followed by a semicolon, the other herbaria are arranged
alphabetically, separated by commas. Authors should
indicate by means of an exclamation mark (!) which of
the types have been personally examined. If only a pho-
tograph or microfiche was seen, write as follows: Anon.
422 (X, holo.— BOL, photo.!). Lectotypes or neotypes
should be chosen for correct names without a holotype.
It is not necessary to lectotypify synonyms. When a
lecto- or a neotype are newly chosen this should be indi-
cated by using the phrase ‘here designated’. If reference
is made to a previously selected lectotype or neotype, the
name of the designating author and the literature refer-
ence should be given. In cases where no type was cited,
and none has subsequently been nominated, this may be
stated as ‘not designated’ .
In notes and brief taxonomic articles
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/countries (typed in capitals) should be cited
in the following order: SWA/Namibia, Botswana,
Transvaal, Orange Free State, Swaziland, Natal,
Lesotho, Transkei and Cape. Grid references should be
cited in numerical sequence. Locality records for speci-
mens should preferably be given to within a quarter-de-
gree square. Records from the same one-degree square
are given in alphabetical order, i.e. (—AC) precedes
(-AD), etc. Records from the same quarter-degree
square are arranged alphabetically according to the col-
lector’s names; the quarter-degree references must be
repeated for each specimen cited. The relevant interna-
tional 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 procedure:
NATAL. — 2731 (Louwsburg): 16 km E of Nongoma (-DD), Pelser
354 (BM, K, PRE); near Dwarsrand, Van der Merwe 4789 (BOL,
M). 2829 (Harrismith): near Groothoek (- AB), Smith 234\ Koffiefon-
tein (-AB), Taylor 720 (PRE); Cathedral Peak Forest Station (-CC),
Marriot 74 (KMG); Wilgerfontein, Roux 426. Grid ref. unknown:
Sterkstroom, Strydom 12 (NBG).
For records from outside southern Africa authors
should use degree squares without names, e.g.:
KENYA. — 0136: Nairobi plains beyond race course, Napier 485.
If long lists of specimens are given, they should be
dealt with as on p. 149, paragraph 5.
la Leaves closely arranged on an elongated stem; a submerged 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; receptacles 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:
Bothalia 19,1 (1989)
In 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 distribu-
tion maps for all relevant taxa and that the distribution
has been described briefly in words in the text. Under the
heading ‘Vouchers’ no more than 5 specimens should be
cited, indicating merely the collector and the collector’s
number (both underlined). 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 semicolon. The
purpose of the cited specimens is not to indicate distribu-
tion but to convey the author’s concept of the taxon in
question.
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: Fisher 840 (NH, NU, PRE); Flanagan 831 (GRA,
PRE); 840 (NH, PRE); Marloth 4926 (PRE, STE); Schelpe 6161,
6163, 6405 (BOL); Schlechter 445 1 (BM, BOL, GRA, K, PRE).
All specimens studied by the author should be listed
together at the end of the article under the heading Speci-
mens examined. They are arranged alphabetically 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 infraspecific taxa
by a small letter. If more than one genus is dealt with in a
given article, the first species of the first genus men-
tioned is indicated as 1.1. This is followed by the inter-
national herbarium designation. Note that the name of
the collector and the collection number are underlined:
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 . Burn 789
(2.6) B, KMG, STE.
Synonyms
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. Illegitimate names are designated by nom.
illeg. after the reference, followed by non with the
author and date, if there is an earlier homonym. Nomina
nuda (nom. nud.) and invalid names are excluded unless
there is a special reason to cite them, for example if they
have been used in prominent publications. Note that in
normal text Latin words are italicized, but in the
synopsis of a species Latin words such as nom. nud. are
not italicized.
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. When a generic
name is repeated in a given synonymy it should be
abbreviated to the initial except where intervening refer-
ences to other genera with the same initial could cause
confusion.
149
Description and example of species treatment
Descriptions of all taxa of higher plants should, where
possible, follow the sequence: Habit; sexuality; under-
ground parts (if relevant). Indumentum (if it can be
easily described for the whole plant). Stems/branches.
Bark. Leaves : arrangement, petiole absent/present,
pubescence; blade: shape, size, apex, base, margin;
midrib: above/below, texture, colour; petiole; stipules.
Inflorescence : type, shape, position; bracts/bracteoles.
Flowers : shape, sex. Receptacle. Calyx. Corolla. Disc.
Androecium. Gynoecium. Fruit. Seeds. Chromosome
number. Figure (word written out in full) number. As a
rule shape should be given before measurements. 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. Language
must be as concise as possible, using participles instead
of verbs. Dimension ranges should be cited as in the
example below. 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; 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 lanceolate’); it is produced on a
typewriter by typing 2 hypens next to each other; and 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 on a
typewriter by typing 3 hypens next to one another. The
use of ‘ ±’ is preferred to c. or ca when describing shape,
measurements, dimensions, etc.
Example:
1 . Bequaertiodendron magalismontanun (Sond.) Heine & Hemsl.
in Kew Bulletin: 307 (1960); Codd: 72 (1964); Elsdon: 75 (1980).
Type: Transvaal, Magaliesberg, Zeyher 1849 (S,
holo.-BOL, photo.!).
Chrysophyllum magalismontanum Sond.: 721 (1850); Harv.: 812
(1867); Engl.: 434 (1904); Bottmar: 34 (1919). Zeyherella
magalismontanum (Sond.) Aubrev. & Pelegr.: 105 (1958); Justin:
(1973).
Chrysophyllum argyrophyllum Hiem: 721 ( 1850); Engl.: 43 ( 1904).
Boivinella argyrophylla (Hiem) Aubrev. & Pellegr. : 37 (1958); Justin:
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.) Aubrev. & Pellegr.:
39 (1958); Justin: 99 (1973). Type: Transvaal, Magoebaskloof, Wilms
1812 (B, holo.; K! , P!, lecto., designated by Aubrev. & Pellegr.: 38
(1958), PRE!, S! W!, Z!).
Bequaertiodendron fruticosa De Wild.: 37 (1923), non Bonpland:
590 (1823); Bakker: 167 (1929); Fries: 302 (1938); Davy: 640 (1954);
Breytenbach: 117 (1959); Clausen: 720 (1968); Palmer: 34 (1969).
Type: Transvaal, Tzaneen Distr., Granville 3665 (K, holo.!; G ! , P!,
PRE!, S!).
Bequaertiodendron 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
3— 10 (—23) x 1,0— 1 ,5( — 4,0) mm, linear to oblanceo-
late, obtuse, base broad, half-clasping. Heads heteroga-
mous, campanulate, 7—8x5 mm, solitary, sessile at tip
of axillary shoots; involucral bracts in 5 or 6 series, inner
150
Bothalia 19,1 (1989)
exceeding flowers, tips subopaque, white, very acute.
Receptacle nearly smooth. Flowers ± 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.
In — 22. Figure 23 B.
New taxa
The name of a new taxon must be accompanied by at
least a Latin diagnosis. Authors should not provide full-
length Latin descriptions unless they have the required
expertise in Latin at their disposal. It is recommended
that descriptions of new taxa be accompanied by a good
illustration (line drawing or photograph) and a distribu-
tion map.
Example:
109. Helichrysum jubilatum Hilliard, sp. nov. H.
alsinoidei DC. affinis, sed foliis ellipticis (nec spatula-
tis), inflorescentiis compositis a foliis non circumcinctis,
floribus femineis numero quasi dimidium hermaphrodi-
torum aequantibus (nec capitulis homogamis vel floribus
femineis 1-3 tantum) distinguitur.
Herba annua e basi ramosa; caules erecti vel decum-
bentes, 100—250 mm longi, tenuiter albo-lanati, remote
foliati. Folia plerumque 8-30 x 5-15 mm, sub capi-
tulis minora, elliptica vel oblanceolata, obtusa vel acuta,
mucronata, basi semi-amplexicauli, utrinque cano-la-
nato-arachnoidea. Capitula heterogama, campanulata,
3,5 -4,0 X 2,5 mm, pro parte maxima in paniculas
cymosas terminales aggregata; capitula subterminalia in-
terdum solitaria vel 2— 3 ad apices ramulorum nudorum
ad 30 mm longorum. Bracteae involucrales 5-seriatae,
gradatae, exteriores pellucidae, pallide stramineae,
dorso lanatae, seriebus duabus interioribus subaequali-
bus et flores quasi aequantibus, apicibus obtusis opacis
niveis vix radiantibus. Receptaculum fere laeve. Flores
± 35-41. Achenia 0,75 mm longa, pilis myxogenis
praedita. Pappi setae multae, corollam aequantes, apici-
bus scabridis, basibus non cohaerentibus.
TYPE. — Cape, Namaqualand Division, Richtersveld,
± 5 miles E of Lekkersing on road to Stinkfontein, kloof
in hill south of the road, annual, disc whitish, 7 xi 1962,
Nordenstam 1823 (S, holo.; E, NH, PRE).
PROOFS
Only galley proofs are normally sent to authors. They
should be corrected in red ink and be returned to the
editor as soon as possible.
REPRINTS
Authors receive 100 reprints free. If there is more than
one author, this number will have to be shared between
them.
DOCUMENTS CONSULTED
Guides to authors of the following publications were
made use of in the compilation of the present guide:
Annals of the Missouri Botanic Garden, Botanical Jour-
nal of the Linnean Society, Bothalia, Flora of Australia,
Smithsonian Contributions to Botany, South African
Journal of Botany (including instructions to authors of
taxonomic papers). South African Journal of Science.
ADDRESS OF EDITOR
Manuscripts should be submitted to: The Editor,
Bothalia, Botanical Research Institute, Private Bag
X101, Pretoria 0001.
Bothalia 19,2: 151-156 (1989)
Studies in the southern African species of Justicia and Siphonoglossa
(Acanthaceae): palynology
K.L. IMMELMAN*
Keywords: Acanthaceae, Justicia , palynology, Siphonoglossa, southern Africa, taxonomy
ABSTRACT
The gross morphology of the pollen of all southern African species and subspecies of Justicia and Siphonoglossa was
investigated, as well as that of eight tropical African species of Justicia. The following pollen types were found in the southern
African species of Justicia: two- or three-colporate, each with the margocolpus either entire or broken up into areolae, and
two-porate areolate pollen. One tropical African species had the colpus very short, and in another tropical African species
it was replaced by an extra row of areolae. All Siphonoglossa species had two-colporate pollen with areolae and long colpi.
The southern African species of Justicia could be separated from Siphonoglossa on pollen characters, and some sections
of Justicia could also be distinguished on the same basis.
UITTREKSEL
Die algemene morfologie van die stuifmeel van alle spesies en subspesies van Justicia en Siphonoglossa in Suider-Afrika,
asook die van agt Justicia- spesies uit tropiese Afrika, is ondersoek. Die volgende stuifmeeltipes is by die Justicia-spesies
van Suider-Afrika gevind: twee- of driekolporaat, elk met die margokolpus of gaaf of opgebreek in areole, en tweeporate
areolere stuifmeel. By een spesie uit tropiese Afrika was die kolpus baie kort, en by ’n ander uit tropiese Afrika is dit deur
’n ekstra ry areole vervang. Alle Siphonoglossa- spesies het tweekolporate stuifmeel met areole en lang kolpusse gehad.
Die Justicia- spesies van Suider-Afrika kon op grond van stuifmeelkenmerke van Siphonoglossa geskei word, en sommige
Justicia- seksies kon ook op dieselfde basis onderskei word.
INTRODUCTION
The gross morphology of the pollen of all southern
African species and subspecies of Justicia and
Siphonoglossa was investigated, as well as that of eight
tropical African species of Justicia. In order to place the
studies into perspective, brief reference is made to findings
of other researchers.
Radlkofer (1883) was the first to point out the potential
value of the pollen in classification of the Acanthaceae,
but it was Lindau (1895) who first attempted to use the
pollen systematically in its classification.
The ‘typical representatives’ of the subtribe Justiciinae,
according to Bremekamp (1965), have prolate, dorsi-
ventrally flattened pollen, with the pore in the middle of
a trema area which is studded with circular ‘insulae’
(areolae).
Steam (1971), after discussing the pollen type of a
number of Jamaican species of Justicia, concluded that
‘The palynology of the group as a whole must certainly
have a significant part in its reclassification. Pending that
reclassification, Justicia is probably best accepted in a
broad sense’.
Gibson (1972) widened the circumscription of Justicia
and, if her delimitation is accepted, the range of pollen
types present in the genus will be greatly increased, and
will include 3- and 4-porate pollen and pollen with the
areolae scattered across the face of the grain.
* Botanical Research Institute, Department of Agriculture and Water
Supply, Private Bag X101, Pretoria 0001.
MS. received: 1988.07.16.
Balkwill & Getliffe Norris (1988), who reappraised the
tribal and subtribal limits of Acanthaceae in southern
Africa, considered that in southern Africa each tribe had
a characteristic pattern of pollen ‘against which aberrant
genera can be contrasted’. They considered prolate
tricolporate grains with entire margocolpi as being the
primitive state.
Graham (1989) surveyed a number of species of Justicia
and related genera worldwide, and greatly widened the
circumscription of the genus to include a number of
formerly segregate genera, including Siphonoglossa. She
divided the genus into a number of sections and subsec-
tions which have been followed here, and considered
pollen, in combination with other characters, to be of
major taxonomic importance. She considered the South
African species of Siphonoglossa to belong in Justicia,
but not in the same section as the South American spe-
cies of Justicia. She described 10 pollen types, but did
not confirm the 4-aperturate pollen types reported for a
few New World species.
METHODS AND MATERIALS
In this study the pollen of all southern African species
and subspecies of Justicia and Siphonoglossa was
investigated with the SEM. Taxa covered were all those
occurring in southern Africa as defined by the Flora of
southern Africa. Eight tropical African species were also
investigated for purposes of comparison, but were not
measured.
Material was obtained from three sources: herbarium
sheets, fresh material grown in the greenhouse at the
Botanical Research Institute, Pretoria, and material
preserved in alcohol (gathered in the field).
152
Bothalia 19,2 (1989)
Fresh material of two of the species, J. flava (Vahl) Vahl
and J. protracta (Nees) T. Anders, subsp. protracta, was
also acetolysed (following Erdtman 1971, as quoted by
Taylor, Hollingsworth & Bigelow 1974), before being
mounted and viewed. The acetolysed material was then
compared with that of the same species mounted directly
onto stubs.
One to four specimens of each species and subspecies
of Justicia and Siphonoglossa in southern Africa were
examined. The anthers were macerated onto 15 mm
aluminium stubs covered with double-sided adhesive tape.
Thin tape was used, as the pollen grains tended to sink
partially into the thicker gum of some tapes and became
difficult to photograph well.
After air-drying, the mounted material was glow-
discharge-coated with ± 400 A° of metallic gold in an
Eiko sputter coater. The specimens were then viewed with
an MSM 4 Hitachi-Akashi (desk top model) SEM at kv
15. Selected views were photographed of the profile (face),
side and amb (polar) faces of the pollen grains of each
taxon, using a Mamiya 6x7 camera and Ilford FP4 125
ASA film. The film was developed in Microdol X.
The size of the pollen grains was measured from water-
mounted pollen with a Kontron Image analyser.
The terminology used is illustrated in Figure 1.
stopple
FIGURE 1. — Diagrams of pollen grains in Justicia and Siphonoglos-
sa: A, with entire margocolpus, x 2400; B, with areolae, x
2400.
RESULTS AND DISCUSSION
No differences were observed between pollen taken from
fresh and from dried material when it was viewed.
No significant difference was found between acetolysed
material and material mounted direcdy onto the stubs. This
is similar to the findings of Munday (1980) in Monechma,
except that acetolysed material tended to loose its ‘stopples’
and had a high proportion of damaged grains. Material
was therefore viewed throughout without pretreatment, and
results given are of such pollen.
Shape of pollen grains
The pollen of both Justicia and Siphonoglossa was found
to be basically prolate, with some species, e.g. J. orchioides
L. f. , J. platysepala (S. Moore) P.G. Mey. and J. thymifolia
(Nees) T. Anders., showing a slight median constriction
(Figure 5C).
Sexine pattern
The sexine pattern was lophate or reticulate over most
of the grain with smooth areas (margocolpi) on either side
of the colpi.
These margocolpi were ornamented either with an entire
band of lophate sexine (paracolpoid ridge) (Figure LA) or
this band was broken up into areolae, i.e. circular areas
of lophate sexine (Figure IB). The areolae were in one
or two rows ( Justicia ) (Figure 3A; 4D, G; 5A) or
sometimes in up to three rows ( Siphonoglossa ). The outer
row was frequently partially merged with the main area
of lophate sexine. In one tropical African species, J.
goetzei Lindau, the area normally occupied by the colpus
was occupied instead by a single row of areolae, as well
as the row on either side, i.e. a total of three rows of areolae
in the pair of margocolpi.
In the case of pollen with entire paracolpoid ridges, the
colpoid streak (which appears as a furrow between the
paracolpoid ridge and the main area of lophate sexine)
would sometimes continue to the poles. This, however,
may be variable (see also Balkwill & Getliffe Norris 1985),
and has not been used in the classification. The
pseudocolpi may actually join at the poles, for instance
in J. campylostemon and J. glabra among the three-
colporate species (Figure 2D).
Pores and colpi
The pollen grains have either two (Figures 4 & 5) or
three pores (Figures 2 & 3). Each may lie within an
elongate colpus running to near each pole. It is sometimes
difficult to establish with certainty whether there is a
colpus or not. Two species, however, J. anselliana (Nees)
T. Anders, and J. anagalloides (Nees) T. Anders. , were
definitely without colpi (Figure 4G). An intermediate stage
was seen in one of the tropical species, J. elegantula S.
Moore, where the colpi were present but short. Another
tropical African species, J. goetzei, had a single row of
areolae where the colpus would normally be. In all species
a granular stopple was present projecting from each pore
(Figure 4B, C).
Bothalia 19,2 (1989)
153
FIGURE 2. — Pollen of Justicia
species, 3-colporate, margo-
colpi entire. A— C, J. betonica,
Strey 8161 : A, face; B, side; C,
amb, all X 1200. D— E, J.
glabra, Smith 2734\ D, amb
showing pseudocolpi continu-
ous at poles, x 1200; E,
surface detail, x 2900.
Length
The length of the pollen grains (taken between the poles)
covered a wide range in the two genera. In Justicia it varied
from 22 /xm (J. anselliana) to 77 pm [J. petiolaris (Nees)
T. Anders.] and in Siphonoglossa from 58 /xm [S.
leptantha (Nees) Immelman] to 90 /xm [5. linifolia
(Lindau) C.B. Cl.]. The pollen in the genera therefore
varies from small to medium to large according to the
categories of Erdtman (1952).
Size (length) of pollen grain, although used in the key
below, has not been used to reach taxonomic decisions,
as there is a large range in pollen sizes in each species,
even in measurements taken from a single flower. Keys
using pollen sizes would need to be drawn up with caution,
and be based on a number of pollen samples from different
individuals and populations.
Abnormal grains
These are seen occasionally in both genera, and take
the form of a much larger than average, triangular grains
with a trilete colpus (Figure 5F). It is presumed that these
are what Bhaduri (1944) called ‘giant grains’.
FIGURE 3. — Pollen of Justicia
species, 3-colporate, margo-
colpi areolate. A & C, J.
kirkiana, Muir 1090 : A, face,
x 1400; C, amb, x 10. B, J.
petiolaris subsp. incerta,
Meeuse 9100 , side, X 1000.
D & E, 7. flava, Werdermann
& Oberdieck 1744, surface de-
tail showing areolae, X 4000.
154
Bothalia 19,2 (1989)
FIGURE 4. — Pollen of Justicia species, 2-colporate. Margocolpi entire: A— C, J. guerkeana, Barnard 122: A, face, X 1200; B, side, x
1200; C, amb, x 1600. Margocolpi areolate, D— I. D— F, J. capensis, Acocks 20100: D, face, x 900; E, side, X 900; F, amb, x
1400. G— H, J. anagalloides, Hafstrom & Acocks 1390: G, face, X 1800; H, side, X 1800. I, J. anselliana, De Winter 2518 , face, x 3000.
Description of pollen of species seen (number in brackets
is the mean length, n = 20):
Justicia
anagalloides (Nees) T. Anders. : two-porate, areolate, one row areolae,
length 28 (31,1) 36 /im ( Hafstrom & Acocks 1390) (Figure 4G, H).
anselliana (Nees) T. Anders.: two-porate, areolate, one row areolae,
length 22 (31,3) 29 /im ( De Winter 2518) (Figure 41).
betonica L.: three-colporate, margocolpus entire, length 36 (46) 58 pm
(Strey 8161) (Figure 2A, B, C).
bolusii C.B. Cl.: three-colporate, margocolpus entire, length 34 (46) 58
pm ( Immelman 376).
campyloslemon (Nees) T. Anders. : three-colporate, margocolpus entire,
length 45 (49,8) 53 pm (McClean 126).
capensis Thunb.: two-colporate, areolate, one row areolae, length 48
(51,2) 60 pm (Acocks 20100) (Figure 4D, E, F).
caerulea Forssk.: two-colporate, areolate, one row areolae, length not
measured (Polhill & Paulo 1022).
cordata (Nees) T. Anders.: two-colporate, margocolpus entire, length
not measured (Greenway & Kanuri 11357).
crassiradix Burkill & C.B. Cl.: two-porate, areolate, one row areolae,
length unknown (Killick & Leistner 3417).
cuneata Vahl (all three subspp.): two-colporate, maigocolpus entire, length
(subspp. cuneata & longebracteata only) 45 (50,9) 61 pm (subsp.
cuneata: Esterhuysen 5550; subsp. longebracteata: Dahlstrand
3157\ subsp. hoerleiniana: Dinter H. 60401).
dinteri S. Moore: two-colporate, areolate, 1—2 rows areolae, length 32
(37.2) 43 pm (Edwards 4338, Smith 1339).
elegantula S. Moore: shortly two-colporate, areolate, one row areolae,
length not measured (Salubeni 778).
extensa T. Anders.: three-colporate, margocolpus entire, length not
measured (Leeuwenberg 6841).
flava (Vahl) Vahl: three-colporate, areolate, one row areolae, length 42
(52.3) 58 pm (Werdermann & Oberdieck 1744) (Figure 3D, E).
glabra Koenig ex Roxb. : three-colporate, margocolpus entire, length 41
(43.4) 50 (im ( Smith 1700, 2734). (Figure 2D, E).
goetzei Lindau: two-porate, areolate, one row areolae on either side of
pore and a single row where colpus would usually be, length not
measured (Semsei 1657).
guerkeana Schinz: two-colporate, margocolpus entire, length 35 (39,3)
45 pm (Barnard 122) (Figure 4A, B, C).
Bothalia 19,2 (1989)
155
interrupta C.B. Cl.: three-colporate, margocolpus entire, length not
measured (R.B. & A.J. Faden 74/337).
kirkiana T. Anders. : three-colporate, areolate, one row areolae, length
47 (54,2) 61 pm ( Muir 1090, Drummond 5329) (Figure 3A, C).
minima A. Meeuse: three-colporate, areolate, two rows areolae, length
30 (33,3) 39 pm (Immelman 145).
montis-salinarum A. Meeuse: three-colporate, margocolpus entire, length
37 (41,8) 45 uni (Van Wyk 5536).
odora (Forssk.) Vahl: two-colporate, areolate, one row areolae, length
37 (41,8) 45 Aim (H. & E. Wanntorp 414).
orchioides L. f. (both subspp.) two-colporate, margocolpus entire, length
30 (39,6) 49 Aim (subsp. orchioides : Hafstrom & Acocks H 905 ;
subsp. glabrata Immelman: Muir 1100).
petiolaris (Nees) T. Anders, (all three subspp.): three-colporate, areolate,
one row areolae, length 56 (67) 77 pm (subsp. petiolaris :
McMurtry 1532; subsp. incerta (C.B. Cl.) Immelman: Meeuse
9100', subsp. bowiei (C.B. Cl.) Immelman: Marriott 3) (Figure
3B).
parvibracteata Immelman: two-colporate, areolate, 1-2 rows areolae,
length 35 (37,9) 42 Aim ( Leistner 1402, 1657).
platysepala (S. Moore) P.G. Mey. : two-colporate, margocolpus entire,
length 32 (37,4) 44 Aim ( Giess , Volk & Bleissner 5863) (Figure
5C).
protracta (Nees) T. Anders, (both subspp.): two-colporate, areolate, 1—2
rows areolae, length 30 (41,9) 48 Aim (subsp. protracta: Hall 4563:
subsp. rhodesiana: Germishuizen 974).
stachytarphetoides C.B. Cl.: three-colporate, margocolpus entire, length
not measured (Tinley 2624).
thymifolia (Nees) C.B. Cl.: two-colporate, margocolpus entire, length
53 (60,6) 68 Aim (Oliver, Tolken & Venter 61).
Siphonoglossa
leptantha (Nees) Immelman (both subspp.): two-colporate, areolate, 2—3
rows areolae, length 58 (64,4) 77 A<m (subsp. leptantha: Galpin
7752, 10856', subsp. late-ovata (C.B. Cl.) Immelman: Fourcade
3724) (Figure 5A, B, D, E, F).
linifolia (Lindau) C.B. Cl.: two-colporate, areolate, 2—3 rows areolae,
length 74 (83,4) 90 Aim ( Meeuse 10093).
nkandlaensis Immelman: two-colporate, areolate, three rows areolae,
length 71 (79,4) 88 Aim (Wells 2495).
Key to pollen of Justicia and Siphonoglossa in southern
Africa
Pollen longer than 58 pm, with 2—3 rows areolae ... Siphonoglossa
Pollen usually shorter than 58 Aim or, if longer, then with
one row of areolae or margocolpus entire Justicia
Key to pollen of sections of Justicia in southern Africa
(Sectional delimitation follows Graham 1989)
la Pollen 2-porate or 2-colporate:
2a Pollen with entire margocolpus sect. Justicia p.p.
(J. orchioides, J. cuneata, J. thymifolia, J. guerke-
ana, J. platysepala)
2b Pollen with areolate margocolpus:
3a Pollen usually 2-porate (in southern African species),
22—36 pm long .. sect. Rostellularia subsect. Ansellia p.p.
(J. anselliana, J. anagalloides, J. crassiradix)
3b Pollen 2-colporate, 34—60 Aim long sect. Hamieria
(J. protracta, J. dinteri, J. parvibracteata, J. capen-
sis, J. odora)
lb Pollen 3-colporate:
4a Pollen with entire margocolpus sect. Raphidospora
(J. glabra, J. campylostemon)
sect. Justicia p.p.
(J. bolusii)
sect. Betonica
(J. betonica, J. montis-salinarum)
4b Pollen with areolate margocolpus:
5a Pollen shorter than 40 Aim
sect. Rostellularia subsect. Ansellia p.p.
(J. minima)
5b Pollen longer than 40 Aim sect. Tyloglossa
(J. flava, J. kirkiana, J. petiolaris)
CONCLUSION
A number of different pollen types were found in the
southern African species of Justicia. Either two or three
colpi were found, and the margocolpus was either entire
or broken up into circular areolae. Most species examined
had an elongated colpus, except for two southern African
species of Justicia where the colpus was lacking, one
tropical African species where it was short, and another
tropical African species where it was replaced by an extra
row of areolae. It was possible to divide the southern
African species of Justicia from Siphonoglossa on pollen
characters, and also to partially divide the southern African
species of Justicia into sections on the same basis.
FIGURE 5. — Pollen of Justicia and Siphonoglossa species, 2-colporate, margocolpi areolate. A, B, D— F, Siphonoglossa leptantha subsp.
leptantha, Galpin 77512: A, face, x 900; B, side, X 900; D, surface detail, X 5200; E, surface detail showing areolae, x 5200;
F, abnormal trilete grain, X 900. C, J. platysepala, Giess, Volk & Bleissner 5863, face, X 1000.
156
Bothalia 19,2 (1989)
The South African species of Siphonoglossa had two-
colporate pollen with areolae and long colpi. On the basis
of the pollen characters the genus would seem to most
resemble Justicia sect. Hamieria.
ACKNOWLEDGEMENTS
This work was carried out as part requirement for a
Ph.D. thesis in the Department of Botany, University of
Natal, Pietermaritzburg. I would like to thank my super-
visor, Dr F. Getliffe Norris, for her help and advice in
the writing up of my thesis, and Mrs S. Perold of the
Botanical Research Institute, Pretoria, for her patient
assistance with the Scanning Electron Microscope and the
photography.
REFERENCES
BALKWILL, K. & GETLIFFE NORRIS, F. 1985. Taxonomic studies
in the Acanthaceae: the genus Hypoestes in southern Africa. South
African Journal of Botany 51: 133—144.
BALKWILL, K. & GETLIFFE NORRIS, F. 1988. Classification of the
Acanthaceae: a southern African perspective. Monographs in
Systematic Botany 25: 503—516.
BHADURI, S. A. 1944. A contribution to the morphology of pollen grains
of Acanthaceae and its bearing on taxonomy. Calcutta University
Journal, Department of Science N.S. 1: 25—38.
BREMEKAMP, C.E.B. 1965. The delimitation and subdivision of the
Acanthaceae. Bulletin of the Botanical Survey of India 7: 21—30.
ERDTMAN, G. 1952. Pollen morphology and plant taxonomy: 18.
Almqvist & Wiksell, Stockholm.
GIBSON, D.N. 1972. Studies in American plants HI. Fieldiana (Botany)
34: 57-87.
GRAHAM, V.A.W. 1989. Delimitation and infra-generic classification
of Justicia (Acanthaceae). Kew Bulletin 43: 551—620.
LINDAU, G. 1895. Acanthaceae. In A. Engler & K. Prantl, Die Natur-
lichen Pflanzenfamilien 4, 3b: 274—354. Engelmann, Leipzig.
MUNDAY, J. 1980. The genus Monechma Hochst. (Acanthaceae tribe
Justicia) in southern Africa. M.Sc. thesis, University of the Wit-
watersrand, Johannesburg.
RADLKOFER, L. 1883. Uber den systematischen Werth der Pollen-
beschaffenheit bei den Acanthaceen. Sitzungsberichte der
Bayerischen Akademie der Wissenschafien 13: 256—314.
STEARN, W.T. 1971. Taxonomic and nomenclatural notes on Jamaican
gamopetalous plants. Journal of the Arnold Arboretum 52:
614-648.
TAYLOR, J., HOLLINGSWORTH, P.G. & BIGELOW, W.C. 1974. Scan-
ning electron microscopy of liverwort spores and elaters. The
Bryologist 77: 281—327.
Bothalia 19,2: 157-160 (1989)
Studies in the genus Riccia (Marchantiales) from southern Africa. 13.
A new species, R. hantamensis , in section Pilifer and a new record for
R. alatospora
S. M. PEROLD*
Keywords: Marchantiales, Riccia, section Pilifer, southern Africa, taxonomy
ABSTRACT
Riccia hantamensis, a new species from the Hantams Mountain, District Calvinia, is described. It is clearly related to
R. alatospora, but is a much larger plant. Like other species with a dorsal epithelium of free-standing cell pillars, it belongs
to section Pilifer Volk. R. alatospora, originally only known from Platklip, Stellenbosch (Volk & Perold 1985), is now also
recorded from Carolusberg, Hester Malan Reserve, Namaqualand.
UITTREKSEL
Riccia hantamensis, ’n nuwe spesie van die Hantamsberg, distrik Calvinia, word beskryf. Dit is duidelik verwant aan
R. alatospora, maar is ’n veel groter plant. Soos die ander spesies met ’n dorsale epiteel van vrystaande selpilare, behoort
R. hantamensis ook tot seksie Pilifer Volk. R. alatospora , oorspronklik slegs bekend van Platklip, Stellenbosch (Volk &
Perold 1985), word nou ook vermeld van Carolusberg, Hester Malan Reservaat, Namakwaland.
R. hantamensis is known from only three collections
from the Hantams Mountain, all in the same vicinity
(Figure 1), but this whole area is undercollected as far as
liverworts are concerned, so it may be more widely
distributed. The specimen, Germishuizen 4034, consists
of only male plants and has been cultivated for 26 months,
growing luxuriantly in a seed tray on soil overlying peat.
Subsequently female plants with ripe sporangia were
collected at ± the same locality on two separate occasions.
The species grows on ± neutral to somewhat alkaline soil,
together with small Crassula spp. and mosses, e.g.
Didymodon ceratodonteus (C. Miill.) Dix. in an area with
predominantly fynbos vegetation. Altitude is ± 1 500 m
above sea level, the annual rainfall is a winter one of less
than 200 mm. The specific epithet refers to the only known
locality.
Riccia hantamensis Perold, sp. nov.
Thallus dioicus, annuus, grandis, in vivo laete viridis
aspectu mtido crystallino, in sicco flavo-virens. Frons
usque ad 10 mm longa, 2,5 -3,8 mm lata, 0,7— 1,2 mm
crassa, 3-3,5 plo latior quam crassa, symmetrice furcata
vel bifurcata, oblonga vel obovata. Anatomia: epithelium
dorsale e cellulis 3( — 4) in columnis liberis, decrescenti-
bus; tela assimilans canalibus aeriis 4—6—8 lateralibus ad
100 ^im latis. Squamae parvae, hyalinae, apicem versus
ventraliter positae. Sporae 60—85 pm diametro, ligno-
brunneae, deltoideo-globulares, polares, ala usque ad 10
pm lata; superficie distale areolis 4 grandis centralibus,
15-20 pm latis, bulla centrali; superficie proximale signo
triradiato distincto, areolis 5-10 pm latis, parietibus ad
nodos prominentibus. Chromosomatum numerus n = 9
(Bomefeld pers. comm.).
TYPE. — Cape Province, 3119 (Calvinia): Hantams
Mountain, Van Rhynshoek Farm, 8 km to FM tower, on
* Botanical Research Institute, Department of Agriculture and Water Sup-
ply, Private Bag X101, Pretoria 0001.
MS. received: 1989.01.25.
clayey soil at streamlet next to road (— BD), 1987.10.03,
S.M. Perold 1830 (PRE, holo.).
Thallus dioicous, annual, bright green, with shiny,
crystalline appearance, in crowded gregarious patches or
scattered, medium-sized to rather large; branches once or
twice symmetrically furcate (Figures 2A; 3A, B), closely
to medium divergent, up to 10 mm long, terminal segments
mostly short, 1,5 — 3,0 mm long, 2,5-3, 8 mm wide,
0,7— 1,2 mm thick, i.e. 3 to 31;2 times wider than thick in
section, shape oblong to obovate, generally widening
distally; apex rounded to truncate (Figure 3C), shortly
emarginate; groove apically deep, disappearing ± midway
along length of thallus; margins rounded, obtuse, over-
hanging, flanks very obliquely sloping (Figure 2G), ventral
surface gently rounded to flat, green (Figure 2G); when
dry, yellowish green, margins raised and incurved, dorsal-
ly flat to slightly concave (Figure 2B). Anatomy, dorsal
FIGURE 1. — Map showing distribution of R. hantamensis and
R. alatospora.
158
Bothalia 19,2 (1989)
FIGURE 2. — Riccia hantamensis . Morphology and anatomy. A, male
thallus with rows of antheridial necks, turgid; B, dry female thallus
with 2 archegonial necks; C, dorsal cells and air pores (hatched
lines) from above; D, transverse section through dorsal epithelium
and assimilation tissue, showing wide air canals (cells with broken
outlines); E, horizontal section through assimilation tissue, with
air canals stippled; F, scale; G, transverse section through thallus
branch. A, C, D, E, G, Germishuizen 4034\ B, F, S.M. Perold
1830. Drawings by J. Kimpton. Scale bars on A, B, G = 1 mm;
C— E = 50 jim; F - 100 ^m.
epithelium ± 135—160 high, in free tapering pillars,
consisting of 3( — 4) thin-walled hyaline cells, 40—58 x
48—80 gm, topmost cells small, rounded to conical, basal
cells wide and bulging at sides (Figures 2C, D; 3E, F),
air pores 3—5-sided, proximally wider and clearly visible
from above; assimilation tissue + 350 gtm thick in section,
i.e. almost 1;2the thickness of thallus, consisting of cells
± 50 x 35 gem, arranged in vertical columns up to 8 cells
high and 1(— 2) cells wide, with abundant chloroplasts,
enclosing 4—6—8-sided, obliquely sloping air canals, up
to 100 gem wide (Figure 2D, E); storage tissue ± 180—200
gem thick, + ]/4 the thickness of thallus, cells roundish,
62 —70 x 55 — 85 gem. Rhizoids arising from ventral
epidermis, mostly smooth, some tuberculate, ± 25 gem
wide. Scales hyaline, small and inconspicuous, 600 x 325
gem, fragile, situated ventrally toward the apex, spaced
(Figures 2F; 3D), cells ± 75 x 50 gem, 4—6-sided.
Antheridia numerous along groove, with conspicuous,
hyaline necks, up to 500 gem long. Archegonia with purple
necks, scattered along length of lobes in female plants.
Sporangia bulging slightly dorsally, up to 700 gem wide,
containing 900-1 200 spores each. Spores (60-)
65— 80(— 85) gem in diameter, pale yellow-brown, semi-
transparent, triangular-globular, polar, with wing up to 10
gem wide, perforated at marginal angles and occasionally
also elsewhere, margin finely crenulate; distal (outer) face
with ± 4 large, central areolae, 15—20 gem wide, some
with central boss (Figure 4B, D, E), outer areolae smaller,
5—12 gem wide, walls extending across wing; proximal
(inner) face with triradiate mark distinct (Figure 4C),
suture ridges + 5 gem high and extending on to wing,
areolae 5—10 gem wide (Figure 4A, C), angular, walls
raised at nodes, often irregular and incompletely separating
the areolae. (The size and ornamentation of the spores
were quite variable when comparing the two gatherings,
S.M. Perold 2338 and 1830, from the same site (Figure
4A, B & 4C— E respectively), the former having been
collected during drought, when the spores were smaller
and more numerous in each sporangium; the latter were
collected during good rains. Chromosome number n =
9 (Bornefeld pers. comm.).
Riccia hantamensis is one of several species which have
a dorsal epithelium consisting of free cell pillars and hence
it is classified in section Pilifer Volk. It is closely related
to, but distinguished from R. alatospora Volk & Perold,
mainly on account of its robust size. In addition, the apical
cells of its free dorsal epithelial pillars are more rounded
and the inconspicuous scales are hyaline and occur
ventrally only near the apex (Figure 3D), in contrast to
the red scales present along most of the length of the flanks
in R. alatospora. The somewhat wider 4—6—8-sided air
canals in the assimilation tissue and the wide-winged
spores with their very distinctive ornamentation, are
characters shared by both species, but the spores of R.
hantamensis are smaller in diameter (Figure 4F), and far
more numerous in each sporangium. The chromosome
number for R. hantamensis is n = 9, in comparison with
n = 8 in R. alatospora, but this may be of less significance
as more than one karyotype is known in some species (Volk
et al. 1988).
Although the spore ornamentation of the two species
is very similar, the differences in thallus morphology
mentioned above, warrant recognition of R. hantamensis
at specific level. Cultures of the two species which were
maintained side by side in a Petri dish for nine months,
retained the differences in size and morphology. Both
species are probably derived from a common ancestor,
Bothalia 19,2 (1989)
159
FIGURE 3. — Riccia hantamensis. Morphology and anatomy. A, dorsal view of thallus, branches once furcate; B, branches twice furcate;
C, apex of thallus from above; D, ventral scales and rhizoids shown from side; E, dorsal cells and air pores from above; F, dorsal
cell pillars partly from the side. A— F, Germishuizen 4034. Scale bars on A— F = 100 /im.
as indicated by similarities in the more conservative
sporophyte.
Specimens examined
CAPE. — 3119 (Calvinia): Hantams Mountain, Van Rhynshoek Farm
(— BD), 1986 October, G. Germishuizen 4034 (PRE); 1987 October, S.M.
Perold 1830 (PRE) (see type); 1988 September, S.M. Perold 2338 (PRE).
A NEW RECORD OF R. ALATOSPORA VOLK & PEROLD
Until now, this species was known only from Platklip,
Stellenbosch, where it was first collected by Duthie in
1929. Two more collections of R. alatospora were recent-
ly made (Figure 1):
CAPE. — 2918 (Gamoep): Carolusberg, Hester Malan Res. (— CA),
S.M. Perold 1425 p.p., 1426 (PRE).
FIGURE 4. — Riccia hantamensis and R. alatospora. Spores. A— Fa, R. hantamensis', Fb, R. alatospora. A, C, proximal face; B, D, distal
face; E, distal face, partly from the side; A-E, SEM micrographs; F, LM (light microscope) micrograph. A, B, Fa, S.M. Perold 2338\
C-E, S.M. Perold 1830\ Fb, Duthie 5004. Photography by S.M. Perold. Scale bars on A-E = 50 pm\ diameter of Fb ± 120 pm.
160
Bothalia 19,2 (1989)
ACKNOWLEDGEMENTS
I wish to sincerely thank Mr G. Germishuizen, BRI,
Pretoria, for collecting the first specimen of this new
species and for proposing the specific epithet; also Dr T.
Bomefeld, Am Reelein 1, Hochberg, F.R.G. for doing the
chromosome counts and Prof, (emer.) Dr O.H. Volk,
Wiirzburg for his advice and fruitful discussions. I am most
grateful to Dr H.F. Glen, BRI, for the Latin diagnosis and
to Mr J. van Rooy for correcting the text.
REFERENCES
VOLK, O.H. 1983. Vorschlag fur eine Neugliederung der Gattung Riccia
L. Mitteilungen aus der Botanischen Staatssammlung, Mtinchen
19: 453-465.
VOLK, O.H. & PEROLD, S.M. 1985. Studies in the genus Riccia
(Marchantiales) from southern Africa. 1. Two new species of the
section Pilifer. R. duthieae and R. alatospora. Bothalia 15:
531-539.
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.
argenteolimbata and R. albomata. Bothalia 18: 155-163.
Bothalia 19,2: 161-165 (1989)
Studies in the genus Riccia (Marchantiales) from southern Africa. 14.
R. concava section Pilifer
S. M. PEROLD*
Keywords: Marchantiales, Riccia concava , section Pilifer, southern Africa, taxonomy
ABSTRACT
A historical account is given of the confusion caused by the application of the name R. concava to various taxa. A new
description of the species as understood by the author is given and a comparison made with related species. Its distribution
and ecology are noted.
UITTREKSEL
’n Historiese oorsig van die verwarring veroorsaak deur die toepassing van die naam R. concava op verskeie taksons word
gegee. 'n Nuwe beskrywing van die spesie, volgens die begrip van die skrywer, word weergegee en ’n vergelyking word
met verwante spesies getref. Die ekologie en verspreiding word ook vermeld.
R. concava was initially named by Bischoff (in MS) and
described by Gottsche, Lindenberg & Nees ab Esenbeck
(1844) from a specimen collected by Krauss in 1838, on
decomposing granite, at the Cape of Good Hope. They
placed it in their group C, 'Subtus Squamatae’, noting that
the fronds were glaucous on both sides, becoming whitish
when dry, fan-shaped, narrowly canaliculate, 2-4-lobed,
occasionally nearly in rosettes, the lobes ovate, obtuse or
emarginate, concave in the dry plant, mussel-shaped, with
raised margins and underneath, toward the apex, with
scales. To them, it was among the biggest species in the
genus, with distinctive lobes in the dry plant. Then there
follows a rather puzzling observation in that, according
to Bischoff (in litt. ), the small scales of the dry plant, when
casually observed, could be taken for cilia. This is difficult
to correlate with the type material as there is no similarity
between these scales and cilia, unless Bischoff possibly
mistook the dorsal cell pillars towards the margins for cilia,
although in the dry state, they would have collapsed and
been less visible. Gottsche et al. compared it with R.
albomarginata and with R. lamellosa, but observed that
in R. concava the lobes were ovate and very concave and
the scales were only visible at the apex.
The type specimens held at G and S are fragmentary,
of poor quality and have probably been pressed. The width
of the widest branch is 3,25 mm, segments are 4 mm long
and the branches up to 7 mm long, the margins are partly
indexed and the flanks here and there have faint purple
colouring; the dorsal cells have collapsed and cannot be
measured; a scale from the apex is 850 x 250 gm, cells
in the body of the scale are oblong-hexagonal, up to 125
x 62 fxm, with smaller cells at the margin. There are no
spores. Enclosed with the isotype specimen (G) and
mounted between two cover-slips, is a dried transverse
section of a thallus branch (by Lindenberg); it is 1,75 mm
wide and 0,65 mm thick, with the dorsal cells collapsed,
the upper surface is ± concave, the flanks ± rounded and
the margins slightly raised and subacute.
* Botanical Research Institute, Department of Agriculture and Water
Supply, Private Bag X101, Pretoria 0001.
MS. received: 1989.01.25.
Stephani (1898) originally placed R. concava in his
Inermes (i.e. without cilia) group IV, but after examining
the 'original' plant, transferred it to his group VII (Frons
Crassa). He, however, expressed doubts that the plant had
been sufficiently studied by Gottsche et al. , because they
had compared it with R. albomarginata which has thin
lobes and with R. lamellosa, which is fleshy. In his leones
hepaticarum (Stephani 1876—1907) (G, M), two widely
different cross sections of the thalli of R. concava are
illustrated, one very thin and slightly concave, with acute,
winged margins and the other thicker and concave, with
obtuse margins. He also described the scales as large and
extending above the thallus margins.
In important aspects, Sim’s (1926) description is not
correct: these plants do not truly grow in rosettes, nor did
he make any mention of the loose dorsal cell pillars, only
noting that there are 'upright pillars of lax, chlorophyllose
cells with a larger globose epidermal cell on each’. Further-
more, he stated that the spores, 80 (not 8) ^im in diameter,
are ‘laxly reticulated with about 5 areolae on the diameter
each way’. Both these characters, viz. a single globose
dorsal cell and only about 5 areolae across the diameter
of the spore, suggest a different species altogether.
Duthie and Garside did not publish anything on R.
concava', a note of Duthie’s was found with a Potts
specimen, CH1010, to the effect that she was not at all sure
of the differences between R. albomarginata and R.
concava. There are, however, several very good Duthie
collections of R. concava (and presumably named by her)
at BOL and S, notably Duthie 5005.
Arnell’s (1963) description may have been based on the
correct species, but he did not cite any specific collections,
so this can only be checked on specimens named by him;
however, here the dorsal cells can no longer be examined
reliably and the spore ornamentation, as stated below, is
somewhat variable. The width at 1,5 mm, which he
reported for the thallus is rather narrow, but the colour,
'glaucous-yellow green', is correct, although also
applicable to other species. The inflated dorsal cells are
not correctly illustrated, as they are longer than wide and
162
Bothalia 19,2 (1989)
[CURE 1 — Riccia concava. Morphology and anatomy. A, fresh thallus, dorsal view; B, fresh thallus, ventral view, C, driea thallus, D1 6,
transverse sections of thalluT branch at different distances from apex to older parts; E, epithelial cells and air pores (hatch**1) from
above F transverse section through dorsal epithelial cell pillars; G, paradermal section through assimilation tissue, H, scale. A B
D SM Sold 143^ C S Perold 1899 ; E, H, Morle} 214, F, S.M. Perold 1447 ; G, Moll 6015. Drawings by J. Kimpton. Scale
bar on A-D = 1 mm; E-G = 50 fim; H = 100 /im.
Bothalia 19,2 (1989)
163
partly collapsed, whereas the ‘hairlike cell pillars’ at the
margin, are the scale cells in cross section; spores with
thick ridges radiating from the centre as drawn by him,
is not a character unique to R. concava. As also concluded
by Volk (1981), it appears highly probable that Sim and
Arnell did not describe the same species.
The specimens Arnell referred to R. concava at PRE
and S have been examined; some have certainly been
correctly determined, but others doubtfully so (see
Specimens examined). It seems likely that he lumped
together as R. concava all those species with loose dorsal
pillars (described by him as velvet-like), which he could
not refer to either R. albomarginata sensu Sim or to R.
villosa. That this is so, is also suggested by the wide
distribution he claimed for it (see below).
Riccia concava Bisch. in Gottsche, Lindenberg &
Nees ab Esenbeck. Synopsis hepaticarum: 604 (1844);
Stephani: 325, 378 (1898); Sim: 12 (1926); Arnell: 22
(1963).
TYPE. — Cape, in Capite Bonae Spei locis humidis,
in saxis graniticis decompositis. Krauss s.n. , 1838 (G8978
in G, iso.!) ex Herb. Musci. Palat. Vindob.; (S, iso.! fide
Grolle 1976 : 226) ex Herb. Lehmannianum.
Thallus monoicous, perennial, in crowded gregarious
patches, not truly in rosettes (Figure 2A); blue-green (to
± yellowish green when actively growing), becoming
whitish along margins, with scales not, or hardly
protruding, except at apex where visible from above, also
when wet and fully expanded (Figure 1A); medium-sized
to large, branches once or twice furcate, rarely simple;
medium to widely divergent, 6-8(-10) mm long, 3-4
mm wide, 0,9— 1,2 mm thick, i.e. 3-4 times wider than
thick in section, broadly ovate to obovate, apex rounded,
emarginate, dorsally deeply grooved toward apex (Figure
2B-D), sides convex, soon becoming flat to slightly
concave proximally, margins acute to subacute, shortly
winged, somewhat recurved, flanks sloping obliquely
upward and outward (Figure 1D1-6), ± arched, pale
mauve to dark purple, covered by scales; ventral surface
rounded, green to purple laterally (Figure IB); when dry,
glaucous to blue-green or mauvish green, scurfy or flaky,
dorsally markedly concave, margins raised and incurved,
partly inflexed (Figure 1C), occasionally almost meeting,
exposing flanks clothed with wrinkled, mosdy dull whitish
to pale cream-coloured scales, basally often with mauve
streaks across or with purple sheen from dark purple flanks
underneath.
Anatomy of thallus : dorsal epithelium consisting of
3(-4)-celled, free-standing, hyaline pillars (Figure 2E, F),
± 180—260 /xm long, top cell smallest, globose or conical,
35—42 x 45—60 /xm, frequently collapsed, second cell
usually wider and inflated, 50-62 /xm long and up to 85
/xm wide, basal cell(s) ± rectangular, 50-75 x 68-75 /xm
(Figure IF); on dorsal face seen from above, cells in apical
parts closely packed, inflated, shiny, like small round glass
beads, in rows, cells in basal parts less orderly arranged
and collapsing; air pores small, 4—5-sided (Figure IE);
assimilation tissue ± 450 /xm thick in section, occupying
almost \ the thickness of thallus and consisting of
vertical columns of 6-8 short- rectangular cells, up to 55
x 43 /xm, enclosing narrow 4—5-sided air canals
(Figure 1G); storage tissue ± 300 /x m thick, ± !/3 (or less)
the thickness of thallus, cells crowded together, round to
angular, 55—62 /xm wide; rhizoids arising from ventral
epidermis ± 20 /xm wide, some smooth, others tubercu-
late. Scales semicircular, imbricate, hyaline, not or hardly
extending beyond thallus margins except at apex, mostly
ventrally situated along concave flanks, 900—1 200 x 600
/xm, cells in body of scale long rectangular, or 5—6-sided,
up to 160 /xm long x 50—65 /xm wide, walls straight, at
scale margin one row of smaller cells, wider than long
(Figure 1H). Antheridia with hyaline necks ± 250 /xm
long, in 2 rows along middle of lobes. Archegonia with
purple necks. Sporangia ± 500 /xm wide, single or in
pairs, each with + 350 spores, bulging dorsally, overlying
tissue disintegrating and liberating spores. Spores
75— 90(— 100) /xm in diameter, triangular-globular, polar,
dark brown, with narrow wing up to 5 /xm wide, angles
notched or with a pore, margin finely crenulate; ornamen-
tation somewhat variable, reticulate to vermiculate, or with
radiating ridges: distal face with 10-14 deep-set areolae
across the diameter, up to 7,5 /xm wide, some with a central
papilla, radial walls thick (Figure 3C), often dusted with
granules (Figure 3D), usually raised at nodes, occasionally
forming short, irregular ridges radiating outwards from
centre (Figure 3E, F); proximal face with triradiate mark
quite prominent, sparsely granular, numerous (30—40)
small round areolae on each facet, walls raised at nodes
(Figure 3A, B). Chromosome number n = 8 (Bornefeld
(1984) on S.M. Perold 470, 485).
R. concava can be distinguished from the other species
in section Pilifer Volk (1983), by its broad thallus, up to
4 mm wide when fully expanded, concave when dry,
glaucous or scurfy blue-green colour, rounded apex,
somewhat overhanging margins mostly obscuring the
scales except those at the apex, and fragile, inflated,
generally wider than long dorsal cells in loose pillars. On
exposure to bright sunlight, it develops a deeper purple
colour at the flanks and ventrally.
In this section, most of the other species, currently
totalling ± 12 species and some still to be described, have
conspicuous hyaline scales, except for R. alatospora Volk
& Perold (1985) and R. hantamensis Perold (1989). In R.
albomarginata sensu Sim [the name has been misapplied
since Sim (1926) — Perold in prep.], the dorsal cell pillars
tend to be more persistent, tall and narrow with all the
basal cells ± equally long; R. villosa is easily recognized
by large, triangular scales and papillose spores; R. parvo-
areolata Volk & Perold (1984) has spores with numerous
small areolae; the rest of the species have tapering or
uniform pillars with cells that are generally longer than
wide. To examine the dorsal cells, living material is
required, as they cannot be reconstituted in long-dried
herbarium material.
Differences in the spore ornamentation between R.
concava, R. albomarginata Bisch. ex G.L. & N., and one
or two new, as yet undescribed species in the section, are
sometimes quite difficult to discern, even on SEM
micrographs, nor are radiating ridges on the distal face
altogether distinctive. Moreover, the spores of R. concava
can be quite variable, even when from the same
sporangium.
R. concava often grows in association with other Riccia
species of section Pilifer and with the moss species
164
Bothalia 19,2 (1989)
FIGURE 2. — Riccia concava. Morphology and anatomy. A, thalli not in rosettes; B, dorsal view of thallus; C, D, apex with dorsal groove;
E, F, dorsal cell pillars. A— F, S.M. Perold 2312. Scale bar on A-C = 1 mm; D— F = 50 /im.
FIGURE 3. — Riccia concava. Spores. A, B, proximal face; C, D, F, distal face; E, radiating ridges on distal face. A, S.M. Perold 2313',
B, S.M. Perold 1791', C, S.M. Perold 1500', D, S.M. Perold 1773', E, F, Garside 6128. Photography by S.M. Perold. Scale bar on A-E
= 50 ^m; diameter of spore on F, ± 90 fim.
Barbula crinita Schultz, Desmatodon convolutus (Brid.)
Grout and Chamaebryum pottioides Ther. & Dix. It prefers
damp, not wet places, being found away from seepages,
on sandy, well drained soil overlying granite outcrops. It
is fairly common in the north-western, western, south-
western and southern Cape Province (Figure 4). Besides
the western Cape, Sim (1931) and Arnell (1963) also
reported it from Natal, Transvaal, S Rhodesia, Portuguese
East Africa, Madagascar and the Canary Islands, but this
has not been verified (see below). (At PRE, a Sim
specimen from Magude, P.E.A. [Mozambique] has been
identified as R. concava, but it has black scales.)
Arnelfs specimens, Arnell s.n., 11.3.59 ( UPS 20635 )
(Lagunetas); 13.3.59 (UPS 20636) (Cueva Grande) and
28.2.59 ( UPS 20637) (La Calzada) from the Canary
Islands, which he named R. concava, do not belong here,
as the shape of the thalli in transverse section, the width
to thickness proportions and the spore ornamentation with
fewer and larger areolae on the dorsal face and granules
Bothalia 19,2 (1989)
165
FIGURE 4. — Distribution map of Riccia concava in southern Africa,
on the proximal face, do not correspond with those of R.
concava. It should be deleted from the species list of the
Macaronesian Islands (Arnell 1961; Dull 1984; Eggers
1982; Sergio 1984). These plants do, however, belong to
section Pilifer Volk, which is unique to southern Africa,
except for the above, and for specimens from lie de l’Est
(Crozet Archipelago), assigned to R. albomarginata by
Jovet-Ast (1986). Unfortunately the identity of Arnell’s
specimens from Gran Canaria is still uncertain and will
probably remain so, unless fresh material can be collected.
Regardless of which species of section Pilifer they belong
to, their original dispersal from southern Africa to
Macaronesia, whether by man or by migrating birds, is
highly probable.
SPECIMENS EXAMINED
CAPE. — 2917 (Springbok): Springbok, opposite Country Club, rock
outcrops (-DB), S.M. Perold 1414, 1415 (PRE); 14 km N of Springbok
at edge of large granite dome (-DB), S.M. Perold 2054, 2057 (PRE);
S of Springbok, 6 km from Kokerboom Motel on road to Kamieskroon
(— DD), S.M. Perold 1438 (PRE). 2918 (Gamoep): Carolusberg, near
gate (-CA), S.M. Perold 1431, 1432 (PRE). 3017 (Hondeklipbaai): 2
km N of Kamieskroon, granitic rock outcrops (-BB), S.M. Perold 1447,
1454, 1455, 2091, 2094 (PRE); Kamiesberg Pass, drier area above seepage
(— BB), S.M. Perold 1604 (PRE); Rietkloof, 14 km S of Kamieskroon,
edge of rock outcrop (-BD), S.M. Perold 2103-2105 (PRE); Brakdam,
31 km S of Kamieskroon, rock outcrops (-BD), S.M. Perold 2113, 2115
(PRE). 3018 (Kamiesberg): 18 km NE of Kamieskroon, on road to
Rooifontein (-AA), S.M. Perold 1460, 1465, 1466 (PRE); 4-5 km along
road to Rooifontein, from Kamieskroon-Leliefontein road (-AA), S.M.
Perold 2148, 2172, 2173 (PRE); Pedroskloof, on road to Rooifontein
(-AA), S.M. Perold 1493 (PRE); Kamassies, large rock outcrop (-AB),
S.M. Perold 1500, 1501 (PRE). 3119 (Calvinia): E of Slagberg, between
Nieuwoudtville and Loeriesfontein, Farm Koringhuis (-AB), S.M. Perold
1795, 1798 (PRE). Nieuwoudtville, sandstone outcrops 2 km SW of town
(-AC), S.M. Perold 2195 (PRE); Nieuwoudtville Falls (—AC), S.M.
Perold 1788, 1791, 1792, 2312, 2313, 2316 (PRE); Nieuwoudtville, Farm
Oorlogskloof (—AC), C.M. van Wyk 1493 (PRE); Van Rhyns Pass, in
ditch on plateau (—AC), S.M. Perold 2185 (PRE); Groothoek, 18 km
on dirt road to Rondekop, Soetlandsfontein River (—AD), S.M. Perold
1773 (PRE). 3219 (Wuppertal): Biedouw Youth Camp, sandstone rock
outcrops (-AA), S.M. Perold 1888 (PRE); Algeria Forest Station, 4 km
S of streambank (—AC), S.M. Perold 2362 (PRE). 3220 (Sutherland):
near Sutherland (-BC), Duthie 5407 (BOL); Montagu, Bath Kloof
(-CC), Arnell 753 (BOL, S); Klein Roggeveld, De Kom, clay soil over
shale (-DA), Oliver 8949 (PRE); Haashoogte, damp E slope (-DA),
Oliver 8957a (PRE); 50 km S of Sutherland, 21 km along road to Wolf-
hoek, Farm Bergsig, streambank (-DA), S.M. Perold 2426, 2427 (PRE).
3318 (Cape Town): Darling, 5 km S of, (-AD), S.M. Perold 485 (PRE);
Lion’s Head above Fresnaye (—CD), Arnell 12a, 67a (BOL); Signal Hill
(-CD), Garside 6128 (BOL); Wellington (-DB), Duthie 5470 (BOL);
Stellenbosch, clayey ground below municipal farm near railway line
(-DD), Duthie 5005 (BOL, S); railway embankment (-DD), Duthie
5417 p.p. (BOL); E end of Stellenbosch Flats near farm, on earth bank
of sloot (-DD), Garside 6108 (BOL); Stellenbosch, Platklip (— DD),
Morley 214 (PRE), S.M. Perold 470 (PRE); Stellenbosch, Papegaaiberg
(— DD), S.M. Perold 478 (PRE). 3319 (Worcester): Tulbagh (—AC),
Duthie 5468 (BOL). 3322 (Oudtshoorn): Meiringspoort, on disturbed
soil at roadside (-BC), S.M. Perold 899 (PRE).
Only specimens which unequivocally could be referred
to R. concava, have been included in the above list. New
collections have all been cultivated and observed over a
period of time. If there was the slightest element of doubt
about the identity of old collections from the western and
south-western Cape Province, that had been named by
Duthie and by Garside, or by Arnell, they have been
excluded.
ACKNOWLEDGEMENTS
The author wishes to thank the curators of BOL, S and
UPS for the loan of specimens; Prof, (emer.) Dr O.H. Volk
of Wurzburg University, W Germany, for numerous
discussions concerning R. concava, its identification and
distribution and Dr (habil.) T. Bornefeld, Am Reelein 1,
Hochberg, W Germany, for the chromosome counts.
Sincere thanks to Mrs A.J. Romanowsky for developing
and printing the micrographs; to the artist, Ms J. Kimpton
and to Mrs M. van der Merwe for typing the manuscript.
REFERENCES
ARNELL, S. 1961. List of the Hepaticae of the Canary Islands. Svensk
botanisk Tidskrift 55: 379—393.
ARNELL, S. 1963. Hepaticae of South Africa. Swedish Natural Science
Council, Stockholm.
BORNEFELD, T. 1984. Chromosomenanalyse der Gattung Riccia von
Slid- und SW-Afrika und allgemeine Bemerkungen zur Zytogene-
tik der Lebermoose. Nova Hedwigia 40: 313-328.
DULL, R. 1984. Bryogeographische Analyse der Makaronesischen Inseln
mit besonderer Beriicksichtigung der Kanarischen Inseln Tenerife
und La Palma. Proceedings of the Third Meeting of the Bryolo-
gists from Central and East Europe, Praha, 14—18 June, 1982 :
177-190. Univerzita Karlova, Praha.
EGGERS, J. 1982. Artenliste der Moose Makaronesiens. Cryptogamie,
Brvologie et Lichenologie 3: 283—335.
GOTTSCHE, C.M., LINDENBERG, J.B.G. & NEES AB ESENBECK,
C.G. 1844. Synopsis hepaticarum. Hamburg. Reprinted 1967.
GROLLE, R. 1976. Verzeichnis der Lebermoose Europas und benach-
barter Gebiete. Feddes Repertorium 87: 171-279.
JOVET-AST, S. 1986. Riccia de l’Archipel des Crozet (lie de L'Est).
Cryptogamie, Bryologie et Lichenologie 7: 479—485.
PEROLD, S.M. 1989. Studies in the genus Riccia (Marchantiales) from
southern Africa. 13. A new species, R. hantamensis in section
Pilifer, and a new record for R. alatospora. Bothalia 19: 157—160.
PEROLD, S.M. in prep. Studies in the genus Riccia (Marchantiales)
from southern Africa. 16. R. albomarginata and R. simii sp. nov.
Bothalia.
SERGIO, C. 1984. The distribution and origin of Macaronesian bryo-
phyte flora. Journal of the Hattori Botanical Laboratory 56: 7— 13.
SIM, T.R. 1926. The bryophytes of South Africa. Transactions of the
Royal Society of South Africa 15: 1-475. Cape Town.
SIM, T.R. 1931. South African Bryophyta. Further notes. Transactions
of the Royal Society of South Africa 20: 15—17.
STEPHANI, F. 1876—1907. leones hepaticarum. Ed. P. Geissler. IDS
Microform Publishers, Zug, Switzerland. Publ. 1986.
STEPHANI, F. 1898. Species hepaticarum. Bulletin de THerbier Boissier
6: 309-411.
VOLK, O.H. 1981. Beitrage zur Kenntnis der Lebermoose (Hepaticae)
aus Siidwestafrika (Namibia). II. Mitteilungen aus der Botanischen
Staatssammlung, Miinchen 17: 245—252.
VOLK, O.H. 1983. Vorschlag fur eine Neugliederung der Gattung Riccia
L. Mitteilungen aus der Botanischen Staatssammlung, Miinchen
19: 453-465.
VOLK, O.H. & PEROLD, S.M. 1984. Studies in the genus Riccia
(Marchantiales) from the south-west Cape. Bothalia 15: 117—124.
VOLK, O.H. & PEROLD, S.M. 1985. Studies in the genus Riccia
(Marchantiales) from southern Africa. 1. Two new species of the
section Pilifer : R. duthieae and R. alatospora. Bothalia 15:
531-539.
Bothalia 19,2: 167-174 (1989)
New taxa, combinations and records of Pteridophyta from southern
and central Africa
J.E. BURROWS*
Keywords: central Africa, new records, pew taxa, Pteridophyta, southern Africa
ABSTRACT
Four new taxa of ferns are described and illustrated from southern Africa: Ophioglossum convexum J.E. Burrows, Mohria
caffrorum (L.) Desv. var. ferruginea J.E. & S.M. Burrows, Marsilea farinosa Launert subsp. arrecta J.E. Burrows and
Asplenium sebungweense J.E. Burrows. The combination of Grammitis rigescens (Bory ex Willd.) J.E. Burrows is made.
Ophioglossum thomasii Clausen, O. rubellum Welw. ex A. Braun, Virtaria ensiformis Swartz and Asplenium buetmeri Hieron.
ex Brause are new records for Zimbabwe, while Hymenophyllum splendidum V.d. Bosch and Asplenium uhligii Hieron.
are new records for Malawi and Zimbabwe. Actiniopteris semiflabellata Pichi-Sermolli is recorded from Namibia and Thelypteris
oppositiformis (C. Chr.) Ching is recorded from the Transvaal.
UITTREKSEL
Vier nuwe varingtaksons van suidelike Afrika word beskryf en geillustreer: Ophioglossum convexum J.E. Burrows, Mohria
caffrorum (L.) Desv. var. ferruginea J.E. & S.M. Burrows, Marsilea farinosa Launert subsp. arrecta J.E. Burrows en Asplenium
sebungweense J.E. Burrows. Die kombinasie Grammitis rigescens (Bory ex Willd.) J.E. Burrows word gemaak. Ophioglossum
thomasii Clausen, O. rubellum Welw. ex A. Braun, Vittaria ensiformis Swartz en Asplenium buettneri Hieron. ex Brause
word die eerste keer in Zimbabwe aangeteken, terwyl Hymenophyllum splendidum V.d. Bosch en Asplenium uhligii Hieron.
die eerste keer in Malawi en Zimbabwe aangeteken word. Actiniopteris semiflabellata Pichi-Sermolli word in Namibie aan-
geteken en Thelypteris oppositiformis (C. Chr.) Ching word in die Transvaal aangeteken.
INTRODUCTION
While carrying out research on southern African
pteridophytes, it became obvious that there were a few
undescribed taxa from, as well as a number of new records
to, the area covered by Flora zambesiaca and Flora of
southern Africa, several being interesting southern
extensions of tropical species. This paper attempts to
update our knowledge of southern African ferns in the light
of recent collections and research.
1. Ophioglossum thomasii Clausen in Memoirs of
the Torrey Botanical Club 19: 152 (1938). Type: Uganda,
Kampala, Kabaka’s Lake, Thomas 1903 (BM, holo.!).
ZIMBABWE.— 2028: Matopo Hills, Besna Kobila Farm, 1 465 m,
Jan. 1956, Miller 3305 (PRE!).
Distribution: Liberia, Ivory Coast, Ghana, Nigeria,
Cameroon, Gabon, Zaire, Uganda, Tanzania and Zambia.
2. Ophioglossum convexum J. E. Burrows, sp. nov.
Rhizoma elongatum, 5—25 mm longum cum vel sine
aliquot basibus petiolorum persistentibus. Radices
proliferae. Folium unum (raro duo), ad angulum 0-30°
portatum vix supra vel terram adpressum. Petiolus 6—25
mm longus, 60-90% longitudinis subterraneus. Lamina
sterilis desuper convexa, ovata vel late ovata, late acuta
vel obtusa et breviter apiculata; basis late cuneata vel
truncata; nervatura obscura. Spica fertilis 30—100 mm
longa, ad vel vix infra basin laminae sterilis inserta;
sporangia paribus 6—15; apex acutus vel breviter
apiculatus.
* Buffelskloof Nature Reserve, P.O. Box 236, Lydenburg 1120.
MS. received: 1988.11.25.
TYPE. — Transvaal, 2530 (Lydenburg): Coromandel
Farm (—AD), Burrows 3683 (PRE, holo.; BOL, K, iso.).
Rhizome elongate, 5—25 mm long with or without a few
persistent petiole bases. Roots proliferous. Leaves one,
rarely two, held at 0-30° from the horizontal, just above
or appressed to the ground. Petiole 6—25 mm long,
60-90% of its length subterranean. Sterile lamina convex
above, ovate to broadly ovate, broadly acute to obtuse and
shortly apiculate, base broadly cuneate to truncate,
venation obscure. Fertile spike 30-100 mm long, inserted
at, or just below, the sterile lamina base, 6—15 pairs of
sporangia, apex acute to shortly apiculate. Figure 1.
ZAMBIA. — 0830: Abercorn District, ± 2,4 km above Sansia Falls,
Kalambo River, Richards 10371 (?K) [Schelpe 1970: 35 as O. rubellum],
ZIMBABWE. — 1731: Ngomakurira Mtn, Chindamora, 1 430 m,
Burrows 3040 (BOL, K, SRGH), 3046 (Herb. Burrows); Mavi 1601
(SRGH).
TRANSVAAL. — 2530 (Lydenburg): Lydenburg, Lisabon State Forest
(-BC), Burrows 3427 (PRE, Herb. Burrows). 2430 (Pilgrim’s Rest):
Pilgrim’s Rest, New Chum Falls (— DB), Roberts 102 (BOL!).
MADAGASCAR. — Grid ref. unknown: Boina, Perrier 7694 (P!) [O.
nudicaule L. f. sensu Tardieu-Blot 1951].
REUNION. — 2155: Chemin des Anglais, Cadet 956, 1203, 1258 (P!)
[all as O. nudicaule L. f.].
Within all the populations seen of O. convexum there
are large numbers of plants that do not bear fertile spikes.
These plants all have the lamina appressed flat on the
ground although those plants that carry a fertile spike
appear to have a petiole that raises the lamina 2—3 mm
above the ground, with the lamina typically held at ± 30°
from the horizontal.
O. convexum is closely related to O. rubellum Welw. ex
A. Braun (Figure 2) but differs from it in having a longer
168
Bothalia 19,2 (1989)
FIGURE 1. — Ophioglossum convexum J.E. Burrows, in natural habitat,
Lisabon State Forest, Lydenburg, Transvaal, Burrows 3427.
4). Proliferating roots as found in this species are not
unusual in the genus, although they are seldom
documented (Chen & Chiang 1972). Due to its proliferous
roots the species tends to form colonies of several square
metres.
3. Ophioglossum rubellum Welw. ex A. Braun in
Kuhn, Filices africanae: 179 (1868). Type: Angola, Pungo
Andongo, Welwitsch 33 (K, holo.!; BM, iso.!).
Icon: Tardieu-Blot: pi. 1, fig. 8 (1953).
Specimens collected on seasonally wet sandy soils in
the Sengwa Wildlife Research Area lack the reddish tinge
that gave the specific name to the type collection, but in
all other characters are identical to Welwitsch’s plants
examined by the author. There are collections from
Zambia, Kenya and Ethiopia that have smaller, single
leaves with a bluish tinge that have been attributed to O.
rubellum but they may in fact be O. convexum J.E. Burrows
(Figure 1), or an undescribed species. (See also notes
under the previous species.)
ZIMBABWE. — 1828: Gokwe, Sengwa Wildlife Research Institute, 0,5
km NE of bridge over the Sengwa Gorge, 880 m, 12.2.1983, Burrows
3019 (BOL, K, PRE, SRGH, Herb. Burrows). Figure 2.
Distribution: Ethiopia, Kenya, Uganda, Tanzania, Zambia
and Angola.
4. Mohria caffrorum (L.) Desv. var. ferruginea J. E.
& S.M. Burrows, var. nov., a var. caffrorum rhachidi
juventute perdense squamis atroferrugineis obtecta differt.
and narrower rhizome, only a single leaf ( O ; rubellum
almost always has two or more), and in having the leaf
appearing convex from above with the margins generally
curving downwards although the midrib region may be
concave, while the leaf of O. rubellum appears folded along
the midrib, sloping upwards to the margin and is never
appressed to the ground, being held at 10-20° from the
horizontal. There are a number of collections of plants
identified as O. rubellum from east and central Africa
which have a single leaf and are stated to have the lamina
lying flat upon the ground which are undoubtedly O.
convexum. The confusion has almost certainly arisen from
a misinterpretation of Welwitsch’s type from Angola which
is in Kew (isotype in BM). On the three sheets (two in
K, one in BM), there are a total of 41 plants, of which
six bear a single leaf, 29 with two and six with three leaves.
Schelpe (1970) states that O rubellum has a single leaf,
quoting and illustrating a plant collected from Zambia
{H.M. Richards 10371 — said to be in Kew but not found
by the author) that is clearly O. convexum. It is likely that
this error has been perpetuated elsewhere in Africa.
O. convexum is also similar to O. nudicaule L. f. (Figure
3). O. nudicaule sens, strict, is, in the author’s opinion,
confined to the Cape Province and has up to five leaves
per plant, each leaf being concave when viewed from
above, with the whole lamina somewhat deflexed. Like
most species of Ophioglossum, O. convexum does not
appear to be closely linked to climate or altitude, occurring
in the Transvaal in montane grassland at altitudes of
between 1 200 and 1 900 metres, but in warmer climates
and at lower altitudes north of the Limpopo River (Figure
TYPE. — Natal, (2929) Underberg: Drakensberg Mts,
Injasuti, below Women Grinding Corn (— AB), Burrows
3670 (BOL, holo.; K, PRE, iso.).
FIGURE 2. — Ophioglossum rubellum Welw. ex A. Braun, in natural
habitat, Sengwa Wildlife Research Institute, Gokwe, Zimbabwe,
Burrows 3019.
Bothalia 19,2 (1989)
169
FIGURE 3. — Ophioglossum nudicaule L. f., ex hort., 14 km from
Grahamstown on Cradock Road, Cape Province, Burrows 3685.
Rhizome erect to procumbent, 5-8 mm in diameter,
with closely packed, tufted, erect fronds. Rhizome scales
rusty brown, linear-lanceolate, attenuate, entire,
concolorous, 2—6 mm long. Stipe 40-120 mm long,
brown basally, stramineous distally when dry, variously
set with dark, reddish brown, subulate, entire scales,
0,5—2 mm long, and scattered scales near the base similar
to those on the rhizome, becoming subglabrous with age.
Lamina 120—300 x 30—60 mm, narrowly elliptic to
narrowly oblanceolate, with the basal pinnae decrescent,
3-pinnatifid. Pinnae ± 25 x 16 mm, ovate to triangular,
bluntly acute. Pinnules oblong, obtuse, deeply pinnatifid
into rounded lobes, margins widely and shallowly serrate-
crenate, involute, glabrous above, with occasional, minute
subulate scales below, both surfaces with scattered,
opaque, linear, gland-like projections ± 0,1 mm long.
Rachis and secondary rachises sulcate above, densely set
with dark, reddish brown, subulate scales similar to those
on the stipe, almost obscuring the rachis when young. Sori
submarginal, partly covered by the involute margins
(Figure 5B).
M. caffrorum var. ferruginea is separated from var.
caffrorum by its thick mat of dark, reddish brown scales
that clothe the rachis when young and by the glabrous to
subglabrous lamina which, in old pressed fronds, turns
a metallic grey. In addition, it appears to be restricted to
wet situations along the margins of mountain streams and
around springs, whereas var. caffrorum is also found on
drier soils and in the shelter of boulder bases, rock cracks,
as well as in scrub on the forest ecotone. Like M. hirsuta
J.P. Roux, it appears to be restricted to high altitudes of
between 1 700 and 2 300 m (Figure 6).
TRANSVAAL. — 2530 (Lydenburg): Lydenburg District, Die Berg
(-AA), Burrows 3677 (BOL, PRE, Herb. Burrows); Lydenburg,
Coromandel Farm (-AD), Burrows 3679 (BOL, J, PRE).
O.F.S. — 2828 (Bethlehem): Golden Gate National Park (— CB),
Roberts 3125, 3277 (PRE).
NATAL. — 2828 (Bethlehem): Mont aux Sources (— DD), Mogg 5303
(PRE). 2829 (Harrismith): Ntonjelane, Mnweni area (— CC), Esterhuysen
14523 (BOL); Lambonja Valley, Cathedral Peak area, Esterhuysen 12894
(BOL, PRE). 2929 (Underberg): Giant’s Cup Trail, Cobham State Forest
(— CB), Nicholas & V.d. Berg 1323 (PRE); Cobham Forest Reserve,
Underberg District (-CB), Hilliard & Burtt 15935 (BOL); Tributary
of Mkhomazi River, Underberg District (-CB), Hilliard & Burtt 15731
(BOL); Bamboo Mountain, Polela District (— CB), Doidge P.8 (PRE).
3029 (Kokstad): Mt Currie, Kokstad (-AD), McLoughlin 771 (BOL,
PRE).
LESOTHO.— 2828 (Bethlehem): Leribe (-CC), Dieterlen 475, 841
p.p. (K, MPU).
CAPE. — 3227 (Stutterheim): Gxulu Mt, Keiskamma Hoek District
(— CA), Story 3502 (PRE).
FIGURE 4. — Distribution of Ophioglossum convexum J.E. Burrows,
H , in the Transvaal and Zimbabwe; Asplenium sebungweense
J.E. Burrows, #, in Zimbabwe.
5. Marsilea farinosa Launert subsp. arrecta J.E.
Burrows, subsp. nov. Differt a subsp. farinosa pedicellis
longioribus et e basi et in axillis stipitis exorientibus
sporocarpiisque ad angulum c. 180° a pedicello feruntur.
TYPE.— Transvaal, (2328) Baltimore: 40 km S of
Groblersbrug on Potgietersrus road (— AA), Burrows 3597
(BOL, holo.; J, K, PRE, Herb. Burrows, iso.).
Differs from subsp. farinosa (Launert 1968, 1983) in
that the pedicels are longer and arise from both the base
of the stipe and the axils of the stipe, and the sporocarps
are held at ± 180° to the pedicels. (Figure 7B).
BOTSWANA.— 2227 (Palapye): ± 10 km N of Martin’s Drift on the
Palapye Road (— DD), Burrows 3715 (PRE, Herb. Burrows). 2425
(Gaberone): 3 miles north of Gaberone (— DB), Mott 314 (BOL, PRE,
UBLS). Grid ref. unknown: Content Farm, Gaberone District, Kelaole
A13 (PRE).
6. Hymenophyllum splendidum V.d. Bosch in
Nederlandsch Kruidkundig Archief 5: 192 (1863). Type:
Fernando Po [Bioko], Mann s.n. (K, ?holo.; L, iso.!).
170
Bothalia 19,2 (1989)
FIGURE 5. — Ophioglossum convexum, Burrows 3683'. Al, various aspects of plants, x 1,5; A2, sterile lamina, view from above and
cross-section, x 1,5; A3, rhizome, x 1,5. Mohria caffrorum var. ferruginea, Burrows 3677 : Bl, plant, x 0,7; B2, pinna, x 1,5;
B3, rachis, x 3,5.
Bothalia 19,2 (1989)
171
FIGURE 6. — Distribution of Mohria caffrorum var. ferruginea JE. &
S.M. Burrows, #, in southern Africa; Marsilea farinosa subsp.
arrecta J.E. Burrows, ■.
Hymenophyllum ciliatum Swartz var. splendidum (V.d. Bosch) C. Chr.:
368 (1906). Sphaerocionium splendidum (V.d. Bosch) Copeland: 31
(1938).
Hymenophyllum plumieri Hooker & Grev. ; t. 123 (1829).
Icon; Tardieu-Blot: pi. IX (1964).
Whereas material recently collected by the author in
both Zimbabwe and Malawi is certainly H. splendidum,
a collection from Gurue in Mozambique, Torre & Cor-
reia 16899 (COI, EA, LISC, LMU, SRGH) has been
determined as H. hirsutum (L.) Swartz. Examination of
the specimen in SRGH revealed a few stellate hairs on the
lamina as well as the margins and veins, making it likely
that it too, is H. splendidum, although the ranges of the
two species overlap through much of tropical Africa.
MALAWI. — 1535: Mount Mulanje, below Lichenya Hut, 1 820 m.
Burrows 3746 (PRE, Herb. Burrows).
ZIMBABWE. — 1832: Nyanga, eastern slopes of Mt Inyangani,
2 040 m, Burrows 2936 (BOL, SRGH, Herb. Burrows), 3698 (PRE,
Herb. Burrows).
Distribution: Cameroon, Equatorial Guinea, Bioko, Sao
Tome & Principe, Gabon, Zaire, Rwanda, Burundi, Kenya
and Tanzania.
7. Vittaria ensiformis Swartz in Gesellsschaft
naturforschender Freunde zu Berlin, Neue Schriften 2:
134, t. 7, fig. 1 (1799). Type: Mauritius, Sonnerat s.n. (P).
Oetosis ensiformis (Swartz) Greene: 103 (1900).
Vittaria plantaginea Bory: 325 (1804). Type: Mascarene Islands.
Icon: Holttum: fig. 359 (1966).
V ensiformis differs from the superficially similar V.
isoetifolia in having broader fronds that arch out from the
vertical substrate on which it grows, rather than hanging
straight down, as does V isoetifolia. In addition, V.
isoetifolia has very distinctive rhizome scales with
thickened cell walls and margins that bear strong,
thorn-like projections, both of which are absent in V.
ensiformis. It has been recorded growing as a low-level
epiphyte on the boles of old trees in tall, semi-deciduous
Newtonia buchananii—Millettia stuhlmannii forest, near
perennial rivers, at an altitude of ± 360 m.
ZIMBABWE. — 1932: Chimanimani District, Ngorima T.T.L., west
bank of the Haroni R., Grid Ref. 016864, Burrows 2737 (BOL, SRGH,
Herb. Burrows); Chimanimani National Park, Mukurupini R., Burrows
2792 (BOL, Herb. Burrows).
8. Actiniopteris semiflabellata Pichi-Sermolli in
Webbia 17: 24 (1962). Type: Ethiopia, Tertale, Pozzi di
El Banno, Corradi 26 (Herb. Pichi-Sermolli, holo.; FI,
iso.).
leones: Pichi-Sermolli: fig. 4 (1962); Lawalree: pi. 1 (1969).
A plant of this fern was first collected by M. Muller
of the Windhoek Herbarium in the Naukluft Mountains
in 1979 and determined as A. radiata (Swartz) Link. Upon
closer examination, it became apparent that the collection
was A. semiflabellata, based upon the homomorphic
fronds (although the fertile fronds are somewhat larger
than the sterile fronds), the two types of rhizome scales
(one concolorous, the other with a dark central stripe) and,
most characteristically, the dried fronds which are only
slightly inclined to one side, whereas in the other three
African species the fan of the dried frond is bent at 90°
or more from the vertical.
A. semiflabellata has, up to now, only been recorded
as far south as Tanzania, Burundi and Zaire, extending
northwards to north Africa and south Asia. This find in
such an isolated situation, therefore, represents an
interesting and puzzling extension for the species, although
the arid habitat of the Naukluft Mountains is very similar
to that in which it occurs throughout much of its range.
NAMIBIA. — 2416 (Maltahohe): Naukluft, Farm Zais, on south-facing
cliffs on border of farm Blasskranz, 1 400 m, (— AA), Burrows 3737 (J,
PRE, Herb. Burrows); M. Muller 995 (WIND).
Distribution: Tanzania, Burundi, Rwanda, Zaire,
Cameroon, Nigeria, Uganda, Kenya, Sudan, Somalia,
Ethiopia, Egypt, Saudi Arabia, Yemen, Socotra, Nepal,
Reunion, Mauritius and Madagascar.
9. Grammitis rigescens (Bory ex Willd.) J.E.
Burrows, comb. nov.
Polypodium rigescens Bory ex Willd. in Species plantarum 4: 183 (1810).
Ctenopteris rigescens (Bory ex Willd.) J. Sm.: 184 (1875). Xiphopteris
rigescens (Bory ex Willd.) Alston: 26 (1956). Type: Reunion, Bory de
St. Vincent s.n. , in Herb. Willdenow no. 19668 (B, holo.; FI, P, iso.).
Grammitis flabelliformis sensu Morton: 57 (1967). Xiphopteris
flabelliformis sensu Schelpe: 217 (1967).
Pichi-Sermolli (1983) has clearly shown that, in terms
of Art. 8 of the Code of Botanical Nomenclature, Morton’s
application of the name Grammitis flabelliformis (Poir.)
Morton (loc. cit.) is unacceptable, and that the plants from
Reunion belong to Polypodium rigescens, described by
Willdenow in 1810, while Poiret’s Polypodium flabelliforme
applies to the central American taxon.
In accordance with current generic concepts in
Grammitidaceae (Morton 1967; Proctor 1985; Stolze 1981),
the recognition of Xiphopteris at generic level is not
upheld, particularly in view of the poor value of degree
of lamina dissection as a distinguishing criterion.
10. Asplenium buettneri Hieron. ex Brause in
Wissenschaftliche Ergebnisse der Deutschen Zentral-
172
Bothalia 19,2 (1989)
FIGURE 7, Asplenium sebungween.se, Burrows 3026: Al, frond and rhizome, x 0,6; A2, pinnule, x 1. Marsilea farinosa subsp. arrecta,
Burrows 3597: Bl, plant, x 0,6; B2, sporocarp, x 2; B3, B4, points of pedicel attachment, x 2.
Bothalia 19,2 (1989)
173
Afrika-Expedition 1907-1908 2 : 23, t.2, fig. 2 (1910).
Type: Togo, Misahohe, Baumann 42 (B, lecto.; P).
Asplenium parablastophorum Braithewaite: 5 (1972). Type: Zimbabwe,
Chimanimani Mts, Mitchell 391 (BOL, holo.; SRGH, iso.!).
leones: Tardieu-Blot: pi. XXXIII (1964); Jacobsen: fig. 270 (1983).
In the course of examining material of Asplenium from
south and central Africa, it became obvious that A.
parablastophorum Braithwaite from south-eastern
Zimbabwe was identical to A. buettneri Hieron. from
neighbouring Mozambique and tropical Africa, and
accordingly the former is hereby sunk into A. buettneri.
Distribution: Ghana, Togo, Nigeria, Gabon, Cameroon,
Zaire, Tanzania, Zambia, Malawi, Mozambique and
Zimbabwe.
11. Asplenium sebungweense J. E. Burrows, sp. nov.
Rhizoma repens, ± 5 mm diametro, frondibus 4-10 mm
distantibus, interdum aspectu caespitosis. Squamae
rhizomae 2,5—4 mm longae, atrobrunneae lineari-
lanceolatae clathratae subintegrae; apex arista longa. Stipes
laminam aequans vel eo longiore, ad 260 mm longus,
castaneus vel fere ater, squamis atrobrunneis clathratis
lineari-lanceolatis ad 2 mm longis modice obtectus,
glabrescens. Lamina 150—250 x 100—140 mm, ovati-
triangularis, bipinnata vel profunde tripinnatifida, pinnae
basales longissimae. Pinnae ovatae vel triangulares.
Pinnulae obcuneatae vel oblanceolatae, 4-14 mm latae,
pinnulae proximales profunde pinnatifidae; margines
distales profunde irregulariterque serrati et incisi,
atrovirides, pagina supera glabra, pagina infera pallidior
et glabra distali aliquot squamis nigris capillaceis ad 1 mm
longis basim pinnularum proximis et secus rhachides
secondarias; nervatura flabellata, prominens in
superficiebus ambabus frondis. Rachis proximali ater,
distali viridescens, squamis et pilis dispersis nigrescentibus
obtectus. Sori numerosi, lineares, secus venas positas;
indusium lineare, integrum, 3—9 x 0,2 mm.
TYPE. — Zimbabwe, tributary of Busi River, 12 km NE
of Lusulu, Grid Ref. NL. 976095, Craig, Mahlangu & Bur-
rows 8 (PRE, holo.; Sengwa Herb., iso.).
Rhizome creeping, ± 5 mm in diameter, with fronds
spaced 4—10 mm apart, sometimes appearing tufted.
Rhizome scales 2,5-4 mm long, dark brown, linear-
lanceolate, clathrate, subentire, with a long hair-tip. Stipe
up to 260 mm long, as long or longer than the lamina,
castaneous to almost black, lightly set with dark brown,
clathrate, linear-lanceolate scales up to 2 mm long,
glabrescent. Lamina 150—250 x 100-140 mm, ovate-
triangular in outline, 2-pinnate to deeply 3-pinnatifid, with
the basal pair of pinnae longer than those above. Pinnae
ovate to triangular. Pinnules obcuneate to oblanceolate,
4-14 mm broad, becoming deeply pinnatifid proximally,
apical margins deeply and irregularly serrate and incised,
dark green, glabrous above, paler below and glabrous dis-
tally with scattered black, hair-like scales up to 1 mm long
near the pinnule bases and along the secondary rachises,
venation flabellate, prominent on both surfaces. Rachis
black proximally, becoming matt-green distally, set with
scattered, blackish scales and hairs. Sori numerous, linear,
set along the veins; indusium linear, entire, 3-9 x 0,2
mm (Figure 7A).
A. sebungweense is closely allied to A. aethiopicum but
is distinguished from the latter by the ovate-triangular
frond, the relatively longer stipe and the thinner and more
widely creeping rhizome. It occurs at altitudes of between
850 and 1 120 m, in hot and dry, deciduous woodland
where it finds a degree of protection in deep ravines in
the sandstone mantle that covers much of north-western
Zimbabwe (Figure 4). A. aethiopicum is not found in this
area and the two species are not known to overlap. A.
sebungweense is remarkably constant morphologically
compared to the highly variable A. aethiopicum.
ZIMBABWE. — 1828 (Kamativi): Gokwe District, Charama Plateau,
near Sai turn-off (— AB), Burrows 3026 (Herb. Burrows); Sengwa Wildlife
Research Institute, Kove River Gorge (-AA), Burrows 2604 (Herb.
Burrows).
ZAMBIA. — 1324: North West Province, Kabompo Gorge, Leach &
Williamson 13469 (PRE).
12. Asplenium uhligii Hieron. in Botanische
Jahrbiicher 46: 374 (1912). Type: Tanzania, Kilimanjaro,
Uhlig 116 (B, holo.; P, iso.).
Icon: Tardieu-Blot: pi. XXVII, fig. 4 (1964).
A. uhligii is a high altitude fern occurring in deeply
shaded recesses among boulders, growing in mats of
Hymenophyllum tunbridgense (L.) Sm. together with
Grammitis rigescens (Bory ex Willd.) J.E. Burrows.
Although smaller than the typical form from central
Africa, the thin, creeping rhizome and the short, ovate,
shiny brown rhizome scales which lack any central cell
wall thickening, are consistent and distinguish it from the
superficially similar A. aethiopicum (Burm. f.) Becherer
and A. linckii Kuhn.
MALAWI. — 1535: Mount Mulanje, Sapitwa, 2 600 m, Burrows 3758
(Herb. Burrows); Wild 6200 (BOL, SRGH!).
ZIMBABWE. — 1832: Nyanga District, Mount Inyangani, 2 560 m.
Burrows 2086 (Herb. Burrows).
Distribution: Togo, Nigeria, Cameroon, Zaire, Uganda,
Kenya and Tanzania.
13. Thelypteris oppositiformis (C. Chr.) Ching in
Bulletin of the Fan Memorial Institute of Biology. Botany
10: 253 (1941). Type: Madagascar, Perrier 7582 (P, holo.).
Dryopteris oppositiformis C. Chr. in Bonap.: 173 (1925).
Amauropelta oppositiformis (C. Chr.) Holtt.: 135 (1974).
Dryopteris tsaratananensis C. Chr.: 45 (1932). Thelypteris
tsaratananensis (C. Chr.) Ching: 255 (1941). Type: Madagascar, Mt
Tsaratanana, Perrier 16455 (P, holo.).
Thelypteris strigosa sensu Schelpe: 193 (1970).
TRANSVAAL. — 2530 (Lydenburg): Die Berg, between Roossenekal
and Lydenburg (— AA), 2 200 mm. Burrows 3709 (PRE, Herb. Burrows).
Distribution: Zimbabwe, Malawi, Tanzania, Uganda,
Kenya, Ethiopia, Sudan, Cameroon, Nigeria.
ACKNOWLEDGEMENTS
I would like to express my gratitude to the staff of the
following herbaria for permission to examine material and
help in various ways: 8M, BOL, GRA, K, L, P, PRE,
SRGH and WIND.
174
Bothalia 19,2 (1989)
REFERENCES
ALSTON, A.H.G. 1956. New African ferns. Boletim da Sociedade
Broteriana , ser. 2a, 30: 26.
BORY DE SAINT-VINCENT, J.B.G.M. 1804. Voyage dans les quatre
principales lies des mers d'Afrique 2: 325. Paris.
BRAITHWAITE, A.F. 1972. Two new species of Asplenium from
southern Africa. Journal of South African Botany 38: 1—7.
BRAUSE, G. 1910. Pteridophyta. In Wissenschaftliche Ergebnisse der
Deutschen Zentral-Afrika-Expedition 1907—1908, 2: 23. Klinkhardt
& Biermann, Leipzig.
CHEN, Y.-S. & CHIANG, Y.-L. 1972. A study of the root of Ophioglos-
sum Hook. Taiwania 17: 92—110.
CHING, R.C. 1941. New family and combination of ferns. Bulletin of
the Fan Memorial Institute of Biology. Botany 10: 235—256.
CHRISTENSEN, C. 1906. Index filicum: 368. Hafniae.
CHRISTENSEN, C. 1925. Fougeres de Madagascar recoltes de M.H.
Perrier de la Bathie. In Le Prince Bonaparte, Notes Pteridolo-
grques 16: 173. Paris.
CHRISTENSEN, C. 1932. The Pteridophyta of Madagascar. Dansk
Botanisk Arkiv 7: 1—253.
CLAUSEN, R.T. 1938. A monograph of the Ophioglossaceae. Memoirs
of the Torrey Botanical Club 19: 152—153.
COPELAND, E.B. 1938. Genera Hymenophyllacearum. Philippine
Journal of Science 67: 31.
GREENE, E.L. 1900. Necker’s genera of ferns I. Pittonia 4: 103.
HIERONYMUS, G. 1912. Polypodiacearum species novae vel non satis
cognitae africanae. Botanische Jahrbiicher 46: 374-375.
HOLTTUM, R.E. 1966. Ferns of Malaya. Flora of Malaya II.
Government Printing Office, Singapore.
HOLTTUM, R.E. 1974. Thelypteridaceae of Africa and adjacent islands.
Journal of South African Botany 40: 123—168.
HOOKER, W.J. & GREVILLE, R.K' 1829. leones filicum: t. 123.
London.
JACOBSEN, W.B.G. 1983. The ferns and fern allies of southern Africa.
Butterworth, Durban.
KUHN, M. 1868. Filices africanae. Engelmann, Leipzig.
LAUNERT, E. 1968. A monographic survey of the genus Marsilea
Linnaeus. Senckenbergiana Biologie 49: 298—300.
LAUNERT, E. 1983. A revised key to and new records of African species
of the genus Marsilea. Garcia de Orta, Serie de Botanica 6: 127.
LAWALREE, A. 1969. Actiniopteridaceae. In Flore du Congo du Rwanda
et du Burundi: 3, fig. 1. Jardin botanique national de Belgique,
Bruxelles.
MORTON, C.V. 1967. Studies of fern types I. Contributions from the
U.S. National Herbarium 38: 57—59.
PICHI-SERMOLLI, R.E.G. 1962. On the fem genus Actiniopteris Link.
Webbia 17: 1-32.
PICHI-SERMOLLI, R.E.G. 1983. Fragmenta pteridologiae — VIII.
Webbia 37: 118-125.
PROCTOR, G.R. 1985. Ferns of Jamaica. British Museum (Natural
History), London.
SCHELPE, E.A.C.L.E. 1967. New taxa of Pteridophyta from south-east
tropical Africa. Boletim da Sociedade Broteriana ser. 2a, 41: 217.
SCHELPE, E.A.C.L.E. 1970. Pteridophyta. In A.W. Exell & E. Launert,
Flora zambesiaca: 193. Crown Agents, London.
SMITH, J. 1875. Historia filicum: 184. London.
STOLZE, R.G. 1981. Ferns and fem allies of Guatemala, Part n. Fieldiana
6: 242-266.
SWARTZ, O.P. 1799. Vittaria, eine neue Farrenkraut-Gattung.
Gesellschaft naturforschender Freunde zu Berlin, Neue Schriften
2: 134.
TARDIEU-BLOr, M.-L. 1951. Ophioglossacees: 3. In H. Humbert, Flore
de Madagascar et des Comores. Museum National d’Histoire
Naturelle, Paris.
TARDIEU-BLOT, M.-L. 1953. Les Pteridophytes de TAfrique
Intertropicale Francaise 28. In Memoires de l ’Institut Francois
dAfrique Noire. Dakar.
TARDIEU-BLOT, M.-L. 1964. Pteridophytes. In A. Aubreville, Flore
du Cameroun. Museum National d’Histoire Naturelle, Paris.
VAN DEN BOSCH, R. 1863. Hymenophyllaceas novas, exposuit.
Nederlandsch Kruidkundig Archief 5: 192.
WILLDENOW, C.L. 1810. In C. Linnaeus, Species plantarum 4: 183.
Berolini.
Bothalia 19,2: 175-178 (1989)
The genera Polygonum and Bilderdykia (Polygonaceae) in southern
Africa: morphology and taxonomic value of the ocrea and fruit
G. GERMISHUIZEN* P.J. ROBBERTSE** and P.D.F. KOK**
Keywords: Bilderdykia , fruits, key, morphology, ocrea. Polygonum , taxonomy
ABSTRACT
The external morphology of the fruit and the ocrea of 16 taxa of Polygonum and Bilderdykia in southern Africa was studied.
Fruits are either lenticular or trigonous. Six types of fruit surfaces were distinguished. Five types of ocreae were found,
characterized by: a green undulating limb; a silvery hyaline sheath; a brown tubular sheath without terminal hairs; a terminal
fringe of short cilia or setae; and a bristly hairy sheath fringed with long rigid setae.
UITTREKSEL
Die uitwendige morfologie van die vrug en die okrea van 16 taksons van die genusse Polygonum en Bilderdykia in suide-
like Afrika is ondersoek. Vrugte is of lensvormig of driehoekvormig. Vrugoppervlakke word in ses groepe verdeel. Vyf
verskillende okrea-tipes word aangetref, gekenmerk deur: ’n groen golwende aanhangsel; ’n silweragtige hialiene skede;
’n bruin buisvormige skede sonder hare aan die punt; ’n fraiing kort siliums of borselhare; en ’n borselharige skede met
’n fraiing lang, stywe borselhare.
INTRODUCTION
The aim of this study was to examine the morphology
of the ocrea and fruit of 16 of the southern African species
of the genera Polygonum and Bilderdykia, and to determine
their taxonomic importance.
MATERIALS AND METHODS
The herbarium voucher specimens mentioned in the
captions of the figures are all kept at the National
Herbarium (PRE), Botanical Research Institute, Pretoria.
Fruits were coated with gold-palladium and studied and
photographed with the aid of a scanning electron micro-
* Botanical Research Institute, Private Bag X101, Pretoria 0001.
** Department of Botany, University of Pretoria, Pretoria 0002.
MS. received: 1989.02.27.
scope (Hitashi-Akashi Model MSM-4). The negatives are
stored in the Botanical Research Institute, Pretoria.
RESULTS
Organography of the fruits
In Polygonum and Bilderdykia the ovary is superior, ses-
sile, 1-locular, with a solitary basal, sessile or stalked
ovule. The fruit is a nut enclosed by the persistent perianth
(Dyer 1975). During dispersal the seed remains enclosed
in the entire fruit wall (Harder & Firbas J965).
Based on the shape of the fruit, two types are recog-
nized (Table 1): 1, trigonous or 3-angled fruits (Figure LA)
and 2, lenticular fruits (Figure IB). In both types the sur-
faces are either concave (Figure LA) or convex (Figure 1C).
Trigonous fruits and lenticular fruits with convex surfaces
TABLE 1. — Fruit shape, surface type and ocrea type found in species of Polygonum and Bilderdykia
176
Bothalia 19,2 (1989)
HP” - '<*
mg } v* / t* *{**? . y^b-v.; .
\ v ** '&&$-* \& <^vv.
Wmmm^
^SiSlIilil
FIGURE 1. — Electron micrographs of Polygonum spp. A, trigonous fruit,
E plebeium, Germishuizen 1386 , x 36; B, lenticular-concave
fruit, E senegalense subsp. senegalense, Germishuizen U69, x
30; C, lenticular-convex fruit, E nepalense, Scheepers 1130, x
54.
are common, whereas lenticular fruits with concave
surfaces are found only in Polygonum lapathifolium and
P. senegalense (Table 1).
The surface of the fruits varies considerably and six
different types of surfaces can be distinguished (Table 1):
1, with protuberances of varying size (Figure 2A). The
fruits have a dull matted appearance;
2, smooth and shiny (Figure 2B);
3, with criss-cross pattern (Figure 2C);
4, ridged (Figure 2D);
5, areolate (Figure 2E), found only in P. nepalense ;
6, warty (Figure 2F), found only in P. amphibium.
Organography of the ocrea
The ocrea (plural: ocreae; sometimes spelled ochreae),
is a tubular sheath formed when the stipules are united
into a hood, which covers the stem apex at first; later it
is ruptured and remains as a membranous tube surrounding
the stem at the nodes (Harder & Firbas 1965).
Ocreae of the southern African species of the genera
Polygonum and Bilderdykia can be divided into five
different types (Table 1):
1, a tubular membranous sheath ending terminally in
a spreading or recurved, green, leaf-like undulating limb
with a strigose margin (Figure 3A). This type occurs only
in P. limbatum.
2, a silvery hyaline sheath, conspicuously veined,
reddish at the base or red all over, lacerating easily (Figure
3B). This type is found in five species.
3, a brown membranous tubular sheath, conspicuously
veined without or rarely with short terminal hairs (Figure
3C). This type occurs in six species.
4, a tubular membranous sheath fringed with short rigid
cilia or setae (Figure 3D). This type occurs in three
species.
5, a brown tubular sheath, bristly hairy and fringed with
numerous long rigid setae and tearing readily on one side
(Figure 3E). This type is found only in Polygonum
pulchrum. Ocreae can be smooth and hairless (Figure 4 A)
or pubescent with multicellular trichomes (Figure 4B).
DISCUSSION AND CONCLUSION
From Table 1 a key has been compiled using only fruit
shape, fruit surfaces and ocreae found in species of
Polygonum and Bilderdykia.
la Nut trigonous:
2a Ocrea silvery hyaline:
3a Nut smooth, shiny E plebeium
P. kitaibelianum
3b Nut with protuberances E aviculare
3c Nut with criss-cross pattern E maritimum
2b Ocrea not silvery hyaline:
4a Ocrea with short rigid setae E salicifolium
4b Ocrea without terminal hairs:
5a Nut surface with protuberances B. convolvulus
5b Nut surface with criss-cross pattern E meisnerianum
lb Nut lenticular:
6a Nut with concave surface:
7a Nut surface smooth, shiny E lapathifolium
7b Nut surface ridged E senegalense
6b Nut with convex surface:
8a Ocrea with green spreading limb E limbatum
8b Ocrea without spreading limb:
9a Nut surface smooth, shiny:
10a Ocrea silvery hyaline E undulatum
10b Ocrea bristly hairy E pulchrum
9b Nut surface ridged, areolate or warty:
11a Ocrea without terminal setae:
12a Nut surface areolate E nepalense
12b Nut surface warty E amphibium
lib Ocrea with short rigid terminal setae; nut surface
ridged E hystriculum
P. hydropiper
Bothalia 19,2 (1989)
177
FIGURE 2. — Electron micrographs of fruit surfaces of Polygonum and Bilderdykia species. A, surface with protuberances, B. convolvulus,
Jacobsz 196, x 860; B, surface smooth, P. salicifolium, Germishuizen 1291, x 4000; C, surface with criss-cross pattern, P. maritimum,
Taylor 4897, x 850; D, surface ridged, P. hystriculum, Germishuizen 1357, x 520; E, surface areolate, P nepalense, Scheepers 1130,
x 650; F, surface warty, P amphibium, Nelson 226, x 220.
178
Bothalia 19,2 (1989)
FIGURE 3. — Schematic representation of ocrea types found in some
taxa of the genus Polygonum. A, P. limbatum , x 4; B, P.
kitaibelianum, x 2; C, P meisnerianum, x 2; D, P. hydropiper ,
x 4; E, P. pulchrum , x 4.
In their treatments of the family Polygonaceae, C.H.
Wright in Flora capensis (1912) and R. A. Graham in Flora
of tropical east Africa (1958) make some use of the fruit
shape and ocrea type in their species keys, and they make
reference to them in their descriptions.
In the Flora of southern Africa (Vol. 9,1 in prep.) fruit
shape, fruit surface and ocrea type play a major role in
the keys. In this paper it is the first time that these features
have been illustrated. In the genera Polygonum and
Bilderdykia the fruit and the ocrea are the two most
important organs for distinguishing between the taxa.
FIGURE 4. — Electron micrographs of a portion of the ocrea surface
of certain members of the genus Polygonum. A, P. aviculare,
Germishuizen 1265, x 360; B, P. pulchrum, Germishuizen 1240,
x 102; C, P. limbatum, Germishuizen 1361, x 204.
GRAHAM, R.A. 1958. Polygonaceae. In W.B. Turrill & E. Milne-
Redhead, Flora of tropical east Africa: 1—40. Crown Agents for
Overseas Government and Administrations, London.
REFERENCES
DYER, R.A. 1975. Polygonaceae. The genera of southern African
flowering plants 1: 59. Department of Agricultural Technical
Services, Pretoria.
HARDER, R. & FIRBAS, F. 1965. Classification of the plant kingdom.
Strasburger’s textbook of botany. Longmans, London.
WRIGHT, C.H. 1912. Polygonaceae. In W.T. Thiselton-Dyer, Flora
capensis 5 : 463 —472. Reeve, London.
Bothalia 19,2: 179-182 (1989)
A re-examination of the genus Amphibolia (Mesembryanthemaceae)
H.E.K. HARTMANN* and M. DEHN*
Keywords: Amphibolia, lectotypification, Mesembryanthemaceae
ABSTRACT
The holotypes of all five species placed in the genus Amphibolia L. Bol. ex Toelken & Jessop have been examined. Only
two of them agree with the generic diagnosis, but they are not congeneric. The type species chosen by Herre is shown to
be in serious conflict with the protologue and a new lectotype (A. maritima L. Bol. ex Toelken & Jessop) is therefore designated.
The other four species are excluded from Amphibolia.
UITTREKSEL
Die holotipes van al vyf spesies wat in die genus Amphibolia L. Bol. ex Toelken & Jessop geplaas word, is ondersoek.
Slegs twee van hulle stem ooreen met die diagnose van die genus, maar hulle is nie kongeneries nie. Daar word aangetoon
dat die tipespesie wat deur Herre gekies is, emstig met die protoloog in stryd is, en daarom word ’n nuwe lektotipe (A.
maritima L. Bol. ex Toelken & Jessop) aangewys. Die ander vier spesies word uit Amphibolia uitgesluit.
INTRODUCTION
In the course of a comprehensive study of the subtribe
Ruschiinae (M. Dehn) and the preparation of a survey of
the genera of the Aizoaceae (H.E.K. Hartmann), the genus
Amphibolia L. Bol. ex Herre (Toelken & Jessop 1976) has
been re-examined. The results and taxonomic conse-
quences are presented.
Amphibolia was first described by Bolus (1965). It was
lectotypified and thus validated by Herre (1971). Toelken
& Jessop (1976) reviewed the nomenclature and supplied
correct names for all five species included in the genus
at that time.
The history of the genus started earlier, though, when
Bolus (1962) noticed that the two species she had described
in the genus Stoeberia Schwant. (S. hallii, S. littlewoodii)
did not conform with the generic characters of that genus,
‘a new genus is therefore required for them . . .’ (l.c. p. 14).
The diagnosis of that new genus, namely Amphibolia
(Bolus 1965), gives three characters: perennial, fruits with
winged valves (like in Lampranthus ) and closing bodies
(like in Ruschia and Stoeberia). In addition Bolus stated
that the capsules differ from those of Stoeberia insofar as
they open spontaneously and do not close again. No further
features were mentioned, but Herre (1971) provided a short
description.
RESULTS
Since the diagnosis of Amphibolia stresses fruit
characters, the present re-examination also concentrates
on capsules. The holotypes of all five species placed in
the genus were examined. Three different combinations
of the relevant character expressions could be distinguished
(Table 1). A. maritima and A. littlewoodii possess valve
wings and closing bodies (CB), as well as closing rodlets
* Institut fur Allgemeine Botanik und Botanischer Garten, Universitat
Hamburg, Ohnhorststrasse 18, D-2000 Hamburg 52.
MS. received: 1989.02.10.
(CR) at the distal end of the covering membranes (Figure
IB). The remaining three species lack closing bodies, but
two of them [A. hallii and A. gydouwensis ) possess closing
rodlets (CR in Figure ID). A. stayneri shows no closing
device at all.
It seems that in the last three species mentioned the
prominent free end of the placenta (PL in Figure ID) has
been understood to represent a closing body (compare data
from descriptions in Table 1). This interpretation, however,
has to be rejected. The relevant placental structure is not
in direct contact with the tangential endocarp, as would
be necessary for an endocarpal closing body (Hartmann
& Liede 1986). Furthermore, funicles occur up to the tip
of the placenta (F in Figure 2), precluding, for lack of
space, the formation of a placental tubercle (as described
for Pleiospilos subgenus Pleiospilos by Hartmann & Liede
1986).
Remarkable, even though not mentioned in all
descriptions, is the existence of closing rodlets in four of
the five species (Table 1, CR in Figure IB, D).
Examination of additional characters shows distinct
differences between the two species with closing bodies.
Capsules of A. maritima break off easily and regularly
after ripening, leaving the persistent peduncle on the plant,
thus forming a spine; capsules and peduncles are light-
coloured, almost white, like all stems and branches of the
plant; the margins of the valves are only moderately raised
and, as a consequence, the valves open so completely that
the tips touch the base of the fruit. Other distinguishing
features of A. maritima are short, inflated, roundish leaves,
small flowers and petals barely reaching the tips of the
calyx lobes. In contrast, capsules of A. littlewoodii remain
on the peduncles and they are dark reddish brown from
numerous tanniniferous idioblasts, like all stems and
branches. The margins of the valves are raised to high rims
preventing the valves from opening further than 180°.
Other characteristic features of A. littlewoodii are long,
slender leaves and larger flowers with the petals exceeding
the tips of the calyx lobes.
180
Bothalia 19,2 (1989)
FIGURE 1. — Longitudinal sections of capsules. A, B, Amphibolia maritima (Hall 2885 , BOL): locule (LO) blocked apically by closing body
(CB) protruding from inner wall (W) of capsule; placenta (PL) torn to its base; in dry state (seen here) closing body touches expanding
keel (EK) but not covering membrane (CM); closing rodlet (CR) and adjacent bend of outer part of covering membrane (OCM) pulled
out from between valve proper and expanding keel at first opening of capsule. C, D, Amphibolia hallii ( Hall 1741, BOL): placenta
(PL) separates apically from endocarp and bears funicles (F) to its very tip; thickening of outer fruit wall (TW) above placenta (similar
to condition in Lampranthus- type fruit; Hartmann 1988), distal end of covering membrane (CM) bent against expanding keel (EK),
resembling position in unopened fruit. RCM, reflexed part of covering membrane; S, seed. Scale in mm.
Bothalia 19,2 (1989)
181
TABLE 1. Comparison of fruit characters of Amphibolia, Ruschia and Lampranthus as observed in the holotypes and as given in original description
( ) = Rare expression of characters.
CONCLUSIONS
In the light of recent findings in fruit typology
(Hartmann 1988) and in generic delimitations in the wider
relationship of the Ruschiinae (Dehn 1989, and in prep.),
the groups described above can be allied to different fruit
types and, consequently, to different genus groups.
Dehn (1989, and in prep.) has been able to show that
valve wings may occur within the Ruschia- type of fruit.
The possession of these structures can therefore not be
used to exclude taxa from that group. Essential character
expressions are the possession of small endocarpal closing
bodies and of closing rodlets at the covering membranes.
A. maritima and A. littlewoodii agree with these require-
ments, but they differ in other characters not considered
by Bolus (1965) in the delimitation of Amphibolia. Seeing
that fruit characters are presently considered to be of great
taxonomic importance in Mesembryanthema (e.g.
Hartmann 1983; Dehn 1989; Ihlenfeldt & Bittrich 1985),
the two species have to be understood as belonging to two
different genera within the group characterized by the
Ruschia- type of fruit. Whereas A. maritima shows several
similarities to the genus Eberlanzia (valve wings, small
closing body, widely opening valves, inflated short leaves,
white stems, pedicels becoming spines; all data after
Stiiber unpublished), A. littlewoodii can be placed
within the close relationship of the genus Ruschia (small
closing body, narrow valve wings opening into an upright
position, long slender leaves, dark reddish stems; Dehn
in prep.).
Of the remaining three species, A. hallii and A.
gydouwensis lack closing bodies but possess valve wings
and closing rodlets (Table 1). This combination places
them closest to the Lampranthus- type fruit (Hartmann
1988) which is characterized by valve wings and closing
ledges in the absence of closing bodies; characteristic also
is a thickened tissue within the outer wall above the zone
where the placenta ends (similar to TW in Figure ID).
Preliminary investigations in the genus Lampranthus
indicate that the taxon is probably heterogeneous and will
have to be divided up, A. hallii and A. gydouwensis
agreeing with a certain part of the material.
Superficially, A. stayneri may be placed in the vicinity
of the Lampranthus- type fruit as well, but the complete
absence of any closing device and the formation of rather
fragile capsules indicates a closer relationship to the
Drosanthemum- type of fruit (Hartmann 1988). Within this
group, the nearest relative seems to be the genus
Mestoklema (this hypothesis is supported by the formation
of a remarkable storage root in A. stayneri, a feature com-
mon in Mestoklema, but further characters will have to
be examined).
FIGURE 2. — Amphibolia hallii :
distal opening of locule of
capsule shown in Figure 1C,D
seen from above after removal
of valve, showing end of
placenta (PL) which can be
mistaken for a closing body.
CM, covering membrane; EK,
expanding keel; F, funicle; PL,
placenta; RCM, reflexed part
of covering membrane; W,
wall. Scale in mm.
182
Bothalia 19,2 (1989)
The results show that only two species, A. maritima and
A. littlewoodii , conform with the generic diagnosis. This
implies that the lectotype species (A. halin') selected by
Herre (1971) is in serious conflict with the protologue
(Greuter et al. 1988, art. 8.2, b) and has to be replaced
by one of the two species named above. A. maritima is
selected as the new lectotype species because its valve
wings are particularly broad, thus conforming best with
the genus Stoeberia considered to be closest to Amphibolia
by Bolus (1965) when she established the latter genus.
The taxonomy of the genus Amphibolia, as given below,
reflects the present state of knowledge.
TAXONOMY
Amphibolia L. Bol. ex Herre, The genera of Mesem-
bryanthemaceae: 70 (1971); L. Bol.: 169 (1965); Toelken,
H. R. & Jessop, J.P. : 64 (1976). Lectotype species (here
designated): A. maritima L. Bol. ex Toelken & Jessop.
This lectotypification supercedes the lectotypification of
Herre (1971), A. hallii, because the character expressions
of that species are in serious conflict with the protologue.
A. maritima L. Bol. ex Toelken & Jessop in Bothalia
12: 64 (1976); L. Bol.: 169 (1965). Lectotype species.
Holotype: Hall 2885 (BOL!). Nomenclature! synonym:
Ruschia maritima Rowley: 62 (1978b).
Species excluded
I. Species close to the genus Ruschia Schwant.:
A. littlewoodii (L. Bol.) L. Bol. ex Toelken & Jessop
in Bothalia 12: 64 (1976); L. Bol.: 170 (1965). Basionym:
Stoeberia littlewoodii L. Bol.: 162 (1960). Holotype:
Littlewood KG 522/59 (BOL!). Nomenclatural synonym:
Ruschia mutata Rowley: 7 (1978a) non Ruschia littlewoodii
L. Bol.
2. Species close to the genus Lampranthus N.E. Br. :
A. hallii (L. Bol.) L. Bol. ex Toelken & Jessop in
Bothalia 12: 64 (1976); L. Bol.: 161 (1960). Basionym:
Stoeberia hallii L. Bol.: 161 (1960). Holotype: Hall 1741
(BOL!). Nomenclatural synonym: Ruschia amphibolia
Rowley: 7 (1978a) non Ruschia hallii L. Bol.
A. gydouwensis (L. Bol.) L. Bol. ex Toelken &
Jessop: 64 (1976), L. Bol.: 306 (1967). Basionym:
Lampranthus gydouwensis L. Bol.: 13 (1963). Holotype:
Leipoldt 4801 (BOL!). Nomenclatural synonym: Ruschia
gydouwensis Rowley: 7 (1978a).
3. Species perhaps close to Mestoklema N.E. Br. :
A. stayneri L. Bol. ex Toelken & Jessop in Bothalia
12: 64 (1976); L. Bol.: 126 (1966). Holotype: Stayner KG
258/65 (BOL!). Nomenclatural synonym: Ruschia
dissimilis Rowley: 62 (1978b) non Ruschia stayneri L. Bol.
ACKNOWLEDGEMENTS
We are indebted to the Director of the Bolus Herbarium
for the loan of all type sheets of all species of Amphibolia
and for permission to examine the material.
Our work on Mesembryanthemaceae has been made
possible through grants from the Deutsche Forschungs-
gemeinschaft and financial support from the National
Botanic Gardens. For both we are most grateful. Work on
fruits has been funded also by the Cactus and Succulent
Society of America through a Research Grant. We should
like to thank all donors.
We are also thankful to Horst Golling for help with
SEM-graphs of fruits.
REFERENCES
BOLUS, H.M.L. 1960. Notes on Mesembryanthemum and allied genera.
Journal of South African Botany 26: 155—164.
BOLUS, H.M.L. 1962. Corrections and additional notes (notes on
Stoeberia hallii and 5. littlewoodii). Journal of South African Bota-
ny 28: 14.
BOLUS, H.M.L. 1963. Notes on Mesembryanthemum and allied genera.
Journal of South African Botany 29: 11-19.
BOLUS, H.M.L. 1965. Notes on Mesembryanthemum and allied genera.
Journal of South African Botany 31: 169-174.
BOLUS, H.M.L. 1966. Notes on Mesembryanthemum and allied genera.
Journal of South African Botany 32: 125-128.
BOLUS, H.M.L. 1967. Notes on Mesembryanthemum and allied genera.
Journal of South African Botany 33: 305—309.
DEHN, M. 1989. Untersuchungen zum Merkmalsbestand und zur
Stellung der Gattung Antimima N.E. Br. (Mesembryanthema-
ceae). Mitteilungen aus dem Institut fur Allgemeine Botanik
Hamburg 22: 195-232.
DEHN, M. in prep. Untersuchungen zum Verwandtschaftskreis der
Subtribus Ruschiinae (Mesembryanthemaceae).
GREUTER, W. et al. 1988. International Code of Botanical
Nomenclature. Regnum Vegetabile 118.
HARTMANN, H.E.K. 1983. Untersuchungen zum Merkmalsbestand
und zur Taxonomie der Subtribus Leipoldtiinae (Mesembryan-
themaceae). Bibliotheca Botanica 136: 1—67.
HARTMANN, H.E.K. 1988. Fruit types in Mesembryanthema. Beitrage
zur Biologie der Pflanzen 63: 313—349.
HARTMANN, H.E.K. & LIEDE, S. 1986. Die Gattung Pleiospilos s.
lat. (Mesembryanthemaceae). Botanische Jahrbiicher 106:
433-485.
HERRE, H. 1971. The genera of Mesembryanthemaceae. Tafelberg, Cape
Town.
IHLENFELDT, H.-D. & BITTRICH, V. 1985. Morphologie, Gliederung
und Abgrenzung der Gattung Psilocaulon N.E. Br. s.l. (Mesem-
bryanthemaceae). Botanische Jahrbiicher 105: 289-322.
ROWLEY, G.D. 1978a. Reunion of some genera of Mesembryan-
themaceae. National Cactus and Succulent Journal 33: 6-9.
ROWLEY, G.D. 1978b. Reunion of some genera of Mesembryan-
themaceae — a postscript. National Cactus and Succulent
Journal 33: 62.
TOELKEN, H R. & JESSOP, J.P. 1976. Mesembryanthemaceae.
Nomenclature of the genus Amphibolia. Bothalia 12: 64.
Bothalia 19,2: 183-188 (1989)
A contribution to the pteridophyte flora of Transkei
C.T. JOHNSON* and A. HUTCHINGS**
Keywords: analysis, checklist, ecology, Pteridophyta, Transkei
ABSTRACT
An analysis of the data is presented in three tables and a comparison is made with that of adjacent areas. Lygodium kerstenii***
and Dryopteris dracomontana are recorded from Transkei for the first time. The 129 species, with their subspecies and
varieties (137 taxa in all), in the 51 genera of Transkei are listed. Coded distribution ranges are given for each taxon.
UITTREKSEL
’n Analiese van die data word in drie tabelle gegee en ’n vergelyking word getref met naasliggende areas. Lygodium kerstenii***
en Dryopteris dracomontana word vir die eerste keer vir Transkei aangeteken. Die 129 spesies, met hulle subspesies en
varieteite (137 taksons altesaam), in die 51 genusse van Transkei word gelys. Vir elke takson word ’n gekodeerde verspreid-
ing gegee.
INTRODUCTION
Prior to this publication, information on the pterido-
phytes of Transkei was limited (Sim 1915; Roux 1982;
Jacobsen 1983; Anthony & Schelpe 1985; Schelpe &
Anthony 1986). The aim of this research project was to
establish a herbarium of the pteridophyte flora of Transkei,
to determine the distribution of the taxa within Transkei,
to compile a floristic analysis of the pteridophytes and to
prepare a checklist of them.
METHODS
The data for this paper was collected over a period of
six years (1981-1986) and presents a summary of an
unpublished report by Johnson & Hutchings (1986).
During the course of this survey extensive collections were
undertaken and the herbarium specimens are housed at
KEI and BOL. The following herbaria were visited in
order to record additional collections: BOL, J, NBG, NU,
SAM, GRA, PRE, UPR.
Transkei comprises a number of quarter degree grids.
For each taxon the grid references are given in which it
was collected in Transkei (Figure 1). The arrangement and
spelling of the families and genera are according to Schelpe
& Anthony (1986). Southern Africa is delimited as in the
Flora of southern Africa.
RESULTS AND DISCUSSION
From the floristic analysis (Tables 1 & 2) it is apparent
that Transkei is relatively rich in pteridophytes. Twenty
three pteridophyte families are present with 51 genera and
129 species. Only five of the 28 families listed for southern
Africa (Anthony & Schelpe 1985) are not found in
Transkei. A comparison of the 23 families (Table 1)
* University of the Western Cape, Private Bag X17, Bellville 7530, South
Africa.
** University of Transkei. Present address: University of Zululand, Private
Bag X1001, Kwadlangezwa 3886, South Africa.
*** The identity of the material is still being investigated. /Die identiteit
van die materiaal word nog ondersoek.
MS. received: 1988.10.12.
common to Transkei and southern Africa reveals a
difference of 16 genera and 114 species.
The four largest pteridophyte families in Transkei are
Dryopteridaceae with seven genera, Adiantaceae and Poly-
podiaceae with six genera and Schizaeaceae with four
genera. In addition Dennstaedtiaceae and Davalliaceae
have three genera, Aspleniaceae, Athyriaceae, Blechna-
ceae, Thelypteridaceae and Osmundaceae are represented
by two genera each, while the remaining 12 families have
one genus each.
The two largest genera are Asplenium and Cheilanthes
with 22 and 10 species respectively. Lycopodium has seven
species followed by Thelypteris with six species. Blech-
TABLE 1. — Comparison of number of taxa per family for Transkei and
southern Africa. Comparative data for southern Africa from
Schelpe & Anthony 1986
Transkei Southern Africa
Genera Spp. Genera Spp. & va s
184
Bothalia 19,2 (1989)
FIGURE 1. — Map showing geo-
graphical location and grid
references of the study area.
num, Polystichum, Pteris and Elaphoglossum each have
five species. Selaginella and Dryopteris have four species
each; Ophioglossum, Trichomanes, Adiantum and
Microsorium have three species each; Gleichenia,
Schizaea, Marsilea, Cyathea, Blotiella, Polypodium and
Pleopeltis have two species each and there are 30 genera
with one species each.
During this survey Lygodium kerstenii* and Dryopteris
dracomontana , previously not recorded for Transkei, were
collected. The gap in the distribution range of Dryopteris
dracomontana, from Natal to the Cape Province (Schelpe
& Anthony 1986), was closed with the discovery of this
species. The distribution range of Elaphoglossum con-
forme, which was previously assumed to be confined to
the southern and south-western Cape, has now been
extended to the Transkei. Mohria cajfrorum and Equisetum
ramosissimum are species with wide distribution and were
recorded in 60% of the degree grids.
It must be emphasized that Adiantum raddianum,
Nephrolepis exaltata and Macrothelypteris torresiana are
introduced species (Jacobsen 1983) which are widely
cultivated and have now become established in nature.
Comparing the statistical data (Table 2), one finds that
the Transkei flora comprises 68,9% (51) and 51,6% (129)
of the southern African genera or species respectively. The
decrease in the number of taxa from Natal to the southern
Cape must be attributed to the differences in the climatic
conditions along this gradient.
Information about density and species distribution can
be drawn from the distributional analysis (Table 3) and
* The identity of the material is still being investigated. /Die identiteit
van die materiaal word nog ondersoek.
TABLE 2. — Comparison of the pteridophyte floras of various regions.
Comparative data from Schelpe & Anthony 1986
* Quarter degree not in study area.
the floristic analysis (Figure 2). The first impression is
that species density is at its highest in grid 31°S 29°E (95
taxa: 69,3% of total number in Transkei) and 31°S 28°E
(65 taxa: 47,4%). Within these degree squares, the most
concentrated collecting occurred around the towns of Port
St Johns and Umtata. Closer examination shows that no
record exists for grid 32°S 27°E. Since the collecting
Bothaiia 19,2 (1989)
185
NUMBER OF GR®S
l\\J Quarter Degrees V//A Degrees
FIGURE 2. — Species density in the study area.
intensity for the entire area is very low (Gibbs Russell et
al. 1984), it is impossible to state whether Pteridophyta
are completely absent from these areas.
ECOLOGY
Pteridophytes are distributed across a wide range of
habitats in Transkei, occurring from the coastal regions
to the highest elevations of over 2 700 m. The greatest
variety of species, habit and development is found in the
forest, where they grow along streams, on the forest floors,
on boulders, or on the stems of trees. Roux (1982) divided
the pteridophyte flora of Transkei into forest and non-forest
types.
High-level epiphytes are found growing high up on the
stems and even branches of trees in humid forest. These
include Pyrrosia africana, Microsorium punctatum,
Pleopeltis macrocatpa and Polypodium polypodioides
subsp. ecklonii. Asplenium sandersonii, A. prionitis and
Microgramma lycopodioides are low-level epiphytes that
are found growing low down on the stems of trees.
Lithophytes that occur in wet coastal forest with high
humidity and deep shade are Asplenium rutifolium and
Microsorium punctatum. True lithophytes are Lycopodium
gnidioides, L. verticillatum, Selaginella mittenii, Tricho-
manes pyxidiferum var. melanotrichum, Microgramma
lycopodioides, Oleandra distenta, Asplenium gemmiferum,
Rumohra adiantiformis and Vittaria isoetifolia.
Occurring as ground cover in forests are Selaginella
kraussiana, Marattia fraxinea var. salicifolia, Pteris
buchananii, P. catoptera var. horridula, Cheilanthes
bergiana and Blechnum punctulatum var. krebsii. Micro-
sorium scolopendrium occurs on the landward side of dune
forests.
Few ferns are found in open grassland and when they
do it is always in association with rocky outcrops, stream
banks, ditches and along forest margins. Selaginella
dregei, Pityrogramma calomelanos var. aureoflava, and
Cyrtomium caryotideum var. micropterum were collected
from exposed rocky outcrops. In a similar habitat but
slightly protected, Selaginella cajfrorum, Cheilanthes
inaequalis var. buchananii, Elaphoglossum macropodium,
* The identity of the material is still being investigated. /Die identiteit
van die materiaal word nog ondersoek.
Blechnum punctulatum var. punctulatum and Lycopodium
gnidioides occur.
Lycopodium clavatum, L. cemuum, L. carolinianum var.
carolinianum,Equisetum ramosissimum, Osmunda regalis,
Todea barbara, Mohria cajfrorum, Gleichenia poly-
podioides, Cyathea dregei, Hypolepis sparsisora, Pteridium
aquilinum, Adiantum capillus-veneris, A. raddianum,
Pteris vittata, Nephrolepis exaltata, Thelypteris confluens,
T. totta, Blechnum capense and B. tabulare are found
growing on wet banks and forest margins. The only aquatic
species are Marsilea macrocarpa and M. capensis.
Selaginella dregei, Schizaea pectinata, Cheilanthes
involuta var. involuta and Pellaea calomelanos are found
in fully exposed dry habitats.
CONSERVATION
Although the primary concern of this manuscript does
not relate to the problems of conservation, it is necessary
to sound a warning on the threat that exists to this element
of Transkei’s flora. Of the 60 species that are regarded
as rare for southern Africa (Jacobsen 1983), Asplenium
simii, Elaphoglossum angustatum, E. hybridum, Lycopo-
dium saururus, Microsorium ensiforme, Polystichum
luctuosum, Psilotum nudum, Trichomanes reptans and
Woods ia montevidensis var. burgessiana were recorded in
Transkei. With the discovery of Lygodium kerstenii* along
the south-facing slope of Mt Sullivan in Port St Johns, the
rare species count was increased to 10. This figure is prob-
ably still too low as Transkei is undercollected.
ACKNOWLEDGEMENTS
The authors wish to express their gratitude to the
Council of the University of Transkei for their financial
assistance. The success of this research project was high-
ly dependant on the co-operation received from the
curators of BOL, J, NBG, NU, SAM, GRA, PRE, and
UPR for allowing us access to their material. The authors
are most grateful for the help we received from Mrs Nicola
Anthony and Mr J.P. Roux with the determination of
species. We acknowledge with gratitude the work of earlier
collectors, without whom we could not have achieved our
goal.
REFERENCES
ANTHONY, N.C. & SCHELPE, E.A.C.L.E. 1985. A checklist of the
Pteridophytes of the Flora of southern Africa region. Bothaiia
15: 541-544.
GIBBS RUSSELL, G.E.. RETIEF, E. & SMOOK, L. 1984. Intensity
of plant collecting in southern Africa. Bothaiia 15: 131—138.
JACOBSEN, W.B.G. 1983. The fern and fern allies of southern Africa.
Butterworth, Durban and Pretoria.
JOHNSON, C.T. & HUTCHINGS, A. 1986. Survey of the wild flowers
of Transkei. Phase 2. Pteridophytes of Transkei. Department of
Botany, University of Transkei. Unpublished.
ROUX, J.P. 1982. A preliminary survey of the Pteridophyta of Transkei,
southern Africa. Fern Gazette 12,4: 191—196.
SCHELPE, E.A.C.L.E. & ANTHONY, N.C. 1986. Pteridophyta. InO.A.
Leistner, Flora of southern Africa. Government Printer, Pretoria.
SIM, T.R. 1915. The ferns of South Africa, 2nd edn. Cambridge University
Press.
186
CHECKLIST OF THE PTERIDOPHYTES OF TRANSKEI
(♦information from Schelpe & Anthony 1986)
Bothalia 19,2 (1989)
PSILOTALES
PSILOTACEAE
Psilotum Swam
nudum (L. ) Beauv., 3129 BD, Johnson 896 (KEI); 3129 DA, Schelpe
350 (NU); 3228 CB, Pocock 30134 (GRA)
LYCOPODIALES
LYCOPODIACEAE
Lycopodium L.
saururus Lam., 3030 CC, Johnson 2116 (KEI)
verticillatum L. f. , 3029 DA, Getliffe 105 (NU); 3128 BC, Hutchings
& Johnson M2 (KEI); 3129 DA, Johnson 138 (KEI); 3228 AD,
Pegler 821 (BOL)
dacrydioides Bak. , 3128 CA, Cawe 709 (KEI); 3129 DA, Mcloughlin
s.n. (BOL)
gnidioides L. f , 3029 DA, Roux 630 (NBG); 3030 CC, Roux 651 (NBG);
3030 AA, Strey 8646 (BOL); 3128 BC, Hutchings & Johnson
M3 (KEI); 3128 CA, Cawe 709 (BOL); 3129 DA, Mcloughlin
s.n., 809 (BOL)
cernuum L. , 3029 DD, Johnson 168 (KEI); 3030 CC, Hilliard 1122
(NU); 3129 BC, Ward 5794 (NU); 3129 BD, Cawe s.n. (KEI);
3129 DA, Hutchings & Plumstead 2058 (KEI); 3130 AA, Ward
205 (NU); 3129 BD, Johnson s.n. (KEI); 3129 DA, Johnson
88, 168 (KEI); 3229 AA, Johnson 81, 758 (KEI)
clavatum L. , 3128 BA, Bolus 10374 (BOL); 3129 AB, Roux 618 (NBG)
carolinianum L. var. carolinianum, 3030 CC, Roux 662 (NBG); 3128
BC, Hutchings & Plumstead 1627 (KEI); 3129 BD, Cawe 110
(KEI); 3129 DA, Schelpe 259 (NU); 3130 AA, Ward 206 (NU)
SELAGINELLALES
SELAGINELLACEAE
Selaginella Beauv.
dregei (Presl) Hieron., 3128 DB, Hutchings 2284 (KEI); 3129 BC,
Leighton s.n. (BOL); 3129 AB, *Bachman 9 (B); 3129 BD, Cawe
342 (KEI)
caffrorum (Milde) Hieron., 3028 AA, Roux 1343 (NBG); 3127 AA,
Britten s.n. (GRA); 3129 DA, Hardcastle 289 (NBG)
kraussiana (Kunze) A. Br. , 3127 AA, Royffe 140 (GRA); 3128 BC, Cawe
479 (KEI, BOL); 3128 CB, Baur 13 (SAM); 3129 DA, Hutch-
ings & Johnson 2239 (KEI); 3228 AD, Mcloughlin 728 (BOL);
3228 BD, Johnson students s.n. (KEI)
mittenii Bak. , 3030 CC, Roux 670 (NBG); 3228 AD, Pegler 893 (GRA)
EQUISETALES
EQUISETACEAE
Equisetum L.
ramosissimum Desf, 3028 BD, Tyson 1622 (BOL, SAM); 3030
CC, Johnson 2050 (KEI); 3127 AA, Pegler & Kolbe s.n. (BOL);
3127 CA, Hutchings & Johnson 1670 (BOL, KEI); 3128 DB,
Schonland 3813 (GRA); 3129 DA, Hardcastle 290 (NBG); 3228
AD, Pegler 553 (BOL)
OPHIOGLOSSALES
OPHIOGLOSSACEAE
Ophioglossum L.
polyphyllum A. Br. , 3128 AA, Roux 1194 (NBG)
reticulatum L. , 3129 AC, Hutchings 957(KEI); 3129 DA, Schelpe 355
(NU)
vulgatum L., 3029 DA, Hilliard & Burtt 7706 (NU); 3128 DB,
Hutchings 2281 (KEI)
MARATTIALES
MARATTIACEAE
Marattia Swam
fraxinea J.E. Sm. exJ.F. Gmel. var. salicifolia (Schrad.) C. Chr. , 3128
BC, Cawe 736 (BOL); 3128 CB, Baur s.n. (GRA); 3128 DB,
Mcloughlin s.n. (BOL); 3129 BC, Meaker s.n. (NU); 3129 BC,
Sims 2451 (GRA); 3129 BD, Roux 614 (NBG); 3129 DA, Bolus
10372 (BOL)
FIL1CALES
OSMUNDACEAE
Osmundia L.
regal is L. , 3029 DD, Schelpe 5070a (BOL); 3030 CC, Abbott 2661
(KEI); 3128 AD, Hutchings 1264 (BOL, KEI); 3128 CB, Baur
s.n. (GRA, SAM)
Todea Willd.
barbara (L.) T. Moore , 3030 CC, Roux 656 (NBG); 3129 BC, Mcloughlin
722 (BOL); 3129 BC, Cawe 30 (KEI); 3129 DA, Schelpe 361
(NU)
GLEICHENIACEAE
Gleichenia J.E. Sm.
polypodioides (L.) J.E. Sm., 3029 BB, Sr. Wheeler s.n. (NU); 3127
DB, Mcloughlin 121 (BOL); 3128 CB, Hutchings 2205 (KEI,
NH); 3129 BD, Roux 611 (NBG); 3129 DA, Jacot Guillarmod
s.n. (GRA)
umbraculifera (Kuntze) T. Moore, 3128 BC, Hutchings 2036 (BOL, KEI);
3128 CB, Baur 10 (SAM), 178 (GRA); 3129 BC, Cawe 79 (KEI);
3130 AA, Taylor 2618 (NBG)
SCHIZAEACEAE
Schizaea J. E. Sm.
pectinata (L.) Swam, 3129 BD, Johnson 176 (KEI)
tenella Kaulf, 3127 DB, Flanagan 2755 (PRE)
Anemia Swam
dregeana Kunze, 3030 CC, Abbott 2457 (KEI); 3128 CA, Hutchings
& Plumstead 2076 (BOL, KEI); 3129 CC, Wells 3540 (GRA);
3129 DA, Hutchings & Johnson 2247 (KEI); 3228 BD, Roux
551 (NBG); 3228 CB, Flanagan 285 (GRA)
Mohria Swam
caffrorum (L. ) Desv. , 3027 DB, Roux 1207 (NBG); 3028 AD, Johnson
713 (KEI); 3029 AD, Taylor 5487 (NBG); 3028 DD, Barker
6158 (NBG); 3029 AB, Schelpe 5062 (BOL); 3127 CB, Schelpe
5831 (BOL); 3128 BC, Hutchings 125 (KEI); 3128 CA, Cawe
839 (KEI); 3129 AC, Hutchings 1010 (BOL. KEI); 3129 DA,
Hutchings & Johnson 2243 (KEI); 3228 BD, Johnson 270 (KEI)
Lygodium Swam
kerstenii Kuhn, 3129 DA, Hutchings 2192 (BOL, E, KEI, NH). The
identity of this material is still being investigated.
MARSILEACEAE
Marsilea L.
macrocarpa Presl, 3127 AA, Pegler & Kolbe 1528a (BOL); 3127 DB,
Pegler 1528 (BOL); 3127 AD, Kolbe & Pegler 193 (GRA); 3129
DA, Hardcastle 317 (NBG); 3228 AB, Roux 543 (NBG)
capensis A. Br. , 3129 DA, Hardcastle 317
CYATHEACEAE
Cyathea J. E. Sm.
dregei Kunze, 3030 BC, Roux 664 (NBG); 3128 BC, Hutchings 1067
(KEI); 3128 CB, Baur 161 (GRA, SAM); 3129 BB, Roux 612
(NBG); 3129 BC, Hutchings 1997 ( KEI); 3130 AA, Griffen s.n.
(GRA)
capensis (L. f.) J.E. Sm. , 3128 CB, Baur 194 (SAM)
HYMENOPHYLLACEAE
Trichomanes L.
reptans Swam, 3129 DA, Schelpe 5043 (BOL)
rigidum Swam, *grid unknown, Drege s.n. (BM)
pyxidiferum L. var. melanotrichum (Schlechtd.) Schelpe, 3030 BC, Roux
648 (NBG); 3129 BC, Strey 8581 (BOL); 3129 DA, Hutchings
1177 (BOL, KEI)
DENNSTAEDTIACEAE
Blotiella Tryon
natalensis (Hook.) Tryon, 3129 BC, Sim s.n. sub CH 3744 (PRE)
glabra (Bory) Tryon, 3129 DA, Bolus 10368 (BOL); *grid unknown,
Drege s.n. (K, BM)
Pteridium Gled. ex Scop.
aquilinum (L.) Kuhn subsp. aquilinum, 3029 AB, Schelpe 5084 (BOL);
3129 BD, Cawe 81 (KEI); 3129 CB, Roux 567 (NBG); 3129
DA, Schelpe 359 (NU); 3130 AA, Schelpe 5084 (BOL)
Hypolepis Bemh.
sparsisora (Schrad.) Kuhn, 3030 CC, Roux 666 (NBG); 3129 BC, Cawe
229 (KEI); 3129 CB, Roux 566 (NBG); 3129 DA, Hutchings
& Johnson 2238 (KEI)
VITTARIACEAE
Vittaria J.E. Sm.
isoetifolia Bory, 3030 CC, Roux 657 (NBG); 3129 BC, Strey 8868 (BOL);
3129 DA, Mcloughlin 776 (BOL); 3129 DA, Hutchings 1175
(BOL, KEI); 3228 CB, Flanagan 1809 (GRA)
Bothalia 19,2 (1989)
187
ADIANTACEAE/PTERIDACEAE
Acrostichum L.
aureum L. , 3130 AA, Ward 911 (NU, KEI)
Pityrogramma Link
calomelanus (Swartz) Link var. aureoflava (Hook.) Weath. ex Bailey ,
3029 AB, Schelpe 5056 (BOL); 3129 DA, Hutchings & John-
son 2246 (KEI)
Adiantum L.
capillus-veneris L. , 3127 AA, Esterhuysen 29208 (BOL); 3127 BC,
Schelpe 5837 (BOL); 3128 BC, Hutchings 2216 (KEI); 3129 DA,
Hutchings & Johnson 2235 (KEI); 3129 CC, Hoffman 014 (KEI)
poiretii Wikstr. , 3028 DD, Barker 6166 (NBG); 3029 DA, Strey 6393
(BOL); 3128 CA, Hutchings & Plumstead 1092 (KEI)
raddianum Presl, 3129 DA, Hutchings & Plumstead 2056 (KEI)
Pteris L.
vittata L. , 3029 CC, Hutchings 1354 (BOL, KEI); 3128 AB, Baur 225
(SAM); 3129 CB, Roux 569 (NBG)
cretica L. , 3028 DD, Barker 6170 (NBG); 3029 DA, Law s.n. (NU);
3127 AA, Roy ffe 240 (GRA); 3128 BC, Hutchings & Johnson
M8 (KEI); 3128 CA, Johnson 331 (BOL, KEI)
buchananii Bak. ex Sim , 3129 DA, Hardcastle 282 (NBG); 3129 CC,
Hoffman 16 (KEI)
dentata Forssk., 3030 CC, Roux 667 (NBG); 3127 DB, Cawe 842
(BOL, KEI); 3128 CA, Wells 3599 (GRA); 3129 DA, Giffen
s.n. (GRA); 3228 BD, Johnson students s.n. (KEI)
caloplera Kunze, 3029 AB, Schelpe 5064 (BOL); 3029 DA, Taylor 5325
(NBG); 3128 BC, Cawe 451 (KEI, BOL); 3129 DA', Schelpe
360 (NU); 3228 BD, Johnson students 2 (KEI)
Cheilanthes Swartz
eckloniana (Kunze) Men. , 3029 AB, Mcloughlin 726 (BOL); 3029 DD,
Schelpe 5076 (BOL); 3127 AA, Pegler 1636 (BOL); 3128 CA,
Cawe 759 (BOL, KEI); 3128 BD, Roux 563 (NBG)
inaequalis (Kunze) Mettx ar. buchananii (Bak.) Schelpe, 3029 AD,
Taylor 5486 (NBG); 3030 CC, Roux 679 (NBG); 3128 DB, Roux
563 (NBG); 3129 DA, Flanagan 2580 (SAM)
parviloba (Swartz) Swartz , grid unknown, Young 3675 (PRE)
hirta Swartz, 3029 CC, Hutchings 1383 (BOL, KEI); 3127 AA, Schelpe
5840 (BOL); 3127 CA, Hutchings & Johnson 1660 (KEI); 3128
CA, Cawe 897 (BOL, KEI); 3129 AB, Hutchings 1353 (KEI);
3129 DA, Roux 595 (NBG)
involuta (Swartz) Schelpe & N.C. Anthony
var. involuta, 3128 DB, Hutchings 96 (KEI)
var. obscura (N.C. Anthony ) N.C. Anthony, 3227 DB, Flanagan 1240
(GRA)
viridis (Forssk.) Swartz
var. viridis, 3128 BC, Lewis 2853 (SAM); 3129 DA, Roux 597 (NBG);
3228 BB, Hutchings 650 (KEI)
var. macrophylla ( Kuntze ) Schelpe & N.C. Anthony, 3029 AD, Taylor
5221 (NBG); 3030 CC, Strey 8718 (BOL); 3129 BC,
Hutchings & Johnson 2251 (KEI); 3129 DA, Hutchings 1176
(BOL, KEI); 3228 DB, Hutchings 650 (KEI); 3228 BC, NBG
365/73 (BOL); 3228 BD, Roux 553 (NBG); *grid unknown,
Drege s.n. (K)
var. glauca (Sim) Schelpe & N.C. Anthony, 3030 CC, Roux 661 (NBG);
3127 DB, Cawe 232 (KEI); 3129 DA, Roux 597 (BOL); 3130
AA, Schelpe 5078 (BOL)
quadripinnata (Forssk.) Kuhn, 3028 AD, Roux 1377 (NBG); 3028 DD,
Barker 6168 (NBG); 3029 AD, Taylor 5483a (NBG); 3029 DA,
Taylor 5324 (NBG); 3127 AA, Pegler 1634 (BOL); 3127 BC,
Schelpe 5843 (BOL); 3127 DB, Esterhuysen 29146 (BOL); 3128
AB. Johnson & Hutchings 1077 (BOL, KEI); 3128 BC,
Hutchings 2167 (BOL, KEI)
multifida (Swartz) Swartz
subsp. multifida, 3127 AA, Schelpe 5838 (BOL); 3129 DA, Mcloughlin
s.n. (BOL)
subsp. lacerata N.C. Anthony & Schelpe, *grid unknown, Drege s.n.
(LZ, B)
bergiana Schlechtd., 3029 AD, Taylor 5230 (NBG); 3029 DA, Taylor
5324 (NBG); 3030 CC, Roux 652 (NBG); 3128 BC, Hutchings
23 (KEI); 3128 CB, Bear 11, 42 (SAM); 3129 DA, Schelpe 352
(NU); 3228 BD, Johnson 14 (KEI)
concolor (Langsd. & Fisch.) R. & A. Tryon, 3030 CC, Roux 691 (NBG);
3128 CA, Cawe 737 (BOL, KEI); 3129 DA, Schelpe 349 (NU);
3228 BD, Johnson students 12 (KEI); 3229 AA, Johnson 753
(KEI)
Pellaea Link
calomelanos (Swartz) Link, 3029 AD, Taylor 5484 (NBG); 3127 CB,
Schelpe 5883 (BOL); 3127 CA, Hutchings 1607 (BOL, KEI); 3128
DB, Johnson 36 (KEI); 3129 DA, Schelpe 356 (KEI); 3228 AD,
Compton 17731 (NBG)
POLYPODIACEAE
Pyrrosia Mirb.
africana (Kunze)Ballard, 3129 BC, Strey 8583 (BOL); 3129 DA, Roux
571 (NBG); 3228 AD, Ward 5710 (NU); 3228 BB, Wells 3566
(GRA); 3228 BD, Johnson students 5a (KEI)
Loxogramme (Blume) Presl
lanceolata ( Swartz ) Presl, 3128 CB, Baur 1 (GRA) s.n. (SAM); 3129
DA, Hardcastle 267 (NBG)
Polypodium L.
vulgare L. , 3029 CB, Tyson 1642 (SAM); 3128 BC, Cawe 625 (BOL,
KEI)
polypodioides (L.) Hitchc. subsp. ecklonii (Kunze) Schelpe, 3128 BC,
Hutchings 47 (KEI); 3129 AB, Roux 619 (NBG); 3128 CB, Baur
s.n. (SAM)
Pleopeltis H.B.K. ex Willd.
macrocarpa (Bory ex Willd.) Kaulf, 3029 AD, Taylor 5228 (NBG);
3128 CB, Baur s.n. (BOL); 3129 DA, Hutchings & Johnson
2242 (KEI)
schraderi (Men.) Tardieu, 3028 DD, Barker 6/6 7( NBG); 3128 BA, Cawe
624 (KEI, BOL); 3128 CA, Hutchings & Plumstead 2078 (KEI)
Microgramma Presl
lycopodioides (L.) Copel., 3130 CC, Johnson 2075 (KEI)
Microsorium Link
punctatum (L.J Copel., 3130 CC, Johnson 2077 (KEI); 3129 DA,
Hutchings & Johnson 2234 (KEI)
scolopendrium (Burm. f. ) Copel. , 3129 DA, Hutchings & Johnson 2231
(KEI); 3130 AC, Hutchings 2215 (KEI)
ensiforme (Thunb.) Schelpe, 3129 DA, Bunt Davy 3895 (PRE)
DAVALLIACEAE
Nephrolepis Schott
exaltata (L. ) Schott, 3129 DA, Hutchings & Johnson 2229 (KEI)
Oleandra Cav.
distenta Kunze, 3129 DA, Roux 591 (NBG)
Davallia J.E. Sm.
chaerophylloides (Poir. ) Steud. , 3030 CC, Roux 678 (NBG); 3129 BC,
Leighton s.n. (BOL); 3129 BD, Hutchings 2024 (KEI); 3129
DA, Schelpe 5030 (BOL); 3130 AA, Schelpe 5076 (BOL)
ASPLENIACEAE
Asplenium L.
anisophyllum Kunze, 3129 BC, Strey 8871 (BOL)
prionitis Kunze, grid unknown, Drege s.n. ; 3030 CC, Roux 649 (NBG);
3128 BC, Hutchings 44 (KEI); 3129 DA, Hutchings 1174 (KEI);
3228 BD, Johnson students 9 (KEI)
boltonii Hook, ex Schelpe, 3029 BB, Schlechter 6229 (GRA); 3128 BC,
Hutchings s.n. (BOL, KEI); 3129 DA, Mcloughlin s.n. (BOL)
gemmiferum Schrad. , 3030 CC, Roux 650 (NBG); 3127 DB, Mcloughlin
s.n. (BOL); 3129 DA, Schelpe 346 (NU); 3228 BD, Roux 549
(NBG)
gemmiferum Schrad. x flexuosum Schrad. , 3129 DA, Mcloughlin s.n.
(BOL)
protensum Schrad. , 3029 BB, Schlechter 6635 (GRA); 3128 AD, Cawe
673 (BOL, KEI); 3128 AC, Cawe 800 (BOL, KEI); 3129 DA,
Mcloughlin 755 (BOL)
sandersonii Hook. , 3228 BD, Roux 545 (NBG)
stoloniferum Bory, 3127 AA, Pegler 1637 (BOL); 3128 AB, Johnson
& Hutchings 1079 (BOL, KEI)
trichomanes L. , 3128 AD, Roux 1379 (NBG)
platyneuron (L.) Oakes, 3027 DB, Roux 1208 (NBG)
monanthes L. , 3028 DD, Barker 6169 (NBG); 3127 AA, Royffe 240
(GRA); 3128 AD, Cawe 665 (BOL, KEI)
lunulatum Swartz, 3030 CC, Roux 653 (NBG); 3128 BC, Cawe 454
(BOL), 456 (BOL, KEI); 3128 BC, Hutchings 119 (KEI); 3129
DA, Hutchings & Johnson 2240 (KEI); 3228 BD, Johnson
students 13 (KEI)
erectum Bory ex Willd. var. erectum, 3127 DB, Cawe 868 (BOL, KEI);
3128 AC, Cawe 799 (BOL, KEI); 3128 BA, Cawe 623 (BOL,
KEI); 3128 BC, Cawe 489, Hutchings 40, 119 (BOL, KEI); 3129
DA, Mcloughlin 783 (BOL)
inaequilaterale Willd., 3129 DA, Schelpe 5042 (BOL)
dregeanum Kunze, 3129 DA, Johnson 108 (KEI); *grid unknown, Drege
s.n.
theciferum (H.B.K.) Mett. var. concinnum (Schrad.) Schelpe, 3029 AD,
Taylor 5229 (NBG)
rutifolium (Berg.) Kunze, 3030 CC, Roux 646 (NBG); 3127 DB,
Mcloughlin 1 (BOL); 3129 BC, Hutchings & Johnson 2248
(KEI); 3129 CB, Wells 3489 (GRA); 3129 DA, Hutchings 1174a
(KEI); 3228 BD, Johnson students 11 (KEI)
188
Bothalia 19,2 (1989)
lobatum Pappe & Rawson, 3128 AD, Cawe 670 (BOL, KEI)
varians Wall, ex Hook. & Grev. subsp. fimbriatum (Kunze) Schelpe,
3129 DA, Hardcastle 255a (NBG)
adiantum-nigrum L. var. solidum (Kunze) J.P. Roux, 3228 CB, Flanagan
146 (PRE)
splendens Kunze, 3030 CC, Roux 647 (NBG); 3128 BC, Hutchings 122
(KEI); 3128 BC, Hutchings 670 (BOL, KEI); 3128 CA,
Hutchings 388 (KEI); 3129 CC, Wells 3547 ( GRA); 3129 DA,
Hutchings & Johnson 1137 (KEI); 3228 BD, Johnson students
U (KEI)
simii Braithwaite & Schelpe, 3130 AA, Bayliss 2223 (BOL, KEI)
aethiopicum (Burnt, f.) Becherer, 3127 AA, Royffe 197 (GRA); 3127
AD, Schelpe 5832 (BOL); 3127 DB, Flanagan 2580 (SAM);
3127 DB, Cawe 865 (BOL, KEI); 3128 AC, Cawe 805 (BOL);
3128 BC, Cawe 484, Hutchings (BOL, KEI)
Ceterach D.C.
cordatum (Thunb.) Desv., 3121 BC, Schelpe 5842 (BOL); 3128 CA,
Cawe 892 (BOL, KEI); 3129 DA, Mcloughlin s.n. (BOL), 794
(BOL); 3228 BD, Roux 556 (NBG)
THELYPTERIDACEAE
Thelypteris Schmidel
madagascariensis (Fee) Schelpe, 3128 BA, Cawe 734 (KEI, BOL)
interrupta (Willd.) K. Iwats. , 3128 BC, Cawe 726 (KEI): 3129 BD, Cawe
80 (KEI); 3129 CB, Wells 3479 (NBG); 3129 DA, Roux 602
(NBG); 3229 AA, Johnson 756 (KEI)
pozoi (Lagasca) Morton, 3029 BB, Cawe 1003 (BOL); 3127 DC, Cawe
901 (BOL, KEI)
dentata (Forssk.) E. St. John, 3127 DB, Cawe 898 (BOL, KEI); 3129
DA, Schelpe 5033 (BOL); 3130 AA, Schelpe 5075 (BOL); 3228
BD, Moll 5747 (BOL); 3229 AA, Johnson 756 (KEI)
gueinziana (Mett.) Schelpe, 3029 AB, Schelpe 5058a (BOL); 3129 DA,
Hutchings & Johnson 2249 (BOL, KEI)
bergiana (Schlechtd.) Ching, 3029 CB, Schlechter 6512 (NBG); 3129
DA, Roux 581 (NBG); 3228 BD, Roux 555 (NBG)
Macrothelypteris (H. Ito) Ching
torresiana (Gaud.) Ching, 3129 DA, Roux 1971 (NBG)
ATHYRIACEAE
Athyrium Roth
schimperi Moug. ex Fee, 3029 BC, Schlechter 6585 (GRA)
Cystopteris Bemh.
fragilis (L.) Bemh., 3029 BB, Schlechter 6638 (GRA); 3029 BC,
Schlechter 6585 (GRA)
LOMARIOPSIDACEAE
Elaphoglossum Schott
hybridum (Bory) Brack, 3127 DB, Mcloughlin 106 (PRE)
macropodium (Fee) T. Moore, 3129 DA, Roux 586 (NBG); 3129 BC,
Sim 2461 (PRE)
conforme (Swartz) J ■ Sm. , 3129 DA, Roux 586 (NBG); grid unknown,
Taylor 2602 (NBG)
acrostichoides (Hook. & Grev.) Schelpe, 3128 BC, Hutchings 1977 (KEI);
3128 CA, Cawe 871 (BOL)
angustatum (Schrad.) Hieron. , 3129 BC, Cawe 231 (KEI)
ASPIDIACEAE/DRYOPTERIDACEAE
Woodsia R. Br.
montevidensis (Spreng.) Hieron. var. burgessiana (Gerr. ex Hook. &
Bak.) Schelpe, 3129 CB, Schlechter 6526 (GRA)
Dryopteris Adans.
squamiseta (Hook.) Kuntze, 3028 AD, Johnson 709 (KEI)
athamantica (Kunze) Kuntze, 3029 AB, Hutchings 2133 (KEI, NU); 3029
CB, Schlechter 6556 (GRA); 3128 CA, Cawe 767 (BOL, KEI);
3128 DB, Fr. Abbott 29 (GRA)
inaequalis (Schlechtd.) Kuntze sens, lat., 3029 AD, Taylor 5482 (NBG);
3029 BC, Schlechter 6512 (GRA); 3129 AB, Schelpe 5063
(BOL); 3129 CB, Roux 565 (NBG); 3129 DA, Hutchings 571
(BOL, KEI); 3228 BD, Johnson students 8 (KEI)
dracomontana Schelpe & N.C. Anthony, 3128 AD, Roux 1376 (NBG)
Cyrtomium Presl
caryotideum (Wall, ex Hook. & Grev. ) Presl var. micropterum (Kunze)
C. Chr. , 3128 AD, Flanagan s.n. (SAM); 3029 BB, Schlechter
6635 (GRA); 3127 DC, Cawe 900 (KEI, BOL); 3128 BC, Cawe
327 (BOL, KEI); 3129 DA, Hutchings & Johnson 2232 (KEI)
Polystichum Roth
transkeiense Jacobsen, 3029 DA, Taylor 5227 (NBG); 3129 BC, Strey
8869 (BOL, NU); 3129 DA, Jacobsen 4301 (BOL)
pungens (Kaulf.) Presl, 3128 AB, Hutchings & Johnson 1081 (BOL,
KEI); 3128 BC, Cawe 443, Hutchings 118 (BOL, KEI); 3129
DA, Schelpe 357, 358 (NU)
luctuosum (Kunze) T. Moore, 3128 AC, Cawe 777 (BOL, KEI); 3128
AD, Cawe 660 (BOL, KEI); 3128 BC, Hutchings 118 (KEI)
transvaalense N.C. Anthony, 3129 DA, Wager s.n. sub CH 2905 (PRE);
3127 DB, Mcloughlin s.n. sub CH 7677 ( PRE); 3127 DA, Young
s.n. sub TM 511c (PRE)
monticola N.C. Anthony & Schelpe, 3129 BC, Cawe 228 (KEI)
Arachniodes Blume
foliosa (C. Chr. ) Schelpe , 3029 CD, Adams 164 (UN)
Rumohra Raddi
adiantiformis (G. Forst.) Ching, 3129 DA, Roux 593 (NBG); 3228 AD,
Ward 5715 (NU)
Ctenitis (C. Chr.) C. Chr. ex Tardieu
lanuginosa (Willd. ex Kaulf. ) Copel. , 3128 BA, Cawe 735 (BOL, KEI);
3128 CB, Baur 221 (GRA); 3129 DA, Mcloughlin 785 (BOL);
*grid unknown, Drege s.n. (LZ)
BLECHNACEAE
Blechnum L.
giganteum (Kaulf.) Schlechtd. , 3127 DB, Cawe 867 (BOL, KEI); 3128
BC, Hutchings & Johnson M7 (BOL, KEI); 3129 DA,
Hardcastle 288 (NBG)
capense (L.) Bunn, f, 3129 BD, Johnson 179 (KEI); 3130 AA, Schelpe
5074 (BOL)
tabulare (Thunb.) Kuhn, 3029 AB, Schelpe 5061 (BOL); 3029 CC, Cawe
945 (BOL); 3128 BC, Hutchings 2165 (KEI, NH); 3129 AB,
Roux 520 (NBG)
australe L. var. australe, 3028 BA, Matthews 957 (NBG); 3127 AA,
Royffe 90 (GRA); 3128 AB, Hutchings & Johnson 1082 (BOL,
KEI); 3127 DC, Cawe 899 (BOL, KEI); 3128 AD, Cawe 666
(BOL, KEI); 3128 CA, Cawe 840 (BOL, KEI); 3128 CB, Baur
s.n. (GRA)
punctulatum Swartz
var. punctulatum, 3129 AD, Johnson 710 (KEI); 3128 CB, Cawe 680
(BOL, KEI); 3129 DA, Williams 102 (NU)
var. atherstonei (Pappe & Raws.) Sim, 3128 CB, Baur 640 (SAM)
var. intermedium Sim, 3128 AC, Taylor 2599 (NBG); 3129 BC, Taylor
2632 (NBG)
var. krebsii (Kunze) Sim, 3129 DA, Mcloughlin 800 (BOL); 3129 DA,
Roux 583 (NBG); 3129 BC, Taylor 2632 (NBG)
Stenochlaena J. Sm.
tenuifolia (Desv.) T. Moore, *grid unknown, Drege s.n. (LZ)
Bothalia 19,2: 189 - 207 (1989)
Taxonomy and leaf anatomy of the genus Ehrharta (Poaceae) in
southern Africa: the Ramosa group
G.E. GIBBS RUSSELL* and R.P. ELLIS*
Keywords: Capensis, culm anatomy, Ehrharta , Fynbos, leaf anatomy, Poaceae, taxonomy
ABSTRACT
The Ramosa species group in the genus Ehrharta is distinguished morphologically by small spikelets with sterile lemmas
similar to each other, with tips rounded, truncate or mucronate, with sides glabrous, scabrous or shortly hairy, and with
bases appendaged and usually bearded. The rectangular intercostal long cells with sinuous walls, the dome-shaped stomata
with a raised rim surrounding the pore aperture, the absence of epicuticular wax and the microhairs without a tapering distal
cell are diagnostic anatomically. The Ramosa group is composed of two species: E. ramosa (Thunb.) Swartz subsp. ramosa ;
subsp. aphylla (Schrad.) Gibbs Russell and E. rehmannii Stapf subsp. rehmannii ; subsp . filiformis (Nees) Gibbs Russell;
subsp. subspicata (Stapf) Gibbs Russell. All taxa are linked by intermediates to one or two others in the group. The closest
relationship of the Ramosa group is to the Calycina species group, on the basis of both morphological and anatomical characters.
UITTREKSEL
Die Ramosa-spesiegroep in die genus Ehrharta word morfologies onderskei deur klein blompakkies met steriele eenderse
lemmas, met punte gerond, afgeknot of stekelpuntig, met kante onbehaar, skurf of kortharig, en met basisse met aanhangsels
en gewoonlik bebaard. Die reghoekige tussenribbige lang selle met gekartelde wande, die koepelvormige huidmondjies met
’n verhewe rand random the porie-opening, die afwesigheid van epikutikulere was en die mikrohare sonder ’n spitslopende
distale sel is anatomies kenmerkend. Die Ramosa-groep bestaan uit twee spesies: E. ramosa (Thunb.) Swartz subsp. ramosa;
subsp. aphylla (Schrad.) Gibbs Russell en E. rehmannii Stapf subsp. rehmannii; subsp. filiformis (Nees) Gibbs Russell;
subsp. subspicata (Stapf) Gibbs Russell. Alle taksons word deur tussenvorms aan een of twee ander taksons in die groep
gekoppel. Die naaste verwantskap van die Ramosa-groep is met die Calycina-spesiegroep, op grand van morfologiese sowel
as anatomiese kenmerke.
INTRODUCTION
Previous papers in this series have outlined the seven
provisional species groups of Ehrharta in southern Africa,
and have dealt with the Setacea, Villosa and Dura groups
(Gibbs Russell & Ellis 1987, 1988; Ellis 1987a; 1987b;
Gibbs Russell 1987a, 1987b). This paper examines the
Ramosa group, which is distinguished from the other
Ehrharta species groups by the small spikelet size (4—9
mm long), sterile lemmas similar to each other, with tips
rounded, truncate or mucronate, with sides glabrous,
scabrous or shortly hairy, and with bases appendaged and
usually bearded (Figure 1). The rectangular intercostal
long cells with sinuous walls, the dome-shaped stomata
with a raised rim surrounding the pore aperture, the
absence of epicuticular wax and the microhairs without
a tapering distal cell are diagnostic anatomically. In
contrast to the species groups covered already, in which
taxa are relatively easy to define and may be distinguished
by clear-cut characters, the five taxa comprising the
Ramosa group intergrade and few consistently reliable
characters adequately separate the species and subspecies.
Therefore, the taxa accepted here must be considered as
noda in a continuum of intra-group variability. Despite the
difficulty, it is desirable to treat the Ramosa group taxa
formally for the following reasons: 1, most specimens fall
within the noda, as here defined, and only a comparative
* Botanical Research Institute, Department of Agriculture and Water
Supply, Private Bag X101, Pretoria 0001.
MS. received: 1988.10.27.
few are intermediates; 2, the great differences between
the ‘ends’ of the continuum, both morphological and
ecological, require a means of distinguishing between,
e.g., the robust bladeless subshrub of rocky places
common at high altitudes (E. ramosa subsp. ramosa ) and
the delicate trailing herb with thin soft leaves from moist
sheltered sites (E. rehmannii subsp. filiformis)', 3, each of
the taxa has a distinct geographical range.
The two taxa that are common in the Fynbos Biome are
large plants of widespread distribution, E. ramosa subsp.
ramosa and subsp. aphylla. They have been recognized
from very early times in the study of the southern African
flora (Thunberg 1794). The smaller, less common plants
with more limited distribution were first treated as separate
species by Stapf (1897, 1900). Previous to his classification,
specimens of these taxa had been attributed to E. ramosa
and E. aphylla.
As a result of the confusing pattern of variation and the
inadequate early classification of the Ramosa group, its
nomenclature has also been somewhat difficult. For each
of the five taxa in the group a lectotype or neotype was
designated.
METHODS
Methods adopted for previous papers of this series were
followed here (Gibbs Russell & Ellis 1987). The
descriptions and keys were prepared through the DELTA
computer system (Dallwitz 1984).
//a ) ] b
TAXONOMY
Key to species in the Ramosa group
Plants robust, strongly suffrutescent; leaf blades absent or
reduced; sterile lemmas usually mucronate E. ramosa
Plants not robust, usually herbaceous; leaf blades present,
expanded; sterile lemmas usually muticous E. rehmannii
1. Ehrharta ramosa (Thunb.) Swartz in Transactions
of the Linnean Society 5: 49 (1802). Thunb.: 336 (1818);
Thunb. ed. Schultes: 335 (1823); Schrader: 2077 (1821);
Schrader in Schultes: 1370 (1830); Trinius: 25 (1839);
Nees: 205 (1841); Steudel: 7 (1855); Stapf: 677 (1900);
Chippindall 39 (1955); Smook & Gibbs Russell: 55 (1985).
Type: Thunberg (sheet 8855), UPS, lecto. here designated
— IDC microfiche 1036 in PRE!
Melica ramosa Thunb.: 21 (1794). Ehrharta digyna Thunb.: 66 (1794).
Perennial, tufted, erect or straggling, long-rhizomatous,
often robust, suffrutescent. Rhizomes woody, branched,
naked or with cataphylls. Cataphylls glabrous, thin, or
thickened. Culms many, erect or geniculate, woody or wiry
and persistent, hollow, crowded at the ends of the
rhizomes, branched at base and branching above, usually
with many fascicled branches, the internodes sometimes
with globose orange gall-like swellings at their base. Young
shoots intravaginal . Leaves not basally aggregated. Culm
leaves with sheaths not overlapping, blades absent or
reduced, sheaths held away from the culm. Basal sheaths
loose, deciduous, or persistent, papery, or membranous,
grey, not bearing blades. Ligule a membrane fringed with
hairs, about 1 mm long.
Inflorescence a raceme or a panicle, open or contracted,
of several to many appressed, spreading or nodding
spikelets, closely subtended or enveloped below by
uppermost leaf sheath or exserted from it, main axis
sinuous around appressed spikelets. Spikelets pedicellate,
distinctly compressed laterally, (5,5— )6— 7,5(— 9) mm long,
about 2—3 mm across. Pedicels usually with short stiff
hairs. Glumes keeled, subequal, slightly shorter than rest
of spikelet to considerably longer than rest of spikelet,
yellowish or green, somewhat spreading or appressed to
lemmas at maturity. Lower glume 5— 7-nerved, acute.
Upper glume 3— 5-nerved, acute. Florets with lemmas
decidedly firmer than the glumes, keeled. Sterile lemmas
similar in shape and texture, laterally compressed, sides
flat. Lower sterile lemma about 3/4 length of upper sterile
lemma to about equalling length of upper sterile lemma,
with keel and margin parallel; base not stipitate, with
PRE): a, whole spikelet; b,
glumes; c, first sterile lemma; d,
fertile lemma; e, second sterile
lemma; f, palea.
auriculate appendages, bearded; sides glabrous or
sometimes scabrous, often short-hairy on the margins,
dull, with 2—3 fairly distinct to faint longitudinal ribs and
rarely with up to 12 faint transverse corrugations; tip
truncate or slightly oblique, usually mucronate, but
sometimes aristate or sometimes muticous. Upper sterile
lemma similar to lower sterile lemma, but often with
transverse corrugations, substipitate, longer than lower
sterile lemma. Fertile floret with lemma differing from
sterile lemmas, strongly laterally compressed and sides
unornamented, not exceeding upper sterile lemma or
sometimes slightly longer than both sterile lemmas,
7— 9-nerved, sides glabrous or scabrous on nerves; tip
truncate, midnerve sometimes submucronate. Palea thinner
than lemma, 3/4 or more as long as lemma. Lodicules 2,
fleshy at base, and membranous above, ciliate at margins,
or glabrous. Stamens 6. Anthers 2,5—4 mm long, yellow.
Ovary glabrous. Stigmas white. Caryopsis not seen
(spikelets shed early).
Both subspecies of E. ramosa inhabit Mountain Fynbos,
growing in xerophytic to mesophytic open habitats.
This species differs from E. rehmannii in its robust,
consistently suffrutescent habit, bladeless or nearly
bladeless leaves, and sterile lemmas with usually
submucronate to shortly aristate tips. Although the break
between the five taxa in the Ramosa group is greatest
between the two species as based on the above characters,
the separation is not complete, for a number of
intermediates are known. The specimens are discussed in
more detail under particular subspecies.
Although the majority of E. ramosa specimens can be
identified to subspecies, there are a number which show
intermediate states for the characters that separate the taxa:
the size of the plant can vary with age, and in any case
may not be apparent on a poor herbarium specimen; the
openness of the inflorescence can vary with developmental
stage, with young inflorescences of subsp. aphylla showing
erect spikelets; glume length is variable in subsp. ramosa ,
being either slightly longer or shorter than the sterile
lemmas. The difficulty of separating the subspecies has
led to their treatment in synonymy as early as Trinius
(1839), or as infraspecific taxa (Gluckmann ex Adamson
1942). The two taxa are treated here as subspecies because
each has a distinct geographical range. Intermediate speci-
mens are discussed under subsp. aphylla.
Bothalia 19,2 (1989)
191
There are three sheets of E. ramosa in Thunberg’s
herbarium. Sheet 8855 was chosen as the lectotype
because it is the most complete, showing typical culm size
and branching, leaf sheaths and several inflorescences.
Key to subspecies
Plants very robust, culms to 5 mm across; glumes usually
slightly shorter than sterile lemmas; inflorescence usually
contracted, pedicels erect to ascending; leaves bladeless
la. E. ramosa subsp. ramosa
Plants robust, culms to 2,5 mm across; glumes slightly to
considerably longer than sterile lemmas; inflorescence
usually open, pedicels spreading to reflexed; leaves
rarely with small blades lb. E. ramosa subsp. aphylla
la. Ehrharta ramosa (Thunb.) Swartz subsp. ramosa
Culms markedly robust, woody, 300—1 000 mm long,
to 5 mm across, erect, or geniculate. Culm leaves with
blades absent, at most the blades occasionally represented
by an apiculate inrolled tip to the sheath. Inflorescence
a contracted raceme 30—140 mm long with appressed
spikelets. Spikelets pedicellate, the pedicels held erect.
Glumes 4,5—8 mm long, usually slightly shorter than rest
of spikelet or sometimes slightly longer than rest of
spikelet. Lower sterile lemma about 3/4 length of upper
sterile lemma to about equalling length of upper sterile
lemma' tip strongly truncate, usually purple-tinged for
about ;4 of lemma length. Fertile floret not exceeding
upper sterile lemma (shorter or subequal).
Subsp. ramosa is distinguished from subsp. aphylla by
its more robust habit and very thick culms, its narrow
inflorescence with erect appressed spikelets, glumes that
are often shorter than the lemmas, and the markedly
truncate sterile lemmas with purple-tinged ends. It is
characteristic of Mountain Fynbos although it extends into
Grassy Fynbos and occurs at low altitudes in the eastern
part of its range. Its distribution is shown in Figure 2.
Subsp. ramosa has the widest range of the five taxa in the
Ramosa species group, although subsp. aphylla extends
slightly further to the north and E. rehmannii subsp.
rehmannii extends slightly further to the east. It is unusual
in Ehrharta for a taxon to have both a western and an
eastern distribution and yet be absent from the Cape
Peninsula-Caledon area. Subsp. ramosa grows in sandy
or stony Table Mountain Sandstone (TMS) or lateritic
soils, often in rocky places, at altitudes of 90 to 2 200 m;
and is common at high altitudes. Flowering occurs from
October to January.
FIGURE 2. — Distribution of E. ramosa subsp. ramosa.
lb. Ehrharta ramosa (Thunb. ) Swartz subsp. aphyl-
la (Schrad.) Gibbs Russell, stat. nov.
Ehrharta aphylla Schrad. in Gottingische gelehrte Anzeigen 1821: 2077
(1821); Schrad. in Schultes: 1369 (1830); Nees: 207 (1841); Steud.: 7
(1855); Stapf: 678 (1900). Ehrharta ramosa Thunb. var. aphylla (Schrad.)
Gluckmann ex Adamson: 273 (1942); Chippindall: 39 (1955). Type: Hesse,
apparently lost; neotype here designated: Ecklon 914 (specimen in K with
suffrutescent culms and bladeless leaves!).
E. ramosa Thunb. var. aphylla (Schrad.) Gluckmann ex Adamson:
273 (1942).
E. aphylla Schrad. var . fasciculata Stapf: 679 (1900). Type: tops of
mountains, Baviaans Kloof, Burchell 7725 (K, holo.!; photo and fragment
in PRE!).
Culms not robust to robust, woody or wiry, 300—800
mm long, to 2,5 mm across, geniculate or decumbent at
base. Culm leaves with blades usually absent, but
occasionally present on upper leaves. Leaf blades when
present lanceolate; to 1 mm across, to 30 mm long, rolled,
erect, herbaceous, glabrous, gradually tapering at the tip.
Inflorescence a raceme or a panicle with 1 or 2 branches,
usually open or sometimes contracted, 20-90 mm long,
with spikelets spreading to nodding. Spikelets pedicellate,
the pedicels spreading to reflexed. Glumes 4,5—9 mm
long, usually slightly or considerably longer than rest of
spikelet but sometimes slightly shorter. Lower sterile
lemma slightly shorter than upper sterile lemma, rarely
purple-tinged and then only at the tips, with up to 12
corrugations; tip oblique or truncate. Upper sterile lemma
usually with marked transverse corrugations. Fertile floret
sometimes exceeding upper sterile lemma.
Subsp. aphylla differs from subsp. ramosa in its
generally smaller size, open inflorescence with spreading
to reflexed spikelets, long glumes and sterile lemmas with
truncate to oblique tips that are uncommonly purple-tinged
only at the tips. Its distribution is shown in Figure 3. Subsp.
aphylla occurs only up the western mountains, and is
absent from the eastern part of the range of the Ramosa
group. It grows in Mountain Fynbos on sandy TMS-
derived soil between rocks, often in dry microhabitats, and
is reported 1-4 years after fire, from altitudes of
230—1 500 m. Flowering occurs from (July) September
to January.
A number of specimens are intermediate between subsp.
aphylla and subsp. ramosa. The specimens that occur in
the area of sympatry of the subspecies fall into two groups:
those with the long glumes of subsp. aphylla and the
192
Bothalia 19,2 (1989)
contracted inflorescence and erect spikelets of subsp.
ramosa (Du Toit 1747; Esterhuysen 3930; Uebenberg 4331;
The Forester 8226) and those with the open inflorescence
of subsp. aphylla but with the short glumes and often with
the dark sterile lemmas of subsp. ramosa ( Andreae 705;
Loxton 221; Stokoe (SAM No. ) 54502). A number of
specimens from the Clanwilliam area fall into this second
group of intermediates, and are leafy, often erect, and with
remarkably short glumes ( Esterhuysen 14978, 32217;
Uebenberg 4308; Stokoe 7809). Within the range of subsp.
ramosa only, there are a few specimens with typical
ramosa spikelets and robust culms but the open
inflorescence of subsp. aphylla ( Gibbs Russell 5683, Kensit
Jan. 1914).
Although the leaves of subsp. aphylla are usually
bladeless or with very' reduced blades, a few specimens
show well developed blades (Du Toit 1425; Ellis 648, 4673;
Gibbs Russell 5668, 5672; Hanekom 1294; Richardson
62). This may be natural or phenological variability within
the taxon, or these specimens may represent a link across
the species line to E. rehmannii subsp. rehmannii, although
the two taxa are allopatric. Such a population at Kirsten-
bosch was collected in 1918 ( Page s.n.) and 1955 ( Salter
8665). Gluckmann in 1938 annotated the Page specimen:
‘I have examined clusters of this growing in Kirstenbosch
and found long-leafed forms intermingled with leafless
forms’.
A second possible link across the species line is
represented by specimens from the Ceres area
(Esterhuysen 28110, 28396; Stokoe 2652). They have
typical aphylla spikelets, but the plants tend to be slender
and leafy with only about 5-10 spikelets per inflorescence,
characteristics of E. rehmannii subsp. filiformis , which
does not occur so far to the north.
The Hesse specimen on which Schrader based the name
aphylla has not been found in GOET, LE, B or P, the
herbaria where it was most likely to have been preserved.
Pending its lucky discovery elsewhere, a lectotype or
neotype must be selected. In his treatment of Ehrharta
in Schultes (1830), Schrader cited an additional specimen,
Ecklon 914. Nees (1832) also cited Ecklon 914 as the type
of E. aphylla var. filiformis. The descriptions of the two
taxa differ, Schrader’s referring to a suffrutescent plant
with no leaf blades, while Nees describes a specimen with
soft flat leaves and a much reduced inflorescence. Stapf
(1900) states that there are two specimens marked Ecklon
914 at Kew, one representing subsp. aphylla and one
representing E. rehmannii var. filiformis. I have seen these
specimens, annotated by Stapf, and propose that they serve
as neotypes for these taxa. They cannot be lectotypes be-
cause there is no indication that either was seen by
Schrader or Nees. It would be futile to attempt to find
unequivocal lectotypes for the following reasons: 1,
specimens labelled Ecklon 914 represent more than one
taxon; 2, Ecklon collected a number of Ramosa group
specimens from different places in the Kleinrivier area
and his duplicates are numerous and widely distributed,
many without collection numbers; 3, the whereabouts of
the personal herbaria of Schrader and Nees is not known;
4, no Ehrharta specimens of historical importance exist
any longer at B, which is the most likely location of Nees’s
and Schrader’s lost herbaria.
2. Ehrharta rehmannii Stapf in Kew Bulletin: 288
(1897); Stapf: 677 (1900); Chippindall: 39 (1955). Type:
Outeniqua Mountains, Montagu Pass, Rehmann 74 (K,
lecto.!, here designated; photo and fragment in PRE!).
Perennial, tufted, erect or straggling, long-stoloniferous,
delicate or neither distinctly delicate nor robust, not
suffrutescent but culms sometimes woody below even in
small plants. Rhizomes branched, naked or with cataphylls.
Culms several to numerous, erect to prostrate, woody
below or wiry or herbaceous, crowded at ends of rhizome,
branched at base and branching above, solid or hollow,
the intemodes rarely with gibbous gall-like swellings at
their base. Young shoots intravaginal. Leaves not basally
aggregated. Culm leaves with sheaths not overlapping,
blades well developed. Leaves auriculate, the auricles with
spreading glassy hairs, produced from top of sheath and
base of blade. Basal sheaths loose, persistent, papery or
membranous, grey, yellowish or whitish, bearing blades
or not bearing blades. Ligule a membrane fringed with
hairs, 0,3—2 mm long. Leaf blades deciduous or persistent
on basal sheaths, persistent on culm sheaths, expanded,
linear or lanceolate, erect or spreading, soft or herbaceous,
glabrous, scabrous or hairy, with marginal vein not
thickened or barely thickened, tip gradually tapering.
Inflorescence a raceme or a panicle, reduced or not
reduced, open or contracted, not overtopping leaves to
considerably overtopping leaves, of 1 to many appressed,
spreading or nodding spikelets, closely subtended or
enveloped below by uppermost leaf sheath or exserted from
uppermost leaf sheath, secund or not secund. Spikelets
pedicellate, distinctly compressed laterally, 4,0-8, 5 mm
long. Pedicels usually with short stiff hairs. Glumes
keeled, subequal, slightly shorter to slightly longer than
rest of spikelet, yellowish or green, gaping widely at
maturity or appressed to lemmas at maturity. Lower glume
3— 7-nerved; acute to apiculate. Upper glume 1-5-nerved,
acute to apiculate. Florets with lemmas decidedly firmer
than the glumes, keeled. Sterile lemmas similar in shape
and texture, laterally compressed, sides flat. Lower sterile
lemma about equalling or somewhat shorter than upper
sterile lemma, with keel and margin parallel; base not
stipitate, with auriculate appendages, bearded or not
bearded; sides glabrous, scabrous or with short hairs, dull,
smooth and unornamented or with 0—3 fairly distinct to
faint longitudinal ribs or with 0-6 faint transverse
corrugations; tip truncate or rounded, usually muticous.
Upper sterile lemma similar to lower sterile lemma but
with distinct corrugations in the upper half, substipitate,
longer than lower sterile lemma. Fertile floret with lemma
differing from sterile lemmas, strongly laterally
compressed and sides unomamented, not exceeding upper
sterile lemma, 5 -7-nerved; tip truncate. Palea thinner than
lemma, \ or more as long as lemma. Lodicules 2, fleshy
at base, or membranous above, ciliate at margins or
glabrous. Stamens 6. Anthers 2,2— 2,6 mm long. Caryopsis
not seen (spikelets shed early).
This species usually grows in wetter habitats than E.
ramosa, and often at lower altitudes. It differs from E.
ramosa in its herbaceous to wiry culms, leaves with well
developed blades and muticous sterile lemmas. There are
three subspecies, which intergrade through a number of
intermediates. These specimens are discussed below under
the appropriate subspecies.
Bothalia 19,2 (1989)
193
Although the two species differ in the characters
mentioned above, the separation is not absolute, and a
number of specimens show intermediate character
conditions between E. ramosa subsp. aphylla and both E.
rehmannii subsp. rehmannii and subsp. filiformis. These
are discussed in more detail under E. ramosa subsp.
aphylla above.
Stapf (1900) is responsible for the recognition of the
herbaceous leafy element in the Ramosa species group,
and the three subspecies treated here were all described
(subsp. rehmannii and subspicata) or separated from the
woody taxa (subsp. filiformis) by him. However, it is
evident that his concepts of the divisions within the leafy
taxa changed during the course of his work. In his original
description of E. rehmannii, Stapf (1897) quotes seven syn-
types. Three of these are also quoted as syntypes of E.
subspicata (Stapf 1900), showing that his original concept
of E. rehmannii included both taxa.
The studies reported here, based on far more specimens
and a wider geographical representation than were
available to Stapf, confirm his opinion that there are three
herbaceous leafy taxa in the Ramosa species group.
However, the boundaries between the taxa in this treatment
differ from those of Stapf s as indicated by the specimens
he cited. In particular, several of the narrow-leaved
specimens with glabrous sterile lemmas he cited as E.
rehmannii (including the contentious Ecklon 914) belong
in subsp. filiformis rather than subsp. rehmannii. Several
of the specimens with branched inflorescences and
herbaceous glumes that Stapf cited as E. subspicata belong
in subsp. rehmannii. The lectotypes and neotype were
chosen from among the several specimens cited by Stapf
to reflect the present concepts of each taxon.
Key to subspecies
la Inflorescence contracted, pedicels and spikelets erect;
glumes subcoriaceous 2c. E. rehmannii subsp. subspicata
lb Inflorescence open, pedicels spreading to reflexed and
spikelets spreading to nodding; glumes membranous:
2a Inflorescence of fewer than 20( — 25) spikelets, leaf
blades narrower than 4 mm; sterile lemmas glabrous on
sides 2b. E. rehmannii subsp. filiformis
2b Inflorescence of more than 20 spikelets; leaf blades to
6 mm across; sterile lemmas usually hairy on sides
or tips and/or strongly scabrous on keels
2a. E. rehmannii subsp. rehmannii
2a. Ehrharta rehmannii Stapf subsp. rehmannii
Tufted, erect; neither distinctly delicate nor robust.
Culms several to many, 300—1 000 mm long, to 2,5 mm
across, usually hard, (but to 5 mm across in rare specimens
with spongy culms), wiry or herbaceous, branches from
upper nodes single or few. Leaf blades linear, to 6 mm
across, 60—300 mm long, usually spreading but some-
times erect, herbaceous, scabrous on margins and keel.
Inflorescence a raceme or a verticillate panicle (the lowest
whorls sometimes with a few branches), not reduced,
open, 45—120 mm long, considerably overtopping leaves,
of more than 20 spreading or nodding spikelets, exserted
from uppermost leaf sheath, secund. Spikelets (5— )6— 8
mm long about 2 mm across laterally above the glumes.
Pedicels spreading to reflexed. Glumes membranous,
5,5—8 mm long, gaping widely at maturity. Lower sterile
lemma usually scabrous or with short hairs on ribs, keel,
tips or margins, rarely glabrous, with 3 fairly distinct
longitudinal ribs and sometimes with up to 6 faint trans-
verse corrugations, tip rounded.
Subsp. rehmannii is distinguished from the other two
subspecies by the long broad leaves, the open paniculate
inflorescence with more numerous spreading to reflexed
spikelets, and the sterile lemmas with scabrous to shortly
hairy keel, ribs and tip. Its distribution is shown in Figure
4. This subspecies grows on streambanks and in rocky
places on mountain slopes and sometimes under trees,
400—660 m, flowering from August to December.
A particularly tall, long-leaved form with thick but soft
culms and numerous short spikelets (5,5—6 mm long)
occurs in forests and on rocky ground at George and
Knysna ( Compton 23076; Eourcade 5529; Ofsowitz 29;
Palmer s.n.).
A putative link between leafy specimens of E. ramosa
subsp. aphylla and this subspecies has been mentioned
under that taxon. A further link may be the hairiness of
the sterile lemmas in subsp. rehmannii, which could be
an elaboration of their strongly scabrous condition
common in E. ramosa subsp. aphylla. The sterile lemma
hairs of subsp. rehmannii are short and appressed and
restricted to the nerves, margins and tips, thus differing
considerably from the longer spreading hairs on the sides
of the sterile lemmas in the E. calycina complex.
2b. Ehrharta rehmannii Stapf subsp. filiformis
(Stapf) Gibbs Russell, stat. nov. Type: Ecklon 914 (in part)
neotype here designated (specimen in K with thin culms
and small leaf blades!).
Ehrharta rehmannii Stapf wax. filiformis Stapf: 677 (1900). Chippindall:
39 (1955); Smook & Gibbs Russell: 55 (1985).
Ehrharta aphylla Schrad. war. filiformis Nees: 334 (1832). Nees: 207
(1841). Kleinriviersberge zwischen Zaunen, Ecklon.
Ehrharta filiformis Mez: 292 (1921). Type: Kapgebiet, Kleinrivier,
Ecklon & Zeyher 85B.
Tufted, erect or straggling, delicate or neither distinctly
delicate nor robust, often growing in dense masses. Culms
many to numerous, 120—800 mm long, 0,3— 0,5 mm
across, erect to decumbent or prostrate in smaller forms,
usually herbaceous or sometimes woody toward base of
culm, branching at upper nodes varies from seldom-
branched with single branches to fascicled with many
branches. Leaves borne all along the culms to just
= j '
14s 16» # To5 55s 54s 56°
FIGURE 4. — Distribution of E. rehmannii subsp. rehmannii.
194
Bothalia 19,2 (1989)
FIGURE 5. — Distribution of E. rehmannii subsp. filiformis.
Tufted erect, neither distinctly delicate nor robust.
Culms several, 300—600 mm long, to 2 mm across, erect
or decumbent at base, herbaceous, branching above, the
branches few and single. Leaf blades linear or lanceolate,
4-8,5 mm across and 30— 120(-170) mm long, flat or
rolled from both margins, erect, herbaceous, glabrous or
margins scabrous. Inflorescence a narrow contracted
raceme, 25 —70 mm long, somewhat overtopping leaves,
of 12—36 erect, appressed spikelets, usually closely
subtended or enveloped below by uppermost leaf sheath.
Spikelets 6—8,5 mm long. Pedicels erect. Glumes
subcoriaceous, nerves obscure; lower glume 6—8,5 mm
long; upper glume 5,5-8 mm long. Lower sterile lemma
glabrous on sides, smooth and unornamented or with
transverse corrugations; 0—3 ribs; 0—4 corrugations, faint;
tip truncate or rounded, rarely submucronate.
below the inflorescence. Leaf blades linear, usually small,
thin and soft, to 4 mm across but most leaves very fine,
15-100 mm long, spreading, sparsely glabrous. Inflores-
cence a reduced open raceme, 10-100 mm long, not or
only a little overtopping leaves, of 1 — 15( — 24) spreading
to nodding spikelets, exserted from uppermost leaf sheath.
Spikelets 4— 6,5(— 8) mm long. Pedicels spreading to
reflexed. Glumes membranous, 4—6,5 mm long, gaping
widely at maturity. Lower sterile lemma glabrous on sides,
with 2—3 fairly distinct to faint longitudinal ribs and rarely
with up to 6 faint transverse corrugations, tip rounded,
muticous. Lemma of fertile floret rarely with very faint
corrugations.
Subsp. filiformis is distinguished from the other two
subspecies by the often delicate habit, the small, soft, thin
leaf blades and the inflorescence with few spikelets borne
on spreading to reflexed pedicels. Its distribution is shown
in Figure 5. It grows on sandy (TMS) and humic soils,
at streamsides, in moist places, and in the shade of rocks,
from altitudes of 25—2 000 m, flowering from October
to February. It is most commonly collected after fire.
A few specimens occur that are intermediate between
subsp. filiformis and both other subspecies. Gibbs Russell
5674 is a typical subsp. filiformis and was collected within
its range, but the sterile lemma margins are hairy, a
characteristic of subsp. rehmannii. The intermediates to
subsp. subspicata are discussed under that subspecies.
As discussed above under E. ramosa subsp. aphylla, this
taxon must be typified by a specimen of Ecklon 914 with
thin culms, soft leaves and few spikelets. There is no
evidence that the Kew specimen of this description was
seen by Nees, so it is designated as a neotype rather than
a lectotype. A fragment of a since-destroyed specimen
from Berlin, allegedly annotated by Nees, exists in PRE
but it lacks a complete spikelet so therefore its identity
cannot be unequivocably determined. Mez (1921) quoted
Nees’s (1841 not 1832) epithet as the basionym when
raising the taxon to species rank, but he cited a different
specimen from those cited by Nees in either publication.
2c. Ehrharta rehmannii Stapf subsp. subspicata
(Stapf) Gibbs Russell, comb, et stat. nov. Type: stream
from Retreat to Muizenberg, Wolley Dod 3519 (K, lecto.!,
here designated; BOL, photo and fragment in PRE!).
Ehrharta subspicata Stapf: 676 (1900), Chippindall: 39 (1955).
Subsp. subspicata is distinguished from the other two
subspecies by the broad erect leaves and narrow inflores-
cences of erect spikelets with subcoriaceous glumes. Its
distribution is shown in Figure 6. It grows in sandy or
gravelly soil in moist places such as streamsides and
seepage areas, near sea level, flowering from October to
December.
Several specimens from limestone areas near the coast
appear to be intermediates between subsp. filiformis and
subsp. subspicata ( Acocks 24130', Ellis 1295, 4660', Gibbs
Russell 5644, 5646). They have the somewhat delicate
branched culms that are woody below and the small
spikelets of subsp. filiformis but the narrow inflorescence
and erect spikelets with subcoriaceous glumes of subsp.
subspicata. Their distribution is not in the area of sympatry
of the two subspecies, but lies to the east of the range of
subsp. filiformis and in the middle of the range of subsp.
subspicata (Figure 6). The limestone area where these
specimens were collected is noted for fts distinctive local
flora (Bond & Goldblatt 1984). However, because of the
general difficulty of delimiting taxa in the Ramosa group
and the paucity of specimens, no formal taxonomic ranking
is applied at this time to the entity represented by these
specimens.
A single specimen. Von Breitenbach 60, appears to be
intermediate between subsp. subspicata and subsp.
rehmannii. It has the erect leaves and narrow inflorescence
with erect spikelets of subsp. subspicata but the mem-
branous glumes and scabrous-keeled sterile lemmas of
subsp. rehmannii.
stone’ intermediates to subsp. filiformis are shown by open dots.
I:
■ 1
• ft
%
Bothalia 19,2 (1989)
195
LEAF BLADE ANATOMY
The leaf blade anatomy of the species of the Ramosa
group is distinctive and distinguishes this group from the
other seven species groups in Ehrharta (Gibbs Russell &
Ellis 1987). However, the leaf anatomy of the five taxa con-
stituting the Ramosa group exhibits no clear structural
disjunctions between the taxa themselves. A large
proportion of the specimens studied have anatomy
intermediate between adjacent taxa and it appears from
this voucher sample as if the taxa recognized only on
morphology reflect trends of specialization in this species
group. This is confirmed by the fact that a number of
herbarium specimens are intermediate between the taxa
as defined and a continuous and reticulate variation pattern
is also evident in external morphological characters, in
accord with the anatomical evidence.
The identifications of the anatomical voucher specimens
naturally ' also reflect this variation pattern with a
disproportionately large number of intermediates. This
situation is further exacerbated by taxa with a suffrutescent
habit, usually with caducous leaves which hamper leaf
anatomical studies. In these taxa only atypical
morphological specimens possessing leaf blades were
suitable for examination. The anatomical sample was,
therefore, not fully representative of all the taxa,
particularly as far as anatomical vouchers closely matching
type specimens is concerned.
For these reasons it is obvious that detailed anatomical
descriptions and diagnoses of individual taxa are
impractical, particularly as the taxa are barely separable
morphologically, being recognized as two species with five
subspecies. It has, therefore, been decided to give a full
anatomical description for the Ramosa group as a whole
followed by brief comments on the anatomical trends in
each of the constituent taxa.
Transverse section
Ribs and furrows may be absent (Figures 11E, F; 13A,
B; 14A-F; L5A, B) or slight ribs may be present over the
bundles (Figures 8B; 9B, E; 11A-D; 15E). Both
conditions appear to occur in all taxa. The mesophyll
generally accommodates a well developed, centrally
located bulliform cell group, the central cell of which may
be large and shield-shaped (Figures 11D, F; 13B; 14F).
The chlorenchyma is of the compact type with small to
medium, isodiametric or angular cells (Figures 9B, E; 11B,
D, F; 13B; 14B, D, F; 15B). These cells are as large as
or slightly larger than the parenchyma sheath cells with
the chloroplasts often peripherally located. The cell walls
are not straight but there is a definite tendency in all taxa
for some of the walls to have slight indentations,
particularly in the abaxially situated cells (Figures 9B, E;
11D, F; 13B). These resemble small invaginations of the
walls but do not appear to be homologous with arm cells.
They require further study but present technical difficulties
as perfect fixation is needed for their elucidation.
The chlorenchyma tissue sometimes displays a some-
what radiate arrangement in thicker leaves with 4-6
chlorenchyma cell layers (Figures 9B, E; 11D, F; 13B).
This pattern is not evident in thinner leaves of 2—3 cell
layers (Figures 14B, D, F; 15B). The thicker leaves tend
to have elongated, angular chlorenchyma cells whereas in
thinner leaves they are more equidimensional.
Abaxial epidermis
Costal and intercostal zones are always clearly differen-
tiated (Figures 8C, E; 12A, C; 13C; 14G; 15C, F). The
intercostal long cells are elongate-rectangular with the
anticlinal walls sinuous but not appearing inflated (Figures
8D, F; 9C; 12B, D; 13D). There is a tendency towards
a fusiform, hexagonal shape in a few specimens (Figure
14H) but this species group can be characterized by the
rectangular shape of the long cells and their very sinuous
walls (Figures 10A, C; 16A, C, E, F).
Successive long cells in a file are separated from one
another by single short cells, short cell pairs, hooks, micro-
hairs or stomata, and very seldom adjoin one another. This
arrangement along a given file alternates with that of
adjacent long cell files resulting in a distinct brickwork
pattern which is particularly conspicuous with the SEM
(Figure 16C, E). Cell size and shape is uniform across
the width of each intercostal zone.
Two files of stomata usually occur in each interstomatal
zone, the stomata in a file being separated by single
interstomatal long cells. The stomata are clearly dome-
shaped and their structure is consistent throughout the
group (Figures 8D; 12B, D; 13D; 14H). With the SEM
the stomata are seen to possess a distinct, raised rim
surrounding the pore aperture and they are not associated
with wax deposits (Figures 10B, D; 16D, F). Epicuticular
wax is also not typical of this group but may be present
as very fine platelets (Figure 10D).
Costal silica bodies are variable in shape, being of the
irregular dumbbell-shaped type (Figures 8D, F; 12B, D;
13 D). Costal zones are narrow, seldom exceeding three
cells in width.
Prickle hairs occur in all taxa but vary in form and
frequency of occurrence. They vary continuously between
those with very short barbs (Figures 8F; 9C; 13D; 15F)
to types with elongate barbs which resemble macrohairs
(Figures 8D; 12C; 15C). These latter types are unicellular
and do not have specialized epidermal cells associated with
their bases and, therefore, are not macrohairs by definition.
Both types may be either costal or intercostal although the
costal prickles on any given specimen are larger than the
intercostal hooks.
The leaf blade is always loosely inrolled from both
margins (Figures 8A; 9A; 11A, C; 13A; 14 A, C; 15A, D)
and there is no evidence of infolding. The margins
themselves range from rounded and blunt to tapering to
the pointed projection of Figure 15E. The midrib is
consistently a median vascular bundle only, without
associated parenchyma development (Figures 8A; 9A, D;
11A, C, E; 13A; 14A, C, E; 15A, D). Successive first order
bundles are separated by 2-5 smaller bundles but this
pattern or arrangement differs in different specimens of
the same taxon or even on opposite halves of the same
blade, as in Figure 15A where either two or four third order
bundles are located between the midrib and the first lateral
first order bundle on either side.
196
Bothalia 19,2 (1989)
FIGURE 7. — Culm anatomy of Ehrharta ramosa. A-C, E. ramosa subsp. ramosa, transverse section: A, culm (c) and surrounding leaf
sheath (s), x 100; B, detail of culm and leaf sheath vasculature and mesophyll tissue, x 250; C, culm epidermis showing stomatal
files (s) in the intercostal zones between successive sclerenchyma girders, x 250. A, B, Ellis 634\ C, Ellis 5525. D-F, E. ramosa
subsp. aphylla, Ellis 4634 , transverse section: D, culm outline, x 100; E, detail of mesophyll, vasculature and pith, x 250; F, epidermal
structure illustrating alternating costal and intercostal zones, the latter with the stomatal files (s), x 400.
Microhairs are common and with the aid of the SEM
are seen to be of a characteristic type (Figures 10B, D;
16B, D, F, H). The basal and distal cells are almost equal
in length. The distal cell is deciduous and has a blunt apex
which does not taper but the hair has a consistent width
throughout its length. This type of microhair occurs in all
taxa of this species group but nowhere else in Ehrharta.
This leaf anatomy is diagnostic for the Ramosa species
group and serves to distinguish it from the other species
groups in the genus. A definite anatomical trend is evident
within the group, from specimens with relatively thick leaf
blades, in which the chlorenchyma cells are rather
elongated with cell wall indentations and displaying a
somewhat radiate arrangement, to very thin leaves. These
have isodiametric chlorenchyma cells without wall
indentations or radiate-type arrangement. This continuum
is paralleled by a trend for the margins to become more
tapering and for the epidermal prickles to become smaller
and fewer in number. However, these anatomical trends
do not appear to be reflected in the classification of this
species group and are assumed to be of no taxonomic
relevance.
E. ramosa
Leaf blades in this species are absent or minute and,
if present, are short-lived and caducous. As a result very
few specimens possess leaf blade material and, conse-
quently, are unsuitable for comparative anatomical study
Bothalia 19,2 (1989)
197
of the leaf blade. All specimens with typical morphology
were devoid of leaf blades and only culms were available
for study. The results are briefly described below and
illustrated in Figure 7.
The culm may be encircled by a leaf sheath which has
overlapping margins (Figure 7A), a very thick cuticle,
stomata, vascular bundles and chlorenchyma only in the
outer cell layers (Figure 7B). Often the bladeless sheaths
are also dehiscent and absent (Figure 7D).
The culm outline is circular (Figure 7A, D) with a
simple epidermis. The stomata are adjacent to the
chlorenchyma zones with the subsidiary cells being flush
with the epidermal cells (Figure 7B). No prickles or other
epidermal appendages are present.
A discontinuous ring of chlorenchyma is present beneath
the epidermis. Each chlorenchyma zone consists of 5— 8
radially arranged columns of tabular cells, 2-3 cells deep
(Figure 7B, E). These zones are separated by the scleren-
chyma girders of the peripheral vascular bundles and are
bounded internally by a continuous ring of sclerenchyma
and collenchyma in which the peripheral vascular bundles
are embedded (Figure 7B, E). This sclerenchymatous ring
encloses the central parenchymatous pith which may or
may not contain vascular bundles near the periphery
(Figure 7A, D).
The vascular bundles in the culms of E. ramosa subsp.
ramosa are arranged in two rings, a ring situated in the
pith tissue and a ring embedded in the sclerenchymatous
ring (Figure 7A). In subsp. aphylla there is only a single
ring associated with the sclerenchyma (Figure 7D). This
FIGURE 8.— Leaf blade anatomy of Ehrharta ramosa subsp. ramosa. A, B, transectional anatomy: A, inrolled outline of leaf blade with
undifferentiated keel (k), x 100; B, anatomical detail with many prickles (p) in both epidermides, x 250. C-F, abaxial epidermis:
C, cell arrangement, x 160; D, detail of costal and intercostal prickle hairs (p), intercostal long cells and stomata, x 400; E, cell
arrangement and numerous prickle hair bases, x 160; F, detail of costal and intercostal prickles (p), long cells, silica bodies and stomata,
x 400. A-D. Ellis 4694; E, F, Ellis 1632.
198
Bothalia 19,2 (1989)
FIGURE 9. — Leaf blade anatomy of Ehrharta ramosa subsp. aphylla. A, B, leaf transverse section: A, loosely inrolled leaf outline and
absence of keel (k), x 100; B, chlorenchyma and vascular bundle detail; note slight invaginations of chlorenchyma cell walls and
prominent abaxial prickle hair base (p), x 400; C, abaxial epidermis showing rectangular, sinuous long cells and large costal prickle
hairs, x 400. D, E, leaf in transverse section: D, outline with median vascular bundle only (k), x 100; E, chlorenchyma detail, x
400. F, abaxial epidermis showing costal prickles and intercostal hooks, x 250. A-C, Ellis 2235; D-F, Ellis 4673.
is a reflection of the difference in size and robustness of
these subspecies. The vascular bundles have a clearly
developed mestome sheath but the parenchyma sheath is
not clear and appears to be replaced by the parenchyma
or sclerenchyma ground tissue (Figure 7B, E). This was
also noted by De Wet (1960).
The culm epidermis reflects this internal anatomy with
alternating bands of stomatal files overlying the
chlorenchyma zones, and thickened epidermal cells overly
the sclerenchyma girders (Figure 1C, F). The form of the
costal cells overlying the girders appears to differ between
E. ramosa subsp. ramosa (Figure 1C) and subsp. aphylla
(Figure 7F) and this may be a taxonomically important
difference between these two taxa. However, a larger study
sample is needed before this can be substantiated.
E. ramosa subsp. ramosa
Two anatomical voucher specimens (Ellis 1632, 4694)
have been identified as belonging to this taxon although
they both deviate from the taxon diagnosis in possessing
leaf blades. They are both suffrutescent and woody but
also possess leaf blades and have erect pedicels. They,
therefore, resemble E. rehmannii subsp. subspicata in the
latter two characters but this taxon is not generally
suffrutescent. These specimens, therefore, are not typical
subsp. ramosa specimens but were the only plants found
which also possessed leaves and, consequently were
suitable for this leaf anatomy study. They appear to
represent very young flowering plants with recent
vegetative growth, perhaps in response to physical damage.
Nevertheless, they do not represent typical examples of
this subspecies and the anatomical results must be
evaluated accordingly.
The anatomy of subsp. ramosa conforms closely to that
given for the Ramosa species group (Figure 8). The leaf
in transection exhibits all the characteristic features of this
species group and even slight cell wall invaginations are
Bothalia 19,2 (1989)
199
visible (Figure 8B). The numerous prickle hair bases
evident in the epidermis are a notable feature (Figure 8B).
The abaxial epidermis is dominated by the large number
of prickle hairs present (Figure 8C-F), both in the costal
and intercostal zones. These may be either typical prickles
with short barbs (Figure 8F) or macrohair-like with much
longer barbs (Figure 8D). The abundance of these prickle
hairs appears to be a feature of this species with reduction
in their size and number in several specimens of E.
rehmannii. The silica bodies in subsp. ramosa are
elongated dumbbell-shaped and tend to be shorter than in
E. rehmannii. The microhairs are typical of those so
diagnostic of the Ramosa species group (Figure 10B).
Specimens examined
CAPE. — 3319 (Worcester): Hex River Mts, Milner Peak (—AD), Ellis
5525 (culm only). 3321 (Ladismith): Langeberge, Cloete's Pass (-DC),
Ellis 634 (culm only), Ellis 4694 (leaves with reduced blades). 3323
(Willowmore): Uniondale Dist., Uniondale Poort (— CA), Ellis 1632 (short
leaf blades).
E. ramosa subsp. aphylla
Typical material of this taxon is also without leaf blades,
as in Ellis 4634. The voucher material which was assigned
to this taxon once again presented problems with identifi-
cation as all plants found with leaf blades were either
sterile or only had very young inflorescences. Positive
identification was, therefore, not possible because the
position of the pedicels (whether reflexed or not) was not
yet visible.
The anatomy of Ellis 2235 is virtually identical to that
of subsp. ramosa, both in transection (Figure 9A, B) and
the abaxial epidermis (Figure 9C). The large prickle hair
bases enchance this resemblance as do the slight cell wall
indentations (Figure 9B).
Another specimen, Ellis 4673, which is intermediate to
E. rehmannii subsp. rehmannii in spikelet characters,
resembles the anatomy of E. rehmannii subsp. rehmannii
more closely than that of subsp. ramosa. This is
particularly evident in the adaxial ribs and furrows in the
leaf section (Figure 9E) which generally is very similar
to that of Ellis 4697 and 4699 (Figure 11B, D), E.
rehmannii subsp. rehmannii. However, the epidermal
features of this specimen do not corroborate this
resemblance with E. rehmannii but, then, neither do they
suggest affinities with other E. ramosa specimens.
Although the very small hooks are unusual, the micro-
FIGURE 10. — Scanning electron micrographs of the abaxial epidermis of Ehrharta ramosa. A, B, E. ramosa subsp. ramosa , Ellis 4694 :
A, intercostal prickle hairs, stomata and rectangular long cells, x 200; B, microhair with distal cell not tapering and stoma (s) with
distinct rim and without associated wax deposits, x 1 000. C, D, E. ramosa subsp. aphylla, Ellis 4673: C, inflated, rectangular inter-
costal long cells, stomata and hooks; costal zones with prickles and silica bodies, x 200; D, microhair with exuding distal cell and
stoma with distinct rim, x 1 000.
200
Bothalia 19,2 (1989)
FIGURE 11. — Transectional leaf blade anatomy of Ehrharta rehmannii subsp. rehmannii. A, B, Ellis 4699: A, inrolled outline without keel
(k) and associated parenchyma, x 100; B, vascular bundle and mesophyll arrangement and chlorenchyma cell detail, x 400. C, D,
Ellis 4697: C, outline with undifferentiated keel (k) and tapering margin, x 100; D, chlorenchyma cell detail showing slight cell wall
invaginations, x 400. E, F, Ellis 618: E, outline showing median vascular bundle without associated parenchyma tissue (k), x 100;
F, compact chlorenchyma cells with slight wall indentations, x 400.
hairs and stomata (Figure 10D) are typical of those of the
Ramosa group.
Specimens examined
CAPE. — 3219 (Wuppertal): Clanwilliam Dist., Pakhuis Pass (-AA),
Ellis 4634 (culm only). 3318 (Cape Town): Jonkershoek State Forest,
Jakkalsvlei (-DD), Ellis 2235 (reduced blades present). 3418
(Simonstown): Hottentot Hollands Mts, Sir Lowry’s Pass (— BB), Ellis
4673 (few, reduced blades).
E. rehmannii
Although this species is characterized by herbaceous
culms and blade-bearing leaves, several anatomical
voucher specimens have suffrutescent culms and reduced,
caducous blades. Examples are Ellis 648 which is
morphologically intermediate between subsp. rehmannii
and E ramosa subsp. aphylla and Ellis 1295 and 4660
which are morphologically intermediate between subsp.
filiformis and subsp. suhspicata. However, no anatomical
intermediacy is evident in these specimens, making their
interpretation difficult.
No clear anatomical discontinuities exist between any
of the three subspecies of E. rehmannii or with E. ramosa.
Instead there is a general trend from E. ramosa through
E. rehmannii subsp. rehmannii to subsp. filiformis along
which leaf thickness decreases, the margins tend to become
more tapered and the prickles become smaller and fewer
in number. Several exceptions to this pattern exist, so the
trend may represent an ecological gradient rather than a
phylogenetic lineage.
E. rehmannii subsp. rehmannii
All specimens have relatively thick mesophyll tissue
displaying a tendency to the radiate condition (Figure 11B,
D, F). The chlorenchyma cells are not isodiametric but
more elongated with slight cell wall indentations. The leaf
margin is more acute than in E. ramosa and may taper
to a definite point (Figure 11C). The adaxia! surface may
be flat (Figure 11F) or ribbed (Figure 11B, D).
The epidermal structure is remarkably uniform through-
out the sample (Figure 12 A— D) with rectangular long cells
being conspicuous. Only Ellis 4697 (Figure 12C, D) has
Bothalia 19,2 (1989)
201
a few intercostal hooks, these with very long barbs. All
have a few costal prickles with short barbs. The silica
bodies are variable in shape but generally are shorter
dumbbell-shaped than in E. ramosa.
Ellis 4697 is noteworthy in that it differs in having a
tapering margin (Figure 11C), has the thickest leaf with
definite ribs (Figure 11D) and possesses intercostal prickles
(Figure 12C). This deserves further comment as it was
collected together with Ellis 4698 and 4699 from an
actively growing and spreading population recovering after
a fire. The latter two specimens differ from Ellis 4697 in
having more abrupt margins, thinner blades (Figure 11A,
B) and no prickles (Figure 12A). Thus, within a single
population, considerable anatomical variation is evident,
even in those characters which show a trend through this
species group. This supports the decision not to attach any
taxonomic importance to these anatomical differences.
Although anatomical differences may, therefore, not be
reflected in the morphology, the reverse is also true. Ellis
648 is morphologically intermediate between E. rehmannii
subsp. rehmannii and E. ramosa subsp. aphylla but
anatomically virtually identical to Ellis 618, subsp.
rehmannii. Reference to Figures HE, F; 12B and 13A— D
illustrate this resemblance clearly, both specimens with
distinctive thick, compact mesophyll and well developed
bulliform groups, without adaxial ribs and without large
prickles in the epidermis. These two specimens are more
alike in leaf anatomy than they are to all the other
specimens in the Ramosa group. It therefore appears that
there is little or no congruence between leaf anatomy and
morphology in this species group, a situation which differs
from that in the other groups of Ehrharta.
Specimens examined
CAPE. — 3320 (Montagu): Barrydale Dist., Tradouw’s Pass (—DC),
Ellis 648 (intermediate between subsp. rehmannii and E. ramosa subsp.
aphylla). 3322 (Oudtshoom): Outeniqua Mts, Robinson’s Pass (-CC),
Ellis 4697, 4698 and 4699. 3323 (Willowmore): Groot River Pass between
Plettenberg Bay and Storms River (—DC), Ellis 618.
E. rehmannii subsp. filiformis
This taxon is placed at one end of the continuum of
anatomical variation and has the thinnest leaves, the most
tapering margins and few, small prickles. The leaf blades
are thin and inrolled from both margins (Figure 14A, C,
E). The chlorenchyma beneath the bulliform cells is only
two to four cells deep and no radiate tendency is evident
(Figure 14B, D, F). The margins are also distinctly tapering
(Figure 14A, C, E). The silica bodies are short dumbbell-
shaped and the prickles are usually few in number and
small and hook-like (Figure 14G, H). The stomata and
microhairs are typical of those of the Ramosa species
group (Figure 16H). The intercostal long cells are
distinctly hexagonal in outline (Figure 14G, H), a feature
not apparent in any other specimens.
This anatomy is consistent with the small, soft, thin leaf
blades of this subspecies and the leaf anatomy is uniform
throughout the study sample. The anatomy is also fairly
FIGURE 12. — Abaxial leaf blade epidermis of Ehrharta rehmannii subsp. rehmannii. A, Ellis 4699, with sinuous, rectangular intercostal
long cells, stomata and costal prickles, x 250. B, Ellis 618, illustrating costal and intercostal cell detail and arrangement, x 400. C,
D, Ellis 4697. C, showing costal and intercostal zone configuration and elongated prickle hairs, x 250; D, detail showing costal prickles
and silica bodies and intercostal long cells, stomata and hooks, x 400.
202
Bothalia 19,2 (1989)
FIGURE 13, — Leaf anatomy of specimen morphologicallly intermediate between Ehrharta rehmannii subsp. rehmannii and E. ramosa subsp.
aphylla, Ellis 648. A, B, leaf transverse sections: A, inrolled outline and median bundle without asociated parenchyma tissue, x 100;
B, detail of chlorenchyma and vascular bundles, x 400. C, D, abaxial epidermis: C, epidermal zonation, x 250; D, elongated, rectangular
intercostal long cells, dome-shaped stomatal subsidiaries, irregular silica bodies and costal prickles, x 400.
distinct from that of the other taxa in the Ramosa species
group and subsp. filiformis appears to represent a more
clear-cut entity than do the other taxa of the group.
Once again, intermediate specimens tend to confuse
these patterns, in this case specimens intermediate between
subsp. filiformis and subsp. subspicata from the limestones
of the De Hoop area. Ellis 1295 and 4660 both diverge
considerably in both morphology and leaf anatomy from
the rest of the sample representing this taxon (Figure
15A— F). The leaf blade of Ellis 1295 has a tapering margin
(Figure 15A), is very thin (Figure 15B), has hexagonal long
cells, elongated dumbbell-shaped silica bodies and very
numerous, conspicuous macrohair-like prickles (Figure
15C). This anatomy is distinct and does not closely
resemble any other Ramosa species group specimen. Ellis
4660 also has very distinctive anatomy, the leaf being very
narrow (Figure 15D) with the tapering margins exaggerated
into a pointed projection (Figure 15E). The abaxial
epidermis of this specimen does not have macrohair-like
prickles but has numerous normal prickles with short
barbs (Figure 15F). Both these specimens differ greatly
from all the others examined and also differ considerably
from one another although they come from virtually the
same locality.
These intermediate specimens from this specialized
limestone habitat are abnormal in that they are woody and
suffrutescent with very few leaves but they are small plants
and their spikelets are similar in form to subsp. subspicata
but fall within the size range of subsp. filiformis. They
are retained in subsp. filiformis even though their leaf
anatomy does not correspond closely to that of either
subsp. filiformis or subsp. subspicata.
Specimens examined
CAPE. — 3318 (Cape Town): Jonkershoek (-DD), Ellis 4676. 3418
(Simonstown): Hottentot Hollands Mts, Sir Lowry’s Pass (— BB), Ellis
2288, 2289. 3419 (Caledon): Hermanus Dist., Olifantsberg (-AD), Ellis
4671. 3420 (Bredasdorp): between Malgas and Wydgelegen (—AD), Ellis
1295, 4660 (intermediate to subsp. subspicata).
E. rehmannii subsp. subspicata
No authentic material of this subspecies was freshly
collected in the field for this study. Consequently,
comparative studies were not possible but herbarium
material ( Taylor 7667) was examined in order to gain an
understanding of the anatomy of this taxon. Indications
are that it belongs closest to E. ramosa along the trend
which is evident in this species group. The leaf appears
to be rather thick with abrupt margins but the epidermis
is without numerous well developed prickles. This must
be confirmed from freshly fixed material.
DISCUSSION AND CONCLUSIONS
The spikelet morphology and leaf anatomy of the five
taxa of the Ramosa group is diagnostic and defines them
as a group separate from the other species groups in
Ehrharta. No spikelet characters are unique to the Ramosa
group alone, but all taxa in the Ramosa group may be dis-
tinguished by the following combination of characters:
small spikelets less than 9 mm long, with the sterile lem-
mas similar in shape and size and about as long as the
fertile lemma, and having the bases appendaged and usual-
ly bearded, the sides glabrous, scabrous or shortly hairy
and the tips rounded, truncate or mucronate. In addition,
all species are perennial and may be suffrutescent, leaf
Bothalia 19,2 (1989)
203
blades may be absent, glumes are two-thirds as long to
longer than the lemmas and the upper sterile lemma is
not stipitate.
Anatomically, the Ramosa group taxa share the follow-
ing unique characters: mesophyll cells compact with slight
cell wall invaginations, stomata dome-shaped with a rim
surrounding the aperture and lacking associated wax
deposits, and the microhairs with the distal cells not
tapering. Other characters which together differentiate the
group include: leaf blades (when present) inrolled, midrib
lacking parenchyma, ribs and furrows absent or only
slightly developed, costal and intercostal zones differen-
tiated, intercostal long cells rectangular with sinuous walls,
silica bodies tending to dumbbell shape; prickle hairs
always present, varying from small and hook-like with
short barbs to macrohair-like with elongated barbs, and
epicuticular wax usually absent. Characters which are
particularly consistent throughout the group are inrolled
leaf blades and the midrib lacking parenchyma. Characters
of ribs, chlorenchyma, silica bodies and prickle hairs tend
to be more variable.
Ik i ^
FIGURE 14. — Leaf blade anatomy of Ehrharta rehmannii subsp . filiformis. A— F, transectional anatomy: A, inrolled blade with tapering
margins and median bundle only, x 100; B, chlorenchyma and vascular bundle structural detail and adaxial prickles, x 400; C, loosely
inrolled outline without keel and with tapering margins, x 100; D, vascular bundle and mesophyll structure and arrangement, x 400;
E, leaf blade outline showing inrolled margin and median bundle only, x 100; F, thin mesophyll layer but cells with slight wall
indentations, x 400, interference contrast. G-H, abaxial epidermis: G, showing epidermal zonation and prickle hairs, x 160; H, with
fusiform long cells, stomata and costal prickles, x 400. A, B, Ellis 2288; C, D, Ellis 2289; E, F, H, Ellis 4676; G, Ellis 4671.
204
Bothalia 19,2 (1989)
FIGURE 15. — Leaf anatomy of specimens intermediate btween Ehrharta rehmannii subsp. filiformis and subsp. subspicata. A— C, Ellis 1295 :
A, leaf outline showing thin inrolled lamina with prominent prickles and without a keel, x 100; B, detail of abaxial and adaxial
macro hair-like prickles and few chlorenchyma cell layers, x 400, interference contrast; C, costal and intercostal prickles which resemble
macrohairs in surface view, x 160. D— F, Ellis 4660 : D, outline showing narrow blade with undifferentiated keel and markedly tapering
margins, x 100; E, detail of narrow projecting margin, x 250; F, abaxial epidermis showing elongated, rectangular long cells, and
costal and intercostal prickles with short barbs, x 250.
The Ramosa group is most closely related to the
Calycina group for both morphological and anatomical
features. Auriculate appendages at the base of the second
sterile lemma occur only in these two groups. Also, the
truncate or tapering mucronate sterile lemma tips of both
the E. ramosa subspecies are similar to those of E.
calycina, and the hairs on the sides of the sterile lemmas
in most specimens of E. rehmannii subsp. rehmannii tend
toward the longer hairs on the lemma sides of all members
of the Calycina group. The anatomical relationship
between the two species groups is not so marked. The long
cells of the Ramosa group are usually rectangular, but a
few specimens of E. rehmannii have fusiform long cells,
an elongated hexagonal shape which is characteristic of
the Calycina group.
Other characters show apparent similarities between the
Ramosa group and other species groups, but closer ex-
amination shows that they are not closely related.
Suffrutescent, bladeless species occur in both the Ramosa
and Villosa groups: E. ramosa is similar in habit to E.
thunbergii and E. villosa in the Villosa species group.
While this might indicate a relationship between the
groups, it is also possible that the bladeless suffrutescent
condition, with photosynthetic culms that persist from year
to year, is an adaptation to conserve plant parts in the low
nutrient soils in the Mountain Fynbos and seaside dunes
where these species occur.
The indentations in the walls of the chlorenchyma cells
present in many of the specimens in the Ramosa group
may be taken to resemble the invaginations characteristic
of arm cells. Engelbrecht (1956) reported arm cell-like in-
vaginations in representatives of the Ramosa group ( E .
ramosa, E. rehmannii and E. subspicata ) based on prepa-
rations from herbarium material. However, the present
study does not confirm Engelbrecht’s observations because
closer examination of well preserved young leaf material
shows that these structures are not inward projections of
cell walls but merely the wavy oudines of these walls.
Bothalia 19,2 (1989)
205
FIGURE 16. — Abaxial epidermal ultrastructure of Ehrharta rehmarmii. A— F, E. rehmannii subsp. rehmannii'. A, rectangular long cells, stomata
with rims and dumbbell-shaped silica bodies, x 200; B, detail of bicellular microhair without tapering distal cell, x 1 000; C, elongated,
rectangular long cells and costal prickles, x 200; D, microhair with distal cell not acutely tapering, x 1 000; E, rectangular long cells
with brickwork type of arrangement and small costal prickles, x 200; F, microhair with intact distal cell and stoma with distinct rim
(s), x 1 000. E. rehmannii subsp. filifonnis: G, long cells elongate rectangular with thick, sinuous anticlinal walls, stomata with rims
and prickles with short barbs, x 200; H, detail of microhair with collapsed but truncated distal cell and thickened, sinuous anticlinal
walls of long cells, x 1 000. A. B, Ellis 4697 ; C, D, Ellis 4698\ E, F, Ellis 4699, G, H, Ellis 4671.
206
Bothalia 19,2 (1989)
Nevertheless, these loose folds of the walls are unique to
and diagnostic for the Ramosa group. They do not appear
to be homologous with arm cells and therefore do not
indicate affinities with the Setacea group or with the Bam-
busoideae.
Individual specimens from the far north-western
distribution limit of the Ramosa Group also show inter-
group relationships. A population from Van Rhyn’s Pass
in the Bokkeveld Mountains shows the spikelet shape and
hairiness of E. calycina, but the plants are suffrutescent,
growing in bushy clumps similar to E. rehmannii ( Ellis
1147, 4625, Gibbs Russell 5588). The anatomy of these
specimens resembles that of the Ramosa group, with
inrolled blades, absence of a midrib and ribs, compact
mesophyll and numerous prickles. However, the stomata
are not of the Ramosa type and the silica bodies are not
dumbbell-shaped. The microhairs appear to be
intermediate, being narrower than the Ramosa type but
not tapering to a point as in the Calycina group. Long cell
shape is variable, with some specimens having rectangular
Ramosa-type cells and others showing fusiform Calycina-
type cells. Specimens of this population appear to establish
a clear morphological and anatomical link between these
two species groups and possibly represent a hybrid
between them.
intermediates for both morphological or anatomical
characters have been recorded (Figure 17); second, some
of the most useful characters for separating taxa depend
on character states that may vary during the life cycle of
an individual. It is particularly difficult to distinguish an
open inflorescence from a contracted one early in the
flowering season, and it appears from field observations
that plants which become suffrutescent and bladeless when
mature may be herbaceous and conspicuously leafy when
young. Long-term phenological studies related to fire
frequency are necessary to understand the true relation-
ship between the infraspecific taxa in the Ramosa group.
ACKNOWLEDGEMENTS
The following herbaria are thanked for the now long-
standing loans of Ehrharta specimens: BOL, J, JF, NBG
& SAM, STE. Specimens from K, P and LE were
examined at Kew. We thank the Keeper, Mr G. Lucas, and
Deputy Keeper, Dr W.D. Clayton, for visitor’s facilities
and especially for the generous gift of many colour
photographs of specimens; the Director of B for visitor’s
facilities; A. Romanowski for photography, S. Perold for
scanning electron microscopy, G. Condy for the spikelet
drawing and H. Ebertson, A. Botha and W. Roux for
technical assistance.
A single specimen (Ellis 5511) from Sneeukop Peak in
the Cedarberg area probably represents a new species. It
has the open inflorescence, long glumes and appendaged
mucronate glabrous sterile lemmas of E. ramosa subsp.
aphylla, but its culms are herbaceous and the base of the
plant is extraordinarily leafy, with long erect blades similar
in number and position to those of E. dura , but with leaf
anatomy similar to E. capensis. Its correct species group
is therefore presently unknown.
Apart from this single specimen, the Ramosa group
shows affinities only with the Calycina group, otherwise
occupying an isolated position within the genus. The
anatomy of the group is distinct, with several unique
features.
The imprecision of defining the taxa in the Ramosa
group is in contrast to other species groups (e.g. Villosa,
Dura, Capensis) where the species and infraspecific taxa
are more easily delineated. There are two sources of doubt
in identifying specimens in the Ramosa group: first, many
ramosa subsp. m ramosa subsp.
aphylla ramosa
FIGURE 17. —The occurrence of intermediates between subspecies in
the Ramosa group is shown by a line between taxa. The symbol
‘m’ indicates morphological intermediates and the symbol ‘a’
indicates anatomical intermediates.
REFERENCES
ADAMSON, R.S. 1942. Some changes in nomenclature. III. Journal
of South African Botany 8 : 273.
BOND, P. & GOLDBLATT, P. 1984. Plants of the Cape Flora. Journal
of South African Botany Supplementary Volume 13.
CHIPPINDALL, L. 1955. A guide to the identification of grasses in
South Africa. In C. Meredith, The grasses and pastures of South
Africa. Central News Agency, Cape Town.
DALLWITZ, M.J. 1984. User’s guide to the DELTA system— a general
system for coding taxonomic descriptions. CSIRO Division of
Entomology Report No. 13.
DE WET, J.M.J. 1960. Culm anatomy in relation to anatomy. Bothalia
7: 311-316.
ELLIS, R.P. 1987a. Leaf anatomy of the genus Ehrharta (Poaceae) in
southern Africa: the Setacea group. Bothalia 17: 75 — 89.
ELLIS, R.P. 1987b. Leaf anatomy of the genus Ehrharta (Poaceae) in
southern Africa: the Villosa group. Bothalia 17: 195—204.
ENGELBRECHT, A. H P. 1956. n Morfologiese studie van die genus
Ehrharta Thunb. M.Sc. thesis. University of Pretoria.
GIBBS RUSSELL, G.E. 1987a. Taxonomy of the genus Ehrharta
(Poaceae) in southern Africa: the Setacea group. Bothalia 17:
67-73.
GIBBS RUSSELL, G.E. 1987b. Taxonomy of the genus Ehrharta
(Poaceae) in southern Africa: the Villosa group. Bothalia 17:
191-194.
GIBBS RUSSELL, G.E. & ELLIS, R.P. 1987. Species groups in the
genus Ehrharta (Poaceae) in southern Africa. Bothalia 17: 51-65.
GIBBS RUSSELL, G.E. & ELLIS, R.P. 1988. Taxonomy and leaf
anatomy of the genus Ehrharta (Poaceae) in southern Africa: the
Dura group. Bothalia 18: 165—171.
LINDER, H.P. 1985. Conspectus of the African species of Restionaceae.
Bothalia 15: 387-503.
NEES AB ESENBECK, G.C. 1832. Plantae ecklonianae. Linnaea 7:
273 - 339.
NEES AB ESENBECK, G.C. 1841. Florae africae australioris illustra-
tiones monographicae . Prausnitzianus, Cracow.
MEZ, C. 1921. Gramineae novae vel minus cognitae, IV. Repertorium
novarum specierum regni vegetabilis 17: 291—303.
SCHRADER, H.A. 1821. Analecta ad floram capensem Sect. II,
Cyperaceae, Gramineae. Gottingische gelehrte Anzeigen 1821:
2065-2079.
SCHRADER, H.A. 1830. Systema vegetabilium. Vol. 7, Part 2, J.A.
Schultes (ed.). Cottae, Stuttgart.
SMOOK, L. 8 GIBBS RUSSELL, G.E. 1985. Poaceae. In G.E. Gibbs
Russell et al.. List of species of southern African plants, edn
2, part 1: 45-70. Memoirs of the Botanical Survey of South Africa
No. 51.
Bothalia 19,2 (1989)
207
STAPF, O. 1897. Ehrharta rehmannii. In Diagnoses africanae. Kew
Bulletin 1897: 288.
STAPF, O. 1900. Gramineae. In W.T. Thiselton-Dyer, Flora capensis
7: 310—750. Reeve, London.
STEUDEL, E.G. 1855. Synopsis plantarum graminearum. Metzler,
Stuttgart.
SWARTZ, O. 1802. The botanical history of the genus Ehrharta.
Transactions of the Linnean Society 6: 40—64.
THUNBERG, C.P. 1794. Prodromus plantarum capensium. Edman,
Uppsala.
THUNBERG, C.P. 1818. Flora capensis Vol. 2. Hafniae [Copenhagen],
Gerhardum Boniacum.
THUNBERG, C.P. 1823. Flora capensis 2nd edn. J.A. Schultes (ed.).
Cottae, Stuttgart.
TRINIUS, C.B. 1839. Phalaridea. Memoires de l Academie imperial des
Sciences de St-Petersbourg, ser. 6,5: 12—26 (of reprint).
SPECIMENS EXAMINED
Acocks 19908 (la) K, PRE; 21178 (2c) PRE; 22484 (2b) K, PRE; 24130
(2b/2c) K, PRE; Acocks & Hafstrom 13.11.1938 (lb) PRE. Adamson 597
(lb) PRE; 675 (2b) PRE; 1250 (lb) BOL; 1253 (2b) BOL; 2427 (2b)
BOL; 2696 (2c) BOL; 2734 (lb) BOL; 3701 (2c) BOL, PRE; 3969 (lb)
JF. Andreae 30 (lb) PRE; 705 (la/lb) PRE; 1298 (la) PRE. Archibald
3480 (la) BOL, PRE.
Balkema 48 (lb) STE. Barnard (la) SAM (No. 28386). F. Bolus Nov.
1913 (lb) PRE; Oct. 1913 (2b) NBG; Jan. 1913 (2b) PRE; Nov. 1913 (2b)
K, PRE; Dec. 1913 (2b) PRE; Dec. 1915 (2b) BOL; Jan. 1915 (2b) BOL,
K. H. Bolus 4733 (lb) BOL; 14670 (lb) K; 14740 (2) K. Bond 1613 (2a)
PRE; s.n. (2a) NBG. BoshoffP302 (la) STE. Boucher 1456 (2b) JF, STE;
1661 (lb) JF, PRE, STE; 1724 (lb) PRE, STE; 2380 (lb) PRE, STE;
2646 (2b) STE; 4202 (la) PRE, STE. Burchell 543 (lb) K, PRE; 4648
(la) K; 5974 (2) K; 6712 (2) K, PRE (fragment). 7011 (la) K; 7312 (la)
K; 7725 (lb) K, PRE (photo and fragment). Burtt Davy 15121 (la) K.
Carmichael s.n. (lb) PRE. Cleghom 2020(26) PRE; 2491 (2c) K, PRE;
2498 (2) K; 3103 (2c) PRE, STE; 3143 (2b) K, PRE, STE; 3164 (lb)
STE, PRE. Compton 8245 (2b) NBG; 12870 (la) BOL, NBG; 16962
(2b) NBG; 18595 (lb) NBG; 20228 (lb) NBG; 23076 (2a) K, NBG. Crook
1021 (lb) K, BOL, NBG, PRE; 1035 (2b) BOL, K, NBG, PRE.
Dahlstrand 2970 (la) J, PRE, STE. Durand 299 (la) JF. Drege (la) SAM
(No. 19639); Du Toit’s Kloof (la) K; Giftberg (lb) K; Paarl Mts (lb) K;
Table Mtn (lb) K; s.n. (la) P. Du Toit 1345 (lb) PRE, STE; 1425 (lb/2a)
PRE, STE; 1747 (la/2a) STE.
Ecklon 914 (in part) (lb) K; 914 (in part) (2b) K; s.n. (la) P. Ecklon &
Zeyher s.n. (la) B. Ellis 618 (2a) PRE; 634 (la) PRE; 648 (lb/2a) PRE;
1295 (2b/2c) PRE; 1632 (la) PRE; 2235 (lb) PRE; 2288 (2b) PRE; 2289
(2b) PRE; 4634 (lb) PRE; 4660 (2b/2c) PRE; 4671 (2b) PRE; 4673 (lb)
PRE; 4676 (2b) PRE; 4694 (la) PRE; 4697 (2a) PRE; 4698 (2a) PRE;
4699 (2a) PRE; 5511 (?) PRE; 5525 (la) PRE. Esterhuysen 580 (2b) PRE;
632 (lb) NBG; 3930 (la/lb) BOL; 6690 (la) K, BOL, PRE; 6751 (2a)
BOL, K, NBG, PRE; 8343 (la) K, BOL, NBG, PRE; 8605 (lb) BOL;
9840 (la) BOL; 10607 (la) BOL, PRE; 10815 (la) PRE; 11317 (la) PRE.
11785 (lb) PRE; U786 (2b) PRE; 11787 ( 2b) BOL; 12348 (lb) BOL, PRE;
13505 (lb) BOL, PRE; 13581 (2a) BOL, NBG, PRE; 14978 (la/lb) NBG;
27314 (2a) BOL, PRE; 27426 (la) PRE; 280U (la) BOL, PRE; 28U0
(lb/2b) BOL, PRE; 28396 (lb/2b) BOL, NBG, PRE; 28651 (2b) BOL,
PRE; 32217 (la/lb) BOL; 32499 (lb) BOL; 33393 (2b) BOL, PRE; 33720
(2c) BOL, K, PRE; 34075 (2c) BOL; 34468 (2c) BOL; 35541 (2b) BOL,
K; 35856 (la) BOL.
Fairale 284 (2c) PRE; 286 (lb) PRE; 297 (2c) PRE. Forsyth 354 (lb)
JF. Fourcade 921 (2a) BOL; 1589 (2a) BOL; 2478 (la) BOL, PRE; 2815
(2a) BOL, K, PRE, STE; 4259 (la) K, STE; 5529 (2a) STE; 6496 (la)
BOL, STE; s.n. STE. Fries, Norlindh & Weimarck 611 (la) PRE.
Galpin 149 (la) STE; 165 (la) K, PRE, STE. Geldenhuys 286 (2a) PRE.
Gericke 1881950 (2c) PRE; s.n. (2a) PRE. Gibbs Russell 5602 (lb) PRE;
5633 (la) PRE; 5644 (2b/2c) PRE; 5646 (2b/2c) PRE; 5651 (2b) PRE;
5654 (2b) PRE; 5657 (lb) PRE; 5659 (2b) PRE; 5660 (lb) PRE; 5662
(lb) PRE; 5664 (2b) PRE; 5668 (lb/2a) PRE; 5672 (lb/2a) PRE; 5674
(la/lb) PRE; 5679 (la) PRE; 5683 (la/2a) PRE; 5687 (la) PRE; 5688
(la) PRE; 5695 (2a) PRE; 5697 (2a) PRE; 5698 (la) PRE; 5699 (la)
PRE. Gluckman 11.10.38 (lb) J. Grobler 31.10.62 (2b) K, STE.
Hanekom 1294 (lb/2a) K, PRE; 2045 (la) PRE. Harvey 318 (2) K; 329
(2b) K; 335 (2b) K. Haynes H460 (lb) JF, K, PRE, STE; 596 (2b) JF,
PRE; 597 (lb) JF, PRE. Henderson 1278 (lb) NBG. Howes 202 (lb) K.
Jacot Guillarmod 8478 (2a) K, PRE.
Kensit Nov. 1913 (lb) K; Jan. 1914 (la/2a) BOL, PRE; Jan. 1914 (2a)
BOL; s.n. (lb) J. Kerfoot & Haynes 8 (lb) JF. Kruger 340 (lb) JF, PRE.
Laughton s.n. (2a) BOL (No. 22542); Levyns 9816 (2b) BOL; s.n. (lb)
BOL. Liebenberg 3805 (2a) K; 3808 (2a) K; 4011 (2a) PRE; 4064 (lb)
PRE, STE; 4308 (la/lb) PRE; 4331 (la/lb) K, STE; 5589 (lb) PRE, STE.
8358 (2a) PRE. Loubser 3219 (2b) STE. Loxton 221 (la/lb) PRE.
MacGillivray 392 (lb) K. MacOwan 1692 (lb) K, SAM. McCollum
Webster N59a (lb) K. Marloth 3051 (la) PRE, STE; 3062 (lb) PRE; 3065
(lb) PRE, STE; 3074 (la) PRE. Meebold 13862 (lb) PRE; 13864 (lb)
PRE. Moss 7641 (lb) J. Muir 3802 (la) PRE. Mund & Maire (la) K,
PRE (photo).
Ofsowitz 29 (2a) PRE.
Page Feb. 1918 (lb/2a) BOL. Palmer s. n. (2a) K, PRE. Parker 4035 (2b)
K, NBG, PRE; 4681 (lb) BOL, K, NBG, PRE; 4931 (2c) BOL, K, NBG,
SAM. Pearson 3222 (lb) K, J; 3533 (lb) K; 5133 (la) BOL, K. Phillips
49 (la) NBG; 1382 (lb) SAM; 1686 (la) SAM; s.n. NBG. Pocock S123
(la) PRE. Prior Sept. 1846 (lb) K, SAM; Apr. 1903 (2a) K.
Rehm 267 (lb) B, K; s.n. (2b) B. Rehmann 74 (la) K, PRE (photo and
fragment). Richardson 63 (lb/2a) JF. Ronassen Dec. 1943 (lb) NBG.
Rycroft 14.12.1945 (2b) JF.
Salter 9648 (2b) BOL; 9665 (lb/2a) BOL, PRE. Sandwith 51 (lb) K;
128 (lb) K, PRE. Scharf 1050 (la) K, PRE; 1594 (la) PRE; 1705 (la)
PRE. Schlechter 7285 (2b) BOL, P, PRE; 9180 (lb) BOL, K, P, PRE;
9417(26) BOL, K, P, PRE; 9873 (la) K, BOL, PRE. Smook 3697 (lb)
PRE; 4083 (2a) K, PRE. Stokoe 2652 (lb/2b) BOL, SAM (No. 49380,
No. 67676); 7809 (la/lb) BOL; 7811 (2b) BOL, SAM (No. 67673, 67674);
8639 (lb) K; SAM No. 54502 (la/lb) SAM. Story 2389 (la) PRE.
Taylor 3636 (2b) PRE, STE; 4235 (la) PRE, STE; 4502 (2b) PRE, STE;
4590 (2b) STE, PRE; 5223 (lb) K, PRE, STE; 7667 (2c) PRE. The
Forester 8266 (la/lb) PRE. Thompson 2253 (la) PRE, STE.
Van Daalen 137 (2a) PRE. Van der Merwe 26-02 (lb) STE; 936 (lb) PRE,
STE; 1794 (lb) STE; 2106 (lb) STE. Van Rensburg U (lb) STE; 212 (lb)
PRE, STE; 214 (2b) K, PRE, STE. Van Wyk, Fellingham & O’Callaghan
432 (la) STE. Von Breitenbach 60 (2a/2c) PRE.
Wells 3236 (la) PRE. Williams 3152 (2b) K, PRE. Wolley Dod 2385 (2b)
K; 3118 (2b) BOL; 3121 (2b) BOL; 3477 (2b) BOL, K, PRE; 3519 (2c)
BOL, K, PRE (photo and fragment). Wright s.n. (lb) K.
Zeyher 85 (lb) BOL; 293 (la) BOL, NBG, PRE, SAM, STE. 4510 (la)
BOL; 45U (la) PRE; 4571 (lb) SAM; Oct. 1830 (la) PRE; s.n. (la) BOL,
P; (lb) SAM (No. 40069). Zinn Feb. 1940 (lb) SAM.
Note that specimens from B, K, LE and P are not included on the distri-
bution maps.
Bothalia 19,2: 209-215 (1989)
Notes on African plants
VARIOUS AUTHORS
ACANTHACEAE
SIPHONOGLOSSA AND AULOJUmCIA IN SOUTHERN AFRICA
The genera Siphonoglossa Oerst. and Aulojusticia
Lindau were recently revised for the southern African
region. As a result it is necessary to describe a new species
and make two new combinations. Aulojusticia is sunk
under Siphonoglossa which is now represented in southern
Africa by three species which are keyed out below. It
should be noted that corolla lengths are measured from
the base of the tube to the apex of the upper lip throughout.
Key to species
la Corolla (35— )40— 58 mm long; leaves 2—6 mm broad;
restricted to Barberton area 3. S. linifolia
lb Corolla 13— 35(— 45) mm long; leaves usually broader
than 6 mm; Natal, Transkei, eastern and southern Cape:
2a Corolla 34-45 mm long, lower lip 0,5 -0,6 times as long
as tube 2. S. nkandlaensis
2b Corolla 13— 31 mm long, lower lip 0,2— 1,2 times as long as
tube 1. S. leptantha
1. Siphonoglossa leptantha (Nees) Immelman, comb,
nov. Type: Cape, 3326 (Grahamstown): Uitenhage,
Olifantshoek, by Boesmans River (-C or -D), Ecklon
s.n. (S, iecto.!).
Gendarussa leptantha Nees in Linnaea 15: 372 (1841). Adhatoda
leptantha (Nees) Nees: 390 (1847). Justicia leptantha (Nees) Lindau:
349 (1893).
Two subspecies are recognized:
Corolla 13-22 mm long, lower lip (0,3-)0,6-l,2 times as
long as the tube, tube 1,5—2 mm in diameter; Knysna-
Tsitsikamma area, rarely as far east as Grahamstown
lb. S. leptantha subsp. late-ovata
Corolla 15—31 mm long, lower lip 0,2— 0,5 times as long as
the tube, tube 0,5— 1,5 mm in diameter; from Grahams-
town through the Transkei to Ngoye Forest (Natal)
la. S. leptantha subsp. leptantha
la. subsp. leptantha
Adhatoda tubulosa Nees: 392 (1847). Justicia tubulosa (Nees) T.
Anders.: 41 (1864). Siphonoglossa tubulosa (Nees) Benth. ex Lindau:
38 (1893); Benth. in Benth. & Hook. f. : 1110 (1896); Clarke in Thiselton-
Dyer: 74 (1901). Syntypes: Natal, 3129 (Port St Johns): Umzimvubu,
500 ft. (-CB), Drege s.n. (G— DC, microfiche in PRE!); S. Africa
(Olifantshoek?), Ecklon & Zeyher s.n. (MEL!).
Siphonoglossa nummularia S. Moore: 40 (1880); C.B. Cl. in Thiselton-
Dyer: 75 (1901). Type: Cape, 3227 (Stutterheim): Keiskamma Hoek
(-CA), Cooper 370 (not found).
lb. subsp. late-ovata (C.B. Cl.) Immelman, comb,
nov. Types: Cape, 3325 (Port Elizabeth): on the rocks of
Swartwaterpoort (— BB), Burchell 3405 (K, lecto.!, here
designated); Burchell 3364 (K!).
Justicia pulegioides subsp. late-ovata C.B. Cl. in Thiselton-Dyer, FI.
Cap. 5,1: 62 (1901).
In the length of the corolla tube this subspecies
resembles Justicia, but it has the pollen and non-recurving
stamens of Siphonoglossa (Table 1).
2. Siphonoglossa nkandlaensis Immelman, sp. nov.,
5. leptantha subsp. leptantha affinis, sed floribus
longioribus distributioque differt.
Herba perennis; rami costati, nodis incrassatis. Folia
lanceolata vel ovata, 33-44 x 15-20 mm, venae paginae
inferae et margo pilis indutae, apex extensus obtusus, basis
cuneata; petioli graciles, 6—17 mm longi, hirsuti. In-
florescentiae cymis remotis axillaribus compositae.
Bracteae foliis similes sed minores. Calyx lobis anguste
lanceolatis acuminatis hirsutis. Corolla (tubus et labium
superum) 34—45 mm longa, purpurea vel flavovirens,
labium inferum 0,6-0,6-plo longitudo tubi; stigma
bilobatum. Antherae thecis elongatis angustatis, ad
altitudines dissimiles insertis, uterque breve mucronatis;
pollen bicolporatum, lenticulare, exino reticulato, area
laevi utrinque colpi areolis elevatis triseriatis. Capsulam
non vidi.
TYPE. — Natal, 2831 (Nkandla): Nkandla (-CD),
Wood 9000 (NH, holo.!).
Perennial herb, branches ribbed, swollen at nodes.
Leaves lanceolate to ovate, 33—44 x 15—20 mm, hairs
present along veins of undersurface and on margin, apex
drawn out into a long blunt tip, base cuneate; petiole
slender, 6—17 mm long, hirsute. Inflorescence of axillary
cymes. Bracts similar to leaves but smaller. Calyx lobes
narrowly lanceolate, acuminate, hirsute. Corolla 34—45
mm long (tube and upper lip), purple or yellow-green,
lower lip 0,5— 0,6 times as long as the tube; stigma bilobed.
Anther thecae elongate, narrow, inserted at different
_r_
FIGURE 1. — Distribution of Siphonoglossa spp. in South Africa: S.
leptantha subsp. leptantha, O; S. leptantha subsp. late-ovata,
#; S. nkandlaensis, I; S. linifolia, ■.
210
Bothalia 19,2 (1989)
TABLE 1.— Comparison of Siphonoglossa linifolia with other Siphonoglossa spp. and Justicia spp.
heights, both shortly mucronate; pollen 2-colporate,
lenticular, exine reticulate with the smooth area on either
side of the colpi studded with 3 rows of raised areoles.
Capsule not seen.
Five specimens of this species have been seen, two from
the eastern Cape and three from Zululand (Figure 1). This
distribution is similar to that of a species of the closely-
related genus Justicia L. , J. capensis Thunb. , which also
occurs in Zululand and the eastern Cape, but has never
been recorded in the intervening areas of southern Natal
and the Transkei. 5. nkandlaensis is found associated with
forest, either on its margin or along forest paths.
NATAL. — 2831 (Nkandla): Nkandla Forest Reserve, in grassveld next
to forest (— CA), Venter 3486 (PRE); Nkandla, forest path (— CA), (in
high forest near road, just before turnoff to Nkandla Forest Station — fide
Wells pers. comm.). Wells 2495 (PRE).
CAPE. — 3227 (Stutterheim): Kei Bridge (— DB), Schonland s.n.
(GRA); Komgha (-DB), Flanagan 675 (GRA).
The very long flowers and the length of the lower lip
relative to the tube distinguish this species from 5.
leptantha subsp. leptantha. The flowers are most similar
to S. linifolia which, however, is quite different
vegetatively, as well as in habitat and distribution.
3. Siphonoglossa linifolia (Lindau) C.B. Cl. in
Thiselton-Dyer, Flora capensis 5,1: 75 (1901). Type:
Transvaal, 2531 (Komatipoort): Barberton, Saddleback
Mountain, 1 300-1 700 m (— CC), Galpin 825 (Bf; PRE,
lecto.!, here designated).
Aulojusticia linifolia Lindau: 325 (1897); Lindau: 209 (1908).
Siphonoglossa linifolia, when originally described, had
been placed in a monospecific genus, Aulojusticia Lindau.
C.B. Clarke later removed the species to Siphonoglossa.
After careful examination of the specimens in the major
South African herbaria, the present author has decided
to follow Clarke because no significant differences were
noted in the pubescence of the corolla, in the stigma or
in the pollen, all of which have at various times been used
to distinguish the genera.
In some respects, e.g. leaves, habit, habitat and anthers,
Siphonoglossa linifolia resembles Justicia rather than
Siphonoglossa. However, the differences in leaves and
habit are probably an adaptation to its more exposed
habitat, and it was decided that S. linifolia would be better
placed in Siphonoglossa. The similarities and differences
considered significant are tabulated in Table 1.
This note is based on a thesis presented for the degree
of Ph.D. at the University of Natal, Pietermaritzburg.
REFERENCES
ANDERSON, T. 1864. An enumeration of the species of Acanthaceae
from the continent of Africa and the adjacent islands. Journal
of the Linnean Society of London (Botany) 7: 13—54.
BENTHAM, G. 1896. Siphonoglossa. In G. Bentham & J.D. Hooker,
Genera plantarum: 1110. Cramer, Weinheim.
CLARKE, C.B. 1901. Acanthaceae. In W.T. Thiselton-Dyer, Flora
capensis 5,1: 1—92. Reeve, London.
LINDAU, G. 1894. Siphonoglossa. In H.G.A. Engler & K.A.E. Prantl,
Die Naturlichen Pflanzenfamilien 4,3b: 338. Engelmann, Leipzig.
LINDAU, G. 1897. Acanthaceae africanae IV: Aulojusticia. Botanische
Jahrbucher fur Systematik, Pflanzengeschichte und Pflanzen-
geographie 24: 324—325.
LINDAU, G. 1908. Nachtrage zu Teil IV Abteilung 3b. In H.G.A. Engler
& K.A.E. Prantl, Die Nattirlichen Pflanzenfamilien, Nachtrdge
III: 309. Engelmann, Leipzig.
MOORE, S. LE M. 1880. Alabastra diversa 4. Journal of Botany, British
and Foreign, London 18: 40.
NEES, C.G. VON E. 1841. Acanthaceae Africae Australioris ab Ecklonio
collectae adiectis nonnullis Dregeanis: Gendarussa. Linnaea 15:
366 -373.
NEES, C.G. VON E. 1847. Acanthaceae. In A.P. De Candolle, Prodromus
systematis naturalis regni vegetabilis 11: 46—521. Treutel & Wiirtz,
Paris.
K.L. IMMELMAN
MS. received: 1988.07.16
POLYGONACEAE
OXYGONUM ALT1SS1MUM, A NEW SPECIES FROM CENTRAL SOMALIA
Oxygonum altissimum Germishuizen, sp. nov. O.
buchananii et O. tristachyo affinis, sed foliis ovatis valde
minoribus, basin cuneatis, in petiolum longum decrescen-
tibus, atque ocrea sine setis longis differt.
TYPE. — Central Somalia, 0346 (Aadan Yabaal
District): 30 km S of Aadan Yabaal (— CA), J.B. Gillett
& J.J. Beckett 23264 (K, holo.; EA, MOG). Figure 2.
Slender, erect, much branched shrub, up to 3 m tall.
Branches glabrous, covered with bloom; older branches
grey or red, with bark peeling off in longitudinal flakes,
revealing reddish wood beneath. Ocreae truncate, up to
10,0 mm long, membranous, white, glabrous, entire or with
a few, short, brown, rigid setae on edge. Leaves simple,
alternate, grey-green, smooth, covered with bloom,
narrowly to broadly ovate, cuneate at base, acuminate at
Bothalia 19,2 (1989)
211
apex, (13,0— )20, 0—28,0 x 10,0—17,5 mm, entire or
marginally pubescent with small, white scales, midrib
ventrally visible, larger leaves towards stem base. Inflores-
cence a long lax thyrse with fascicles of up to 3 flowers
in the axils of brown cuspidate membranous bracts; axis
up to 110 mm long. Perianth 5-lobed, pinkish white; lobes
oblong, up to 5 mm long. Stamens 8, included; filaments
up to 5 mm long; anthers up to 1 mm long. Styles 3, up
to 4 mm long, joined for two-thirds of the way; stigmas
capitate. Fruit immature.
CENTRAL SOMALIA.— 0346: 20 km WSW of Aadan Yabaal
(—AC), Kuchar 17291 (K; PRE); 30 km S of Aadan Yabaal (— CA), Gillen
& Beckett 23264 (EA; K; MOG).
Oxygonum altissimum is endemic to the sand plain area
of Aadan Yabaal District of central Somalia. Found in soft
yellowish orange, level sand in Acacia-Commiphora-
Loewia glandulosa bushland. O. altissimum flowers during
May and June. It is reported that the leaves of O.
altissimum are untouched by stock, but occasionally eaten
by camels when hungry.
The specific epithet altissimum is the Latin word
meaning the tallest , and is used on recommendation of
J.B. Gillett, an allusion to the tall habit these shrubs attain.
ACKNOWLEDGEMENTS
The author would like to thank the Director and staff
(particularly J.B. Gillett) of Kew Herbarium for the loan
of specimens and the opportunity to describe this new
taxon. Dr H.F. Glen and A. Romanowski of the Botanical
Research Institute, Pretoria are also thanked for their
valuable assistance.
G. GERMISHUIZEN
FIGURE 2. — Oxygonum altissimum Germishuizen, holotype. MS. received: 1989.02.27
POLYGONUM HYDROPIPER IN SOUTHERN AFRICA
During the course of a revision of the genera Polygo-
num L., Bilderdykia Dumort and Reynoutria Houtt. in
southern Africa, it was found that numerous herbarium
specimens filed under Polygonum salicifolium Willd. were
wrongly identified.
In P. salicifolium the perianth is eglandular and pink
to purple and the nut is always trigonous. In contrast, all
the wrongly identified specimens under P. salicifolium had
a green, glandular punctate perianth and a lenticular nut.
Initially these specimens were thought to belong to a new
taxon, but further studies showed that they belong to P
hydropiper L.
Studies were undertaken to establish whether other
characters could be used to distinguish between the two
taxa.
In all the investigated material it was found that P
hydropiper possesses a glandular perianth and lenticular
fruit and P. salicifolium an eglandular perianth with a
trigonous fruit (Figures 3 & 4). Only three other charac-
teristics may be of some value in separating the two
species. The fruit is no longer than 3 mm in P. salicifolium
but always longer than 3 mm in P hydropiper (Figures
3 & 4). The leaves of P salicifolium are usually no wider
than 2 mm whereas those of P. hydropiper are mostly
broader than 2 mm (Figure 4). The width/length ratio of
P salicifolium is mostly less than 0,18 and that of P.
hydropiper more than 0,19.
Polygonum hydropiper L. , Species plantarum 1: 361
(1753); Meisn.: 109 (1856); Benth. & F. Muell.: 269 (1870);
Steward: 58 (1930); Webb & Chater: 79 (1964); Ohwi: 411
(1965); Lai: 271 (1976). Type: from Europe (collector and
herbarium unknown).
Persicaria hydropiper (L.) Spach: 536 (1841); Britton & Brown: 670
(1913), Spach non Opiz.
Erect or basally decumbent slender annual, up to 1 m
tall; stems simple or branched, glabrous. Ocreae tubular,
membranous, brown, up to 20 mm long, thinly covered
with close ascending, bristly hairs and terminally fringed
with short erect-patent stiff bristles, 10,0-20,0 mm long
(Figure 5B). Leaves subsessile; blade lanceolate,
(50 — )60 — 120( — 150) x (5 — )14 — 27( — 32) mm, apically
212
Bothalia 19,2 (1989)
FIGURE 3. — A scatter diagram based on the width of the leaves in mm
(X axis) and length of fruit in mm (Y axis) of Polygonum salicifo-
lium, <$; and P. hydropiper, 4. • = glandular perianth, o =
eglandular perianth, 4 = lenticular fruit, 6 = trigonous fruit.
acute, cuneate at the base (Figure 5A), glandular-dotted
on the lower surface, more or less scabrous at the margins.
Inflorescence a slender terminal thyrse up to 120 mm long,
often nodding at the ends of the stem branches. Bracts
glabrous, truncate or truncate-rounded, reddish brown,
with a terminal fringe of rigid bristles about 1 mm long.
Perianth 4-5-lobed; lobes greenish white, 3-5 mm long
(Figure 5C & D), brownish glandular-dotted. Stamens 5.
FIGURE 4. — A scatter diagram based on the width and length ratio
of the leaves (X axis) and length of fruit in mm (Y axis) of Poly-
gonum salicifolium, 6 and P. hydropiper, 4.9= glandular
perianth, O = eglandular perianth, 4 = lenticular fruit, 6 =
trigonous fruit.
FIGURE 5. — Polygonum hydropiper L. A, habit, x 1; B, ocrea, x 1;
C, flower, x 5; D, flower, longitudinal section, x 5; E, fruit,
x 6.
Styles 2, united for half their length. Nut lenticular, dark
brown, finely granulate, 3,0-4,5 mm long (Figure 5E).
A native of Europe, it is now widely distributed in the
temperate and subtropical regions of the world. In southern
FIGURE 6. — Distribution of Polygonum hydropiper L.
Bothalia 19,2 (1989)
213
Africa this species is found in the south-eastern Trans-
vaal, Natal and south-western Cape Province (Figure 6).
It occurs in damp places, often growing in water, and on
stream and river banks and on the edge of dams.
The disjunct distribution may be attributed to the fact
that seeds could have been transported to the main
harbours of southern Africa, and spread from there. It
flowers from December to April.
Voucher specimens: Codd 6944 (PRE); Esterhuysen 20155 (BOL);
Germishuizen 1640, 1858 (PRE); Ward 5830 (NU, PRE).
ACKNOWLEDGEMENTS
The authors would like to thank the Directors and staff
of BOL, NH, NU, PRE, PRU and STE-U for the loan
of specimens.
REFERENCES
BAKER, J.G. & WRIGHT, C.H. 1909. Polygonaceae. In W.T. Thiselton-
Dyer, Flora of tropical Africa 6,1: 99—111. Reeve, London.
BENTHAM, G. & VON MUELLER, F. 1870. Polygonaceae. Flora aus-
traliensis: a description of the plants of the Australian territory
5: 266—269. Reeve, London.
BRITTON, N. & BROWN, A. 1913. Illustrated flora of the United States
1,2 : 670. New York.
LAI, MING-IOU 1976. Polygonaceae. Flora of Taiwan 2: 271. Epoch
Publishing Co., Taipei.
LINNAEUS, C. 1753. Species plantarum. (Facsimile edn 1957) 1: 361.
Ray Society, London.
MEISNER, C.F. 1856. Polygonaceae. In A.P. De Candolle, Prodromus
systematis naturalis regni vegetabilis 14: 109. Paris.
OHWI, I. 1965. Polygonaceae. Flora of Japan : 411. Smithsonian Insti-
tution, Washington DC.
SPACH, E. 1841. Phanerogames. Histoire naturelle des vegetaux 10: 536.
Paris.
STEWARD, A.N. 1930. The Polygonaceae of eastern Asia. Contribu-
tions from the Gray Herbarium of Harvard University 88: 58.
Cambridge, Massachusetts..
WEBB, D.A. & CHATER, A.O. 1964. Polygonaceae. In T.G. Tutin et
al.. Flora europaea 1: 79. Cambridge University Press.
WRIGHT, C.H. 1912. Polygonaceae. In W.T. Thiselton-Dyer, Flora
capensis 5,1: 459-482. Reeve, London.
G. GERMISHUIZEN* P.D.F. KOK** and PI. ROBBERTSE**
* Botanical Research Institute, Private Bag X101, Pretoria 0001.
** Department of Botany, University of Pretoria, Pretoria 0002.
MS. received: 1989.02.27.
ARECACEAE
NEW BOTANICAL PERSPECTIVES ON THE ORIGIN OF THE RAPHIA PALMS AT MTUNZINI
The natural distribution of raphia palms in South Africa
is limited to the Kosi Bay area in the extreme north-east
of Zululand. However, early this century a grove of raphia
palms was established at Mtunzini about 250 km further
south on the Zululand coast (Figure 7).
There has been much speculation as to exactly when,
why and how the first raphia palms were introduced to
Mtunzini. Secondary sources, past and present, verbal and
ngn-verbal, (Austen 1953; C.C. Foxon Jrpers. comm. 1987;
Harrison 1986; Oberholster 1972; Palmer & Pitman 1972)
differ somewhat with respect to the ‘when’ and ‘why’.
The original supply of seed was in fact sent from Pretoria
to Mtunzini by the then Secretary for Justice and Direc-
tor of Prisons, Jacob de Villiers Roos (1916), who wrote
to the magistrate at Mtunzini on the 21st July 1916, that
it would be economical to grow the raphia palm in the
marshy Government ground there, to supply the fibre for
the Prison brush and broommaking industry, instead of
importing it from West Africa via London.
This directive from Roos to C.C. Foxon, who was magis-
trate at Mtunzini from 1905 to 1921, clearly fixes the date
and the reason for the establishment of raphia palms at
Mtunzini. The origin of the seed supplied by Roos is not
stated. However, on the 19th July 1916, A.M. Bottomley
(1916) of the Division of Botany in Pretoria, wrote to Roos
stating, ‘I am forwarding you herewith the seed of Rafia
(sic) vinifera which we obtained from Zululand’. It seems
likely but not certain, that this was the seed, which, two
days later, was forwarded to Foxon by Roos.
Two further parcels of raphia seed were sent to Foxon
on behalf of the Director of Prisons, Pretoria, for planting.
The covering letter (Director of Prisons 1917a) for the
second batch, dated 10th January 1917, gives no indication
whatever of the source of the seed.
It may also have been Zululand or perhaps Portuguese
East Africa, but it is not unreasonable to consider the
possibility that the seeds may even have come from West
Africa, since raphia produce was imported from that
source on a regular basis.
However, the covering letter (Director of Prisons 1917b)
for the third (and apparently final) batch, dated 20th
January 1917 states, ‘I am forwarding under separate cover
a consignment of the abovementioned seed (R. vinifera)
which has been received from Portuguese East Africa’.
This batch may have been R. australis, a species described
only in 1969 (see discussion below) or R. vinifera, both
of which occur in what is today Mozambique.
It is clear that in no single instance can the identity of
the original batches of seed be established beyond
reasonable doubt. Obviously, at least one batch must have
contained R. australis in order to account for the existence
of the present population at Mtunzini. At the time it was
believed that all these seeds were of the species R. vinifera,
but this was to be disproved later, as we shall see.
The raphia palms indigenous to northern Zululand and
southern Mozambique are geographically isolated from
other giant raphia palms in Africa. Despite this they were
originally presumed to be of the species R. vinifera which
occurs commonly in West Africa. An indication to the con-
trary was recorded when specific differences between the
fruits of the Kosi Bay palms and those of R. vinifera, were
noted by Aitken & Gale (1921), who concluded that ‘either
214
Bothalia 19,2 (1989)
FIGURE 7. — Raphia palms at Mtun-
zini, Zululand coast.
this palm is a variety of R. vinifera, or it may be a new
species of the genus’. King (1925) also referred specifically
to ‘the true Raffia Palm’, R. vinifera, and ‘the giant palms
from Kosi Bay’, clearly believing them to be different
species.
Much later, 1967, two palms at the Botanic Station in
Durban, which were grown from seed collected at
Mtunzini by Dr V.A. Wager, developed large (±3 m),
erect inflorescences growing centrally above the crown of
the leaves. It was then realized that a previously
undescribed species was involved since, amongst others,
the inflorescence of R. vinifera is pendulous. Obermeyer
& Strey (1969) described the new species and named it
R. australis. Despite the inclusion of ‘Historical Notes’
in their paper, Obermeyer & Strey make no reference to
the observations of Aitken & Gale and King, whose earlier
reports appear to have been overlooked.
The very uncertain identity of the original seed and the
proven ease with which the true identity of established
raphias may be overlooked, suggest the strong possibility
that one or even more raphia species other than R. aus-
tralis were also introduced to Mtunzini. Further support
for this possibility is contained in a letter addressed to the
Conservator of Forests, Pietermaritzburg, by King (1925),
who stated that, ‘The seeds supplied by Mr de V. Roos
were of the true Raffia Palm. In addition to these certain
seed of the giant palms from Kosi Bay was planted’. Yet
today only R. australis is known to grow in the Mtunzini
area. It could be speculated that the exotic species died
out because they were less well adapted to local conditions.
The increase of Raphia trees in the immediate vicinity
of Mtunzini was facilitated in the 1940’s by L.E. Davis
who collected seeds from the original grove and planted
them while visiting the surrounding swamplands in the
course of his duties as malaria control officer (Palgrave
1977; Wicht 1969).
In 1942 the original grove was declared a national monu-
ment (Government Gazette 1942). The grove is of no great
historic significance and the procedures which led to the
declaration were somewhat unorthodox. In spite of this,
Mtunzini’s raphia palms joined about 17 other botanical
specimens which form about 5 % of the list of the country’s
official national monuments (Oberholster 1972). The grove
at the original site was virtually destroyed in 1948 when
a spark from a passing locomotive set the trees ablaze
(Oberholster 1972). The remaining trees were able to
regenerate and a fine grove of palms may be seen at the
site today.
Over the last three decades Mr Ian Garland has made
a substantial contribution to the population of raphia palms
by planting and distributing almost a thousand R. australis
on, and in the vicinity of his farm ‘Twin Streams’, which
borders Mtunzini (I.F. Garland pers. comm. 1987).
In recent years the Mtunzini Town Board has played an
active role in promoting the raphia palm (Figure 7), which
along with the closely associated palm nut vulture,
Gypohierax angolensis, has been incorporated into the
official town emblem (Government Gazette 1981).
Even though it is doubtful whether the exact origin of
Mtunzini’s raphia palms will ever be known their present
status is secure and improving.
REFERENCES
AITKEN, R.D. & GALE, G.W. 1921. Botanical Survey of South Africa,
Memoir No. 2: Botanical Survey of Natal and Zululand: 17-18.
Government Printer, Pretoria.
AUSTEN, W.M. 1953. Palm-nut vultures (Gypohierax angolensis) in
Raphia palms at Mtunzini, Zululand. The Ostrich 24: 98-102.
BOTTOMLEY, A.M. 1916. Letter dated 19 July. C. A., JUS237, 3/569/16.
DIRECTOR OF PRISONS 1917a. Letter dated 19 January. Natal Archives
(N.A.), 1/MTU3/4/2/7.
DIRECTOR OF PRISONS 1917b. Letter dated 20 January. N.A.,
1/MTU3/4/2/7.
Bothalia 19,2 (1989)
215
GOVERNMENT GAZETTE 1942. Proclamation No. 2685, dated 21
December.
GOVERNMENT GAZETTE 1981. Proclamation No. 7673, dated 13
March.
HARRISON, E. 1986. The Raphia’s of Mtunzini. The Palm Enthusiast
3 : 22 -27.
KING, R.F. 1925. Letter dated 17 November. C.A., NTS5984:74/321.
OBERHOLSTER, J.J. 1972. The historical monuments of South Africa.
The Rembrandt van Rijn Foundation for Culture, Cape Town.
OBERMEYER, A. A. & STREY 1969. A new species of Raphia from
northern Zululand and southern Mozambique. Bothalia 10:
29-37.
PALGRAVE, K.C. 1977. Trees of southern Africa. Struik, Cape Town.
PALMER, E. & PITMAN, N. 1972. Trees of southern Africa, Vol. 1.
Balkema, Cape Town.
ROOS, J. DE V. 1916. Letter dated 21 July. N.A., 1/MTU3/4/2/7.
WICHT, H. 1969. The indigenous palms of southern Africa. Howard
Timmins, Cape Town.
G.D. PECKHAM* and F.A. VAN JAARSVELD*
* University of Zululand, Private Bag X1001, KwaDlangezwa 3886.
MS. received: 1988.08.08.
.
Bothalia 19,2: 217-223 (1989)
Leaf anatomy of the South African Danthonieae (Poaceae): XIX. The
genus Prionanthium
R.P. ELLIS*
Keywords: Arundineae, leaf anatomy, linear glands, Poaceae, Prionanthium , taxonomy
ABSTRACT
The leaf blade anatomy of Prionanthium dentatum (L. f.) Henr., P. ecklonii (Nees) Stapf and P. pholiuroides Stapf is
described and illustrated. The transectional anatomy is non-Kranz with diffuse chlorenchyma. The adaxial epidermis has
dome-shaped stomata, dumbbell-shaped to nodular silica bodies and elongated microhairs. The three species differ in the
presence or absence of macrohairs, abaxial stomata and marginal linear, sessile glands. The latter character is shared with
Pentaschistis triseta (Thunb.) Stapf and its allies, and relationships with Pentaschistis are indicated rather than with any
other extant arundinoid genus.
UITTREKSEL
Die blaaranatomie van Prionanthium dentatum (L. f.) Henr. , P. ecklonii (Nees) Stapf en P. pholiuroides Stapf word beskryf
en geillustreer. Die anatomie in dwarssnee is nie-Kranz met eweredig verspreide chlorenchiem. Die abaksiale epidermis
het koepelvormige huidmondjies, murgbeenvormige silikaliggaampies en langwerpige, vingeragtige mikrohare. Die drie spesies
verskil ten opsigte van die voorkoms van makrohare met geen gespesialiseerde basale epidermisselle, abaksiale huidmondjies
en lineere, steellose kliere op die blaarrand. Soortgelyke kliere kom ook by Pentaschistis triseta (Thunb.) Stapf en sy verwante
spesies voor en verwantskappe met Pentaschistis word aangedui, eerder as met enige ander teenwoordigers van die tribus
Arundineae.
belongs with the primitive arundinoid genera such as
Tribolium, Urochlaena, Elytrophorus and others from
Australasia. The phylogenetic implications of the leaf
anatomy of Prionanthium will be considered in this paper
in order to shed more light on this problem. The purpose
of this paper is, therefore, to describe and illustrate the
leaf blade anatomy of Prionanthium and to compare this
structure with that of the other South African danthonioid
species, as well as with earlier published results for Pri-
onanthium.
Little detailed attention has been given to the leaf
anatomy of Prionanthium in the published literature but
the information available does indicate typical arundinoid
anatomy. The anatomy is non-Kranz with a double bundle
sheath, non-radiate mesophyll with a maximum lateral cell
count greater than four, bulliform cell groups not
associated with colourless cells, finger-like microhairs
with tapering distal cells, domed subsidiary cells and
dumbbell-shaped to nodular silica bodies (Hansen &
Potztal 1954; Renvoize 1986; Watson etal. 1986). Renvoize
(1986) notes two unusual features for the subfamily: the
chlorenchyma is not interrupted by sclerenchyma girders
above and below the vascular bundles, and the epidermal
cells are large and thin-walled. De Wet (1956) notes that
the epidermis of Prionanthium is panicoid (with linear
microhairs) but that the anatomy is festucoid with the
chlorophyll tissue being uniformly distributed between the
bundles.
MATERIALS AND METHODS
Specimens of all three species of Prionanthium were
collected in the south-western Cape. Herbarium voucher
specimens were prepared for verification by the staff of
the National Herbarium (PRE) where they are housed.
INTRODUCTION
The genus Prionanthium is a small genus of three
species all endemic to the south-western Cape Province.
It includes small, ephemeral or annual plants which are
exceedingly difficult to locate and are considered to be
rare and/or endangered (Davidse 1988). Taxonomically the
genus does not pose any particular difficulties and it
contains three morphologically distinct species: P.
dentatum (L. f.) Henr. (= P. rigidum Desv.), P. ecklonii
(Nees) Stapf and P pholiuroides Stapf. These three species
are recognized by most previous workers (Chippindall
1955; Clayton & Renvoize 1986; Watson et al. 1986;
Davidse 1988) although all these workers did not have
access to material of all three taxa.
The recent revision by Davidse (1988) included all the
anatomical voucher specimens used for the present study
and the anatomical data reported on here is complementary
to the morphology and cytology discussed in this revision.
The present study is the first in which comparative leaf
anatomy of all three species has been undertaken.
Although there is little difficulty with the delimitation
of taxa within Prionanthium, the generic relationships of
the genus are puzzling. Most authors now agree with the
placement of the genus in the Arundinoideae, usually in
the tribe Arundineae (= Danthonieae). Davidse (1988)
convincingly argues the case for arundinoid relationships
and, furthermore, demonstrates that Prionanthium and
Pentaschistis are probably related. This conflicts with the
suggestion of Clayton & Renvoize (1986) that the genus
* Botanical Research Institute, Department of Agriculture and Water
Supply, Private Bag X101, Pretoria 0001.
MS. received: 1989.02.15.
218
Bothalia 19,2 (1989)
FIGURE 1. — Leaf blade anatomy of Prionanthium dentatum. A-D, leaf in transverse section: A, outline of lateral part of lamina showing
undifferentiated midrib (m) and narrow, projecting margin, x 100; B, detail of margin showing sessile gland cells (g) in the extreme
margin and the diffuse nature of the chlorenchyma, x 250; C, outline of lateral part of blade without midrib (m) and with tapering
margin, x 160; D, anatomical detail of chlorenchyma, vascular bundles, adaxial ribs and sclerenchyma strands (s), x 250. E— H, abaxial
epidermal structure as seen in surface view: E, macrohairs with single modified basal cell, microhairs and nodular silica bodies, x‘
250; F, interference contrast of macrohairs and silica bodies, x 250; G, lateral part of leaf blade with glandular marginal cells with
persistent nuclei, x 250; H, sessile linear gland in leaf margin, x 250. A, B, G, Ellis 5416\ C, F, Ellis 2452\ D, H, Ellis 5773\ E,
Ellis 5417
These same specimens were also examined by Davidse for
a revision of the genus (Davidse 1988).
Central segments of leaf blades were removed and
immediately fixed in FA A. Transverse sections and
epidermal scrapes of these blade segments were prepared
following the methods outlined in previous papers in this
series (e.g. Ellis 1988).
The standardized terminology of Ellis (1976, 1979) was
used for the anatomical descriptions. The following
abbreviations are used in the descriptions:
vb/s — vascular bundle/s
l’vb/s — first order vascular bundle/s
2’vb/s — second order vascular bundle/s
3’vb/s — third order vascular bundle/s
ibs - inner bundle sheath; mestome sheath
obs — outer bundle sheath; parenchyma sheath
Specimens examined
Prionanthium dentatum
CAPE. — 3119 (Calvinia): 15-16 km SE of Nieuwoudtville on
Oorlogskloof road to Clanwilliam (—AC), Ellis 2452 (16.10.1975), 5416,
5417 (20.9.1987), 5773 (11.10.1988).
Bothalia 19,2 (1989)
219
FIGURE 2. — Leaf anatomy of Prionanthium ecklonii, Davidse 34018. A-C, leaf in transection: A, outline with undifferentiated midrib and
adaxial ribs, x 100; B, detail of gently tapering left hand margin with few glandular cells in extreme margin (g), x 250; C, detail
of other margin, also with slightly enlarged epidermal cells constituting a sessile gland, x 250. E-F, abaxial epidermal surface anatomy:
E, detail of sinuous intercostal long cells, dome-shaped subsidiary cells, microhairs and silica bodies, x 250; F, long cell, stomata,
silica body and microhair structure, x 250.
/WvW\rA Ajuv/- '
-vc/v^v/.'Vy_rv
/v t/u.'Vx
#"WVvv -
y V/-^5-iAaA; • ^UVVW/ V
w -A/VW^TiA
' v'V' =A/Sy/VVyV 'va a ^ /
AfVta^W^r
VAA/V v
(.Vft, SA,yy<J\nTJ'\
I E VU Www
Vi/VUA' VOvA/l
Prionanthium ecklonii
CAPE.— 3218 (Clanwilliam): 45 km N of Citrusdal (— BD), Davidse
34018, Ellis 5784.
Prionanthium pholiuroides
CAPE. — 3318 (Cape Town): Darling Dist., 3 km E of Mamre Road,
Rondevlei Farm (— BC), £7//s 5433, 5434, 5435. 3420 (Bredasdorp): 6
km N of Struisbaai (-CA), Davidse 34053.
LEAF ANATOMY OF THE GENUS PRIONANTHIUM
Leaf blade in transverse section
Outline : open, expanded, flat in P. dentatum and P.
ecklonii (Figures 1A, C; 2A) but slightly inrolled to
infolded in P. pholiuroides (Figure 3A, C, G). Ribs and
furrows : medium to deep, open to narrow and cleft-like
furrows present between all vbs. Similar rounded adaxial
ribs associated with all vbs (Figures LA— D; 2A— C;
3A-D, G). No abaxial ribs or furrows but slight
undulations may be associated with the vbs (Figure 2A).
Median vascular bundle : not structurally distinct from
lateral l’vbs. Vascular bundle arrangement : tends to vary
across the lamina width with more 3’ vbs between
successive l’vbs in the region of the median bundle than
laterally; 3 (P. ecklonii) or 2 (P. dentatum and P.
pholiuroides) 3 ’vbs between consecutive l’vbs in region
of median bundle but only 1 laterally in all species; 3 ( P.
pholiuroides), 5 (P. ecklonii and P. dentatum) and rarely
7 (P. dentatum) l’vbs in leaf section. No 2’vbs. All vbs
located in centre of blade. Vascular bundle description :
3 ’vbs elliptical with xylem and phloem distinguishable,
l’vbs round to elliptical in shape, phloem adjoins the ibs,
metaxylem vessels very narrow with diameters much less
than those of the obs cells (Figures IB, D; 2B, C; 3B, D)
with thin walls. Vascular bundle sheaths: double; round
to slightly elliptical; obs often incomplete, particularly
abaxially (Figure 3B, D, G); no extensions; obs cells
inflated, rounded, the cells smaller than the chlorenchyma
cells; walls thin; without chloroplasts or with few, small
chloroplasts; ibs entire around all bundles; of small
lignified cells with inner tangential and radial walls
thickened. Sclerenchyma: adaxial sclerenchyma in the
form of strands, either minute (Figure 2B, C), small
(Figure IB, D; 3B, D) or conspicuous (Figure 3G) in the
Struisbaai population of P pholiuroides-, abaxial strands
smaller and often not developed in association with the
3’vbs (Figure 3B, D, G). Fibres not lignified. Minute cap
in margin but margin distinctly tapering in P dentatum
and P ecklonii but not P. pholiuroides. No sclerenchyma
present between bundles. Chlorenchyma : irregular, with
no pattern in the cell arrangement; cells of different size
and shape with conspicuous intercellular air spaces;
occupy the entire area between the adaxial and abaxial
epidermides, even the space above and below the vbs
(Figures IB, D; 2B, C; 3B, D, G); no arm cells or fusoids.
Colourless cells : absent. Adaxial epidermal cells: small
to medium-sized bulliform groups at bases of furrows in
P. dentatum and P. ecklonii (Figures IB, D; 2B, C), occupy
less than \ leaf thickness, may be completely absent as
in P pholiuroides (Figure 3B, G); cuticle thin; epidermal
appendages absent; no papillae although epidermal cells
220
Bothalia 19,2 (1989)
mwip&i
vn fvttfu
*/\0/V>A/V>AA
nfinfvm)
un/W-
/~l /> aAO A/VftA /"'rv.
/|^«Uwv\tvVl
rUUWU'
'WVWWWUUWu
'Wviv
»U
j$g5*
FIGURE 3. Leaf anatomy of Prionanthium pholiuroides . A— F, specimens from the Mamre Road population. A— D, leaf blade transections:
A, leaf outline showing vascular bundle arrangement and median vascular bundle not structurally differentiated from first order bundles,
x 100; B, detail of leaf margin with no sessile linear gland cells, x 250; C, blade outline without midrib, x 100; D, detail of leaf
margin with adaxial ribs and furrows, sclerenchyma strands, vascular bundles and chlorenchyma — note the absence of a narrow
pointed margin with sessile glandular cells, x 250. E— F, abaxial epidermal preparations: E, intercostal long cells with thin, slightly
sinuous walls, microhair basal cells and absence of stomata, x 250; F, long cells with sinuous walls, microhairs, stomatal absence
and nodular silica bodies, x 400. G— H, examples of specimens from the Struisbaai population: G, leaf outline illustrating infolded
nature of blade and well developed adaxial sclerenchyma strands, x 250; H, abaxial epidermis with sinuous long cell walls, micro-
hairs, no stomata and variable, nodular silica bodies, x 250. A, B, E, Ellis 5435\ C,F, Ellis 5433; D, Ellis 5434; G, H, Davidse 34053.
Bothalia 19,2 (1989)
221
tend to be inflated. Abaxial epidermal cells', bulliform cells
absent; similar in size to adaxial epidermal cells but all
cells of uniform size; cuticle thin; no epidermal
appendages present. Slightly enlarged sessile glandular
cells present in the extreme margin in P. dentatum and
P. ecklonii but absent in P. pholiuroides (Figures IB; 2C);
probably constitute part of marginal linear glands which
appear to vary from 3—7 cells wide and are located on
a distinctly narrow and projecting margin. These structures
are also visible in surface view (Figure 1G, H) where the
persistent nuclei are clearly visible.
Abaxial epidermis in surface view
Intercostal long cells: elongate, rectangular (Figures IF,
H; 2D, E; 3H) to fusiform (Figures IE, G; 3E, F); walls
virtually straight to moderately or deeply sinuous (Figure
4A, C, E); end walls vertical; walls unthickened to slightly
thickened. Cell shape and size consistent across intercostal
zones; long cells adjoin one another or separated by short
cells; no bulliform cells. Stomata: usually absent in P.
pholiuroides (Figure 3E, F, H) but present in P. dentatum
and P. ecklonii (Figures IE— H; 2D, E); low dome-shaped
to dome-shaped subsidiary cells; 1—3 rows of stomata per
intercostal zone; 1 elongate, rectangular interstomatal cell
separates individual stomata in a file. Intercostal short
cells: presence variable, either absent or irregularly
present; when present usually paired; square to tall and
narrow in shape. Papillae absent. Prickles: absent. Micro-
hairs: present on all specimens; bicellular, but variable
with basal cell either shorter than distal cell (Figures 3F;
4B, D) or the two cells equal in length (Figure 4F); hairs
longer than stomatal length; distal cell deciduous and
thin-walled. Macrohairs: usually absent but always present
in P dentatum (Figure 1E-H); rarely present in P.
pholiuroides', unicellular, soft hairs with single inflated,
hemispherical cell associated with the superficial base.
Silica bodies: angular, irregular dumbbell-shaped to
nodular; horizontally elongated; exclusively costal in very
narrow (1—3 files) costal zones; separated by costal short
cells.
DISCUSSION AND CONCLUSIONS
Differences between the species of Prionanthium
The three species of Prionanthium can readily be
distinguished from one another on anatomical criteria as
summarized in Table 1. P dentatum is unique in the genus
in possessing macrohairs but it otherwise resembles P.
ecklonii very closely in leaf anatomy: both species have
similar outlines, ribs and furrows, abaxial stomata and
microhairs. They also share the very unusual marginal,
sessile glands which are currently only known in four or
five species of Pentaschistis. In addition to the macrohairs,
P. dentatum and P. ecklonii also differ slightly in the
arrangement of the vascular bundles and in the extent of
the development of the sclerenchyma strands. P dentatum
and P ecklonii, therefore, appear to be more closely related
to each other than either is to P. pholiuroides. This agrees
with the morphological evidence, particularly that of the
glume glands where P. dentatum and P ecklonii share
similar stalked multicellular glands, whereas in P.
pholiuroides the glands are sessile (Davidse 1988).
P. pholiuroides is distinct in the genus on the basis of
a number of features: the leaf outline is inrolled to
infolded; the adaxial furrows are narrow and cleft-like;
there are fewer large vascular bundles in the leaf
transection which is, consequently, narrower; the outer
bundle sheath is often not entire but interrupted abaxially;
the sclerenchyma strands may be conspicuous; the margin
is not tapering and lacks the linear glands and abaxial
stomata are absent or very rare. P. pholiuroides can,
therefore, be readily separated from both P. dentatum and
P. ecklonii on leaf anatomy and this study fully supports
the recognition of these three species in the genus.
The results presented here must be interpreted with
caution as only single populations of both P. dentatum and
P. ecklonii were examined in this study due to the rarity
of all the species. In the case of P. dentatum the same
population was sampled on three different occasions — in
the spring of 1975, 1987 and 1988. All these specimens are
virtually identical in leaf anatomy and very little variation
was exhibited. However, two populations of P. pholiuroides
were included in the study and they exhibit obvious ana-
tomical variation as illustrated in Figure 3. The Mamre
population has more open leaves with smaller adaxial
sclerenchyma strands than the Struisbaai population.
Although these differences are visually rather striking, they
do not represent significant differences, and characters
such as vascular bundle number and arrangement, the
cleft-like furrows and the absence of abaxial stomata re-
main consistent. It is of relevance to note that the Struis-
baai population also differed cytologically from the Mamre
population in the occurrence of a small B chromosome
and this population was also the most variable in spikelet
morphology (Davidse 1988). Nevertheless, it appears like-
ly that most of the anatomical features separating the spe-
cies will hold even when more material becomes available.
TABLE 1. — Anatomical differences between the three species of Prionanthium
Outline
Adaxial furrows
No. l’vbs in section
No. 3’vbs between l’vbs
Outer bundle sheath
Adaxial strands
Bulliform cells
Margin in transverse section
Marginal glands
Abaxial stomata
Macrohairs
P. dentatum
P. ecklonii
P. pholiuroides
open, expanded
shallow, open, wide
5 or 7
2
entire
small
medium
projecting, narrow
present
present
present
open, expanded
shallow, open, wide
5
3
entire
minute
small
tapering
present
present
absent
inrolled
deep, narrow, cleft-like
3
2 or 1
absent opposite phloem
small or conspicuous
small or absent
abruptly pointed
absent
absent
absent
222
Bothalia 19,2 (1989)
FIGURE 4.— SEM micrographs of the abaxial epidermis of different species of Prionanthium. A— B, P. dentatum, Ellis 5416'. A, general
epidermal features with macrohairs, stomata and inflated, slightly sinuous long cells, x 204; B, bicellular microhair with tapering distal
cell, x790. C— D, P. ecklonii, Davidse 34018 : C, thickened, very sinuous long cells, stomata present (but not illustrated), x 210; D,
bicellular microhair with distal cell longer than basal cell, x 810. E— F, P. pholiuroides, Ellis 5433 : E, long cells slightly inflated and
walls not undulating, stomata and macrohairs absent, x 210; F, microhair with basal cell longer than distal cell, x 780.
Subfamilial and tribal classification
Numerous characters of the leaf anatomy support the
classification of Prionanthium in the Arundinoideae,
confirming the reports of previous workers (Hansen &
Potztal 1954; De Wet 1956; Renvoize 1986; Watson et al.
1986). The anatomy is C3 with diffuse mesophyll, a
double bundle sheath and without colourless cells
accompanying the bulliform cells. This structure
eliminates the possibility of panicoid or chloridoid
relationships and the absence of fusoid and arm cells rules
out a bambusoid alliance. The presence of microhairs,
dome-shaped stomata and dumbbell to nodular silica
bodies are all characters which are unequivocally
Bothalia 19,2 (1989)
223
arundinoid and exclude pooid affinities. The anatomical
evidence is, therefore, in full agreement with all other
indications (Davidse 1988) and Prionanthium appears best
placed in the Arundineae of the Arundinoideae.
Within the Arundineae the generic relationships of
Prionanthium are subject to differing interpretations.
Clayton & Renvoize (1986) are of the opinion that
Prionanthium, and in particular P. dentatum, suggests a
relationship with Tribolium and the other primitive
arundinoid genera. Leaf anatomical characters can be used
to substantiate such a relationship, particularly as all the
primitive South African arundinoid taxa display similar
mesophyll characteristics — which will here be termed the
orthophyllous or mesic anatomical type. This differs
significantly from the sclerophyllous type found in genera
such as Merxmuellera and Pentameris. These two basic
arundinoid anatomical types are discussed in Ellis (1988a)
and Prionanthium shares many anatomical similarities with
primitive genera such as Tribolium and Urochlaena (Ellis
1988a). However, other arundinoid genera not included
in the heterogeneous group of primitive genera (Clayton
& Renvoize 1986) also have this orthophyllous type of
anatomy. Chaetobromus (Ellis 1988b), Schismus,
Karroochloa and some species of Pentaschistis are
examples and, therefore, the possession of the mesic type
of arundinoid anatomy does not necessarily appear to
indicate affinity.
Davidse (1988) proposes a distant relationship between
Prionanthium and Pentaschistis based on several shared
spikelet characters, the most important being the occur-
rence of well differentiated multicellular glands on the
glumes of all three species of Prionanthium and on the
glumes, pedicels and leaves of several species of
Pentaschistis (Linder et al. in prep.). Although stalked
multicellular glands do not occur on the leaves of
Prionanthium, the presence of sessile, linear glands serves
to strengthen the argument of affinities between these two
genera.
The leaf glands do not appear to represent equivalent
structures to the sessile glands of the glumes of P.
pholiuroides which appear to be similar to the multicellular
glands but lack the stalks. The sessile glands on the leaves
are linear structures with the glandular tissue not
aggregated into raised, cylindrical cones as is the case with
the glume glands. Prionanthium is, therefore, unique in
the possession of two gland types. In Pentaschistis both
these gland types occur (Linder et al. in prep.) but not
on the same plants or species and are characteristic of two
different species groups. On the basis of this character
alone Prionanthium, therefore, appears to be intermediate
between the Pentaschistis species groups represented by
P triseta and P thunbergii (Kunth) Stapf. Affinities with
Pentaschistis are supported by the anatomical evidence,
but a critical revision of Pentaschistis is needed before the
phylogenetic links of Prionanthium become more lucid.
ACKNOWLEDGEMENTS
The co-operation and suggestions of G. Davidse
contributed substantially to this paper. Mrs A. Botha is
gratefully thanked for technical assistance, Mrs A.
Romanowski for the photography, Mrs S. Perold for the
electron microscopy and Mrs S. Brink for typing the
manuscript.
REFERENCES
CHIPPINDALL, L.K.A. 1955. A guide to the identification of grasses
in South Africa. In D. Meredith, The grasses and pastures of
South Africa. Central News Agency, Cape Town.
CLAYTON, W.D. & RENVOIZE, S.A. 1986. Genera graminum. Kew
Bulletin Additional Series 13: 1—389.
DAVIDSE, D. 1988. A revision of the genus Prionanthium (Poaceae:
Arundineae). Bothalia 18: 143—153.
DE WET, J.M.J. 1956. Leaf anatomy and phylogeny in the tribe
Danthonieae. American Journal of Botany 43: 175—182.
ELLIS, R.P. 1976. A procedure for standardizing comparative leaf
anatomy in the Poaceae. I. The leaf blade as viewed in transverse
section. Bothalia 12: 65—109.
ELLIS, R.P. 1979. A procedure for standardizing comparative leaf
anatomy in the Poaceae. II . The epidermis as seen in surface view.
Bothalia 12: 641-672.
ELLIS, R.P. 1988a. Leaf anatomy of the South African Danthonieae
(Poaceae). XVI. The genus Urochlaena. Bothalia 18: 101—104.
ELLIS, R.P. 1988b. Leaf anatomy of the South African Danthonieae
(Poaceae). XVII. The genus Chaetobromus. Bothalia 18:
195-209.
HANSEN, I. & POTZTAL, E. 1954. Beitrage zur Anatomie und
Systematik der Leptureae. Botanische Jahrbiicher 76: 251—270.
LINDER, H.P., THOMPSON, J.F., ELLIS, R.P. & PEROLD, S.M.
in prep. The occurrence, anatomy and systematic implications
of multicellular glands in Pentaschistis and Prionanthium
(Poaceae — Arundinoideae — Arundineae) .
RENVOIZE, S.A. 1986. A survey of leaf anatomy in grasses VIII.
Arundinoideae. Kew Bulletin 41: 323—342.
WATSON, L., DALLWITZ, M.J. & JOHNSTON, C.R. 1986. Grass
genera of the world: 728 detailed descriptions from an automated
database. Australian Journal of Botany 34: 223—230.
Bothalia 19,2: 225-235 (1989)
Observations on plant usage in Xhosa and Zulu medicine
A. HUTCHINGS*
Keywords: ethnobotany, pharmacognosy, practitioners, medicine, Xhosa, Zulu
ABSTRACT
The holistic concept of Xhosa and Zulu traditional medicine and some differences from Western orthodox practice are
briefly outlined. The transmission of herbal knowledge within various social groups is outlined. The background, training
and some procedures followed by five of the informants are discussed. Plant characteristics that may be seen, felt, smelled
or tasted are considered as possible determinants of usage. The form of plant parts accounts for some usage in the more
magically orientated medicines whereas colour, texture or the production of froth may signal the presence of medicinally
active components such as tannin, mucilage and saponin. The role of plants producing a milky latex is discussed. Vesicant
or irritant properties are utilized in septic or inflammatory conditions. Aromatic plants are used for respiratory or digestive
disorders and pungent-smelling plants are used in the treatment of catarrh and some stress-related disorders. Bitter or sour-
tasting plants may be used as an aid to digestion or serve a deterrent function. Parallel usage of some related plants in African
and European herbal practice indicates that appropriate usage may be widely determined by easily discerned plant characteristics.
Two herbal medicinal recipes recorded by the author and a list of medicinal plants collected in Transkei are presented.
UITTREKSEL
Die holistiese konsep in die tradisionele geneeskunde van die Xhosa en die Zoeloe, en enkele verskille van ortodokse
Westerse gebruike word in hooftrekke beskryf. Die oorlewering van kennis omtrent kruie in verskillende sosiale groepe
en die opleiding, agtergrond en prosedures wat gevolg is deur vyf van die verskeie beoefenaars met wie die onderhoude
gevoer is, word bespreek. Plantkenmerke wat gesien, gevoel, geruik of geproe kan word, word beskou as moontlik bepalend
vir gebruik. Die vorm van plantdele gee aanleiding tot hulle gebruik in die meer magies georienteerde geneesmiddels, terwyl
kleur, tekstuur, of die vorming van skuim ’n aanduiding kan gee van die aanwesigheid van medisinaal aktiewe komponente
soos tannien, plantslym en saponien. Die rol van plante met melksap word bespreek. Blaartrekkende of irriterende eienskappe
word aangewend in gevalle van septiese toestande en ontsteking. Aromatiese plante word gebruik vir respiratoriese of
spysverteringongesteldhede en plante met ’n skerp, prikkelende geur word gebruik vir die behandeling van katar en sommige
spanningstoestande. Plante met ’n bitter of suur smaak kan as hulpmiddel by spysvertering of as afskrikmiddel dien. Parallelle
gebruik van sekere verwante plante in kruiegeneeskunde in Afrika en Europa dui aan dat gepaste gebruik algemeen bepaal
word deur maklik onderskeibare kenmerke. 'n Lys van plante wat in Transkei versamel is, is opgestel en twee kruiegeneeskundige
resepte wat deur die outeur opgeteken is, word gegee.
CONTENTS
Introduction 225
Traditional Xhosa, Zulu and Western orthodox me-
dicine
A background to Xhosa and Zulu traditional me-
dicine 226
Some differences between traditional Xhosa,
Zulu and Western orthodox medicine 226
Transmission of herbal knowledge
1 Categories of practitioners 226
1.1 Villagers 226
1.2 Herbalists 227
1.3a Diviners 227
1.3b Traditional doctors 227
1.4 Homeopath/herbalists 227
2 Informants — background and practise 227
Plant characteristics as determinants of usage 228
1 Characteristics that may be seen
a. Suggestive forms 228
b. Colour 229
c. Plants that froth in water 229
d. Mucilage 229
e. Milky latex 229
2 Characteristics that may be felt 229
3 Characteristics that may be smelled 230
* Zulu Folk Medicine Research, Department of Biochemistry, University
of Zululand, Private Bag X1001, KwaDlangezwa 3886.
MS. received: 1988.10.12.
4 Characteristics that may be tasted 230
5 Conclusion 230
Plant usage in Transkei 230
Recipes for herbal medicines
Recipe 1: medicine for swollen glands 231
Recipe 2: medicine for ipleyiti 231
Appendix 232
Acknowledgements 235
References 235
INTRODUCTION
This study is an attempt to answer the question ‘How
do people know which plants to use?’ in traditional Xhosa
and Zulu medicine. Both the transmission of herbal
knowledge and the apparent role of easily discernible plant
characteristics as determinants of usage are considered.
Botanical fieldwork, focussed mainly on the establishment
of a herbarium at the University of Transkei, presented
the author with an opportunity to collect and observe many
of the plants known to be used in traditional medicine.
Plant species listed in this paper were collected or observed
in the company of an informant and further usage was
recorded from interviews. Informants from Transkei
included villagers, herbalists, traditional healers and
homeopath/herbalists. The author attended two meetings
with a group of ten Xhosa traditional healers at St
Elizabeth’s hospital in Lusikisiki, Transkei, in 1986 and
was also present at a meeting held in Umtata, attended by
226
Bothalia 19,2 (1989)
a very large group of healers from many parts of Transkei.
They met to discuss the possibility of joining the SA
Traditional Healers Council. One meeting with a group
of ± 30 traditional Zulu healers was attended at Valley
Trust in Natal in 1987. Communal cultivation of medicinal
plants in short supply, potential problems in the use of toxic
plants, possible means of co-operation with the relevant
institutions and some plant usage were among the topics
discussed at these meetings.
TRADITIONAL XHOSA, ZULU AND WESTERN
ORTHODOX MEDICINE
A background to Xhosa and Zulu traditional medicine
The first written records of Xhosa and Zulu medicinal
plant usage were published as early as 1885 (Smith 1895)
and 1909 (Bryant 1966). Smith (1895:6) refers to the age-
old oral transmission of herbal knowledge as the ‘heritage
of experience’. In order to understand something of this
heritage a few points on the underlying concepts of disease
and its treatment need to be made, as these differ from
the modern Western orthodox approach to medicine. A
holism, involving both the relationship between body and
mind in the individual and the relationship between the
individual and his social and physical environment, is to
be found in traditional Xhosa and Zulu attitudes to health
and disease. Bryant (1966:16) refers to the recuperative
powers of the Zulu as being possibly attributable to: ‘his
possession of a mind working in more perfect harmony
with the requirements of the body’. Ngubane (1977)
distinguishes between diseases believed by the Zulu to be
caused by natural, biological factors and those believed
to be caused by environmental factors. Traditional belief
in the presence of the ancestors, sorcery, evil spirits and
mystical forces that produce pollution are presented by her
as being part of the perceived social and physical environ-
ment. These are the means by which a sense of moral order
and of community is fostered. Although the ancestors are
believed to cause some disease themselves when offended
by a failure to carry out proper rites, their function is
primarily a protective one. Through the medium of dreams
they call the diviners to their profession and, through them,
reveal the cause and nature of an illness and also direct
its cure. Bryant (1966), Ngubane (1977) and Broster (1982)
all refer to the high moral standing of the diviners within
their society. The sense of responsibility healers feel
towards the community and the realisation of the potential
effects of one person’s illness on the community was
demonstrated by one of the healers interviewed (Mr
V.M. — see informant 3). He said that if any member of
a patient’s family became ill during treatment, that person
was treated by him without further charge. He also paid
the medical costs of any one of his patients he felt needed
the help of a Western-trained doctor while undergoing
treatment. Traditional healers referred, on more than one
occasion, to their custom of taking patients to stay in their
own homes while undergoing treatment.
Some differences between traditional Xhosa, Zulu
and Western orthodox medicine
The differences in the understanding of the cause of
disease may account for the following differences from
Western medicinal practice in forms of administration in
traditional Xhosa and Zulu practice: 1, the wider use of
emetics and enemas; 2, the use of snuff in stress-related
disorders; 3, the rubbing of powdered medicines into
scarifications on the joints; 4, the use of charms.
Ngubane (1977) refers to the extensive use of emetics
and enemas to cleanse the body from harmful substances.
Snuff-taking would have the same effect of cleansing the
nasal passages and one group of healers, who passed snuff
around before a meeting, said they were taking it ‘to clear
their minds’.
The rubbing of powdered medicines into freshly cut
scarifications on the joints is attributed by Ngubane (1977)
to the vulnerability of these areas to evil elements.
Powdered medicines are also used for the relief of pain
or as anti-inflammants. One healer (Mr V.M.) used this
form of treatment for an ailment he referred to as
rheumatism. He said that this ailment was caused by evil
spirits.
Charms are used to ward off evil and also to procure
the goodwill or affection of others. In Xhosa and Zulu
practise, plant material may be taken, inhaled, bathed with,
sprinkled, worn or simply grown. The use of charms may
appear magical in the sense of being founded on belief
in the supernatural rather than on observed effects that can
be scientifically accounted for, but their function is a
psychological, reassuring one. The status of this sort of
categorization is liable to change, being dependent on the
state of knowledge at any particular time. For example,
the widespread use of love-charm emetics and various
fertility cures or medicines taken to procure a given gender
in the baby certainly appears to be more magically than
medicinally based. It is possible, however, that constituents
such as steroidal saponins, which are known to be present
in many of the plants used, do affect the sex hormones.
TRANSMISSION OF HERBAL KNOWLEDGE
1 Categories of practitioners
Transmission of herbal knowledge takes place within
social groups. Most of the fieldwork for this study was
undertaken in the rural areas of Transkei and the
practitioners observed could be roughly categorized into
the following groups, which are also applicable to Zulu
culture:
1.1 Villagers gather the plants used for various common
minor ailments and some charm remedies for themselves.
These plants are called by locally known common names
and are often recognized by their leaves and used before
flowers or fruit are present. In one village visited a very
young child was sent out to collect a well known purge.
He returned very quickly with a freshly dug up Ledebouria
sp. which he had identified correctly from the Xhosa
name. Some common ailment remedies such as the
influenza and cold cure Artemisia afra Jacq. ex Willd.
(Hutchings & Johnson 1986 — see also the Appendix)
appear to be used in all parts of Transkei and also in
Zululand. It is known by the same name, umhlonyane, in
both Xhosa and Zulu. Within the village, more specialised
knowledge is common to smaller groups, such as the
grandmothers who, from information apparently passed
on through the family, collect the necessary plants and
Bothalia 19,2 (1989)
227
prepare and administer medicines for their grandchildren.
These medicines include the purges deemed necessary for
the cleansing of the newly-born or weaning child from
impurities believed to be passed on by the mother. They
are known as isicakathi and iyeza-lamasi in Xhosa. Small
boys know the plant charms which may be placed in the
mouth or hair against the wrath of a teacher or father.
Older men often know which plants to use to cure or
prevent disease in stock animals.
1.2 Herbalists gather and sell or prescribe herbal
medicines and may be either men or women. As
herbalists’s children frequently gather herbs with or for
their parents, they grow up well informed in local plant
lore and often become herbalists or healers themselves.
Some herbalists support themselves by their trade, dealing
with healers or selling directly to the public, often from
street stalls. Some practise the craft for the benefit of their
family or neighbours and earn their living by other means.
1.3a Diviners (known as amgqirha in Xhosa and
izangoma in Zulu), keep contact with the ancestors, divine
the causes of misfortune and illness and may treat patients
themselves. They may also refer patients to specialist
traditional doctors. Diviners are usually, but not always,
women. They invariably receive a strong vocational calling
which they themselves refer to as a sickness, known in
Xhosa as ukuthwasa. They receive their training from
practising diviners but never from a member of their own
family, although the calling frequently comes through the
medium of a departed relative, often the diviner’s grand-
mother, who was herself a diviner. The period of training
of the ten diviners interviewed varied from eight months
to five years.
1.3b Traditional doctors (known as amaxwhele in Xhosa
and izinyanga in Zulu) sell and prescribe herbal remedies
for various ailments but do not usually divine the causes
of an illness. According to Ngubane (1977) an inyanga is
a male practitioner and a man who wants to become an
inyanga is normally apprenticed to a practising inyanga
for a period of not less than a year and the skill may be
passed on to one of his sons. The men who attended the
meeting in Umtata introduced themselves in English, using
the title ‘dtfctor’. Most of the women present wore the
traditional head gear of a diviner.
There is an overlap between traditional doctors and
diviners — many of the practitioners at the Valley Trust
meeting described themselves as being both an inyanga
and a sangoma and those interviewed in Transkei belonged
to associations of ‘traditional healers’ which issued
certificates of membership in English. The term ‘healer’
is used in this paper for both traditional doctors and
diviners unless a distinction is necessary. The locally
formed associations of traditional healers follow their own
strict codes of ethics.
Knowledge of plant usage is taught in the field by
trainers. At the meetings held at St Elizabeth’s Hospital
and Valley Trust specific plants were called by their local
names, and their usage was quite openly discussed. Some
individual forms of treatment used by diviners are revealed
to them by their ancestors through the medium of dreams.
Although some of these forms were discussed openly, the
author was also requested to discuss others privately.
Illnesses are treated with herbal remedies which may
consist of only one part of a plant or a mixture of various
parts of one or more plants. Medicinal plants may be used
fresh or may be dried in the sun and then stored in glass
containers or hung from the rafters of huts. Roots and bark
may be ground after drying. Sometimes insects or parts
of animals are used and patent medicines may also be used.
1.4 Homeopath/herbalists undergo various correspond-
ence courses in both herbalism and homeopathy. They
frequently come from families of herbalists or traditional
healers. They do not consult the ancestors and are the only
group who appear to make use of published information.
They may use either herbal or homeopathic remedies and
study subjects such as human anatomy as well as diag-
nostic techniques such as reflexology or iridology. In
Transkei they are referred to as ‘Ooquira’, the same term
that is used for conventionally Western-trained doctors.
One Transkeian homeopath running a correspondence
course from Butterworth claimed to have over 400
students.
2 Informants — background and practise
The five informatns described below were all inter-
viewed on more than one occasion and provided much of
the information on plant usage referred to in this paper.
It is difficult to assess how representative this small group
is of the ancient traditional practise. However, many of
the plants they use and customs they referred to have been
recorded in the early literature. The vocational calling of
diviners through the medium of dreams is well established
(Krige 1950; Broster 1982).
1, Mrs S.M., a 75 year-old retired teacher whose parents
had both been herbalists. She had acquired an expertise
from them which she practised for the benefit of her large
extended family and neighbours. When asked how she
knew that a plant she described had been used by the Bush-
men, she said simply ‘through the ancestors’. She was in-
troduced to the author by her son, a colleague in the Botany
Department at the University of Transkei. At the time of
the interviews she had charge of six grandchildren in a
remote rural village. She was interviewed at her home
where she provided recipes for herbal medicine, including
the recipe for swollen glands recorded in this paper (see
Recipe 1). She also accompanied the author on a plant
collecting expedition to Camana Forest near Cofimvaba.
2, Mr C.M. , a middle-aged traditional healer and owner
of herbalist shops in Lusikisiki and Flagstaff. The son of
a herbalist, he had received a calling in a dream while
working as a builder in Durban. He walked to Zululand
and found his trainer waiting for him, although there had
been no written or verbal communication. His training
took five years, during which time he had no communi-
cation with his family. He then returned to his home in
Lusikisiki, where he set up his business. He attended the
first meeting at St Elizabeth’s Hospital and was later
interviewed in one of his shops. He had started his business
by collecting all his own plants but he now purchases plants
from other collectors. He cultivates some in pots, including
Aloe aristata Haw. , which he uses for a variety of illnesses.
He trains his own assistants to dispense medicines. He said
that many of the illnesses his patients suffered from were
believed by them to be caused by umfufunyane (an evil
spirit) and thus needed to be seen to be treated by stronger
228
Bothalia 19,2 (1989)
spirits, which he claimed to keep in beaded calabashes.
This appeared to be a psychological ploy for coping with
hysterical complaints. He had a special interest in the
treatment of venereal diseases and also claimed an
expertise in the treatment of difficult and delayed con-
finements.
3, Mr V.M. , a middle-aged traditional healer who had
been practising for two years after a training of eight
months. He gave up his job as a transport manager on a
mine in order to become a healer. He was called in a
dream, in which he saw the face of his trainer, a woman
living in Ladysmith in the Orange Free State. He attended
both meetings at St Elizabeth’s Hospital and was also
interviewed twice, once at his own home. He described
how he had been trained to feel in his own body, by
concentration, the symptoms experienced by his patient.
His special interest was that of his trainer — what he
termed ‘mental’ illnesses. He said that he received a lot
of help and advice from his neighbour, a more experienced
healer who also attended both meetings. He cultivated a
few medicinal plants in his garden among which were two
Chenopodium spp. and Artemisia afra Jacq. ex Willd. He
collected others in the field and had to buy some such as
iqwili ( Alepidia amatymbica Eckl. & Zeyh.) because, he
said, it grows only in the mountains, and umavumbuka
(probably Sarcophyte sanguinea Sparrm.) because that,
he said, could only be found in Port St Johns.
Umavumbuka he used not only for diarrhoea but also
because he found it an important plant for what he
described as ‘bringing out the illness in a patient’ (see
Appendix and Recipe 1). He indicated that many of his
patients were suffering from diseases caused by sorcery
or evil spirits because of a disturbed and changing life style
and he felt that some of the children’s ailments he saw were
caused by lack of adequate parental care.
4, Mr A. B., a middle-aged labourer of mixed Black and
White parentage was the only healer interviewed who had
not been formally trained. He claimed to have been
instructed by his grandmother in dreams as to which plants
to use. He attended the first meeting at St Elizabeth’s
Hospital and was obviously well known by the group of
healers present. He was interviewed the following day at
the hospital. He brought the author a bottle of medicine,
the recipe and the plant Drimiopsis maculata Lindl.
( Hutchings 2225 KEI) used for the infant disease known
as ipleyiti, discussed later in this paper.
5, Mr F.N. a 40 year-old homeopath-herbalist whose
father had been a herbalist and taught him much traditional
usage. He later studied herbalism and home-nursing by
correspondence. He then completed a four year course in
homeopathy with a correspondence college in England.
The author met him in the University of Transkei
herbarium where he came with a query about Hypoxis spp.
He was subsequently interviewed at his surgery a number
of times and accompanied the author on a brief collecting
expedition in the vicinity of his surgery at Nqeleni in
Transkei. He uses either herbal or homeopathic remedies
but does not mix them. He employs iridology and
reflexology in diagnosis. He has a trained hospital nurse
working for him and he dispenses his own medicines.
Many of the plants he uses grow in his garden. He sends
his gardener out to collect others when needed and he buys
some from herbal vendors. He has a special interest in
cancer. He sometimes uses Hypoxis spp. corms to treat
cancer and also, for uterine tumours, the young root of
a Phytolacca sp. He attributed most of the illness he sees
in babies and adults to malnutrition and also said that he
frequently has patients suffering from hysteria caused by
the belief that they have been bewitched. In the treatment
of hysteria he often uses a tea made from Viscum anceps
E. Mey. ex Sprague (see Appendix). He said an overdose
could cause drowsiness, which passed in time, and he was
careful to avoid using the plant when in fruit.
PLANT CHARACTERISTICS AS DETERMINANTS OF USAGE
Usage of related groups of plants for similar ailments
is recorded in the literature and has been observed by the
author. While transmitted knowledge obviously determines
usage it seems that easily discernible plant characteristics
were probably the original determinants. Some of these
characteristics are outlined below with a few added
examples from cultures other than Xhosa and Zulu.
1 Characteristics that may be seen
a. Suggestive forms
Some evidence of the role of suggestive forms in plant
parts in Zulu folk medicine has been documented, mainly
in connection with procreation-related conditions. A
traditional healer’s claim to be able to cure barrenness by
the use of a corm resembling the female genitalia was
recorded and published in 1927 (Bayer & Lebzelter). The
same plant, Gloriosa superba L., has been recorded by
Bryant (1966) and Gerstner (1939) as being given to parents
wishing to have a baby of a particular gender and also as
being used as an aphrodisiac, while Hulme (1954) records
its use as a love charm emetic. The closely related
Sandersonia aurantiaca Hook, and Littonia modesta
Hook. , with corms of a similar shape, are also recorded
as aphrodisiacs (Gerstner 1939). Crocosmia and Gladiolus
spp. are aptly described by Gerstner (1941: 375) as having
a ‘string of corms grown together’ and recorded by him
as being used to treat barren women. He compares the
Zulu name of the medicine, uNdwendweni to
uDwendwe — the wedding procession. He also records
that various other Iridaceae species are carried during
planting as charms to bring fertility to the crops. Hulme
(1954: 10) records that a man, suspecting that his girl’s
love is waning, gives her an infusion of Cyrtorchis arcuata
(Lindl.) Schltr. so that ‘she will cling to him as the orchid
clings to the tree’. The epiphytic orchids frequently sold
in herbalists shops as love charm emetics or aphrodisiacs
for men are likely also to be used on account of the form.
A parallel European example is that of the ‘Mandrake’
which was known as male or female according to the form
of its roots. Desmond (1986) records that Mandragora
offtcinarum L. (Solanaceae) was frequently illustrated in
herbals, one of the earliest records known being in the
Anglo Saxon Herbal (± 1 200). The roots of the plant were
thought to resemble a human being and if pulled out were
said to emit such a scream as to cause instant insanity or
death in the collector. This could be avoided by ritual
incantations or by having the roots pulled out by a dog,
who would then go mad. The fearsome plant was highly
valued as a powerful aphrodisiac. Early parallel usage of
Orchidaceae is also known. Richter (1965) points out that
Bothalia 19,2 (1989)
229
the origin of the name orchis is from the original Greek
word for testicle. He claims that in medieval times, when
the medical Doctrine of Signatures was adhered to,
preparations from the tubers of certain orchids were
regarded as sexual stimulants and also that a child of the
required gender could be produced by using a tuber of
the right age, the younger ones being thought to procure
a male child.
b. Colour
While colour as a determinant has not been documented
in the literature surveyed for this study, the author
accompanied Mr F.N. in a search for the plant he knew
as umavumbuka which differed from the author’s
description of Sarcophyte sanguined Sparrm. He was
looking specifically for a red material and eventually
found, just below the surface of the ground, a very large
reddish swelling on the root of an Acacia karroo Hayne
tree that was neither of the two red parasites called
‘Umavumbuka’ by both the Zulu and Xhosa. These are
Sarcophyte sanguined Sparrm. and Hydnora africana
Thunb. and both are used for diarrhoea and dysentery. The
plant material was sent away for identification but no con-
clusion has yet been reached. A. karroo itself has a red
root bark and inner bark and is used as an astringent medi-
cine (Bryant 1966). It seems likely that the colour here
does play a determinant role and possibly signals the
presence of tannin, which is often present in the plants
used for sore throats or diarrhoea and dysentery, and would
be effective on account of its protein-precipitating, barrier-
forming property. Bryant describes the inner bark of the
‘uNgazi’ tree as crimson and the roots of Elephantorrhi-
za elephantina (Burch.) Skeels = E. burchellii Benth. and
an Indigofera sp. as red. All of these plants are used for
diarrhoea and dysentery. Red streaking has been observed
by the present author on Geraniaceae spp. used for the
same purpose and was observed in the 17th century by Cul-
peper (1826) on ‘Herb Robert’ (= Geranium robertianum
L.). This was used as an astringent ‘to stay the blood’. It
is also refered to by Fliick (1976) as an anti-diarrhoeal and
as used for inflammation of the mucosa of the mouth.
Fliick (1976) refers to the astringent properties and presence
of tannin as well as to the red tinges frequently seen on
the stem of Polygonum hydropiper L.
c. Plants that froth in water
Saponins are widely present in plants and may be
detected by their property of frothing in water, a property
which is made use of in the preparation of emetics.
Mention has already been made of the large number of
emetics used in traditional medicine and they are used for
a wide range of conditions, including nausea, fever,
snake-bite and coughs. They are also taken to induce the
trances needed for divining, as love charms and as
antidotes against bewitchment. Saponins have an irritant
effect on the mucosa, which is why they make effective
emetics. They are well known in the closely related
Caryophyllaceae and Illecebraceae. Silene spp. and
Dianthus crenatus Thunb. (both Caryophyllaceae) are used
as Zulu emetics (Hulme 1954; Gerstner 4666 PRE) while
Saponaria officinalis L. (Caryophyllaceae), commonly
known as ‘soapwort’ in English, is used as a mild laxative
and expectorant in Europe (Fliick 1976).
d. Mucilage
A mucilaginous exudate has been observed by the author
in the bulbs of some Amaryllidaceae species used for
wound healing and rashes and also in the purging medicine
made from the bulbs of Drimiopsis maculata Lindl. This
may be a guide to usage. Mucilage applied externally
would promote healing by forming a barrier to further
irritation. Taken internally, it has a laxative effect which
is attributed by Fliick (1976) to its property of swelling in
water. The closely related Malvaceae and Tiliaceae families
are known to be rich in mucilage (Trease & Evans 1983).
This seems likely to be the constituent utilized in the
Hibiscus and Grewia spp. used in the treatment of urinary
disorders. Bryant (1966) records that the medicine is
directly introduced through the urethral channels. Grewia
cajfra Meisn. and Sida dregei Burtt Davy are also recorded
as being used in the treatment of sores and wounds
(Gerstner 1938, 1939). Fliick (1976) records the use of
Malva neglecta Wallr. (Malvaceae) for abscesses and as
a mild purgative in Europe.
e. Milky latex
The use of Ficus sur Forssk. (= F. capensis Thunb.)
as a bovine galactalogue and Sarcostemma viminale (L.)
R. Br. as a human and bovine galactalogue would appear
to be on account of presence of the milky latex. The
practice has been described as a matter of mimetic magic
(Watt & Breyer-Brandwijk 1962). The milky latex found
in many Euphorbiaceae, Apocynaceae and Asclepiadaceae
may signal their toxicity and may have accounted for the
wide African use of various species as arrow poisons or
snake-bite cures. Gerstner (1939) records the use of
Euphorbia ingens E. Mey. ex Boiss. as a purgative given
in very small amounts. The latex is known to be highly
irritant. Watt & Breyer-Brandwijk (1962) refer to African
emetic and purgative use of Euphorbia pugniformis Boiss. ,
which may have accounted for a reported death following
medicinal administration. Culpeper (1826: 168) writes of
‘petty spurge’ (also Euphorbiaceae) as: ‘The whole plant
is full of a caustic milk, burning and inflaming the mouth
... a strong cathartic ... by reason of its sharp corrosive
quality and therefore ought to be used with caution’.
2 Characteristics that may be felt
Bryant (1966: 57—59) refers to the use of various caus-
tic plants including Ranunculus multifidus Forssk. ( =R .
pinnatus Poir.), Mikania capensis DC. and Cardiosper-
mum halicacabum L. as poultices in the treatment of
venereal sores, stating that they are ‘said to burn away all
the foulness of the ulcerated parts, leaving them clean, and
stimulating them to rapid healing’. He suggests that the
same principle is employed when Croton spp. are inserted
into the womb in cases of uterine inflammation. Vesicant
or caustic properties are known to be present in many
species of Thymelaeaceae, Euphorbiaceae and Ranuncula-
ceae. These properties are likely to account for the usage
of many species from these families for skin complaints,
wounds and sores and for their occasional use as cancer
cures. Counterirritants recorded by Bryant (1966) include
Diospyros villosa (L.) De Winter (= Royena villosa L.)
and Croton spp. The stinging properties of various
Urticaceae spp. would explain their use as sexual irritants
for cattle or men, recorded by both Gerstner (1938) and
230
Bothalia 19,2 (1989)
Bryant (1966). In Europe Ranunculus spp. are referred to
as being rubbed into the skin for rheumatism by Fliick
(1976). Fliick (1976) and Culpeper (1826) refer to the use
of Urtica spp. for rheumatism.
Plants that have been recorded as irritant to the eyes,
nose or mouth and are used for headaches or catarrh
include Andrachne ovalis (Sond.) Muell. Arg. (Gerstner
1941) and Synadenium cupulare (Boiss.) L.C. Wheeler
(Watt & Breyer-Brandwijk 1962).
3. Characteristics that may be smelled
device (pers. comm.) Gerstner (1938) records that a
number of strong-smelling Apiaceae and Lamiaceae are
grown to keep away evil spirits.
Culpeper (1826: 59) writes of ‘common feverfew’
( Chrysanthemum sp.): ‘Its unpleasant foetid smell bespeaks
it useful in hysteric disorders’. The drug Valerian, often
used in Europe as a carminative and antispasmodic in
hysteria and nervous disorders comes from Valeriana
officinalis L. (Trease & Evans 1983) and is commonly
called by English botanists ‘stinking Valerian ’ on account
of the odour that develops when the root is dried.
Scented flowers do not appear to play a role in Zulu or
Xhosa medicine, but sweetly scented or aromatic leaves
or roots are sometimes used as cosmetic or purification
washes. Cymbopogon marginatus (Steud.) Stapf ex Burtt
Davy has an aromatic rootstock and, according to the
Valley Trust group of healers, is used by the sangoma as
a purification wash after funerals and by all women as a
purification wash after menstruation. Hulme (1954)
redords that the lemon-scented Heteropyxis natalensis
Harv. is used as a perfume. The aromatic Achyrocline
stenoptera (DC.) Hilliard & Burtt (= Helichrysum
stenopterum DC.), is used by women to wash away body
odours while Helichrysum cooperi Harv. is used as a wash
by young men wishing to attract women (Watt & Breyer-
Brandwijk 1962). Helichrysum odoratissimum (L.) Sweet
is used by the Sotho to fumigate huts and to make a
pleasantly perfumed ointment (Watt & Breyer-Brandwijk
1962). This plant is burnt by the Xhosa as an incense to
invoke the ancestors and as a purification and protective
charm (Hutchings & Johnson 1986).
The principal causes of aromatic odour in plants are
volatile oils (Fliick 1976), a number of which are known
to have therapeutic or antispasmodic activity (Trease &
Evans 1983). Species from the notably aromatic families,
Rutaceae, Apiaceae, Lamiaceae and Verbenaceae, and
various aromatic Asteraceae spp. are used by the Xhosa
and Zulu for coughs, colds and influenza (see Artemisia
afra Jacq. ex Willd. in Appendix) as well as carminative
purposes. Back (1987) records that the strongly scented
Achillea millefolium L. is used in England for feverish
colds and indigestion or flatulence.
Aromatic or pungent-smelling plants used in stress-
related disorders include Clausena anisata (Willd.) Hook,
f. ex Benth., the pounded roots of which are used in an
emetic for people made ill by evil spirits or the ancestors
(pers. comm.). The peppermint-smelling roots of Monan-
thotaxis caffra (Sond.) Verde, are smoked for hysteria
(Gerstner 1939). The plant is also used as a charm against
bad dreams (Watt & Breyer-Brandwijk 1962). Ocotea
bullata (Burch.) Baill. and Cardiospermum halicacabum
L. are among the strong-smelling plants used to relieve
headaches and catarrh (Hulme 1954; Watt & Breyer-
Brandwijk 1962). A Kaempferia sp. is referred to by Watt
& Breyer-Brandwijk (1962) as good for catarrh, driving
away snakes and warding off lightning. Taken in
mealiemeal, it is supposed to keep away the effects of
drought and heat and also to protect the inyanga from the
dangerous effects of the plants collected. Garlic-smelling
Tulbaghia spp. are grown by both the Xhosa and Zulu to
keep snakes away from the homestead and smeared on the
bodies of Xhosa diviners before dancing as a protective
Strong-smelling plants are also often used by the Xhosa
and Zulu as anthelmintics e.g. Clausena anisata (Willd.)
Hook. f. ex Benth. and Clerodendrum glabrum E. Mey.,
or as insect repellants e.g. Cymbopogon marginatus
(Steud.) Stapf ex Burtt Davy (= Andropogon marginatus
Steud.) and Andrachne ovalis (Sond.) Muell. Arg. (Watt
& Breyer-Brandwijk 1962; Gerstner 1941). The strong-
smelling Chrysanthemum parthenium (L.) Bemh. is an
English insect repellant (Back 1987).
4 Characteristics that may be tasted
The sour- tasting leaves of Embelia ruminata (E. Mey.
ex A. DC.) Mez and a Pavetta sp. are chewed as a tonic
(Gerstner 1938). The roots of Mondia whitei (Hook, f.)
Skeels, chewed for the relief of indigestion, taste first bitter
and then sweet (Gerstner 1941). Bryant (1966) records that
the very bitter Vemonia adoensis Sch. Bip. ex Walp. (=V
woodi O. Hoffm.) is said to be a useful stomachic. A
number of Aloe spp. are recorded by Gerstner (1941) as
being applied to the mother’s breast at weaning. The known
bitter taste of many species would surely account for their
use in discouraging suckling. Any fieldworker who has
collected Asclepiadaceae spp. would understand the use
made of the bitter latex when applied to eggs to deter dogs
from stealing them as recorded by Hulme (1954).
The bitter taste of aloes was recorded by Culpeper
(1826). The purgative properties of Aloe spp. are noted
in the Oxford English Dictionary, with English citations
going back to the 14th century and etymological evidence
going back to Latin and Greek. The use of the bitter
Artemisia absinthum L. and A. vulgaris L. as digestion
stimulants was recorded by Fliick (1976).
5 Conclusion
Jensen & Nielsen (1984) point out that chemistry has
always been used in the classification of plants, exemplify-
ing chemical characters by the colour, taste and smell of
various parts of the plant. It would seem from parallel
usage of related plants in African and European herbal
practice that appropriate usage may be widely determined
by easily discerned plant characteristics.
PLANT USAGE IN TRANSKEI
Medicinal plants were collected and their usage recorded
by the author in Transkei from 1983—1987 (see Appendix).
The categories of informants interviewed include the
following:
Bothalia 19,2 (1989)
231
Villagers: (a) villagers encountered by chance on collecting
trips; (b) some inhabitants of Ndunguniyeni Village in the
Engcobo area of Transkei; (c) staff and students who either
brought the author plants or were with the author when
plants were collected. They recalled plant usage from their
earlier experience in the rural areas.
Herbalists: (a) Mrs S.M. (informant 1 above); (b) a
herbalist who visited the herbarium and discussed various
medicinal plants on display; (c) street vendors from whom
plants were bought by the author.
Healers: (a) Mr C.M. (see informant 2 above); (b) Mr
V.M. (see informant 3 above); (c) healers attending the
St Elizabeth’s Hospital and Valley Trust meetings.
Homeopath/herbalist: Mr F.N. (see informant 5 above).
All the plants in the Appendix, with the exception of
Sarcophyte sanguinea Sparrm. and the Hypoxis spp. , were
collected or observed by the author in the company of an
informant. The anti-diarrhoeal use of Sarcophyte
sanguinea was recorded from many informants and its
Xhosa name was confirmed from a pickled herbarium
specimen ( Johnson 222 KEI). This had originally been
collected for the herbarium by a traditional healer from
Kei Mouth. Hypoxis spp. were shown to Mr F.N. so that
he could confirm his identification of the genus and were
also discussed by their Zulu names at the Valley Trust
meeting. The plants listed in the Appendix were selected
on account of properties which appear to the author to
indicate a possible appropriate usage.
RECIPES FOR HERBAL MEDICINES
Recipe 1: medicine for swollen glands
The following recipe for swollen glands was given to
the author by Mrs S.M. (informant 1 above) in the
presence of her son, who is a botanist. Mrs S.M. said that
the medicine was also effective against cancerous growths.
Unfortunately the four ingredients were recorded during
a drought and not collected. It was possible to make
hypothetical determinations from the Xhosa names and
the plant descriptions. The recipe is included for its
information on method. Although the plants used are
different, the way in which the medicine is prepared and
used is similiar to that described by Bryant (1966) for the
treatment of tumours ascribed by him to scrofula.
Method: mash and boil the roots of isinama and amaselwa
and the rootstock of umavumbuka to make one litre of pulp.
Place the cooking pot while it is still hot on a folded shawl
on the patient’s head for neck glands, chest for armpit
glands and stomach for groin glands. When cool enough
use the mixture directly as a poultice on the affected
glands. When further cooled, add two teaspoons of
imithombo as a ferment and give the patient two spoonfuls
as necessary.This medicine has a bad taste and may be
diluted.
The four ingredients are interesting. Isinama was
described as sticking to clothing and is likely to be the
common weed Achyranthes aspera L. (Amaranthaceae),
which is also called isinama by the Zulu. Watt & Breyer-
Brandwijk (1962) record that the leaf and seed are applied
in India to inflamed and enlarged glands. Oliver-Bever
(1986), quoting Neogi et al. (1970), refers to the diuretic
and slightly anti-pyretic properties of achyranthine, the
betaine derived from the plant. She also tables the
anti-leprosy action of the seeds produced by the oleanolic
glycoside, referring to the work of Gopalachari & Dhar
(1958) and Ojha et al. (1966).
Amaselwa was described as the calabash creeper and
identified by the informant’s son as being either Lagenaria
siceraria (Molina) Standi. orL. sphaerica (Sond.) Naud.
(Cucurbitaceae). A pounded root decoction of L. sphaerica
has been recorded by Hulme (1954) as being used by the
Zulu for treating a swollen body caused by some blood
disorder. Watt & Breyer-Brandwijk (1962) refer to a small
form of L. siceraria which is thought to contain amygdalin
and so to be cyanogenetic. Another member of the family,
Momordica charantia L. is referred to in the following ex-
tract (Sofowara 1982: 208): ‘With alcoholic extracts of the
stems, leaves and fruits Abbott et al. (1966) demonstrated
remarkable anticancerous action on mice with transport-
able 180 tumours. Aqueous extracts of the roots also proved
effective in reducing the tumour’.
Imithombi is a solidified fermented paste made from the
fruit of a cultivated Sorghum sp. The stem pith has been
recorded as being used by the Xhosa to treat tubercular
swellings (Watt & Breyer-Brandwijk 1962). Sorghum is
one of the genera cited as containing cyanogenic
compounds and free HCN by Nartey (1981: 73). He writes:
‘cyanogenic glycosides are reputed to possess some
therapeutic properties against cancer. Their action against
cancer cells produces large amounts of B-glucosidases, so
that HCN produced by the enzymic cleavage of, for
example amygdalin and prunasin, exerts its full inhibitory
influence on the growth of neoplastic cancer cells’.
Umavumbuka, described by the informant as having a
red rough-textured rootstock with red sap and small white
flowers and growing on the roots of old trees, is likely
to be the parasite Sarcophyte sanguinea Sparrm., referred
to in the Appendix as a diarrhoea remedy. While no
chemical research on this plant is known to the author,
another parasite, Hydnora johannis, from the Sudan, has
been found to have a high concentration of phenolic
compounds in the roots, imparting an astringent quality
which would account for its antidiarrhoeal use (Visser &
Musselman 1986). Hydnora africana Thunb. is also known
to the Zulu as umavumbuka and is similar in colour and
habit to S. sanguinea, both being found on the roots of
trees. A species from another parasitic genus, Viscum has
been used in Africa for the removal of warts (Watt &
Breyer-Brandwijk 1962) and it has been recorded that V.
album L. may inhibit the growth of certain tumours if
applied directly on or into the tumour (Fliick 1976).
Recipe 2: medicine for ipleyiti
Ipleyiti is an alleged disease of newly bom and very
young infants described variously by several informants
as ‘producing an old look in the newly bom’, ‘producing
green veins stretching from the arms to the stomach’, ‘the
result of a placenta formed like an enamel plate’,
‘producing much crying and green stools’. It is a condition
frequently treated by traditional healers. It was ascribed
by one healer, Mr V.M. to unsuitable behaviour on the
part of the mother, such as going to too many drinking
232
Bothalia 19,2 (1989)
_ 2
2 '5
I'f
3|
U .£
f 1
C 3
< T3
e c
2 |
< 2
■8 .5*
II
i£3 a
§=c
“ a
-o o
g£
li
1!
II
1.
O -o
3 o
4-. U
O x>
<2 §
is -5
.3 CO
•a g-
•e 1
a e
o
l; ^
<u • -
op S
a -a
>1
Cd
K >3
as
a .o
5,3 :
y
W
”8 — :
"O *3
8 «
2 S
si
2 d
-a 5
c ^
c3
g 8 ^
o
5
I S
S 3
a %
ii
5 “
u §
Go
g
X
-2
5 -~
■S y
f 1
J I
•5 e
03
1S
< $
~ 2
13 00
3 -a
S 8
1 a
•e 60
e >
<L> w
cu'3
o 3
-o
■go
■a s
2 «
-o .5
C W;
O <£>
11
*s o
1 >
■f.fi
ts 3
•2 D.
■a g
■e I
£ o
X j=
^ 8
£
.« < 3
•3 ^
S -6
■8 g.
3 2
« 00
T3 £
a'-a
£ £
M £
S2
a i
'C
DO \£>
1) ^
e
<L> —
£ 3
X **
<U <*>
<3 s
3 £
.a s
X -r
a 6
SI
3 «
ffl y
0)
2 S
D CJ
II
§,£
&
§
> 0)
- >
"3 cS
773 DO
O fa
•° d,
a cs
3 CA
CQ cd
■B &
< '5
CJ CN
<U vO
s a
u
=3 **
O *3
IS
g 1
O o-
a b
D cd
Dm oo
cd _0J
^ ‘3)
O
■8 ^
cd Om
Q cd
•J c/3
? T *
e .a I
Cd <L>
* s
■a 8
S3 §
§ 3 Si
|||
§ X oo
.2 tt « «
s ^ 'S . .
<t
■p <5
2 ^
a <n
"8 o
| ^
2 2
e S >>
s "3
Fo
2 8 3
s |i
111
■a ■£, «
c — .E
y t2 .2
S-S-S
•* as
■g c .
§ o a
g js :s
o 3 S-
_ ^
3 c
2 .2
S (S
O D
.fr'S
0 c
1 g>
i!
^ 00
!i„
s a «
JlE
OS O co
u u
I s
.§ “
si
<D
g D
o *c3
X J§
1?
8 E
E -c
° 1
3 E
i §2
1 oo1*-
1 -a -S t«
! glS
s S3 8.§
a a a
11^^
I -S
Sm^
t: w-
cd «P
(D ^
-C T3
»- c
O ^
it.
3 8 S
g £1
.2 5 2
IJS
s feN
• > >>
£ 3
* Personal communication from Noristan Laboratories.
Bothalia 19,2 (1989)
233
if?
S S
Sic
<u ^
g3^
•53 «u
i- <d
<
E .8
< S
u so
i- oo
>, ON
•i S'
c 00
< 2
c £
5 2
2 2
i s
c »
E- _•
“ I
ao "
—
111
S
s 5. £
•o 5
I g -
§ £ 00
^ oo>
i— 1 CJ —
c _
.2 e
W cd
~X
Ql, cj
O 2
& <j
5 d?
T3 O
4) <U
£ i
O o
w)S:
a <n
■5S CM
T3 ^
<U <50
« -S
.> -r
C B ^
W o
- -O o
£ ^ -ri
U ^
3 0) c
U^3 OC H
||
JD S
O ^
c .5
> -r
>
2 =s
O S:
c &
« -o
CD
I §
c e
i 3
c 0X1
o
g o c
,J) o "§
DC T3 >S
"8 _
3 §
C E
*£ ‘3
Cd <3
E -o
o «
5 £ to
X) _ gv
3 TJ £
•s 8-S
§.s *
2 — «
“><S *S
■n c c3
S o ”0
•r o cd
Q 8-^
E -s
.E S fc
w C cd
c ^
<D c
^ 2
<D <U 4)
Op « g
a «
o 2
2 E
-a "2
u S
3 §
I £
3 JJ
|.E
u "o
T3 c
£ 3
8. g3
■81
O I
> ^
<u
> <D
o .E
O. -j;
u |
Q .5
i I
Q .S
1.3
cC -3
£3 T3
H |
§<§
U .E
a* o
= 1
> j*
.2 ■£
> o.
, V
k y
£
■3 „
^ =3
J g3
5 E
c '3
££
a 2 |
I a J
I? §
^ 3 |
<n O ^
ago
§> g
§ « „
10 3 ^
s® |
£> -r -S
Schinus molle L.; Herbalist Leaf infusion inhaled or drunk for cold and Observed in garden Aromatic, pungent-tasting. Analgesic*; anti-inflammatory*; anti-
Anacardiaceae; pepper influenza Resin, volatile oils* hypertensive*; antidepressant*; anti-
tree arrythmia*
234
Bothalia 19,2 (1989)
J-8
£ s
•r '■o
s 5;
60 °0
1 I
2 "§
5 S:
I ’2
<1
•S'S
II
>- .s
•§ 1
.2 a
•■3 <S
s«
e-1
I a
£ 8 ■
.2 ^
a S3
■SeSj
i|-s I
S -§ I- a
a a 2 §
3 ffl y
S c
vj a
a ^
C3 *
Sfj
a* g
3©
>-v 3
4) •—
.^S
S o3
4) w
o §•;?
8-r= -C
*&y
■S !s?
2. u
4) i— 1
O
« 5
• - a
c =5
;§
o v.
sz ^
.a S oo
55 .5 25
C "S
1 1
o- a
51
•a g.
£ -s
a s-
lit
o jg
o *C3
c -c
'cd T3
D- C
52 'G
E S
2 £
x cd
5 T3
3 OJ
O .>
w) td
73 g? c*
4) 3 4)
r a-c
o
cd cd
S' u
S 3
o
• ed
W o
«0 I-T*
§■>
§ s
a §)
k 2 -Si
a co23
G on ^
•2 *
^ S -2
ir "5
a) i«
<5
11
| -o
1 -a
a 2
§ 3 2
s-^ do
fc •- c
121
\%i
& J a
Bothalia 19,2 (1989)
235
parties before giving birth, or to sorcery. The homeopath/
herbalist, Mr F.N. said that the cases he sees are either
cases of colic or are babies bom to mothers suffering from
malnutrition. One informant from a village said that the
disease had come from Zululand. This belief is also
mentioned by Broster (1982), who confirms the high
incidence, the deformed placenta and the attribution to
sorcery. She states that a baby suffering from the disease
is usually bom prematurely and remains sickly. The
disease is also known among the Zulu, and Ngubane (1977)
suggests that the Zulu term ipleti is used in a manner that
suggests deprivation and starvation.
The recipe for medicine for ipleyiti was provided by Mr
A.B. The plant used is Drimiopsis maculata Lindl. and
the author was able to see the medicine as well as to collect
the plant ( Hutchings 2225 KEI). The medicine is known
as nstwilisa or nomatyuntyuma.
Method : crush the bulbs and add cooled, boiled water (+
4 bulbs to 250—300 ml water, depending on size of bulbs).
Add a pinch of salt to preserve the medicine, which will
last for abojut ten days. The dose varies from one teaspoon
to a tablespoon as required. For older children, a spoon-
ful of Epsom salts may be added.
The medicine was very slimy, indicating the presence
of mucilage. Mr V.M. confirmed that he used the same
plant for treating ipleyiti. The bulbs of Drimiopsis macula-
ta are recorded by Hulme (1954) as being used by the Zulu.
They are steeped in water to make an enema for young
children with stomach trouble.
ACKNOWLEDGEMENTS
The research for this study was partly undertaken
through Zulu Folk Medicine Research, funded by the De
Beers Chairman’s Fund Educational Trust who are thanked
for their support. The author is grateful to the University
of Transkei for enabling her to pursue her interest in
medicinal plants while she was working on the Wild
flowers of Transkei Project (1982—1987). This paper could
not have been written without the help of the informants
and the author is most grateful to them and wishes to thank
them all. Prof. G.J.M. Hutchings, Mrs Annetjie Kemp,
Mrs Gillian Lewis and Prof. S.E. Terblanche of the
University of Zululand are all thanked for their help and
encouragement. Dr T.G. Fourie of Noristan Laboratories
is thanked for permission to use the information on some
of the plants tested by the firm.
REFERENCES
BACK, P. 1987. The illustrated herbal. Octopus, London.
BAYER, F.A.H. & LEBZELTER, V. 1927. Treatment of barrenness and
venereal disease amongst the Zulus. Anthropos 22: 287-289.
BROSTER, J. 1982. Amagqirha. Via Africa, Cape Town.
BRYANT, A.T. 1966. Zulu medicine and medicine-men. Struik, Cape
Town (originally published in 1909 in the Annals of the Natal
Museum).
CHIEJ, R. 1984. The Macdonald encyclopedia of medicinal plants. Mac-
donald, London.
CULPEPER, N. 1826. Culpeper’s complete herbal ... Gleaver, Man-
chester (originally published 1653).
DESMOND, R. 1986. Wonders of creation. The British Library, London.
DREWES, S. & LIEBENBERG, R.W. 1983. Extracts of plants from the
Hypoxidaceae family for treatment of cancer. Eur. Pat. Appl.
EP2226a 26 Oct 1983. From Chemical Abstracts 100(9): 61782 m.
FLUCK, H. 1976. Medicinal plants. Foulsham, London.
GERSTNER, J. 1938. A preliminary checklist of Zulu names of plants
with short notes. Bantu Studies 12: 215—236, 321—342.
GERSTNER, J. 1939. A preliminary checklist of Zulu names of plants
with short notes. Bantu Studies 13: 49—64, 131—149, 307—326.
GERSTNER, J. 1941. A preliminary checklist of Zulu names of plants
with short notes. Bantu Studies 15 : 277- 301, 369 — 383.
GOPALACHARI, R. & DHAR, M.L. 1958. Studies in the constitution
of the saponins from seeds of Achyranthes aspera. Identification
of the sapogenin. Journal of Scientific and Industrial Research,
India 17B: 276 -278.
HULME, M.M. 1954. Wild flowers of Natal. Shuter& Shooter, Pieter-
maritzburg.
HUTCHINGS, A. & JOHNSON, C.T. 1986. Glimpses of a Xhosa Her-
bal. Veld & Flora 72 : 59 -62.
JENSEN, S.R. & NIELSEN, B.J. 1984. Chemical characters. In R.M.T.
Dahlgren, H.T. Clifford, & P.F. YEO. The families of the
Monocotyledons : 17—22. Springer-Verlag, New York.
KRIGE, E. 1950. The social system of the Zulus, 2nd edn. Shuter &
Shooter, Pietermaritzburg.
NARTEY, F. 1981. Toxicological aspects of cyanogenesis in tropical food-
stuffs. In R.L. Smith & E.A. Balocbunni, Toxicology in the
tropics: 53—73. Taylor & Francis, London.
NEOGI, N.C., GARG, R.D. & RATHOR, R.S. 1970. Preliminary phar-
macological studies on achyranthine. Indian Journal of Pharmacy
32: 43-46.
NGUBANE, H. 1977. Body and mind in Zulu medicine. Academic Press,
London.
OJHA, D. , TRIPATH, S.N. & SINGH 1966. Role of an indigenous drug
{Achyranthes aspera) in the management of reactions of leprosy.
Leprosy Review: 37, 115. Indian Council of Medical Research Pub-
lications, 1966.
OLIVER-BEVER, B. 1986. Medicinal plants in tropical West Africa.
Cambridge University Press, Cambridge.
RICHTER, W. 1965. The orchid world. Studio Vista, London.
SMITH, A. 1895. A contribution to South African materia medica, 3rd
edn. Juta, Cape Town.
SOFOWARA, A. 1982. Medicinal plants and traditional medicine in
Africa. John Wiley, Chichester.
TREASE, G.E. & EVANS, W.C. 1983. Pharmacognosy, 12th edn. Bail-
liere Tindall, London.
VISSER, J.H. & MUSSELMAN, L.J. 1986. The strangest plant in the
world. Veld & Flora 72: 109—110.
WATT, J.M. & BREYER-BRANDWUK, M.G. 1962. The medicinal and
poisonous plants of southern and eastern Africa, 2nd edn. Living-
stone, London.
Bothalia 19,2: 237-261 (1989)
Invasive alien woody plants of Natal and the north-eastern Orange Free
State
L. HENDERSON*
Keywords: alien, coastal communities, forest, grassland, KwaZulu, Natal, Orange Free State, savanna, survey, invasive plants (woody)
ABSTRACT
The frequency and abundance of invasive alien woody plants were recorded along roadsides and at watercourse crossings
in 87 % (152/175) of the quarter degree squares in the study area. The survey yielded 130 species of which the most prominent
species (in order of prominence) in roadside and veld habitats were: Chromolaena odorata, Solarium mauritianum, Psidium
guajava, Rubus spp.. Acacia meamsii and Lantana camara. The most prominent species (in order of prominence) in streambank
habitats were: Acacia dealbata, A. meamsii and Salix babylonica.
The greatest intensity of invasion was recorded in the Natal midlands and in the coastal belt of southern Natal, including
the metropolitan areas of Pietermaritzburg and Durban. There was relatively little invasion in the north-eastern lowlands
of Natal but the potential for expansion is great. Little invasion was recorded in the north-eastern Orange Free State except
along some watercourses.
UITTREKSEL
Die frekwensie en volopheid van uitheemse houtagtige indringerplante is langs paaie en by oorgange oor waterlope in
87% (152/175) van die kwartgradevierkante in die studiegebied aangeteken. Daar is 130 indringers aangetref waarvan die
mees prominente (in volgorde van prominensie) Chromolaena odorata , Solanum mauritianum, Psidium guajava, Rubus spp.,
Acacia meamsii en Lantana camara langs paaie en in veldhabitats was. Die mees prominente spesies (in volgorde van
prominensie) langs stroomoewers was Acacia dealbata, A. meamsii en Salix babylonica.
Die ergste indringing is in die Natal middelland en in die kusstreke van suid-Natal aangetref, met insluiting van die stede-
like gebiede van Pietermaritzburg en Durban. Daar was betreklik min indringing in die noordoostelike laagland van Natal
maar die potensiaal vir uitbreiding is groot. Behalwe langs waterstrome, is daar min indringing in die noordoostelike Oranje-
Vrystaat aangeteken.
CONTENTS
Introduction 237
Survey history and objectives 237
The study area 238
Method 239
Sampling method 239
Abundance ratings 240
Sampling level envisaged and achieved 240
Data treatment — formulae used 241
Frequency 241
Prominence value 241
Mean species abundance rating in roadside
and veld habitats 241
Mean abundance of invaders per km in road-
side and veld habitats 241
Results 242
The streambank habitat 242
The whole study area 242
Analysis according to veld type 242
Analysis according to species 242
Frequency 242
Prominence 243
Roadside and veld habitats 243
The whole study area 243
Analysis according to veld type 243
* Plant Protection Research Institute, Department of Agriculture and
Water Supply; stationed at Botanical Research Institute, Private Bag X101,
Pretoria 0001.
MS. received: 1988.11.04.
Analysis according to species 243
Frequency 243
Prominence 243
Patterns of invasion 243
Discussion 243
Sampling 243
Prominent and potentially important species 254
Relation of invasion to environmental factors .... 258
Conclusion 259
Acknowledgements 259
References 259
Appendix 260
INTRODUCTION
Survey history and objectives
This study, which covers Natal and the adjacent north-
eastern Orange Free State is the second of eight regional
surveys which together are designed to reflect invasion by
woody alien plants in the Republic of South Africa as a
whole. The survey method was developed during a study
of the south central region of the Transvaal (Wells et al.
1980) and then used in a survey of the rest of the Transvaal
(Henderson & Musil 1984). This survey of Natal and the
north-eastern Orange Free State was undertaken during
the 1986/87 summer season.
The objectives of the survey are: to produce a checklist
of the major invasive alien woody plants of streambank.
238
Bothalia 19,2 (1989)
| ~| grassland biome
| ~1 SAVANNA BIOME
PMB • PIETERMARITZBURG
,..A INTENSIVE SITE
SURVEY ROUTE
FIGURE 1.— The study area, Grass-
land ,and Savanna Biomes, in-
tensive sites and survey routes.
roadside and veld habitats in the study area; to determine
the pattern of alien woody invasion as a whole and for in-
dividual species; to attempt to relate distribution to
environmental factors and to determine which are the most
prominent and potentially important invaders.
The study area
The study area comprises Natal, including KwaZulu,
which together occupy an area of 86 967 km2
(Macdonald & Jarman 1985) and the extreme north-eastern
Orange Free State which covers an area of approximately
17 400 km2. It lies between latitudes 26° and 31°S and
longitudes 28° and 33°E (Figure 1). From east to west
the altitude rises rapidly, from sea level on the Indian
Ocean coastline of Natal, in a series of terraces more or
less parallel to the coast, to greater than 3 000 m on the
Drakensberg escarpment. The western boundary of Natal
is formed by the Drakensberg mountain range. The
north-eastern Orange Free State lies to the west of the
Drakensberg range at elevations of 1 500 m to 1 800 m
on the highveld plateau.
The landscape of Natal is rugged and dissected by the
valleys of 11 major rivers which arise in or adjacent to
the province. The terrain is level in the north-eastern
coastal region where numerous pans and lakes have de-
veloped.
Rainfall varies from 300 to 600 mm in some of the drier
river valleys and in the north-eastern interior region to over
1 500 mm along the coast and on some of the peaks in
the Drakensberg. Rain falls mostly in summer from
November to March but on the coast up to one third of
the rain falls during the winter period (Haigh & Wilhelmij
1973; Francis 1977).
The climate of the coastal belt and lowland interior is
subtropical to tropical. From east to west, with rising al-
titude, the climate becomes progressively more temper-
ate. The incidence of frost varies from light to moderate
in the mistbelt (600—1 200 m) and moderate to severe in
higher areas [Poynton (1972) — silvicultural map of the
Republic of South Africa]. Snow falls most years at
elevations of over 1 200 m and is a regular winter feature
along the Drakensberg (Haigh & Wilhelmij 1973).
There are two major indigenous vegetation types or
biomes, Savanna and Grassland, in the study area
(Rutherford & Westfall 1986) (Figure 1). Twenty Acocks
Veld Types (Acocks 1988) occur in the study area and have
been grouped into six veld type categories for the purposes
of this survey. These are temperate grassland, moist
subtropical grassland, dry subtropical grassland, mistbelt
grassland, all falling within the Grassland Biome; tropical
bush and savanna, and tropical forest (Table 1 and Figure
2) are classified under the Savanna Biome.
Bothalia 19,2 (1989)
239
TABLE 1. — Veld type categories in the study area and the equivalent Acocks Veld Type groupings and Veld Type numbers
Temperate grassland occupies the highest and coldest
parts of the study area at elevations of 1 500 m to greater
than 3 000 m. Rainfall ranges from 700-1 000 mm per
annum. Moist subtropical grassland occurs on the eastern
slopes and foothills of the Drakensberg from about 1 350
m to 2 150 m. Rainfall ranges from 750-1 500 mm per
annum. Dry subtropical grassland lies at elevations of
between 900 and 1 500 m and rainfall ranges from
600-1 000 mm per annum. This grassland has been
invaded by woody thomveld species especially in its drier
parts (Edwards 1967). Mistbelt grassland occupies rolling
misty country at elevations of between 450 and 1 350 m.
Rainfall ranges from 750—1 300 mm per annum. Tropical
bush and savanna occupies the hot and dry river valleys
and the north-eastern interior lowlands. Rainfall ranges
from 300—800 mm per annum. Tropical forest is the major
vegetation type of the warm and moist coastal belt. Rain-
fall ranges from 900—1 500 mm per annum. Little remains
of the forest between Durban and the Transkei border to
the south, but extensive patches exist on the north coast,
particularly north of Lake St Lucia Estuary.
METHOD
Sampling method
The method used in this survey was basically the same
as that used in the Transvaal (Henderson & Musil 1984)
J MOIST SUBTROPICAL GRASSLAND
mm
TROPICAL BUSH A SAVANNA
FIGURE 2. — The six broad veld type
categories in the study area (af-
ter Acocks 1988).
Idry subtropical grassland
TROPICAL FOREST
240
Bothalia 19,2 (1989)
but with minor changes to the abundance ratings (see next
subheading). The presence and abundance of all
naturalized alien trees, shrubs and conspicuous climbers
were recorded for each veld type category, habitat type
(roadsides and adjoining veld, and streambanks) and
quarter degree square traversed by road. Although the
objective of the survey was to record woody species, other
large non-woody and succulent species were included
rather than lose valuable information.
Recordings of roadside and veld invaders were made
from a moving vehicle whereas recordings of streambank
invaders were made at watercourse crossings. Abundance
estimates of roadside and veld invaders were based on
frequency of encounter within road transects of five to ten
kilometres in length. Abundance estimates of streambank
invaders were based on estimates at specific sites.
The width of road transects and length of watercourses
scanned for invaders varied according to local conditions.
Usually no more than 50 m of veld and 100 m of stream-
bank habitat were scanned on either side of the road for
invaders. Species occurring beyond these ranges and along
watercourses which were not crossed were recorded as
present in the given habitat type and veld type category
but were not included within the formal recordings.
Although Henderson & Musil (1984) suggested the use
of a single road transect length of ten kilometres, a variable
transect length of between five and ten kilometres was
found to be more practical. This applied particularly in
the highly dissected veld type categories in the midlands
and lowlands of Natal. In these regions it was often not
possible to accommodate ten kilometre transects and where
there was much invasion a shorter transect length proved
to be more manageable. Since all road transects were
plotted on maps the survey is easily repeatable despite the
use of variable transect lengths.
Seventeen quarter degree squares were selected for more
intensive surveying (Figure 1). These intensive sites were
selected primarily to ensure that representative parts of
each veld type category and geographical region in the
study area were well sampled. They may also be used at
a later date for a quick resurvey of the study area to assess
any changes that may have taken place. In each of the
seventeen sites abundance recordings of roadside and veld
invaders were made along a total road length of
approximately 30 km (six transects each five km long).
Recordings were made at virtually all watercourse
crossings. Herbarium specimens of all invader species
which were flowering or fruiting, were collected.
Survey routes and road transects were plotted on
1:250000 maps (general survey area) and 1:50000 maps
(intensive sites) before a field trip was undertaken.
Wherever possible two or more road transects were plotted
per quarter degree square. In most instances road transects
were not contiguous but were separated by a distance of
between five and ten kilometres. This approach was
adopted mainly due to time considerations since abundance
estimates along road transects can be very time-consuming
depending on the intensity of invasion. In addition, it was
necessary to have a breathing space between transects not
only to avert eye strain but to free one’s attention from
the immediate roadside and to observe invasion further
afield. Road transects along national roads and other routes
with heavy traffic were kept to a minimum. All road
transects to be sampled were plotted before a field trip was
undertaken to ensure non-selective recording in the field.
Recordings were made at most bridges over watercourses
but some were omitted because of time constraints and
traffic considerations.
Abundance ratings
Minor changes were made to the abundance ratings for
roadside and veld habitats used in the Transvaal survey
(Henderson & Musil 1984). The two ratings below the old
rating T were removed. These ratings became obsolete
as a result of the standardization of road transect length
from 5—10 km. The abundance ratings for roadside and
veld habitats and streambank habitats are given in Table 2.
Sampling level envisaged and achieved
The sampling level envisaged was at least 60% of the
total quarter degree squares at an average of 33 km per
square which was achieved in the Transvaal (Henderson
& Musil 1984). The sampling level achieved in this survey
was 87 % (152 of the total 175 quarter degree squares) at
an average of 38 km travelled per square. An average of
TABLE 2. — Abundance ratings
* Approximate numbers of individuals/groups per 10 km transect.
Bothalia 19,2 (1989)
241
TABLE 3. — Sampling coverage in each veld type category, biome and the study area
* This represents the distance along which abundance recordings were made. Total distance along which observations were made is approximately
twice that given.
17 km of road transects were sampled per quarter degree
square for abundance estimates of roadside and veld in-
vaders.
The veld type coverage in terms of quarter degree
squares and road transects sampled, kilometres travelled
and watercourse recordings made, is given in Table 3.
Data treatment — formulae used
Frequency
The percentage frequency of occurrence of a species x
in veld type category y was calculated as follows:
no. of watercourse recordings/road transects
in veld type y having species x
frequency = x 100
total no. of watercourse recordings/road
transects in veld type y
Prominence value
Prominence is used here in preference to the term
Importance defined by Henderson & Musil (1984). The
prominence value, which has been derived from Curtis’
Importance Value (Mueller-Dombois & Ellenberg 1974),
is a measure of the prominence (in terms of frequency and
abundance) of a species in a vegetation category relative
to all other species in the same category. Other aspects,
such as rate of spread and difficulty of control, which
should be taken into account when assessing species
importance, are not included here, hence the preferred use
of the term prominence.
In streambank habitats the prominence value for a
species x in veld type category y was calculated as follows:
frequency of species x in veld type y
scoring 5, 6 or 7
x 100
sum frequency of all species in veld
type y scoring 5, 6 or 7
prominence value = +
frequency of species x in veld type y
sum frequency of all species in veld
type y
* each abundance rating was expressed in numbers of individuals/groups
recorded per transect (see Table 2). To be both conservative and consistent
the minimum number was used in each instance, e.g. an abundance rating
of 5 over ten kilometres = 50 and an abundance rating of 5 over five
kilometres = 25.
** mean no. of individuals/groups per 10 km converted to rating (see
Table 2).
The selection of abundance rating 5 as the cut-off point
is arbitrary but one at which a species can be regarded
as locally prominent (see definition in Table 2). A formula
using all abundance ratings would be preferable if each
rating could be converted to an absolute value.
In roadside and veld habitats the prominence value for
a species x in veld type category y was calculated as
follows:
total abundance* of a species x in veld
type y
x 100
sum of the abundances* of all species
in veld type y
prominence value = +
frequency of a species x in veld type y
x 100
sum frequency of all species in veld
type y
The highest prominence values in a given category
which add up to approximately 160 points out of a total
of 200 are printed in bold in Tables 6 and 7. The cut-off
point of 160 points is arbitrary but represents 80% of the
summed prominence values.
Mean species abundance rating in roadside and veld
habitats (see Table 7)
The mean abundance rating** of species x in veld type
category y was calculated as follows:
mean no. of total no- °f individuals/groups of species x in
individuals/ ve^ lyPe y lfl
groups per total distance along which species x was rated
10 1™ in veld type y
Mean abundance of invaders per km in roadside and veld
habitats (see Table 5 and Figure 5)
The mean abundance of invaders per kilometre in veld
type category y/quarter degree square z was calculated as
follows:
total abundance* of all species in veld type
y/quarter degree square z
mean abundance =
total kilometres rated for abundance estimates
in veld type y/quarter degree square z
242
Bothalia 19,2 (1989)
TABLE 4. — Streambank statistics for each veld type category, biome and the study area
* 1 or more species scored 5 or more. ** invaders present.
RESULTS
The survey yielded 130 naturalized alien species. These
species are listed in the Appendix together with a further
50 species which were obtained from various literature
and other sources. The distributions and high abundance
areas of 32 of the most prominent species are given in
Figures 6 & 7 (see Appendix).
The streambank habitat
The whole study area
Five hundred and seventy eight watercourse crossings
were sampled in which 90 species were recorded, with
up to 13 species in one sample (Table 4). Invaders were
present at 81.7 % of all crossings and 24,9% of all crossings
were heavily invaded (Table 4).
Analysis according to veld type
More alien species were recorded in the Savanna Biome
than in the Grassland Biome. The most species were
recorded in tropical forest and the least in temperate
grassland.
In the Grassland Biome there was a progressive increase
in the severity of invasion in terms of percentage crossings
heavily invaded with decreasing elevation from the
montane region (temperate grassland) to the mistbelt. The
rivers in mistbelt grassland and tropical forest south of
Durban were the most invaded in terms of percentage
crossings heavily invaded and percentage crossings
invaded. The rivers of tropical bush and savanna were the
least invaded (Table 4).
Analysis according to species
Frequency
Only eight species were recorded at 10% or more
crossings in the whole study area (Table 6). Acacia
dealbata and Salix babylonica were the most frequently
encountered species.
In the Grassland Biome the most frequently recorded
species were Acacia dealbata, Salix babylonica and Acacia
meamsii. Ricinus communis was by far the most frequent
invader in the Savanna Biome followed by Psidium
guajava, Chromolaena odorata, Solanum mauritianum
and Melia azedarach which were almost equally frequent.
The highest percentage frequency recorded was 73 % for
Salix babylonica in temperate grassland.
TABLE 5. — Statistics for roadside and veld habitats in each veld type category, biome and the study area
* 1 or more species scored 5 or more. ** See data treatment — formulae used.
243
Bothalia 19,2 (1989)
Prominence
The most prominent invader in the study area was Acacia
dealbata, followed by A. meamsii and Salix babylonica
(Table 6). A. dealbata was very common to abundant (i.e.
scored a 5, 6 or 7) at 10% (60/578) of all river crossings
in the study area. A. meamsii and Salix babylonica were
very common to abundant at 4% (24/578) and 3% (19/578)
respectively (Table 6).
In the Grassland Biome A. dealbata was by far the most
prominent invader followed by Salix babylonica and A.
meamsii. Salix babylonica was prominent in temperate
grassland, A. dealbata most prominent in moist sub-
tropical grassland and both Acacia spp. were prominent
in dry subtropical grassland and mistbelt grassland.
In the Savanna Biome Chromolaena odorata, Lantana
camara and Ricinus communis were the most prominent
invaders. The first two species were most prominent in
tropical forest and although they were not the most
frequently recorded species they formed dense stands more
often than did any other species. Ricinus communis rarely
formed dense stands, its prominence being largely
attributable to its high percentage frequency. Sesbania
punicea and Melia azedarach were the most prominent
species in tropical bush and savanna.
Roadside and veld habitats
The whole study area
One hundred and fifty two quarter degree squares and
350 road transects were sampled in which 116 species were
recorded. Up to 44 species were recorded per quarter
degree square. Invaders were recorded in 93,7% of all
transects sampled and 30,6% of all transects were heavily
invaded (see Table 5).
Analysis according to veld type
Tropical forest was the most heavily invaded in terms
of species diversity and abundance of invaders (Table 5).
Mistbelt grassland was the next most heavily invaded.
Within these veld type categories the Pietermaritzburg,
Durban and particularly the south coast were the most
heavily invaded (Figures 4 & 5). Temperate grassland was
the least invaded.
Analysis according to species
Frequency
The most frequently recorded species in the study area
were Melia azedarach, Acacia meamsii, Solanum
mauritianum and Ricinus communis (Table 7). Only about
14 spp. were recorded in more than 10% of all transects.
The most frequently recorded invader in the Grassland
Biome was Acacia meamsii, followed by A. dealbata,
Eucalyptus spp., Solanum mauritianum, Melia azedarach,
Prunus persica and Rubus spp. (mainly R. cuneifolius) .
In the Savanna Biome Ricinus communis, Melia
azedarach and Psidium guajava were the most frequently
recorded invaders, followed by Solanum mauritianum,
Chromolaena odorata and Lantana camara. Opuntia ficus-
indica was the most frequent invader in tropical bush and
savanna.
Prominence
Chromolaena odorata, despite the fact that it was almost
entirely confined to the coastal belt, scored the highest
prominence value in the study area (Table 7). It had the
highest mean abundance rating (7) of all species in the
study area (Table 7). The next most prominent species in
order (mean abundance ratings in brackets) were Solanum
mauritianum (5), Psidium guajava (5), Rubus spp., mainly
R. cuneifolius (5), Acacia meamsii (4) and Lantana
camara (5).
In the Grassland Biome Rubus spp., Solanum mauri-
tianum and Acacia meamsii were the most prominent
species. In the Savanna Biome Chromolaena odorata,
Psidium guajava and Lantana camara were the most
prominent species.
Acacia dealbata deserves mention as the second most
prominent invader after Rubus spp. in moist subtropical
grassland. Opuntia ficus-indica rated second most
prominent after Lantana camara in bush and savanna.
Melia azedarach rated second most prominent after Acacia
meamsii in dry subtropical grassland.
Patterns of invasion
Most invasion in terms of species diversity and
abundance of invaders was recorded in the coastal belt and
adjacent midlands. Within this zone most invasion occurs
in and around towns and cities particularly Pietermaritz-
burg and Durban. The coastal belt is heavily invaded south
of Durban as well as in the north around Mtunzini and
Lake St Lucia (Figure 5). The north-eastern coastal plains
and bushveld north of Lake St Lucia have little invasion
except along the perennial rivers such as the Pongola.
A comparison of Figures 3 and 4, indicating the severity
of invasion in streambank and roadside and veld habitats
respectively, shows similar patterns except that in the
uplands there is more severe invasion of the streambank
habitat than of roadside and veld habitats. This pattern of
streambank invasion in the uplands is mainly the result
of Acacia dealbata invasions (Figure 6A) and, to a lesser
extent, that of Salix babylonica (Figure 7K).
DISCUSSION
Sampling
As mentioned previously (Henderson & Musil 1984) the
sampling method has its limitations, such as the under-
sampling of certain habitats which are inaccessible by road,
and the less distinctive species. The results reflect only
the situation along road verges, which are highly disturbed
sites, and a small strip of veld and watercourse visible from
the road. Despite these limitations the method has proved
successful and economical in obtaining information that
otherwise would be unobtainable.
One recommendation with regard to the abundance
ratings is that the seven point scale used for streambank
habitats be revised or replaced with a cover-abundance
scale such as used in the Braun-Blanquet method of
vegetation analysis (see Mueller-Dombois & Ellenberg
1974). The present scale is unsatisfactory as it cannot cope
TABLE 6. — Alien species occurring in streambank habitats
244
Bothalia 19,2 (1989)
If s
c c a
73 |
~ o g 2
8 5 so
12 *
O
•e -a |
3 -a
2 §
<D .3
Oh %
B g
£ 5b
1
'S.-S
•a „
a
a
a
3 :§
3 '1= § -2 I I
-2 5 I §, «« 3
F = % frequency of occurrence; I = % crossings heavily invaded; P = prominence value; * = species occurring in the given category but not included in a formal recording at a watercourse crossing; bold numbers = the
highest prominence values in a given category which add up to ± 80% of the summed prominence values (see text).
TABLE 6. — Alien species occurring in streambank habitats (continued)
Bothalia 19,2 (1989)
245
F = % frequency of occurrence; I = % crossings heavily invaded; P = prominence value; * = species occurring in the given category but not included in a formal recording at a watercourse crossing; bold numbers
highest prominence values in a given category which add up to ± 80% of the summed prominence values (see text).
TABLE 6. — Alien species occurring in streambank habitats (continued)
246
Bothalia 19,2 (1989)
F = % frequency of occurrence; I = % crossings heavily invaded; P = prominence value; * = species occurring in the given category but not included in a formal recording at a watercourse crossing; bold numbers
highest prominence values in a given category which add up to ± 80% of the summed prominence values (see text).
TABLE 6. — Alien species occurring in streambank habitats (continued)
Bothalia 19,2 (1989)
247
F — % frequency of occurrence; I = % crossings heavily invaded; P = prominence value; * = species occurring in the given category but not included in a formal recording at a watercourse crossing; bold numbers
highest prominence values in a given category which add up to ± 80% of the summed prominence values (see text).
TABLE 7.— Alien species occurring in roadside and veld habitats
248
ifg
«- T3
73 S
x> JS
E
oa 5 73 s
-a “ .a -2
C ti g. £
•S O 2 g
$ 5 60
2
O
3 S I
a T3
<3 C
b 42
a. X
E g
[8 &
2 ^
'5.-S
73 _
o o
£ ^
Bothalia 19,2 (1989)
O. O, Q. O. O. Q. o, Q, O.
co co" c4 Tt co" <n C4 04
o o o ^ Or rs
» o h n o
*— i co
O. O. o. Q. O. o,
CO CO 04 ^ CO vH
00 04 00 00 ON
On irf c4 co o
CO 04 CO to
-T cT *-<" O'
O. CD o Q.
04 rj- CO 04
o oi >n h
cT co" ©" cT
o, cd„ o, o„
CO CO 04 co
oo VO *0 "7f
Pi 10 -a* -
04 oo ON
04 *Q * 04
04 ON
h 04 O
oo r-
>ciri O
vo VO
Q 22
O CD.
Mills
F = % frequency of occurrence; A = mean abundance rating; P = prominence value; * = species occurring in the given category but not included in a formal recording in a road transect; bold numbers — the highest
prominence values in a given category which add up to ± 80% of the summed prominence values (see text).
TABLE 7. — Alien species occurring in roadside and veld habitats (continued)
Bothalia 19,2 (1989)
249
F = % frequency of occurrence; A = mean abundance rating; P = prominence value; * = species occurring in the given category but not included in a formal recording in a road transect; bold numbers = the highest
prominence values in a given category which add up to ± 80% of the summed prominence values (see text).
TABLE 7. — Alien species occurring in roadside and veld habitats (continued)
250
Bothalia 19,2 (1989)
3 T3
13 S
X) J2
X> _ -O
3 J g
to
o 8 g
S 00
IIS
I'll
Q oo
s ^
'5.-S
in
1 a
^ 8
8 c
3 G
5 a
a -2 £
§ -5 ^ S
B.-R S '? .g 8 I
a-S-g^i'c o
.a 'c -c
■g g -a s s
. s ■
i.-5 •,
^3 a 3
.s
3 8- s
a .a
c -o
F = % frequency of occurrence; A = mean abundance rating; P = prominence value; * = species occurring in the given category but not included in a formal recording in a road transect; bold numbers = the highest
prominence values in a given category which add up to ± 80% of the summed prominence values (see text).
TABLE 7. — Alien species occurring in roadside and veld habitats (continued)
Bothalia 19,2 (1989)
251
F = % frequency of occurrence; A = mean abundance rating; P = prominence value; * = species occurring in the given category but not included in a formal recording in a road transect; bold numbers = the highest
prominence values in a given category which add up to ± 80% of the summed prominence values (see text).
TABLE 7. — Alien species occurring in roadside and veld habitats (continued)
252 Bothalia 19,2 (1989)
F = % frequency of occurrence; A = mean abundance rating; P = prominence value; * = species occurring in the given category but not included in a formal recording in a road transect; bold numbers — the highest
prominence values in a given category which add up to ± 80% of {he summed prominence values (see text).
TABLE 7. — Alien species occurring in roadside and veld habitats (continued)
Bothalia 19,2 (1989)
253
254
1 OR MORE RIVER CROSSINGS INVADED
1 OR MORE RIVER CROSSINGS HEAVILY INVADED
a
10 OR MORE SPECIES PER 1/4“ SQUARE
FIGURE 3. — Invasion in streambank habitats in terms of the intensity
of invasion of watercourse crossings and species diversity per
quarter degree square.
a 15 OR MORE SPECIES PER 1/4° SQUARE
FIGURE 4. — Invasion in roadside and veld habitats in terms of the in-
tensity of invasion of road transects and species diversity per
quarter degree square.
with all situations and especially where only one species
is present. The term ‘commonest species' has not been
defined but has been subjectively evaluated in terms of
cover and density.
Prominent and potentially important species
Although 130 species were*recorded in the study area,
most invasion can be attributed to the following species:
Chromolaena odorata, Psidium guajava and Lantana
camara in the Savanna Biome and Acacia dealbata, A.
meamsii, Solatium mauritianum and Rubus spp. in the
Grassland Biome.
Chromolaena odorata is the most outstanding invader
in Natal. Its vigorous growth, prolific seed production
(1 300000 seeds annually per plant have been recorded)
and efficient dispersal mechanism have enabled it to
rapidly encroach large areas (Erasmus 1986). It is thought
to have been unintentionally introduced to Durban during
the Second World War (1939-1945). By 1950 it was
conspicuous as a weed and had spread 120 km north of
Durban (Liggitt 1983). By 1962 it was spreading
‘virulently’ along the Natal coast (Erasmus 1986). By 1981
it was recorded from Kosi Bay just south of the
Mozambique border and by the mid 1980’s it was recorded
as far south as Hluleka Nature Reserve on the Transkei
coast (about 300 km south of Durban) and in the Wolkberg
in the north-eastern Transvaal (about 650 km north of
Durban) (Macdonald 1984).
In the study area C. odorata is virtually confined to the
narrow, hot and moist coastal belt of Natal (Figure 6J)
where it forms dense monospecific thickets. It accounts
for 23% of the total invasion recorded in roadside and veld
habitats in the whole study area. From Mtunzini
southwards it is exceedingly abundant, especially along
□
INDIVIDUALS /GROUPS
Mils! >49
FIGURE 5. — Invasion in roadside and veld habitats in terms of the mean
abundance of invaders per kilometre in each quarter degree square.
i it n
Bothalia 19,2 (1989)
255
FIGURE 6. — Distribution and high abundance areas of the most prominent species: A, Acacia dealbatat B, A. decurrens\ C, A. longifolia;
D, A. meamsii; E, A. melanoxylon\ F, A. podalyriifolia ; G, Arundo donax\ H, Caesalpinia decapetala', I, Cassia didymobotrya , J,
Chromolaena o do rata ; K, Eucalyptus spp. : L, Jacaranda mimosifolia\ M, Lantana camara\ N, Leucaena leucoccphala , O, Melia
azedarach; P, Montanoa hibiscifolia. Roadside and veld habitats, □, highest abundance rating of 5 or more; streambank habitat. A,
highest abundance rating of 5 or more; streambank, roadside and veld habitats, O, with the aforementioned values.
roadsides where it forms continuous stands. It frequently
forms a dense margin to bush clumps and forest patches.
North of Mtunzini it is locally abundant on the western
shores of Lake St Lucia. Further north it is relatively
scarce but has been observed at Manzengwenya and at
Manguzi about 15 km south of the Mozambique border
(M.C. Ward pers. comm.). It is absent from dry bush and
savanna except where it has made incursions along water-
courses.
C. odorata is native to a large area which includes
Central and South America and the Caribbean Islands.
According to preliminary findings the Natal plants are
morphologically most similar to plants growing at Manaus
(Amazonian region) in northern Brazil (R.L. Kluge pers.
comm.). Based on these findings and the rampant growth
of C. odorata in the tropical coastal belt of Natal it is
predicted that it has the potential to invade the tropical
East African coast up to Somalia. It is already present in
equatorial West Africa in the Ivory Coast (Erasmus 1986),
Cameroon (National Herbarium, Pretoria), Ghana and
Nigeria (Holm et al. 1979). Its southern limit as a major
weed is likely to be defined by Acocks’s Veld Type, Coastal
Forest and Thornveld (to which it is almost confined in
256
Bothalia 19,2 (1989)
Natal) near East London in the eastern Cape of South
Africa. Further south to Port Elizabeth the coastal
forests become progressively more arid and unsuitable
for growth of C. odorata. Beyond Port Elizabeth the
coastal Knysna Forest, although receiving a high rain-
fall and no frost, is probably too temperate for the likes
of this tropical weed. In Natal this species has virtually
reached the limits of its distribution but it is expected to
increase in density throughout its range especially in the
northern coastal belt unless strict control measures are
exerted.
Psidium guajava was the most frequently recorded
invader of roadside and veld habitats in the coastal belt
and the second most abundant after Chromolaena odorata.
It not only invades roadsides and other highly disturbed
sites, where it can form dense stands, but also coastal
grasslands, bush clumps, forest patches and riverine
vegetation. It is the author’s contention that the importance
of this species in Natal was greatly underestimated at the .
1984 workshop meeting (Macdonald & Jarman 1985). This
survey has shown that it is one of the most prominent
invaders in the very land use types — utility areas and
FIGURE 7. — Distribution and high abundance areas of the most prominent species: A, Moms alba ; B, Opuntia ficus-indica; C, Pereskia
aculeata\ D, Pinus patula\ E, Populus x canescens ; F, Pmnus persica', G, Psidium guajava ; H, Pyracantha spp. ; I, Ricinus communis',
J. Rubus spp.; K, Salix babylonica ; L, S. lasiandra', M, Schinus terebinthifolius', N, Sesbania punicea\ O, Solanum mauritianum', P,
Tithonia diversifolia. Roadside and veld habitats, D, highest abundance rating of 5 or more; streambank habitat, A, highest abundance
rating of 5 or more; streambank, roadside and veld habitats, O, with the aforementioned values.
Bothalia 19,2 (1989)
257
coastal urban open space — where it was considered
unimportant and which significantly lowered its
importance ranking. As could have been expected, the
survey showed that Psidium guajava was not abundant in
the Durban metropolitan area on which the workshop
based their coastal urban open space judgements. It is not
known why this area differs in this respect from the other
urban areas in the coastal zone.
Lantana camara has a wider climatic tolerance than the
previous two species but grows best in the hot and humid
coastal belt where it can form dense thickets. It is abundant
along the coast south of Lake St Lucia but scarce north
of this point (Figure 6M). It is abundant along the Pongola,
Usutu and Ingwavuma Gorges in the aftermath of the
floods caused by cyclone Demoina in January 1984 (M.C.
Ward pers. comm.).
Solanum mauritianum is a widespread invader in the
mistbelt and coastal belt of Natal (Figure 70). It is most
abundant in mistbelt grassland where it forms dense stands
along roadsides and is a major understorey weed of
plantations (Le Roux 1982).
Acacia meamsii is widely naturalized in Natal but is
most abundant in the midlands where it has been
commercially planted on a vast scale (Figure 6D). It is
least abundant in the high grasslands, coastal belt and
tropical bush and savanna. In the whole study area it rated
as the second most prominent riverine invader after A.
dealbata.
Acacia dealbata is a widespread invader in the study
area. Its distribution stretches from the temperate
grasslands of the Orange Free State down to the Natal
Mistbelt and even into the rivers of tropical bush and
savanna (Figure 6A). It is by far the most serious invader
of watercourses in the Grassland Biome. Its massive seed
production and efficient downstream dispersal have
enabled it to penetrate watercourses far from plantings.
It rated equally prominent as A. meamsii in mistbelt
grassland although it has been planted there to a much
lesser degree.
Rubus spp. , mainly R. cuneifolius, are most abundant
in the Natal mistbelt and foothills of the Drakensberg in
southern Natal (Figure 7J). They are major understorey
weeds in Pine plantations (Le Roux 1982) and form dense
stands in open grassland and along road verges.
The aforementioned seven most prominent species in
the study area are not likely to greatly increase their
distribution ranges. Solanum mauritianum can be expected
to invade the coastal plains north of Lake St Lucia.
Chromolaena odorata, Psidium guajava and Lantana
camara are all expected to expand their distributions in
the same region. Further expansion by all seven species
will probably occur in marginally suitable areas. All
species can be expected to increase in density within their
ranges.
Melia azedarach and Ricinus communis are widespread
but most common in the coastal belt (Figures 60 & 71).
They are considerably less abundant than the species
already mentioned and therefore received lower
prominence rankings.
Ricinus communis has always been regarded as an
introduced weed from elsewhere in Africa. However,
recent archaeological diggings in the eastern Cape have
unearthed seeds of this species from a grass-lined storage
pit dating back more than 1 200 years (Brink 1988). This
suggests that, if indeed introduced, primitive hunter-
gatherers were the agents (Brink 1988). This is in sharp
contrast to the majority of our foreign weeds which have
been introduced since the colonization of the Cape 300
years ago. Whether it is classified as an alien or indigenous
plant, R. communis is not generally regarded as an
important weed (e.g. Macdonald 1983) because it is a
pioneer plant, colonizing disturbed sites, and eventually
giving way to longer-lived species.
Melia azedarach on the other hand is a long-lived
tree and can grow to a considerable size (12—23 m in
height and a spreading canopy) (Poynton 1972). Add to
this its production of vast quantities of berries which are
dispersed by birds and water, its high germinability, its
hardiness to drought and cold, rapid growth and its
response to felling by coppicing profusely, then it deserves
recognition as an important invader. It is most important
in streambank habitats where its efficient downstream
dispersal enables it to invade protected areas far from
source areas (Macdonald 1983). In this survey it rated as
the most prominent tree species in tropical bush and savan-
na. It has virtually reached the limits of its distribution
in the study area but can be expected to increase in abun-
dance especially along river banks in the Savanna Biome.
Salix babylonica is a widespread streambank invader in
the upland grasslands (Figure 7K). It is unable to
reproduce sexually as only female trees have been
introduced into South Africa, but reproduces vegetatively
from branches which are torn from trees by floodwaters
and deposited downstream (Poynton 1973). It does not form
dense stands like the Acacia spp. but a single large tree
probably achieves the same basal area as 10, 20 or more
Acacia trees. It can also effectively exclude the growth of
other species under its canopy (pers. obs.). In places such
as along the Umzimvubu River in south-western Natal,
it forms continuous stands which stretch for many
kilometres.
Salix lasiandra, referred to by Immelman (1987), is less
widely distributed (Figure 7L) than S. babylonica but it
too can form continuous stands in places. It is particularly
abundant along the Wilge River near Harrismith. It is more
widely distributed and abundant in southern Natal than
shown in Figure 7L but the data are not available as it was
mistaken for an indigenous Salix species during this part
of the survey.
Sesbania punicea was the fourth most prominent
streambank invader in the study area. It was locally
abundant along watercourses in the interior and
along roadsides in the humid coastal belt (Figure 7N).
Three species of herbivorous weevils have been imported
for biological control of this weed. Already one of the
weevils has made an outstanding contribution to the
biological control of S. punicea and, in combination with
the two other species, is expected to halt the invasive spread
of this plant in South Africa (Hoffmann & Moran 1988).
258
Bothalia 19,2 (1989)
Opuntia spp. are widely distributed in the study area
but occur mainly as scattered plants. Opuntia ficus-indica
(Figure 7B) and O stricta are locally abundant in the
Tugela River valley and elsewhere in tropical bush and
savanna. Opuntia vulgaris occurs as widely scattered plants
in the coastal belt. All these species have been the subjects
of biological control campaigns and their numbers have
been greatly reduced. They are no longer considered a
threat but may still form localized infestations
(Zimmermann et al. 1986).
Cassia didymobotrya (Figure 61) and Caesalpinia
decapetala (Figure 6H) are fairly widespread in the coastal
belt and mistbelt of Natal. They are locally abundant,
particularly in disturbed sites around kraals, villages, along
roadsides and riverbanks. Cassia didymobotrya is a central
African plant and although very abundant in places it is
mainly a pioneer plant. Flowering specimens are invariably
host to a caterpillar which may help to control the spread
of this species. Caesalpinia decapetala is a vigorous-
growing, exceedingly thorny woody shrub or climber that
can form a dense canopy that smothers the existing vege-
tation and excludes other species. It is particularly trouble-
some as an invader of forest margins. It is easily
overlooked during a roadside survey when not flowering
and thus is likely to be more widely distributed and abun-
dant than shown.
Less widespread species which are locally abundant and
can form dense stands include: Arundo donax (Figure 6G),
Pereskia aculeata (Figure 1C), Pinus elliottii (Macdonald
& Jarman 1985), Schinus terebinthifolius (Figure 7M) and
Solanum seaforthianum in the coastal belt; Tithonia
diver sifolia (Figure 7P), Montanoa hibiscifolia (Figure 6P)
and Leucaena leucocephala (Figure 6N) around Durban;
Cardiospermum grandiflorum and Cestrum laevigatum in
the Pietermaritzburg, Durban and coastal areas and
Populus x canescens (Figure 7E) along streambanks in
the Grassland Biome. Ulpx europaeus and Cytisus
scoparius have been known to form dense stands in moist
subtropical grassland at the Highmoor Forest Estate
(National Herbarium specimens collected by D. Edwards
in 1961).
Arundo donax, the giant reed, has invaded streambank
habitats largely unnoticed because it is similar in
appearance to the indigenous reeds ( Phragmites spp.). In
the Natal coastal belt, where it forms dense stands in places
and flowers prolifically, it can be easily recognised by its
large inflorescence with dense ascending branches.
Suitable habitat for this species occurs along river banks
throughout the Savanna Biome. It can be expected to
expand its distribution in this region.
Schinus terebinthifolius is a popular ornamental and
hedge plant in the coastal belt and produces large numbers
of berries. There have been observations of seed predation
by insects which has probably prevented it from becoming
a serious weed (S. Neser pers. comm.). It should
nevertheless be regarded as a potentially important invader
since it has become a pest in other parts of the world, can
thrive at the outer limits of vegetation exposed to salt spray
and has even invaded mangroves in Florida, USA (Morton
1978; Toufexis 1985). Dense stands of this species were
observed in the Umgeni River in Durban North just
upstream of the Beachwood Mangroves Nature Reserve.
If seed is made available S. terebinthifolius may even
threaten the swamp forests of Maputaland in north-eastern
Natal. This habitat is of very limited extent and according
to Macdonald & Jarman (1985) is the least invaded of any
habitat in Natal. However several species are being
cultivated in these swamps, namely Mangifera indica,
Carica papaya, Musa sp. and Ananas sp. M. indica is
naturalized in the swamp forest at Kosi Bay.
Concern has been expressed about the invasiveness of
Litsea glutinosa in the coastal lowlands of Natal
(Macdonald & Jarman 1985). According to records in the
National Herbarium it has been much planted in Durban,
regenerates easily from seed and it was recorded as
naturalized in the Eshowe District as far back as 1937. It
is regarded as a weed on the island of Mauritius (Holm
et al. 1979).
Pereskia aculeata is potentially a very serious invader
of coastal forests. It is a very thorny vigorous-growing
climbing cactus which can smother and kill the trees it
overtops. Its current widespread distribution in the
conservation areas of the coastal lowlands in northern
Natal is mainly thought to be the result of previous
intentional plantings around kraals and burial sites
(Macdonald & Jarman 1985). Natural spread from these
sites of previous introduction has in many localities been
surprisingly limited given its bird-dispersed fruit. But the
rate at which it can be spread by frugivorous animals and
by vegetative reproduction, is potentially rapid (Macdonald
& Jarman 1985). Campbell (1988) reports that the seed
is geared for rapid germination in a range of habitats but
that the soil seed bank is likely to deteriorate rapidly, with
the occurrence of either germination or seed death.
P. aculeata is a difficult weed to control as any part of
the plant which survives treatment can reproduce
vegetatively and restart the thicket. Control methods
involving slashing, poisoning and burning can result in
the total destruction of all vegetation in the affected area
(Bruton 1981). However, in several KwaZulu nature
reserves infestations have been hand-cleared with as little
disturbance as possible. Hand-weeding and herbicide
treatments are necessary in follow-up operations
(Macdonald & Jarman 1985).
Relation of invasion to environmental factors
Alien invasion is related to indigenous veld type
categories and broad climatic factors. There is a general
trend for more invasion in terms of species diversity and
abundance with decreasing elevation from the cold upland
grasslands to the warm coastal belt. Most invasion occurs
in the humid to subhumid coast (tropical forest) and
mistbelt (mistbelt grassland) where there is little or no
incidence of frost. There is considerably less invasion,
particularly in roadside and veld habitats, in the colder
and drier veld types.
The limited invasion of the northern coastal belt relative
to that in the south of the province is interpreted as being
a result of less disturbance and fewer plantings of alien
species. Localized infestations, for example around Lake
St Lucia, indicate that this region is vulnerable to invasion.
According to Liggitt (1983) Chromolaena odorata is
spreading rapidly in the north where up to 2000%
(i Id
ill
ini
lit
St
j ■'*
5ft
(4
anl
iit-j
Bothalia 19,2 (1989)
259
increases in vegetation cover have been recorded in the
Dukuduku plantations during a time span of five years.
The distributions of some species correspond well with
broad climatic zones. For example Chromolaena odorata
is virtually restricted to the frost-free and moist coastal
belt, although, being wind-dispersed, it has a potentially
wide distribution. Members of the Rosaceae, such as
Rubus, Rosa, Prunus, Pyracantha and Cotoneaster spp.,
are most evident in the cold high-lying grasslands. This
may be attributable to a dormancy mechanism in their
seeds which is terminated by cold winter temperatures
(Dean et al. 1986).
Watercourses have played an important role in the
dispersal of species and in particular those which otherwise
have a limited dispersal range. For example Acacia
dealbata, A. meamsii, Sesbania punicea and Caesalpinia
decapetala have rather immobile and hard seeds but which
are readily transported by water. The abrasion which the
seeds receive along their journey may well promote ger-
mination.
Watercourses have also enabled the long-range vegetative
dispersal of species such as Salix babylonica and Opuntia
spp. The spread of suckering species such as Populus x
canescens and Robinia pseudoacacia is also promoted by
stream flow.
Some of the important invader species are dispersed by
birds enabling them to invade relatively undisturbed sites
and far afield from parent plants. Notable species include
Lantana camara, Solarium mauritianum, Melia azedarach,
Pyracantha spp., Psidium guajava, Rosa spp. and Rubus
spp. In the grassland regions bird-dispersed species are
clearly associated with perching sites, such as fence lines,
rocky outcrops, bush clumps and plantations. In the
mistbelt Solanum mauritianum and Rubus cuneifolius form
dense thickets in the understoreys of plantations.
CONCLUSION
The intensity of alien plant invasion is expected to
increase in all parts of the study area and particularly in
the coastal and mistbelts of Natal. Top priority should be
given to the control of invaders, especially Chromolaena
odorata, in north-eastern Natal [see Macdonald & Jarman
(1985), where a possible control strategy is detailed] which
is an important conservation area and where the potential
for expansion is great.
Shortly after completion of this survey, in September/
October 1987, Natal experienced devastating widespread
floods. Most of the major river valleys as well as the
floodplains along the coast were severely affected. Vast
tracts of riverine vegetation that used to grow within the
flood line were swept away. The long-term consequences
of the floods remain to be seen but in the short-term it
can be expected that an explosion of pioneer and other
fast-growing plants will occur.
In the coastal belt Chromolaena odorata can be expect-
ed to rapidly invade floodplains, river banks and any other
exposed land. Ricinus communis is likely to greatly
increase in abundance especially in riverbeds as it did in
the Pongola River after the floods caused by cyclone
Demoina in January 1984. The floods are also expected
to promote the downstream spread of water-dispersed
species such as Acacia dealbata, A. meamsii, Melia
azedarach and Salix babylonica. The floods, together with
the prospect of a wetter climatic cycle ahead (Tyson 1986),
could result in the spread of species into areas which
previously were too dry and inhospitable. In this respect
there is a danger that A. dealbata and C. odorata will
become more widely established and abundant in bush and
savanna.
An assessment of the ecological consequences of
streambank invasion, particularly pertaining to water usage
and soil conservation, is considered to be extremely
important in the motivation for the control of streambank
invaders. Acacia dealbata and A. meamsii, although
declared invaders, have spread uncontrolled throughout the
Grassland Biome from the upland grasslands along the
Drakensberg escarpment down into the mistbelt. In the
case of A. meamsii it has even spread into the coastal belt.
Apart from a few notable exceptions most of the
important alien woody invader species in Natal and the
rest of South Africa (Henderson & Musil 1984; Macdonald
& Jarman 1984; Macdonald et al. 1986; Stirton 1978) have
been cultivated either on a grand scale in plantations, or
as barrier plantings, cover/binders, shelterbelts and
ornamentals in gardens. This raises the issue of screening
alien plant species for potential invasiveness before they
are allowed to be propagated on a grand scale. This applies
particularly to the establishment of plantations of species
new to the region belonging to the genera Acacia and
Pinus, many species of which have become serious
invaders in South Africa.
ACKNOWLEDGEMENTS
I thank Miss C. Craemer of the Plant Protection
Research Institute and Mrs H. Joffe of the Botanical
Research Institute for their assistance in the field. The
assistance and co-operation received from the Director and
Mr M.C. Ward of the Bureau of Natural Resources,
KwaZulu are gratefully acknowledged. Thanks are also
due to the Department of Environment Affairs, Directorate
of Forestry, Zululand and Natal Forest Regions and the
Natal Parks Board for information received.
REFERENCES
ACOCKS, J.P.H. 1988. Veld types of South Africa, 3rd edn. Memoirs
of the Botanical Survey of South Africa No. 57.
BRINK, E. 1988. Interesting record. The Elephant's Child. Newsletter
of the Albany Museum 11,3.
BRUTON, M.N. 1981. Major threat to the coastal dune forest in
Maputaland. The Naturalist 25,1: 26 —27.
CAMPBELL, P.L. 1988. Seed germination of Harrisia martinii and
Pereskia aculeata with reference to their potential spread in Natal.
Applied Plant Science 2: 60—62.
DEAN, S.J., HOLMES, PM. & WEISS, P.W. 1986. Seed biology of
invasive alien plants in South Africa and South West Africa/
Namibia. In I.A.W. MacDonald, F.J. Kruger & A. A. Ferrar, The
ecology and management of biological invasions in southern
Africa. Oxford University Press, Cape Town.
DYER, R.A. 1975. The genera of southern African flowering plants Vol.
1. Department of Agricultural Technical Services, Pretoria.
EDWARDS, D. 1967. A plant ecology survey of the Tugela River basin.
Natal. Memoirs of the Botanical Survey of South Africa No. 36.
ERASMUS, D.J. 1986. Triffid weed/Paraffienbos. Weeds/Onkruid
A17/1986. Farming in South Africa.
260
Bothalia 19,2 (1989)
FRANCIS, L.M. 1977. Guide to tree-planting: Zululand. Pamphlet No.
197. Department of Forestry, Pretoria.
HAIGH, H.H. & WILHELMU, H. 1973. Guide to tree-planting: Natal.
Pamphlet No. 124. Department of Forestry, Pretoria.
HENDERSON, L. & MUSIL, K.J. 1984. Exotic woody plant invaders
of the Transvaal. Bothalia 15 : 297— 313.
HENDERSON, M., FOURIE, D.M.C., WELLS, M.J. & HENDER-
SON, L. 1987. Declared weeds and alien invader plants in South
Africa. Bulletin No. 413. Department of Agriculture and Water
Supply, Pretoria.
HOFFMANN, J.H. & MORAN, V.C. 1988. The invasive weed Sesbania
punicea in South Africa and prospects for its biological control.
South African Journal of Science 84 : 740—742.
HOLM, L., PANCHO, J.V., HERBERGER, J.P. & PLUCKNETT, D.L.
1979. A geographical atlas of world weeds. Wiley, New York.
IMMELMAN, K.L. 1987. Synopsis of the genus Salix (Salicaceae) in
southern Africa. Bothalia 17: 171—177.
LE ROUX, P.J. 1982. Plant invader species in pine plantations in South
Africa. Proceedings of the fourth National Weeds Conference of
South Africa, 1981. Balkfema, Cape Town.
LIGGITT, B. 1983. The invasive alien plant Chromolaena odorata, with
regard to its status and control in Natal. Institute of Natural
Resources, University of Natal, Pietermaritzburg. Rural studies
series, monograph 2.
MACDONALD, I. A.W. 1983. Alien trees, shrubs and creepers invading
indigenous vegetation in the Hluhluwe-Umfolozi Game Reserve
Complex in Natal. Bothalia 14: 949-959.
MACDONALD, LAW. 1984. Infiltration of dreaded weed alarms
experts/Pesplant mik nou ook na Krugerwildtuin. Custos 13,8:
31-35.
MACDONALD, I. A.W. & JARMAN, M.L. (eds) 1984. Invasive alien
organisms in the terrestrial ecosystems of the fynbos biome, South
Africa. South African National Scientific Programmes Report
No. 85. CSIR, Pretoria.
MACDONALD, I.A.W. & JARMAN, M.L. (eds) 1985. Invasive alien
plants in the terrestrial ecosystems of Natal, South Africa. South
African National Scientific Programmes Report No. 118. CSIR,
Pretoria.
MACDONALD, I.A.W., KRUGER, F.J. & FERRAR, A. A. (eds) 1986.
The ecology and management of biological invasions in southern
Africa. Oxford University Press, Cape Town.
MOLL, E. 1981. Trees of Natal. Eco-lab Trust Fund, Cape Town.
MORTON, J.F. 1978. Brazilian pepper — its impact on people, animals
and the environment. Economic Botany 32: 353—359.
MUELLER-DOMBOIS, D. & ELLENBERG, H. 1974. Aims and
methods of vegetation ecology. Wiley, New York.
POYNTON, R.J. 1972. Characteristics and uses of trees and shrubs.
Bulletin No. 39. Department of Forestry, Pretoria.
POYNTON, R.J. 1973. Two hundred selected indigenous and exotic
species: how to recognise and grow them. In W.F.E. Immelman,
C.L. Wicht & D.P. Ackerman, Our green heritage. Tafelberg,
Cape Town.
ROSS, J.H. 1972. Flora of Natal. Memoirs of the Botanical Survey of
South Africa No. 39.
RUTHERFORD, M.C. & WESTFALL, R.H. 1986. Biomes of southern
Africa— an objective categorization. Memoirs of the Botanical
Survey of South Africa No. 54.
SPIES, J.J. & DU PLESSIS, H. 1985. The genus Rubus in South Africa.
1. Chromosome numbers and geographical distributions of
species. Bothalia 15: 591-596.
STIRTON, C.H. (ed.) 1978. Plant invaders — beautiful but dangerous.
Department of Nature and Environmental Conservation of the
Cape Provincial Administration, Cape Town.
TOUFEXIS, A. 1985. The trees are taking over. Time 24: 51.
TYSON, P.D. 1986. Climatic change and variability in southern Afri-
ca. Oxford University Press, Cape Town.
WARD, C.J. 1980. The plant ecology of the Isipingo Beach area. Natal,
South Africa. Memoirs of the Botanical Survey of South Africa
No. 45.
WELLS, M.J., DUGGAN, K.J. & HENDERSON, L. 1980. Woody plant
invaders of the central Transvaal. Proceedings of the third National
Weeds Conference of South Africa, 1979. Balkema, Cape Town.
ZIMMERMANN, H.G., MORAN, V.C. & HOFFMANN, J.H. 1986.
Insect herbivores as determinants of the present distribution and
abundance of invasive cacti in South Africa. In I.A.W.
MacDonald, F.J. Kruger & A. A. Ferrar, The ecology and
management of biological invasions in southern Africa. Oxford
University Press, Cape Town.
APPENDIX
The names of 180 species of naturalized alien trees, shrubs and climbers are listed. Some non-woody species are included. Names and dates in
brackets: literature references. (PRE): cited on National Herbarium specimen labels.
Acacia
dealbata Link, silver wattle
decurrens Willd., green wattle
famesiana (L.) Willd. (Ross 1972), sweet acacia
longifolia (Andr.) Willd., long-leaved wattle
meamsii De Wild., black wattle
melanoxylon R. Br., blackwood
podalyriifolia A. Cunn. ex G. Don, pearl acacia
saligna (Labill.) Wendl. (MacDonald & Jarman 1985), Port Jackson
willow
Agave
americana L., century plant
sisalana Perrine, sisal
Ageratina adenophora (Spreng.) R.M. King & H. Robinson, crofton weed
Ailanthus altissima (Mill.) Swingle, tree-of-heaven
Albizia
lebbeck (L.) Benth. (Ross 1972), lebbeck tree
procera (Roxb.) Benth. (Ross 1972), false lebbeck
Anacardium occidentale L. (Ross 1972), cashew
Anredera
baselloides (H.B.K.) Baill. (Ross 1972)
cordifolia (Ten.) Steenis (PRE), Madeira vine
Antigonon leptopus Hook. & Am., Mexican creeper
Araujia sericifera Brot. (PRE), moth catcher
Arundo donax L. , giant reed
Bambusa balcooa Roxb. ex Roxb., common bamboo
Bambuseae spp. , (two small spp. , one possibly Phyllostachys sp.),
bamboos
Basella paniculata Volkens (Ross 1972)
Bauhinia variegata L., white orchid tree
Caesalpinia decapelala (Roth) Alston, Mauritius thorn
Cajanus cajan (L.) Millsp. (Ross 1972), Congo pea
Canna
edulis L. (MacDonald & Jarman 1985), edible canna
indica L., canna
Cardiospermum grandiflorum Schwartz, balloon vine
Carica papaya L., papaya
Cassia
bicapsularis L.
coluteoides Collad.
corymbosa Lam. (PRE), autumn cassia
didymobotrya Fresen., peanut-butter cassia
floribunda Cav., arsenic bush
hirsuta L.
occidentalis L., wild coffee
Casuarina
equisetifolia G. Forst., horsetail tree
sp. cf. C. cunninghamiana Miq., beefwood
Cedrela odorata L. (PRE), West Indian Cedar
Cedrus deodara (Roxb.) G. Don, deodar
Ceiba pentandra (L.) Gaertn. (Ross 1972), kapok tree
Cereus peruvianus (L.) Mill., queen of the night
Cestrum
aurantiacum Lindl., yellow cestrum
laevigatum Schlechtd., inkberry
Chromolaena odorata (L.) R.M. King & H. Robinson, triffid weed
Cinnamomum camphora (L.) J. Presl, camphor tree
Citrus
aurantium L. (MacDonald & Jarman 1985), sour orange
limon (L.) Burm. f. (MacDonald & Jarman 1985), lemon
reticulata Blanco (MacDonald & Jarman 1985), mandarin
Colocasia esculenta (L.) Schott, taro
Cortaderia sp. , pampas grass
Cotoneaster franchetii Bois, orange cotoneaster
Bothalia 19,2 (1989)
261
Cotoneaster sp., cotoneaster
Crataegus phaenopyrum (L. f.) Medic., Washington thorn
Crotalaria agatiflora Schweinf. subsp. imperials (Taub.) Polhili,
canary-bird bush
Cryptomeria japonica (L. f.) D. Don, Japanese cedar
Cupressus sp. cf. C. arizonica Greene, Arizona cypress
Cupressus spp., cypresses
Cyphomandra betacea (Cav.) Sendtn. (PRE), tree tomato
Cytisus scoparius L., Scotch broom
Dahlia imperialis Roezl. ex Ortg., tree dahlia
Delonix regia (Bojer) Raf., flamboyant
Duranta erecta L. (= D. repens L.) (Ross 1972), forget-me-not-tree
Eriobotrya japonica (Thunb.) Lindl. (MacDonald & Jarman 1985), loquat
Eucalyptus
elata Dehnh. (= E. andreana Naud.) (Ward 1980), river peppermint
camaldulensis Dehnh., red gum
cinerea F.J. Muell. ex Benth., florist’s gum
grandis W. Hill ex Maid. (MacDonald & Jarman 1985), saligna gum
robusta Sm., swamp mahogany gum
Eucalyptus spp., gums
Euphorbia pulcherrima Willd. ex Klotzsch, poinsettia
Gleditsia triacanthos L., honey locust
Gmelina arborea Roxb. (Dyer 1975), white teak
Grevillea robusta A. Cunn., Australian silky oak
Hakea sericea Schrad. (= H. tenuifolia (Salisb.) Domin) (Moll 1981),
silky hakea
Harrisia martinii (Labouret) Britton (Henderson etal. 1987), moon cactus
Hedychium
coronarium J. Konig, butterfly ginger
flavum Roxb.
Hylocereus undatus (Haw.) Britt. & Rose (PRE), night-blooming cereus
Ipomoea
alba L., moon flower
coccinea L. (Ross 1972), red morning-glory
conge sta R. Br.
fistulosa Choisy
nil (L.) Roth (Ross 1972)
purpurea (L.) Roth (Ross 1972), common morning-glory
Jacaranda mimosifolia D. Don, jacaranda
Jatropha curcas L. (PRE), physic nut
Lagerstroemia indica L., pride-of-India
Lantana camara L., lantana
Lespedem cuneata (Du Mont) G. Don (Ross 1972), bush clover
Leucaena leucocephala (Lam.) De Wit, leucaena
Ligustrum ovalifolium Hassk. (PRE), California privet
Litsea glutinosa (Lour.) C.B. Robinson (= L. sebifera Pers.) (MacDonald
& Jarman 1985), Indian laurel
Macfadyena unguis-cati (L.) A. Gentry (PRE), cat’s-claw creeper
Mangifera indica L., mango
Manihot sp., cassava
Melia azedarach L., syringa
Metasequoia glyptostroboides H.H. Hu & Cheng, dawn redwood
Monstera deliciosa Liebm., Swiss-cheese plant
Montanoa
bipinnatifida (Kunth) C. Koch (Moll 1981), tree daisy
hibiscifolia Benth.
Morns
alba L., white mulberry
alba L. var. multicaulis
Musa sp., banana
Nerium oleander L., oleander
Nicotiana glauca R.C. Graft., wild tobacco
Opuntia
dillenii (Ker-Gawl.) Haw. (Henderson et al. 1987), pipestem prickly pear
exaltata A. Berger (Henderson et al. 1987), long-spine cactus
ficus-indica (L.) Mill., sweet prickly pear
imbricata (Haw.) DC. (Henderson et al. 1987), chain-link cactus
spinulifera Salm-Dyck, large round-leaved prickly pear
stricta Haw., pest pear of Australia
vulgaris Mill., cochineal prickly pear
Pandanus spp., screw-pines
Paraserianthes lophantha (Willd.) Nielsen subsp. lophantha (= Albizia
lophantha (Willd.) Benth.) (Ross 1972), stinkbean
Parkinsonia aculeata L. (Ross 1972), Jerusalem thorn
Passiflora
edulis Sims, purple granadilla
foetida L., love-in-a-mist
suberosa L. (Ross 1972)
subpeltata Ortega (Ross 1972), granadina
Pennisetwn purpureum Schumach., elephant grass
Pereskia aculeata Mill., Barbados gooseberry
Persea americana Mill. (PRE), avocado pear
Pinus
canariensis Sweet ex K. Spreng., Canary pine
elliottii Engelm. (MacDonald & Jarman 1985), slash pine
patula Schlechtd. & Cham., patula pine
? taeda L., loblolly pine
sp. cf. P. uncinata Mill, ex Mirb.
Pinus spp., pines
Populus
x canescens (Ait.) J.E. Sm., grey poplar
deltoides Bartr. ex Marsh., match poplar
nigra L. var. italica Muenchh., Lombardy poplar
Prunus
armeniaca L., common apricot
persica (L.) Batsch., peach
Psidium
guajava L., common guava
littorale Raddi var. longipes (O. Berg) Fosb. (= P. cattleianum Sabine)
(Ross 1972), strawberry guava
Punica granatum L., pomegranate
Pyracantha
angustifolia (Franch.) C.K. Schneid., yellow firethorn
fortuneana (Maxim.) H.L. Li
rogersiana (A.B. Jacks.) Bean
sp. cf. P. coccinea M.J. Roem., red firethorn
Quercus robur L., English oak
Robinia pseudoacacia L., black locust
Rosa
eglanteria L., eglantine
odorata (Andr.) Sweet, tea rose
sp. cf. R. multiflora Thunb. ex J. Murr. (PRE), baby rose
Rubus
affinis Wight & Arn., blackberry
cuneifolius Pursh., American bramble
phoenicolasius Maxim. (Spies & Du Plessis 1985), wineberry
Saccharum officinarum L., sugar cane
Salix
babylonica L., weeping willow
caprea L., pussy willow
lasiandra Benth.
Sambucus
canadensis L., American elder
nigra L. (PRE), European elder
Schinus
molle L., pepper tree
terebinthifolius Raddi var. acutifolius Engl., Brazilian pepper tree
Sesbania
bispinosa (Jacq.) W.F. Wight var. bispinosa (Ross 1972), spiny sesbania
punicea (Cav.) Benth., red sesbania
Solanum
hermannii Dun., bitter apple
mauritianum Scop., bug tree
pseudocapsicum L., Jerusalem cherry
seaforthianum Andr., potato creeper
Spathodea campanulata Beauv., African flame tree
Spiraea cantoniensis Lour., Reeves spiraea
Syncarpia glomulifera (Sm.) Niedenzu (MacDonald & Jarman 1985),
turpentine tree
Syzygium cumini (L.) Skeels, Java plum
Thevetia pemviana (Pers.) K. Schum., yellow oleander
Tipuana tipu (Benth.) O. Kuntze, tipu tree
Tithonia
diversifolia (Hemsl.) A. Gray, Mexican sunflower
rotundifolia (Mill.) S.F. Blake (Ross 1972)
Toona ciliata M.J. Roem. (MacDonald & Jarman 1985), toon tree
Ulex europaeus L., gorse
Vitex trifolia L. (Ross 1972)
Yucca sp. cf. Y. aloifolia L., Spanish bayonet
Bothalia 19,2: 263-274 (1989)
Miscellaneous notes
VARIOUS AUTHORS
IMPROVING THE RESOLUTION OF FLORISTIC/HABITAT PATTERN CORRELATIONS
ON PHYTOSOCIOLOGICAL TABLES
INTRODUCTION
One of the aims of causal-analytical vegetation research
is to analyse the reaction of plant groups to a combination
of habitat factors. In particular, it is important to recognize
the factors that are primarily responsible for floristic
differentiation (Mueller-Dombois & Ellenberg 1974).
Tabular portrayal of vegetation-habitat relationships is an
effective means of realizing this objective. Traditionally,
habitat factors in the form of itemized symbols are
appended above the floristic classification without any
attempt at clustering on the basis of similarity
(Bredenkamp 1975; Robinson 1976; Van der Meulen 1979;
McDonald 1983; Van Rooyen 1983; Westfall 1985).
Presentation of habitat data in this format may be termed
‘passive’ since these data require no additional
manipulation. Interpretation in this context, however, can
be difficult and cumbersome. An improved method of
portraying habitat correlations, whereby ‘diagnostic’
groups of habitat factors are juxtaposed above floristic
units, is described here. This involves a more ‘active’ type
of habitat classification.
METHODS
In the Braun-Blanquet phytosoeiological approach, as
described by Westhoff & Van der Maarel (1973), Werger
(1974) and Mueller-Dombois & Ellenberg (1974), the cover-
abundance of plant species in stands of vegetation is
presented in two-way tables with columns representing
stands (releves) and rows representing species. Columns
are re-arranged so as to group releves having similar
species and rows are re-arranged to group species
occupying similar releves.
By treating habitat factors as species on a presence/
absence basis, they can be re-arranged within the
framework of a floristic classification to form diagnostic
groups of habitat/environmental factors.
The PHYTOTAB computer package (Westfall et al.
1982) was used to structure a floristic table for the grass-
lands of the Sabie area, Eastern Transvaal Escarpment.
Thirteen syntaxa (including nine communities) were elicit-
ed from 37 releves. In a separate table, habitat and
structural data rows pertaining to each releve were su-
perimposed on the floristic classification and re-arranged
into groups occupying similar releves. In this way, habitat
groups were highlighted in the same fashion as species
groups (Table 1).
RESULTS AND DISCUSSION
The habitat groups provide a ready means of identifying
and ‘labelling’ the different syntaxa. For example, the
Gladiolus densiflorus—Loudetia simplex Short Closed
Grassland (Community 45) is found on the Escarpment
Lower Slopes in the Transitional Mistbelt between 1 112
m and 1 233 m elevation on level terraces in Land Type
Ab33b (Table 1). Conversely, the Cliffortia
repens -Loudetia simplex Short Open Shrubland
(Community 46) is found on the Escarpment Upper Slopes
on fairly rocky sites of Land Type Ab36a (Table 1). These
examples are taken from Deall (1985).
The habitat groups may also be useful for highlighting
specific factors influencing floristic differentiation. For
example, the Wahlenbergia huttonii— Eragrostis racemosa
Low Closed Grassland (Community 49) is differentiated
by the Acrotome hispida species group (Table 1: 10). Such
differentiation is probably largely influenced by the red
clay soils derived from Oaktree Dolomite (Table 1).
The advantages of ‘active’ as opposed to ‘passive’
classification of habitat for correlation with floristic
classifications are manifold:
1, ‘noise’ is more easily observed;
2, pattern is better defined and therefore interpretability
is enhanced;
3, pattern is not obscured by parameters with small class
intervals;
4, gradients can, if necessary, be indicated by means
of class intervals;
5, tables can, if necessary, be constructed independently
of floristics;
6, there is practically no limit to the number of habitat-
factor/class-interval combinations that can be used.
The portrayal of habitat factors in this manner is a
convenient visual aid, facilitating rapid environmental
interpretation of floristic classifications.
ACKNOWLEDGEMENTS
The authors wish to thank Dr J.C. Scheepers for helpful
comments and Mrs M. van der Merwe for typing the text.
REFERENCES
BREDENKAMP, G.J. 1975. 'n Plantsosiologiese studie van die Suiker-
bosrandnatuurreservaat. M.Sc. thesis, University of Pretoria.
DEALL, G.B. 1985. A plant-ecological study of the Eastern Transvaal
Escarpment in the Sabie area. M.Sc. thesis, University of
Pretoria.
McDONALD, D.I 1983. The vegetation of Swartboschkloof, Jonkershoek,
Cape Province, South Africa. M.Sc. thesis, University of Cape
town.
MUELLER-DOMBOIS, D. & ELLENBERG, H. 1974. Aims and
methods of vegetation ecology. Wiley, New York.
264
Bothalia 19,2 (1989)
TABLE 1. — Floristic classification and habitat correlation in Grassland of the Humid Mistbelt, Sabie Area (from Deall 1985)
(1) Differential species of the Gladiolus densiflorus—Loudetia simplex Short Closed Grassland (Community 45)
Indigofera sp. (Fb)
Gladiolus densiflorus (Fb)
Lopholaena disticha (Fb)
Aster comptonii (Fb)
Selago atherstonei (Fb)
13 3 2
4 12 2 2
12 2 1
2 2
2 2
(2) Differential species of the Cliffortia repens- Loudetia simplex Short Open Shrubland (Community 46)
Lobelia decipiens (Fb)
Cliffortia repens (Sh)
Styppeiochloa gynoglossa (Gr)
Senecio oxyriifolius (Fb)
Helichrysum mimetes (Fb)
Pycreus muricatus (Cy)
3 3
3 3
3 2
3 3
3 3
(3) Differential species of the Tetraselago natalensis—Monocymbium ceresiiforme Low Closed Grassland (Community 47)
(5) Differential species of Communities 45—47
+ Denotes presence. Digits 1-9 in matrix denote Domin-Krajina cover-abundance values. Growth forms: Tr = tree; Sh = shrub; Ln — lianoid;
Fb = forb or herb; Cy = sedge; Gr = grass; Pt = fern.
Bothalia 19,2 (1989)
265
TABLE 1, — Floristic classification and habitat correlation in Grassland of the Humid Mistbelt, Sabie Area (from Deall 1985) (continued)
(6) Differential species of the Rendlia altera- Monocymbium ceresiiforme Low Closed Grassland (Community 48)
Alloteropsis semialata subsp. eckloniana (Gr)
Commelina sp. (Fb)
Senecio erubescens (Fb)
Senecio gerrardii (Fb)
Nidorella auriculata (Sh)
Alhanasia calva (Fb)
Helichrysum sp. (Fb)
Drosera sp. (Fb)
Helichrysum cephaloideum (Fb)
Euphorbia striata (Fb)
Polygala hottentotta (Fb)
(7) Differential species of Communities 47—48
Kyllinga alba (Cy) 2323332 2
Rendlia altera (Gr) 5 3 6 9 2
Tolpis capensis (Fb) 2 1 3 2 1
(8) Differential species of Communities 46-48
Selago muddii (Fb)
Harpochloa falx (Gr)
Koeleria capensis (Gr)
Trachyandra saltii (Fb)
Hypoxis filiformis (Fb)
Stiburus alopecuroides (Gr)
Panicum ecklonii (Gr)
Stachys nigricans (Fb)
(9) Differential species of Communities 45-48
Hemizygia subvelutina (Fb) 23 3433 223
Alhanasia acerosa (Fb) 1 3 3 1 2 1 5 5 4
(10) Differential species of the Wahlenbergia huttonii—Eragrostis racemosa Low Closed Grassland (Community 49)
Acrotome hispida (Fb)
Hyparrhenia hirta (Gr)
Wahlenbergia huttonii (Fb)
Sporobolus ‘complex’ (Gr)
Brachiaria subulifolia (Gr)
Digitaria apiculata (Gr)
Eriosema cordatum (Fb)
Hibiscus aethiopicus var. ovatus
Pearsonia aristata (Fb)
Sonchus integrifolius (Fb)
Helichrysum subulifolium (Fb)
Tristachya leucothrix (Gr)
Senecio latifolius (Fb)
Triumfetta welwitschii var. hirsuta (Fb)
(11) Differential species of the Hypoxis multiceps— Wahlenbergia huttonii—Eragrostis racemosa Variant (49A)
+ Denotes presence. Digits 1—9 in matrix denote Domin-Krajina cover-abundance values. Growth forms: Tr — tree, Sh shrub, Ln lianoid,
FT) = forb or herb; Cy = sedge; Gr = grass; IT = fern.
266 Bothalia 19,2 (1989)
TABLE 1. — Floristic classification and habitat correlation in Grassland of the Humid Mistbelt, Sabie Area (from Deall 1985) (continued)
(12) Differential species of the Parinari capensis subsp. capensis—Wahlenbergia huttonii—Eragrostis racemosa Variant (49B)
Raphionacme elata (Fb)
Sphenostylis angustifolia (Ln)
Piloselloides hirsuta (Fb)
(13) Differential species of Communities 46—49
Becium obovatum (Fb)
Indigofera sanguined (Fb)
2 2
3 5 2 4
2 2
3
3 3
(14) Differential species of Communities 45-49
+ Denotes presence. Digits 1—9 in matrix denote Domin-Krajina cover-abundance values. Growth forms: Tr = tree; Sh = shrub; Ln = lianoid;
Fb = forb or herb; Cy = sedge; Gr = grass; Pt = fern.
ROBINSON, E.R. 1976. Phytosociology of the Namib Desert Park, South
West Africa. M.Sc. thesis, University of Natal, Pietermaritzburg.
VAN DER MEULEN, F. 1979. Plant sociology of the western Transvaal
Bushveld, South Africa. Cramer, Vaduz.
VAN ROOYEN, N. 1983. Die plantegroei van die Roodeplaatdamnatuur-
reservaat. II. Die plantgemeenskappe. South African Journal of
Botany 2: 115—125.
WERGER, M.J.A. 1974. On concepts and techniques applied in the
Zurich-Montpellier method of vegetation survey. Bothalia 11:
309-323.
WESTFALL, R.H. 1985. The plant ecology of the farm Groothoek,
Thabazimbi District. II. Classification. Bothalia 15: 655-688.
WESTFALL, R.H., DEDNAM, G., VAN ROOYEN, N. & THERON,
G.K. 1982. PHYTOTAB — a program package for Braun-
Blanquet tables. Vegetatio 49: 35—37.
WESTHOFF, V. & VAN DER MAAREL, E. 1973. The Braun-Blanquet
approach. In R.H. Whittaker, Ordination and classification of
vegetation. Handbook of Vegetation Science 5 : 617—726. Junk,
The Hague.
G.B. DEALL and R.H. WESTFALL
MS. received: 1988.10.21.
PLANT COLLECTING APPARATUS FOR TAXONOMIC AND ECOLOGICAL STUDIES
1. A LIGHTWEIGHT PLASTIC PLANT PRESS FOR ON-SITE SPECIMEN PRESSING
The advantages of pressing plant specimens on-site,
while fresh, are: 1, a natural layout may be achieved that
can often resemble the original plant habit; 2, the specimen
forms a site record rather than a reconstruction of a site
record; 3, fresh material is often easier to arrange than
wilted material; 4, it increases sampling efficiency because
only sufficient material to fill the required number of
herbarium sheets need be collected, and 5, it reduces
information loss through a reduction in specimen handling.
Factors which can inhibit on-site plant pressing include
the difficulty in carrying plant presses in the field and
pressing plant specimens in even moderate wind. The
following is a description of an economical and portable
plant press that can be used on-site, even in windy con-
ditions.
A rectangular plastic bin, with stacking base design for
rigidity, and dimensions of 490 x 260 x 240 mm forms
the basic unit. These are available commercially. The sides
can be cut to a convenient depth. A depth of 115 mm
provided a reasonable compromise between bulkiness and
capacity. Four slots are cut in the angle between the base
and the long side as illustrated in Figure 1, for press strap
insertion. The two press straps are each 25 x 1 300 mm
nylon straps with double D-rings on one end. The straps
can be secured to the bottom of the bin by rivets so that
the D-ring side of the strap is slightly longer than the bin
FIGURE 1. — Portable press with shoulder strap showing strap layout
and spare specimen folders secured inside lid.
Bothalia 19,2 (1989)
267
FIGURE 2. — Portable press mounted on rucksack frame with additional
pockets for equipment.
depth. Figure 1 shows the inside of the plant press with
press straps and shoulder strap which is attached to the
bottom of the bin by rivets.
The lid which is also available commercially is cut to
fit snugly inside the press. A nylon strap can be riveted
to one face of the lid to form a handle. Two loops of
elasticized straps of 25 mm width are riveted to the lid
for holding spare specimen folders on the face opposite
to that with the handle (Figure 1).
In use, folders are numbered prior to pressing, while
still held down by the elasticized straps as the lid also forms
a good writing surface. If the end of the press strap is
looped before threading through the second D-ring then
the free end becomes a quick-release device to facilitate
press opening. The press can be used either with a
shoulder strap (Figure 1) or it can be mounted on a
rucksack frame (Figure 2). The former is often more
suitable for ecological studies where an abundance of
equipment often necessitates a separate rucksack.
The portable press is not suitable for drying specimens,
therefore drying paper is not required. Specimens should
be transferred to drying presses with drying paper inserted
between each folder as soon as possible. The portable press
protects specimens from moist vegetation and light rain.
The press sides afford protection from wind when laying
out specimens and clothes pegs can be used to keep folder
covers open, if necessary. The sides also ensure that the
specimens are not subjected to undue shifting such as is
often experienced when loading and aligning standard
presses. Pressing of fresh material before transfer to a
standard drying press also facilitates alignment of specimen
folders.
Use of a portable press should improve the quality and
information content of plant specimens.
ACKNOWLEDGEMENTS
The authors thank L. Fish, T.H. Arnold, J.M. van
Staden and J.C. Scheepers for assistance and suggestions.
M.D. PANAGOS* and R.H. WESTFALL*
*Botanical Research Institute, Department of Agriculture and Water
Supply, Private Bag X101, Pretoria 0001.
MS. received: 1988.10.20.
2. COLDAT: A FIELD-DATA CAPTURE PROGRAM FOR COLLECTOR’S DATA AND HERBARIUM LABELS
A plant specimen is rarely worth preserving unless the
place and time of collection and the collector, preferably
with a collecting number, are known. Information
concerning the habitat and plant characteristics, not evident
in the dried specimen, can enhance the value of a specimen
considerably.
Initial recording of observations relevant to herbarium
specimens should be recorded at the place and time of
collection. However, the occurrence of errors and hence
the reliability of the data, can be considerably influenced
by the number of times the data are manually transferred,
from initial recording, to the herbarium label. Writing
herbarium labels for more than one specimen from a
particular collecting site entails much duplication as only
the plant characteristics and specimen numbers change.
It should, therefore, not be necessary to record all data
for each label, at a site, thereby saving on high-cost field
time. Insertion of data at a later stage may lead to errors.
Computerized field-data capture can facilitate on-site
data recording, overcome the problem of duplicating
data and eliminate manual data transfer. The program
RH WESTFALL Coll no: 103
Specimen no: 2288
Grldr-ef- 2428AC Region: Transvaal
Date: 1982/1 J/14
HaJ loci UATERBER6
run loc: UAAL WATER
Prec loc: 242134S280244E FARH WOLUEN
FONTEIN
Altitude: 1188 m Aspect: N£
Life form: Climber
Plant height: 10 cm
Hab i t at
SHORT CLOSED WOODLAND ON SANDY ROCK
Y SOILS. GENTLE SLOPE ON INTERFLUUE.
Plant note*
CREEPER WITH PURPLE FLOWERS AND TUB
ER.
Loc/Rel no: 820041 Bio effect*: 10
Ueg type: 2 Substrate: 2
Hoist reg: 2 Soil type: 2
PLANT NAHE:
FIGURE 3.— Mini label for
collector’s register
with provision for later
insertion of plant
name.
268
Bothalia 19,2 (1989)
BOTANICAL RESEARCH INSTITUTE, SOUTH AFRICA
2428AC Grid ref. Regio Transvaal
RH WESTFALL Legit Anno 1987/11/14
2288 No. Alt. 1180 m
Name
Locality:
WATERBERG VAALWATER
2421 34S280744E FARM W0LWENF0NTEIN
Habitat: Aspect: NE
SHORT CLOSED WOODLAND ON SANDY ROCKY SOILS. GENTLE
SLOPE ON INTERFLUVE.
Notes: Climber 10 cm
CREEPER WITH PURPLE FLOWERS AND TUBER.
Det. I Ref.
Bio.eff. 10 Veg. type 7 Substrate 2
Moisture reg. 2 Soil type 2 Loc.no. 870041
FIGURE 4. — Herbarium label from mainframe printer after
computerized data transer to B7900 computer.
COLDAT, written in BASIC for the Sharp PC1500A pocket
computer is designed for field-data capture of herbarium
label data, data transfer and label production.
Data input provides for the following: collector’s name
and number; region; grid reference; date; major, minor
and precise localities; locality or releve number; aspect;
altitude; PRECIS (Morris & Glen 1978) codes for biotic
effects, vegetation type, substrate, moisture regime and
soil type; habitat notes; specimen number; plant height;
PRECIS (Morris & Glen 1978) life-form code; and plant
characteristic notes. Complete data are entered for the first
label, thereafter, only data that change are entered.
Specimen numbers are allocated automatically after the
first number. Mini labels (Figure 3) can be printed in the
field. They form the original hard copy of the data and
can be pasted into a book to form a collector’s register.
Provision is made on the mini labels (Figure 3) for later
insertion of plant names. Data are transferred to cassette
before the computer capacity of 40 labels is reached.
Data transfer from cassette to a host computer can be
achieved with a suitable program. The program
COLTRAN written in BASIC for the Sharp PC1500A
pocket computer transfers data from cassette to the
Burroughs B7900 mainframe computer. The transfer
format required is the same as that used for printing large
format labels (Figure 4) on the mainframe printer. The
data can then be transferred to a data bank.
This procedure permits minimum data input and
physical data handling with consequent increase in data
reliability. Experience has shown that minimum data input
leads to a greater concentration on plant characteristics
and habitat recording on-site, with a corresponding in-
crease in quality of herbarium label information, compared
to manual methods used previously.
ACKNOWLEDGEMENTS
The author thanks Mr T.H. Arnold and Dr J.C.
Scheepers for comments and suggestions.
REFERENCES
MORRIS, J.W. & GLEN, H.F. 1978. PRECIS the National Herbarium
of South Africa (PRE) computerized information system. Taxon
27: 449-462.
R.H. WESTFALL
MS. received: 1988.10.20.
3. A NEW TOP-LOADING PLANT PRESS FOR OFF-SITE SPECIMEN PRESSING
Correct alignment of specimen folders and drying papers
when pressing plants reduces specimen damage caused by
specimen-folder shift and increases the functional life span
of specimen folders and drying papers through a reduction
of friction and edge-wear. It is, furthermore, often difficult
to close a standard strap-type plant press without causing
folder shift especially when uneven plant specimens are
pressed. The potential for specimen damage is increased
with the insertion of additional specimens, windy
conditions and the necessity for changing drying papers.
The procedure for pressing plants with a standard
strap-type plant press can be laborious and often frus-
trating.
The plant press shown in Figures 5 and 6 minimizes
specimen-folder shift, even with bulky specimens and is
easy to use. The features (Figure 5) include:
1, a top which is hinged by means of two nuts and bolts
(item 7) for loading and access to plant specimens while
in the press;
2, a steel frame (item 1) which provides alignment
guides for correct alignment of specimen folders and dry-
ing paper;
3, quick pressure release with the spring release bar
(item 6) and release mechanism (item 5);
4, rapid opening and closing of the hinged top with
locking pin (item 8); and
5, a simple and effective pressure mechanism, using
modified silicone caulking guns (item 2).
The double pressure action provided by the two caulking
guns and the swivel action of the pressure plate (item 11)
allows for even pressure to be applied to uneven specimens.
Pressure is applied by squeezing the caulking gun handles.
In use, only two fingers need be used on each handle to
exert a pressure of about 80 kg on the plant specimens.
Pressures greater than 80 kg could damage plant
specimens.
The mass of the empty top-loading plant press is about
12 kg which inhibits movement of the press in the vehicle
but can preclude on-site use. The use of a portable press
(Panagos & Westfall 1989a) is recommended whenever
collecting outside the immediate vicinity of the collector’s
vehicle. Specimen folder transfer and drying paper
insertion are extremely simple and quick when a portable
press is used.
String or ribbon can be draped inside the top-loading
plant press prior to loading (Figure 6). The specimen
Bothalia 19,2 (1989)
269
FIGURE 5. — Plan of top-loading plant press with materials required (a) side view; (b) end view and (c) top view.
Use of the top-loading plant press can reduce
information loss through specimen damage and time spent
on pressing specimens, as well as increase the life span
of specimen folders and drying paper. It can even be used
in windy conditions.
ACKNOWLEDGEMENTS
The authors thank J.C. Scheepers and J.M. van Staden
for suggestions and assistance.
REFERENCES
PANAGOS, M.D. & WESTFALL, R.H. 1989a. Plant collecting apparatus
for taxonomic and ecological studies. 1. A lightweight plastic plant
press for on-site specimen pressing. Bothalia 19: 266—267.
FIGURE 6. — Open top-loading plant press showing draped ribbons for
batch removal of plant specimens.
folders can then be tied and removed in a batch when
required, minimizing specimen handling and obviating the
need to remove the top-loading plant press from the
vehicle.
R.H. WESTFALL* P.J. BRITZ** and M.D. PANAGOS*
* Botanical Research Institute, Department of Agriculture and Water
Supply, Private Bag X101, Pretoria 0001.
** Agricultural Engineering, Department of Agriculture and Water Supply,
Private Bag X515, Silverton 0127.
MS. received: 1988.10.20.
4. DRIER-TRANSPORTERS FOR PLANT PRESSES
Extended plant collecting trips necessitate frequent
changes of drying paper to prevent plant specimens
becoming mouldy. This, together with vehicle interior
space limitations, which can lead to frequent plant press
restacking, can cause specimen damage. Changing drying
paper is also time-consuming. Plant presses transported
outside the vehicle can allow better interior space
utilization and the method can include a specimen drying
function not possible inside the vehicle.
A fully enclosed aluminium rack, mounted on the roof
of a NISSAN long wheelbase Ekonovan, is shown in
Figure 7. The roofrack is suitable for 12 standard strap-type
plant presses, filled up to about 350 mm in height. Plant
presses are loaded through four side-flaps and are secured
by straps in guides which ensure airflow around each plant
press. The end-flaps can remain open, even when
travelling, for ventilation. In moist or dusty conditions the
end-flaps can be closed. The exterior is painted black for
heat absorption, to assist the drying process. The drying
process is, however, not even throughout the roofrack as
the four front presses have a markedly faster drying rate
than the others when the vehicle is moving.
270
Bothalia 19,2 (1989)
FIGURE 7. — Roofrack drier-transporter showing open front and side
flaps and stowed canvas awning on awning pole box.
A partitioned 2,12 m trailer with a perspex lid over the
rear partition for light transmission and heat capture is
shown in Figure 8. A large ventilator on each of the sides,
the tailgate and the rear of the lid ensure adequate
ventilation when stationary and, forced ventilation when
moving. The lid and a side ventilator, in the open position,
can be seen in Figure 8. The front partition can be used
to stow camping gear. This configuration is suitable for
six top-loading plant presses (Westfall et al. 1989). Both
partitions have webbed rubber mats for floor protection
and insulation.
In later models the rear lid was made of metal and
painted black for heat absorption as the perspex, apart
from being expensive, is inclined to crack. This did not
influence internal temperatures significantly as
temperatures of about 53°C at 1 700 m altitude were
recorded in both models when cloud cover was absent.
The perspex model did, however, appear to have a faster
rate of heat buildup than the painted model.
The roofrack drier-transporter should dry plant
specimens more efficiently because considerably more air
can pass over the plant presses than in the trailer. However,
in inland summer conditions, the trailer has proved
adequate for specimen drying, eliminating the need to
change drying paper in all but the wettest specimens, such
as geophytes with large, moist storage organs. In
comparison with the trailer, the roofrack has the following
disadvantages: 1, loading and unloading of plant presses
is difficult; 2, plants cannot be pressed with the press in
situ\ 3, noticeably higher fuel consumption, especially with
lid and top-loading plant presses.
speeds in excess of 90 km/h; 4, altered vehicle handling
characteristics; and 5, higher manufacturing cost.
The plant-press securing straps of the roofrack can be
used to take up slack in the plant presses as the material
dries. The roofrack is also equipped with a canvas awning
on an awning pole box, with width almost that of the
roofrack length and length sufficient to pass over the open
door and reach the ground. The awning is rolled and
stowed on the side of the roofrack as shown in Figure 7.
The trailer lid can also offer some protection from
inclement weather when raised, as shown in Figure 8.
The trailer drier-transporter, although possibly slightly
less efficient than the roofrack drier-transporter for drying
plant specimens, can reduce plant-press handling
considerably, thereby saving time and possible information
loss through specimen damage.
ACKNOWLEDGEMENTS
The authors thank Drs B. de Winter, J.C. Scheepers and
Mr J.M. van Staden for comments and assistance.
REFERENCES
WESTFALL, R.H., BRITZ, P.J. & PANAGOS, M.D. 1989. Plant
collecting apparatus for taxonomic and ecological studies. 3. A
new top-loading plant press for off-site specimen pressing.
Bothalia 19: 268—269.
M.D. PANAGOS and R.H. WESTFALL
MS. received: 1988.10.20.
5. A GAS DRIER FOR FIELD DRYING OF PLANT SPECIMENS
The use of a portable specimen drier can overcome the
necessity for changing drying paper either when plant
specimens are too moist for drying in a drier-transporter
(Panagos & Westfall 1989b) or when working in very
humid conditions. Characteristics of a portable specimen
drier should include: (a) robust construction; (b) efficient
drying; (c) suitability for drying specimens in plant
presses; (d) simple and inexpensive construction; and (e)
independence of electrical power. These characteristics
exclude specimen driers such as that described by Botha
& Coetzee (1976), in which gas lamps are used as heat
sources, because of the fragility of the mantles.
Furthermore, aluminium construction is expensive in
terms of both material and labour.
In designing a portable specimen drier, maximum use
was made of commercially available apparatus in order
to reduce material and labour costs. A Cadac Skottelskaar
Braai forms the basic unit. Modification simply entails
the following:
Bothalia 19,2 (1989)
271
FIGURE 9. — Specimen drier showing (a) press support unit, (b) burner
pan, (c) stem, (d) gas cylinder stabilizers, (e) 2,8 kg gas cylinder
and (f) retaining clip handles wedged on the gas cylinder.
1, removing the cooking dish;
2, reducing the stem to an overall length of 155 mm
(excluding jet protrusion) by cutting, for stability;
3, replacing the standard jet with a Cadac no. 31 (0,12
mm) jet for heat reduction;
4, placing rubber tubing over the retaining clip handles
so that the handles can be wedged against the gas cylinder
for extra stability; and
5, drilling five 11 mm diameter holes in the base of the
burner pan in the same contour as the three existing holes
for increased airflow.
To prevent the grid centre from overheating, the flame
is deflected by a 55 mm deep, lipped cone and a 6 mm
thick plate both of fibre-reinforced cement and with a
diameter of 275 mm. These are bolted to the underside
of the grid. A fibre-reinforced cement ashtray lid is suitable
for the cone and can withstand temperatures of up to about
200°C.
A 550 x 1 960 mm canvas skirt, with eyes at 100 mm
intervals along the edge of one side for threading a securing
cord, channels air through the grid and press.
The assembled field drier mounted on a 2,8 kg gas
cylinder is shown in Figure 9. Assembly entails screwing
the stem into the gas cylinder and placing the burner pan
onto the stem and the press support unit onto the burner
pan. The retaining clip handles are then wedged onto the
cylinder. The cylinder can be stabilized by the addition
of gas cylinder stabilizers as shown in Figure 9. Calcula-
tions indicate a burning time in excess of 120 hours with
a 2,8 kg gas cylinder as shown in Figure 9.
After lighting through the large burner pan holes and
adjusting the flame to a low setting, the plant press is
placed with the long side on the press support unit in such
a way that the hot air can rise through the plant press
cardboard ventilators and between the drying papers. The
canvas skirt is wrapped around the press and secured with
the cord as shown in Figure 10. Some excess hot air should
be able to pass between the press and skirt to prevent over-
heating.
v
FIGURE 10. — Specimen drier with plant press showing the canvas skirt
wrapped around the plant press.
Air heating, press support and flame deflection are
provided by a simple unit placed on the burner pan for
supporting the press.
This unit consists of a flat expanded metal grid 520 mm
in diameter with the following specifications: 2,5 mm
strand width; 1,6 mm strand thickness; mass 9,1 kg m~2;
and 21% opening. The grid serves to heat the air and also
prevents gas ignition on the upper side of the grid, as in
a Davy lamp. A ring of 10 mm diameter rod is brazed on
the upper side of the grid around the circumference, to
provide a smooth edge and increase rigidity.
The plant press is supported by a 370 x 370 mm square
of 10 mm diameter rod. Four struts, also of 10 mm
diameter rod, are brazed to the corners of the square and
to the grid circumference rod, to support the plant press
105 mm above the grid.
272
Bothalia 19,2 (1989)
After use a plastic cap should be placed over the jet and
a plug inserted into the open end of the stems, to prevent
jet clogging. The components can then be wrapped in the
canvas skirt for transportation.
The specimen drier is robust, extremely efficient, simple
to construct and easy to transport.
ACKNOWLEDGEMENTS
The authors thank Dr B. de Winter, Mr T.H. Arnold,
Dr J.C. Scheepers and Mr J.M. van Staden for suggestions
and assistance.
REFERENCES
BOTHA, D.J. & COETZEE, J. 1976. A portable drier for plant specimens.
Journal of South African Botany 42: 41—44.
PANAGOS, M.D. & WESTFALL, R.H. 1989b. Plant collecting apparatus
for taxonomic and ecological studies. 4. Drier-transporters for
plant presses. Bothalia 19: 269—270.
M.D. PANAGOS* P.J. BRITZ** and R.H. WESTFALL*
* Botanical Research Institute, Department of Agriculture and Water
Supply, Private Bag X101, Pretoria 0001.
** Agricultural Engineering, Department of Agriculture and Water Supply,
Private Bag X515, Silverton 0127.
MS. received: 1988.10.20.
6. A TRANSPORTABLE MAP CABINET FOR VEHICLE AND OFFICE USE
The simplest reference for collecting or sample site
locality is the grid system of parallels and meridians, used
throughout the world, and is the most common of
co-ordinate systems (Strahler 1975). This grid system is
also used on the South Africa 1:50 000 sheets which,
because of scale convenience, are frequently used for
determination of site locality.
The precision with which site localities are determined
can influence 1, subsequent mapping precision; 2,
re-collecting or re-sampling efficiency; 3, re-sampling
validity; and 4, validity of correlations with other sources.
In field situations, it is often impracticable to determine
positions on a map with greater precision than the nearest
millimetre. This implies a ground error of less than, or
equal to 25 m at 1:50 000 scale. It is, therefore, not
necessary under these circumstances, to record a locality
with a precision greater than the nearest second, which
implies a ground error of less than 16 m. This level of
precision can be considered adequate for much plant col-
lecting and vegetation sampling work. Fluctuating
humidity with consequent stretch and shrinkage of maps
should not significantly alter this level of precision because
of proportionality. However, the method of map trans-
portation, including folding, rolling and subjection to
impacts can cause considerable map distortion, often with
irregular loss of precision.
These causes of precision loss can be overcome with
a vertical map filing cabinet. A multi-prong map-support
system, for use with pre-punched tape on the maps,
facilitates indexing, filing and retrieval. This system is used
in the map cabinet shown in Figure 11. Maps are supported
MATERIALS
FIGURE 11.— Plan of portable map cabinet with list of materials required.
Bothalia 19,2 (1989)
273
FIGURE 12. — Portable map cabinet in vehicle with front flap down
to form a working surface. Note opissometer (a), attached with
spring clips, for measuring distance.
by five metal prongs, three of which are attached to a fixed
flat iron bar and two of which are attached to a removable
bar for map retrieval. Holes drilled in the bars opposite
each prong allow support for the removable bar and
simplify map retrieval and filing. The removable bar is
held in place by the front flap when in the closed position.
The front flap also serves as a large working surface and
is supported by fold-down legs when open.
The map cabinet has a net mass of about 20 kg and a
capacity of over 100 maps. It can be carried by one person
and is suitable for use in a vehicle such as is illustrated
in Figure 12. Map storage and transport as well as site
location and plotting in the field are greatly facilitated with
the use of this map cabinet.
ACKNOWLEDGEMENTS
The authors thank Dr J.C. Scheepers and Mr J.M. van
Staden for suggestions and assistance.
REFERENCE
STRAHLER, A.N. 1975. Physical geography. Wiley, New York.
R.H. WESTFALL* and M.D. PANAGOS*
* Botanical Research Institute, Department of Agriculture and Water
Supply, Private Bag X101, Pretoria 0001.
MS. received: 1988.10.20.
7. A TRANSPORTABLE CAMPING KITCHEN FOR VEHICLE USE
A considerable saving of time and vehicle space can be
effected by making use of efficient provisioning equipment.
A simple, sturdy, transportable camping kitchen suitable
for 8/12-seater light bus is shown, as well as a standing
basin which is very useful in situations where a single
camp site is used (Figure 13).
The upper partitioned area is for food, to which easy
access can be gained through the upper lid (Figure 14).
The lower partitioned area is suitable for 2,0 x 2,8 kg gas
bottles with burners, plastic cooler box and eating, cooking
and cleaning utensils. The front lid also serves as a
working surface.
The capacity of the camping kitchen unit is sufficient
for utensils and food for three to four people, for up to
two weeks. Exterior dimensions are such that the unit can
be positioned behind the rear seat with access through the
rear door of the light bus. The unit is best filled while
in position as the laden mass inhibits movement. The spare
wheel should be removed from the side panel and can be
placed under the rear seat because of restricted access with
the laden unit. The working surface support chains can
be covered with cycle tubes as chain covers (Figure 14)
to suppress rattles when the vehicle is in motion. Sufficient
space exists between the unit and side panel of the vehicle
for a 20f plastic water container. This should be equipped
with an extension tube and tap for easy access to water
during food preparation.
Stainless steel cooking and eating utensils, which
experience has shown are easier to clean than aluminium,
can be obtained economically from mine suppliers. The
FIGURE 13. — The camping kitchen positioned in the rear of a light bus
showing working surface, water container and standing basin.
274
Bothalia 19,2 (1989)
MATERIALS
FIGURE 14. — Plan for transportable camping kitchen listing materials required.
pan unit of a CADAC minibraai was also found to dissipate
heat from the gas burner evenly over thin-bottomed pots,
thereby preventing food from burning. Some protection
from light rain, during food preparation is afforded by the
vehicle’s open rear door. A 12V fluorescent tube with
socket for the vehicle’s cigarette lighter receptacle can
provide sufficient light for meal preparation and also for
work within the vehicle.
ACKNOWLEDGEMENT
The authors thank Dr J.C. Scheepers for comments and
suggestions.
R.H. WESTFALL* M.D. PANAGOS* and J.M. VAN STADEN*
* Botanical Research Institute, Department of Agriculture and Water
Supply, Private Bag X101, Pretoria 0001.
MS. received: 1988.10.20.
Bothalia 19,2 : 275 -294 (1989)
New taxa, new records and name changes for southern African plants
B.C. DE WET, G.E. GIBES RUSSELL, G. GERMISHUIZEN, B.D. SCHRIRE, M. JORDAAN, B.J. PIENAAR,
W.G. WELMAN, C. REID, C.M. VAN WYK, L. FISH, K.L. IMMELMAN, J. VAN ROOY, H.F. GLEN and
N.P. BARKER*
ABSTRACT
Alterations to the inventory of about 24 000 species and infraspecific taxa of bryophytes and vascular plants in southern
Africa are reported for the year 1988. The inventory, as currently maintained in the Taxon component of the PRECIS system,
contains the accepted name for each taxon, synonyms previously in use as accepted names during the past half-century,
and literature references necessary to identify species in each genus and to establish the synonymy. The inventory is updated
as new research affecting plant classification in southern Africa is published. During 1988 there were 744 alterations, affecting
about 3% of the total number of taxa.
UITTREKSEL
Daar word vir die jaar 1988 verslag gedoen van veranderings aan die lys van ongeveer 24 000 spesies en infraspesifieke
taksons van mosse en vaatplante in suidelike Afrika. Die lys, soos dit tans in die Takson-komponent van die PRECIS-stelsel
in stand gehou word, bevat die aanvaarde naam van elke takson, sinonieme voorheen in gebruik as aanvaarde name gedurende
die afgelope halfeeu, en literatuurverwysings wat nodig is om spesies in elke genus te identifiseer en om sinonieme vas
te stel. Die lys word bygewerk soos nuwe navorsing wat plant-klassifikasie in suidelike Afrika raak, gepubliseer word. Gedurende
1988 was daar 744 veranderinge wat ongeveer 3% van die totale aantal taksons verteenwoordig.
INTRODUCTION
This is the fifth in this series that reports annual
alterations to the complete inventory of southern African
plants maintained in the Taxon component of the computer
system PRECIS. References to the first three of these lists
are given in the introduction to the fourth (Gibbs Russell
et al. 1988). The format continues to be that used in
previous lists of the series. The complete and up-to-date
listing of names, literature and useful synonyms for all the
24 000 southern African plants is continuously maintained
in the Taxon component of PRECIS, and the plant
identification service of the Botanical Research Institute
uses the names as currently recorded. The Institute can
supply PRECIS listings of the most recent species
treatment for any family or genus. In addition the data files
for the alterations reported in this series have been available
on floppy disk in MS-DOS or BTOS format since 1988.
This year, as in 1985 and 1988, alterations for all plants
have been included. In 1986 and 1987, while the literature
and synonyms for dicotyledons were being completed,
only cryptogams and monocots could be covered in these
annual lists. A total of 744 changes were recorded during
1988, affecting about 3% of the total number of taxa in
the southern African flora.
The largest number of name changes is to be found in
genera in which active taxonomic research is in progress,
or for which revisions have recently been published. One
of the most extreme examples of this is the genus In-
digofera, where we record changes in 272 records (about
50% change). Because of the extent of the changes in this
genus, we break with the tradition of presenting only name
changes in these lists, and list Indigofera in full.
* All members of staff of the Botanical Research Institute, Private Bag
X101, Pretoria 0001.
MS. received: 1989.04.26.
This list covers name changes recorded since the
previous list went to press, and includes material received
before the end of February 1989. Due largely to postal
delays, therefore, this indicates that name changes
published in overseas journals after November 1988 will
only appear in next year’s list. Over 120 journals are
routinely scanned for relevant papers, and the changes
reported here were published in about 170 papers and
books.
Extensions being tested at present to Specimen- and
Taxon-PRECIS and the curatorial data base (Arnold in
press) mean that in future each name change will need
to be entered into the computer systems of the National
Herbarium only once. The full implementation of these
extensions will mean a drastic reduction in the effort
required to keep PRECIS up to date, and an improvement
in the overall standard of accuracy of the names in our
data bases. We welcome reports of changes from botanists
outside the Botanical Research Institute.
Spermatophyte families and genera follow the order and
numbering of the Englerian classification system as given
by Dyer (1975, 1976), with extensions to accommodate new
genera as described in detail by Morris & Glen (1978).
The only exception is the family Poaceae, which follows
an unpublished system of generic numbering. Bryophytes
follow Crosby & Magill (1981) and Grolle (1983).
Pteridophytes follow Schelpe & Anthony (1986). Species
are listed in alphabetical order: a name in current use
appears in upper case letters with its PRECIS number,
synonyms appear in lower case. Each synonym is entered
twice, once indented below the name for which it is a
synonym and once in its alphabetical place in the genus.
New collection records are indicated by quoting a
specimen and its locality. Naturalized taxa are shown by
an asterisk following the name. A few names have been
included because the author name or publication date have
to be changed; with most of these names the reason for
inclusion is not specifically mentioned.
276
Bothalia 19,2 (1989)
BRYOPHYTA Contributed by J.
HEPATICAE
RICCI AC EAE (H8)
1000
1016
1016
100
225
2 50
300
A 5 0
650
700
800
1050
1550
1750
1800
1850
1875
1950
2050
2150
2200
27 50
2950
32 50
184.
7. JOVET-AST. 1983. CRYPTOGAMIE 4,1: 37.
8. VOLK & PEROLD. 1984. BOTHALIA 15, 1 &
117-124.
9. VOLK & PEROLD. 1985. BOTHALIA 15, 3 &
531-539.
10. VOLK & PEROLD. 1986. BOTHALIA 16,1
33.
11. VOLK & PEROLD. 1986. BOTHALIA 16,2
201 .
12. PEROLD & VOLK. 1988. BOTHALIA 18,1
49.
13. VOLK & PEROLD. 1988. BOTHALIA 18,2
163.
14. PEROLD. 1989. BOTHALIA 19,1.
R. ALATOSPORA VOLK & PEROLD
R. ALBOLIMBATA S. ARNELL
(-R. albosquamata S. Arnell) 14
R. ALBOPOROSA PEROLD
R. ALBORNATA VOLK & PEROLD
R. albosquamata S. Arnell - R. ALBOLIMBATA
R. ARGENT EOLIM BATA VOLK & PEROLD
R. CAMPBELLIANA HOWE
R. canescens Steph. - R. TRICHOCARPA
R. capensis Steph. - R. LIMBATA
R. CONGOANA STEPH.
(-R. rhodesiae S. Arnell) 11
R. DUTHIEAE VOLK & PEROLD
R. HIRSUTA VOLK & PEROLD
R. LIMBATA BISCH.
(-R. capensis Steph.) 12
R. MAMMIFERA VOLK & PEROLD
R. MICROC ILIATA VOLK & PEROLD
R. MONTANA PEROLD
R. NIGRELLA DC.
R. PARVOAREOLATA VOLK & PEROLD
R. PERSSONI I KHAN
(Note spelling correction)
R. rhodesiae S. Arnell - R. CONGOANA
R. ROSEA VOLK & PEROLD
R. SCHELPEI VOLK & PEROLD
R. TRICHOCARPA HOWE
cens Steph.) 7
29-
169-
37-
155-
( ■ R •
MUSCI
POTT I ACE AE (14)
1301
1401
-BARBULA HEDW.
B . t richos tot
ea C. Muell. -
AUSTR ALAS IAE
DI DYMODON
-DESMATODON BRID.
2. TOWNSEND. 1987. LINDBERGI A 12: 67-71.
D. BOGOSICUS C. MUELL.
-DIDYMODON HEDW.
1. MAGILL. 1981. FSA .
2. ZANDER. 1978. PHYTOLOGI A 41,1: 11-32.
3. GUERRA & ROS. 1987. CRYPTOGAMIE 8,1: 64.
D. AUSTR ALAS 1 1 (HOOK. & GREV.) ZANDER
(-Barbula t r icho s t oraac ea C. Muell.) 1
(-Trichostomopsis australasiae (Hook. &
Grev.) Robinson) 2
D. TRIVIALIS (C. MUELL.) GUERRA
(-Trichostomopsis trivialis (C. Muell.)
Robinson) 3
-Tricho
3 P8 i
T. trivialis
s Card. - DIDYMODON
ae (Hook. & Grev.) Robins -
DIDYMODON A USTRALASI I
(C. Muell.) Robins - DIDYMODON
TRIVIALIS
GRIMMIACEAE (15)
-GRIMM I A HEDW.
G. nigrovirlde C. Muell. var.
(C. Muell.) Pa
LAMPROCARPUM
-RACOMITRIUM BRID.
2. OCHYRA ET AL. 1988. BULL. JARD. BOT. NAT.
BELG. 58: 225-258.
R. defoliatura Dix. - R. LAMPROCARPUM
R. LAMPROCARPUM (C. MUELL.) JAEG.
(-Grimmia nigrovirlde C. Muell. var.
robusticulum (C. Muell.) Par.)
2
(-R. defoliatura Dix.) 2
( - R . nigro-viride (C. Muell.) Par.) 2
( -R . pseudoacicular is (C. Muell.) Par.)
2
R. nlgrorvlride (C. Muell.) Par. - R.
LAMPROCARPUM
R. pseudoacicular is (C. Muell.) Par. - R.
LAMPROCARPUM
BRYACEAE (24)
1508
100
-BRYUM HEDW.
B. ALPINUM HUDS. EX WITH.
(Note change in spt
1618 -MACROMITRI UM BRID.
2. MAGILL & VAN ROOY. MS. FSA.
M. schlothe imiaef ormis Par. - CAR DOT I ELLA
SECUNDA
M. secundum C. Muell. - CARDOT I E L LA SECUNDA
500 M. SERPENS (HOOK. & GREV.) BRID.
( -M . tristratosum Dix.) 2
M. tristratosum Dix. - M. SERPENS
1620 -SCHLOTHEIMIA BRID.
2. MAGILL & VAN ROOY. MS. FSA.
S. exrug ulosa C. Muell. - S. RUFOPALLENS
100 S. FERRUGINEA (HOOK. & GREV.) BRID.
(-S. subventrosa Broth. & Bryhn) 2
425 S. RUFOGLAUCA C. MUELL.
450 S. RUFOPALLENS C. MUELL.
(-S. exr ugulosa C. Muell.) 2
S. subventrosa Broth. & Bryhn*- S. FERRUGINEA
1622 -ZYGODON HOOK. & TAYL.
2. MAGT LL & VAN ROOY. MS. FSA.
Z. africanus Sim - Z. EROSUS
300 Z. EROSUS MITT.
(-Z. africanus Sim) 2
500 Z. LEPTOBOLAX C. MUELL.
(-Z. transvaaliensis Rehmann ex Sim) 2
550 Z. PERREFLEXUS C. MUELL.
Z. transvaaliensis Rehmann ex Sira - Z.
LEPTOBOLAX
1623 -CARDOTIELLA VITT
2. MAGILL & VAN ROOY. MS. FSA.
100 C. SECUNDA (C. MUELL.) VITT
(-Macromitr ium schlothe imiaeformis Par.)
2
( -Macromitr ium secundum C. Muell.) 2
RHABDOWEIS IACEAE (42A) 1631
1632 -RHABDOWEIS IA B.S.G.
2. MAGILL & VAN ROOY. MS. FSA.
50 R. CRISPATA (WITH.) LINDB.
POLYTRICHACEAE (83)
1921
1922 -POGONATUM BEAUV.
2. FANSHAWE. 1980. POLYTRICHACEAE. M.SC.
UCT.
50 P. BORGENI I (HAMPE) JAEG.
100 P. CAPENSE (HAMPE) JAEG.
(-Polytrichum transvaal iense C. Muell.)
2
300 P. 0 LI GOD US (C. MUELL.) MITT.
(-Polytrichum natalense Sim) 2
1923 -POLYTRICHUM HEDW.
2. FANSHAWE. 1980. POLYTRICHACEAE. M.SC.
UCTv
100 P. COMMUNE HEDW.
(-P. commune Hedw. var. trichodes (C.
Muell.) Dix.) 2
P. commune Hedw. var. trichodes (C. Muell.)
Dix. - P. COMMUNE
P. formosum Hedw. - POLYTR ICHASTRUM FORMOSUM
P. natalense Sim - POGONATUM OLIGODUS
P. t ransvaaliense C. Muell. - POGONATUM
CAPENSE
1924 -Psilopilura Brid. Southern African species
moved to OLIGOTRICHUM
2. FANSHAWE. 1980. POLYTRICHACEAE. M.SC.
UCT.
P. afrolaevigatum Dix. - OLIGOTRICHUM
AFROLAEVIGATUM
P. wageri Broth, in Dix. - OLIGOTRICHUM WAGER I
1925 -OLIGOTRICHUM LAM. & DC.
1. FANSHAWE. 1980. POLYTRICHACEAE. M.SC.
UCT.
50 0. AFROLAEVIGATUM (DIX.) G.L. SMITH
(-Psilopilura afrolaevigatum Dix.) 1
100 0. CAPENSE SCHELPE & FANSHAWE
200 0. TETRAGONUM SCHELPE & FANSHAWE
300 0. WAGERI (BROTH.) G.L. SMITH
(-Psilopilura wageri Broth.) 1
1926 -POLYTRICHASTRUM G.L. SMITH
1. FANSHAWE. 1980. POLYTRICHACEAE. M.SC.
UCT .
100 P. FORMOSUM (HEDW.) G.L. SMITH
(-Polytrichum formosum Hedw.) 1
PTERI DOPHYTA
A DI ANTACEAE
Contributed by C. Reid
ORTHOTRICHACEAE (42)
350 -DOR YOPTER IS J. SM .
1. SCHELPE & ANTHONY. 1986. FSA.
2. JACOBSEN & JACOBSEN. 1988. BOTHALIA 18:
90.
200 D. PILOSA (POIR.) KUHN
370 -PITYROGRAMMA LINK
300 P. CALOMELANOS (SWARTZ) LINK VAR. AUREOFLAVA
(HOOK.) WEATH. EX BAILEY *
(Note author change)
ASPIDIACEAE 59
8LECHNACEAE
Bothalia 19,2 (1989)
277
690 -BLECHNUM L.
200 B. AUSTRALE L. VAR. AUSTRALE
(Correction of species number)
ANGIOSPERMAE
100 A. PARAD0XA (CHERM.) GOETCHEBEUR & VORSTER
(■Harlscu* paradoxus (Chera.) Chera.) 1
0471000 -PIMBR I ST YLIS VAHL
P. exilis (H.B.K.) Roea. & Schult. -
BULBOSTYLIS HISPIDULA
P0ACEAE
9900890
3360
4970
9901020
95
9901070
150
9901160
800
9901240
300
500
730
MONOCOTYLEDONAE Contributed by C. Reid
and L . Fish
Contributed by L. Fish 9900010
-DIGITARIA HALLER Revision: P.D.F. Kok (PRU).
D. RUKWAE CLAYTON
( East tropical African species collected
in Natal. 2732 (Ubornbo): Lake St. Lucia,
northern end of False Bay (-CD) Ellis
4517)
D. VIOLASCENS LINK *
(Tropical Asian and American species
collected in Natal. 2930 (Pietermaritz-
burg): Pietermaritzburg, Chase Valley
(-CB) Ellis 4416)
-ERI0CHL0A KUNTH
E. MEYER I ANA (NEES) PILG. SUBSP. GRANDIGLUMIS
(STENT & RATTRAY) GIBBS RUSSELL
(Correction of species number and
spelling)
-PASPALUM L.
4. BRUMMITT. 1983. TAXON. 32: 281.
P. DISTICHUM L.
(-P. paspalodes (Michx.) Scribn.) 4
P. paspalodes (Michx.) Scribn. - P. DISTICHUM
-PANICUM L.
P. COLORATUM L. VAR. COLORATUM
(■P. coloratum L. var. maka r i ka r i en s e
Goossens ) 3
P. coloratum L. var. makar ikar ien se Goossens *
P. COLORATUM VAR. COLORATUM
-SACC IOLEPIS NASH
S. CHEVALIERI STAPF
(Tropical African species. Specimens of
this species have been m i s id e n t i f i ed
under S. typhura)
S. cymbiandra auct . , non Stapf - S. INTERRUPTA
S. INTERRUPTA (WILLD.) STAPF
(-S. cymbiandra auct., non Stapf)
(Indian to S.E. Asian species apparently
introduced into Africa, previously
wrongly identified as a West African
s pec ies )
0471010 -BULBOSTYLIS KUNTH
4. GOETCHEBEUR & COUDIJZER. 1985. BULL.
JARD. BOT. NAT. BELG. 55: 207.
770 B. HISPIDULA (VAHL) R. HAINES
( -F labr 1 at yl 1 a exilis (H.B.K.) Roea. &
Schult.) 4
0525000 -CAREX L. Revision: C. Reid (PRE).
2. CLARKE. 1908. KEW BULL. ADD. SER. 8: 74.
C. condensate C.B. Cl. non Nees - C. ZULUENSIS
2300 C. ZULUENSIS C.B. CL.
(-C. condensate C.B. Cl. non Nees) 2
ARACEAE Contributed by C. Reid
684000
0764000 -STYL0CHIT0N LEPR.
#. PRE HERBARIUM PRACTICE, FOLLOWING MAYO
(KEW) .
100 S. NATALENSIS SCHOTT
(-S. obliquine r v is Peter) #
(■S. puberulus N.E. Br.) #
(-S. rogersll N.E. Br . ) #
S. obliquine rvls Peter - S. NATALENSIS
S. puberulus N.E. Br . - S. NATALENSIS
S. rogersii N.E. Br. - S. NATALENSIS
RE STI0NACEAE Contributed by C. Reid 804000
0804140 -CANN0M0IS DESV. (Previous nuaber 0817000)
500 C. SCIRPOI DES (KUNTH) MAST.
(Note author correction)
JUNCACEAE Contributed by C. Reid 930000
0936000 -JUNCUS L.
720 J. CAPENSIS THUNB. X J. LOM ATO P HYLLUS SPRENG.
LILIACEAE Contributed by C. Reid 942000
0969000 -ANDROCYMBIUM WILLD. Revision: U. & D.
Mul ler-Dobl ies (Herb. M-D) .
1870 A. POELTIANUM U. & D. MULLER-DOBL IES
(Note change in species nuaber)
1012000 -ERI OS PERMUM JACQ. EX WILLD.
#. PRE HERBARIUM PRACTICE, FOLLOWING PERRY.
2280 E. DIELSIANUM SCHLTR . EX V. POELLN.
3370 E. LANUGINOSUM JACQ.
9901800 -PRIONANTHIUM DESV.
2. DAVIDSE. 1988. BOTHALIA 18,2: 149.
50 P. DENTATUM (L. F.) HENR.
(■P. rlgldum Desv . ) 2
P. rigidua Desv. - P. DENTATUM
9902860 -ERAGROSTIS WOLF
300 E. ANNULATA RENDLE EX SCOTT ELLIOT
(Note author correction for species)
ZAMIACEAE Contributed by C. Reid
3000
1027000 -GASTERIA DUVAL Revision: E. van Jaarsveld
( NBG) .
HO. PR F. HERBARIUM PRACTICE, FOLLOWING VAN
JAARSVELD.
2100 G. EXCELSA BAK.
4150 G. OBLIQUA (DC.) DUVAL
1081000 -GALTONIA DECNE.
3. HILLIARD & BURTT. 1988. NOTES R. BOT. GDN
EDINB. 45: 95.
200 G. PRIM CEPS (BAK.) DECNE.
0005000
540
560
600
650
-ENCEPHALARTOS LEHM.
2. LAVRANOS & GOODE. 1988. BULL. JARD. BOT.
NAT. BELG. 58: 219.
3. ROBBERTSE ET AL. 1988. S. AFR. J. BOT.
54,4: 363.
4. ROBBERTSE ET AL. 1988. S. AFR. J. BOT.
54,5: 487.
#. PRE HERBARIUM PRACTICE, FOLLOWING REID.
E. DOLOMITICUS LAVRANOS & GOODE
( «E . verrucosus P.J. Vorster et al.) 9
E. DYERI ANUS LAVRANOS & GOODE
(-E. graniticolus P.J. Vorster et al.) #
E. EUGENE-MARAISII VERDOORN SUBSP. EUGENE-
MARAISII
E. EUGENE-MARAISII VERDOORN SUBSP.
MIDDELBURGENSIS LAVRANOS &
GOODE
E. graniticolus P.J. Vorster et al. - E.
DYERI ANUS
E. verrucosus P.J. Vorster et al. « E.
DOLOMITICUS
PINACEAE Contributed by C. Reid
22000
0022000 -PINUS L.
250 P. PINEA L. *
(Exotic species becoming naturalised.
3226 (Fort Beaufort) ( - AD ) , on the road
froa Fort Beaufort to Tarkastad near
Fenella Falls Fara, Henderson 962)
HYDROCHAR ITACEAE Contributed by C. Reid 85000
0087010 -EGERIA PLANCH.
(Note correction of genus nuaber)
CYPERACEAE Contributed by C. Reid 452000
0459030 -MARISCUS GAERTN. Revision: P.J. Vorster
(STE).
M. paradoxus (Chera.) Cherm. » ALINULA
PARADOXA
0459050 -ALINULA J. RAYNAL
1. GOETGHEBEUR & VORSTER. 1988. BULL. JARD.
BOT. NAT. BELG. 58: 457.
1098000
200
210
215
220
700
-LACHENALIA JACQ. F. EX MURRAY
11. DUNCAN. 1988". ANN. KIRSTENB. BOT. GARD.
17.
L. ALOIDES (L. F.) ENGL. VAR. ALOIDES
L. ALOIDES (L.
L. ALOIDES (L.
L. ALOIDES (L.
) ENGL. VAR.
ENGL.
F.) ENGL. VAR.
(JACQ.) ENGL.
F.) ENGL. VAR.
BARKER
AUREA (LINDL.)
QUADRICOLOR
VANZYLIAE W.F.
(Note author change)
F
L.
(Note author change)
BULBIFERA ( CYR. ) ENGL.
(Note author change^)
1109000 -DRACAENA VAND. EX L.
Off. PRE HERBARIUM PRACTICE, FOLLOWING BOS.
150 D. MANNII BAK.
1113000 -ASPARAGUS L.
A. asparagoldes
A. asparagoldes
A . as pa r ago ides
A. asparagoldes
A. asparagoldes
Revision: A. A. Obermeyer (PRE)
sensu Jessop, non (L.) Wight,
p.p. - MYRSIPHYLLUM
ASPARAGOI DES
sensu Jessop, non (L.) Wight,
p.p. - MYRSIPHYLLUM
KRAUSSIANUM
sensu Jessop, non (L.) Wight,
p.p. - MYRSIPHYLLUM
MULTITUBEROSUM
sensu Jessop, non (L.) Wight,
p.p. - MYRSIPHYLLUM OVATUM
sensu Jessop, non (L.) Wight,
p.p. - MYRSIPHYLLUM VOLUBILE
111 3020
200
-MYRSIPHYLLUM WILLD.
1. OBERMEYER. BOTHALIA 15: 77.
M. ASPARAGOIDES (L.) WILLD.
(-Asparagus asparagoldes sensu Jessop,
non (L.) Wight, p.p.) I
M. KRAUSSIANUM KUNTH
(■Asparagus asparagoldes sensu Jessop,
non (L.) Wight, p.p.) 1
M. MULTITUBEROSUM ( R . A . DYER) OBERM.
(■Asparagus asparagoldes sensu Jessop,
non (L.) Wight, p.p.) I
M. OVATUM (SALTER) OBERM.
278
Bothalia 19,2(1989)
(-Asparagus asparagoides sensu Jessop,
non (L.) Wight, p.p.) 1
1200 M. VOLUBILE (THUNB.) OBERM .
(-Asparagus asparagoides sensu Jessop,
non (L.) Wight, p.p.) 1
HYPOXI DACEAE
Contributed by C. Reid
1230000 -HYPOXIS L.
270 H. ANGUSTIFOLIA LAM. VAR. BUCHANANI I BAK. X H.
MEMBRANACEA BAK.
DIOSCOREACEAE
Contributed by C. Reid
1252000 -DIOSCOREA L.
1400 D. MUNDII BAK.
(Note spelling change)
I R I DACEAE
Contributed by C. Reid
1259000
1265000 -MORAEA MILL.
1. GOLDBLATT. 1973. ANN. MISSOURI BOT. GARD.
60: 204.
9. GOLDBLATT. 1986. ANN. KIRSTENBOSCH BOT.
GARD. 14.
1520 M. BREVISTYLA (GOLDBL.) GOLDBL.
( -M . pubi flora N.E. Br . subsp.
brevistyla Goldbl.) 9
2370 M. EXILIFLORA GOLDBL.
3350 M. GAR I PENS IS GOLDBL.
M. mossii N.E. Br. - M. STRICTA
5320 M. NAMAQUAMONTANA GOLDBL.
M. pubiflora N.E. Br . subsp. brevistyla Goldbl.
- M. BREVISTYLA
7300 M. STRICTA BAK.
( -M . mossii N.E. Br . ) 1, 9
( -M . thomsonli sensu Goldbl. 1977, p.p.,
non Bak.) 9
( -M . trita N.E. Br . ) 1, 9
7455 M. THOMSONl I BAK.
M. thonsonll sensu Goldbl. 197.7, p.p., non Bak.
- M. STRICTA
M. trita N.E. Br . - M. STRICTA
8240 M. VERECUNDA GOLDBL.
1273000 -HEXAGLOTTIS VENT.
3. GOLDBLATT. 1987. ANN. MISSOURI BOT. GARD.
74: 542.
50 H. BREVITUBA GOLDBL.
H. flexuosa (L. f.) Sweet - H. LEWISIAE SUBSP.
LEWIS IAE
100 H. LEWISIAE GOLDBL. SUBSP. LEWISIAE
( -H . flexuosa (L. f.) Sweet) 2
120 H. LEWISIAE GOLDBL. SUBSP. SECUNDA GOLDBL.
200 H. LONGI FOLIA (JACQ.) SALISB.
( -H . longifolia (Jacq.) Vent. var.
angustifolia G.J. Lewis) 3
H. longifolia (Jacq.) Vent. var. angustifolia
G.J. Lewis - H. LONGIFOLIA
250 H. NAMAQUANA GOLDBL.
310 H. R I PAR I A GOLDBL.
320 H. VIRGATA (JACQ.) SWEET SUBSP. KAROOICA
GOLDBL.
400 H. VIRGATA (JACQ.) SWEET SUBSP. VIRGATA
( - H . virgata (Jacq.) Sweet var. lata
G. J. Lewis) 3
H. virgata (Jacq.) Sweet var. lata G.J. Lewis -
H. VIRGATA SUBSP. VIRGATA
1301000 -HESPERANTHA KER-GAWL. Revision: O.M.
Hilliard 6 B.L. Burtt (E).
##. PRE HERBARIUM PRACTICE, FOLLOWING
HILLIARD & BURTT.
2650 H. PUBINER VIA HILLIARD & BURTT
(Note spelling change)
2830 H. RUPESTR IS N.E. BR. EX R.C. FOST.
1302000 -I XI A L.
4. DE VOS. 1988. S. AFR. J. BOT. 54: 596.
1270 I. ESTERHUYSENIAE DE VOS
1450 I. FREDER ICKI I DE VOS
3320 I. MOSTERTI I DE VOS
1303000 -DIERAMA K. KOCH Revision: O.M. Hilliard (E).
2. HILLIARD. 1988. NOTES R. BOT. GDN EDINB.
45: 78.
20 D. ADELPHICUM HILLIARD
50 D. AMBIGUUM HILLIARD
D. davyi N.E. Br . - D. INSIGNE
450 D. DISSIMILE HILLIARD
470 D. DRACOMONTANUM HILLIARD
620 D. ERECTUM HILLIARD
650 D. PLOR IPERUM HILLIARD
670 D. PORMOSUM HILLIARD
1000 D. INSIGNE N.E. BR.
( -D . davyi N.E. Br . ) 2
1050 D. JUCUNDUM HILLIARD
1400 D. MEDIUM N.E. BR.
D. medium N.E. Br . var. mossii N.E. Br . - D.
MOSSII
1520 D. MOBILE HILLIARD
1540 D. MOSSII (N.E. BR.) HILLIARD
( -D . medium N.E. Br . var. mossii N.E.
Br . ) 2
1560 D. NEBROWNI I HILLIARD
2650 D. TYRIUM HILLIARD
1306000 -TRITONIA KER-GAWL.
T. flava sensu Bak. non (Ait.) Ker-Gawl. - T.
KAROOICA
1370 T. KAROOICA DE VOS
( -T . flava sensu Bak. non (Ait.) Ker-
Gawl.) 1
T. ma thews lana L. Bol. - CROCOSMIA MATHEWS I ANA
1306010 -CROCOSMIA PLANCH. Revision: M.P. de Vos
( STEU) .
3. DE VOS. 1984. JL S. AFR. BOT. 50: 463.
170 C. X CROCOSMIIFLORA (LEMOINE EX BURBIDGE &
DEAN) N.E. BR.
(Note change in taxon number)
500 C. MATHEW S I ANA (L. BOL.) GOLDBL. EX DE VOS
(-Tritonia mathewsiana L. Bol.) 3
1311000 -GLADIOLUS L.
5. HILLIARD & BURTT. 1983. NOTES R. BOT. GDN
EDINB. 41: 306.
G. cruentus sensu Oberm., p.p., non S. Moore -
G. FLANAGANI I
4850 G. FLANAGANI I BAK.
("G. cruentus sensu Oberm., p.p., non S.
Moore) 5
1312010 -TRITONIOPSIS L. BOL.
1. LEWIS. 1959. JL S. AFR. BOT. 25: 319.
400 T. DO D 1 1 (G.J. LEWIS) G.J. LEWIS
( -Hebea dodii G.J. Lewis) 1
1312011 -Hebea L. Bol. non (Pers.) Hedw. f. -
TRITONIOPSIS
H. dodii G.J. Lewis - TRITONIOPSIS DODII
1315000 -WATSONIA MILL. Revision: P. Goldblatt (MO).
5. GOLDBLATT. 1987. ANN. MISSOURI BOT. GARD.
74: 570.
#. PRE HERBARIUM PRACTICE, FOLLOWING
GOLDBLATT .
450 W. BORRONI C A (POURR.) GOLDBL.
(-W. pyramidata (Andr.) Stapf) 5
2320 W. LEPIDA N.E. BR.
W. pyramidata (Andr.) Stapf - W. BORBONICA
ORCHIDACEAE
Contributed by C. Reid
1389000
1434000 -DISA BERG.
1. LINDER. 1981. CONTR. BOL. HERB. 9.
9. LINDER. 1988. S. AFR. J. BOT. 54: 496.
620 D. CEDARBERGENSIS LINDER
3750 D. OBTUSA LINDL. SUBSP. PICTA (SOND.) LINDER
( - D . picta Sond.) I
D. picta Sond. - D. OBTUSA SUBSP. PICTA
1438000 -PTERYGODIUM SWARTZ
2. LINDER. 1988. S. AFR. J. BOT. 54: 496.
P. rubiginosura Sond. ex H. Bol. - CORYCIUM
RUBIGINOSUM
1350 P. SCHELPEI LINDER
1439000 -CERATANDRA ECKL. EX BAUER
350 C. GRANDIFLORA LINDL. X C. ATRATA DUR. & SCHINZ
1440000 -CORYCIUM SWARTZ
1. ROLFE. 1913. JC 5: 283.
2. STEWART ET AL. 1982. WILD ORCH. S. AFR.
3. OLIVER. 1985. S. AFR. J. BOT. 52: 256.
#. PRE HERBARIUM PRACTICE, FOLLOWING
SCHELPE .
1100 C. RUBIGINOSUM (SOND.) ROLFE
( -Pte rygodium rubiginosum Sond. ex H.
Bol.) 1, #
1561000 -ACROLOPHIA PFITZER
3. SUMMERHAYES & HALL. 1962. TAXON 11: 201.
4. LINDER. 1988. S. AFR. J. BOT. 54: 496.
#. PRE HERBARIUM PRACTICE, FOLLOWING HALL.
##. PRE HERBARIUM PRACTICE, FOLLOWING LINDER.
20 A. BARBATA (THUNB.) LINDER
( -A . lunata (Schltr.) Schltr. & H. Bol.)
4
150 A. CAPENSIS (BERG.) FOURC.
A. capensis (Berg.) Fourc. var. lamellata
(Lindl.) Schelpe - A.
LAMELLATA
A. comosa (Sond.) Schltr. & H. Bol. - A.
CAPENSIS
400 A. LAMELLATA (LINDL.) SCHLTR. & H. BOL.
( - A . capensis (Berg.) Fourc. var.
lamellata (Lindl.) Schelpe) ##
(-Eulophia barbata (Thunb.) Spreng.) 3,
#
A. lunata (Schltr.) Schltr. & H. Bol. - A.
BARBATA
650 A. PANICULATA CRIBB
A. sphaerocarpa (Sond.) Schltr. & H. Bol. - A.
CAPENSIS
A. tristis (L. f.) Schltr. & H. Bol. - A.
CAPENSIS
1565000 -POLYSTACHYA HOOK.
#. PRE HERBARIUM PRACTICE, FOLLOWING LINDER.
300 P. MODESTA REICHB. P.
800 P. TESSELLATA LINDL.
1568000 -ANSELLIA LINDL.
#. PRE HERBARIUM PRACTICE, FOLLOWING LINDER.
20 A. APRICANA LINDL.
( -A . africana Lindl. var. australis
Sunmerh . ) #
( ■ A . gigantea Reichb. f. var. gigantea)
#
(-A. gigantea Reichb. f. var. nilotica
( Bak . ) Summer h . ) #
A. africana Lindl. var. australis Sunmerh. -
A. AFRICANA
A. gigantea Reichb. f. var. gigantea - A.
AFRICANA
A. gigantea Reichb. f. var. nilotica (Bak.)
Summerh. - A. APRICANA
Bothalia 19,2 (1989)
279
1648000 -EULOPHIA R. BR. EX LINDL.
E. barbata (Thunb.) Spreng.
LAMELLATA
ACROLOPHI A
SALICACEAE
Contributed by B.J. Pienaar
0 . PRE HERBARIUM PRACTICE, FOLLOWING
IMMELMAN.
S. capensls Thunb. var . garleplna (Burch.)
Anders. - S. MUCRONATA SUBSP.
CAPENSIS
S. MUCRONATA THUNB. SUBSP. CAPENSIS (THUNB.)
IMMELMAN
(-S. capensls Thunb. var. garleplna
(Burch.) Anders.) #
( -S . raucronata Thunb. var. caffra Burtt
Da vy ) If
(-S. raucronata Thunb. var. Integra Burtt
Davy) #
S. MUCRONATA THUNB. SUBSP. WILMSII (SEEMEN)
IMMELMAN
(-S. woodil Seeraen var. wllmsll (Seemen)
Skan) #
S. mucrcnata Thunb. var. caffra Burtt Davy -
S. MUCRONATA SUBSP. CAPENSIS
S. raucronata Thunb. var. Integra Burtt Davy -
S. MUCRONATA SUBSP. CAPENSIS
S. woodil Seeraen var. wllmsll (Seemen) Skan -
S. MUCRONATA SUBSP. WILMSII
- X. AMERICANA
X. rogersll Burtt Davy - X. AMERICANA
HYDNORACEAE Contributed by B.J. Pienaar 2182
2182000 -HYDNORA THUNB.
3. MUSSELMAN & VISSER. 1987. DINTERIA 19:
77-82.
150 H. JOHANNIS BECCARI
(■H. solmslana Dlnter) 3
H. solmslana Dlnter - H. JOHANNIS
POLYGONACEAE
Contributed by B.J. Pienaar
2184000
Contributed by B.J. Pienaar
1908000
1000 -FICUS L. Revision: C.C. Berg (U); H.
Baljnath (UD-W); J.V. van
Greun lng ( PRU ) .
3. BERG ET AL. 1984. FL. DU GABON 148.
4. BERG. 1986. BULL. MUS. NATN. HIST. NAT.
PARIS 4: 39-40.
5. BERG. 1988. KEW BULL. 43,1: 77-97.
If. PRE HERBARIUM PRACTICE, FOLLOWING
PIENAAR.
50 F. ABUTILI FOLIA MIQ.
(■F. soldanella Warb.) 3
F. burke 1 (Mlq.) Mlq. - F. THONNINGII
F. capensls Thunb. - F. SUR
700 F. CORDATA THUNB. SUBSP. CORDATA
750 F. CORDATA THUNB. SUBSP. SALICIFOLIA (VAHL)
BERG.
(■F. sallcifolla Vahl) 5
F. exasperata Vattl - F. SYCOMORUS
900 F. FISCHERI WARB. EX MILOBR. & BURR.
(-F. kiloneura Hornby) 5
1000 F. GLUMOSA DEL.
( -F . sonderl Mlq.) 5
1200 F. INGENS (MIQ.) MIQ. VAR. INGENS
(-F. Ingens Mlq. var. tomentosa Hutch.)
3
F. ingens Mlq. var. tomentosa Hutch. - F.
INGENS VAR. INGENS
F. kiloneura Hornby - F. FISCHERI
1300 F. LUTEA VAHL
(»F. nekbudu Warb.) 3
(■F. quebeba Welw. ex Fical.) 4
( ■ F . vogelil (Mlq.) Mlq.) 5
1500 F. NATALENSIS HOCHST. SUBSP. NATALENSIS
F. nekbudu Warb. - F. LUTEA
F. petersli Warb. - F. THONNINGII
1600 F. POL I TA VAHL SUBSP. POLITA
F. pretorla^ Burtt Davy - F. CORDATA SUBSP.
SALICIFOLIA
F. quebeba Welw. ex Fical. ■ F. LUTEA
F. sallcifolla Vahl - F. CORDATA SUBSP.
SALICIFOLIA
F. soldanella Warb. ■ F. ABUTILIFOLIA
F. sonderl Mlq. - F. GLUMOSA
2250 F. SUR FORSSK.
(■F. capensls Thunb.) 5
2300 F. SYCOMORUS L.
(-F. exasperata Vahl) #
2450 F. THONNINGII BLUME
( • F . burke 1 (Mlq.) Mlq.) 5
(»F. petersli Warb.) 5
F. vogelil (Mlq.) Mlq. - F. LUTEA
PROTEACEAE
Contributed by B.J. Pienaar
2016000
2031000 -MIMETES SALISB.
2. ROURKE. 1988. JL . S. APR. BOT. 54,6: 636-
639.
250 M. CHRYSANTHUS ROURKE
2035000 -PROTEA L.
4. CHISUMPA & BRUMMITT. 1987. KEW BULL.
42,4: 815-853.
P. bolusll Phill. - P. CAFFRA SUBSP. CAFFRA
1200 P. CAFFRA MEISN. SUBSP. CAFFRA
(-P. bolusll Phill.) 3
(-P. mul t lbrac teata Phill.) 3
(-P. rhodantha Hook, f.) 3
P. mult Ibracteata Phill. ■ P. CAFFRA SUBSP.
CAFFRA
P. rhodantha Hook. f. - P. CAFFRA SUBSP.
CAFFRA
2204000 -OXYGONUM BURCH.
4. GERMISHUIZEN. 1988. BOTHALIA 18,2: 173-
181, 187-188.
If. PRE HERBARIUM PRACTICE, FOLLOWING
GERMISHUIZEN .
50 0. ACETOSELLA WELW.
0. calcaratum Melsn. - O. DREGEANUM SUBSP.
CANESCENS VAR. CANESCENS
0. canescens Sond. - 0. DREGEANUM SUBSP.
CANESCENS VAR. CANESCENS
0. canescens Sond. var. subglabra Schlnz - 0.
DREGEANUM SUBSP. CANES-CENS VAR.
CANESCENS
300 0. DELAGOENSE KUNTZE
(■0. delagoenae Kuntze var. strlctum
C. H. Wr . ) #
(■0. dregeanura Melsn. var. pubescens
(C.H. Wr.) Burtt Davy) #
(•0. dregeanum Melsn. var. strlctum
(C.H. Wr.) R . A . Grah. ) #
0. delagoense Kuntze var. strlctum C.H. Wr . -
0. DELAGOENSE
400 0. DREGEANUM MEISN. SUBSP. CANESCENS (SOND.)
GERMISHUIZEN VAR. CANESCENS
(-0. calcaratum Melsn.) 1
(■0. canescens Sond.) 1
(-0. canescens Sond. var. subglabra
Schlnz) .1
(•0. dregeanum Melsn. ver . canescens
(Sond.) R . A . Grah.) 4
(■0. zeyherl Sond.) 1
430 0. DREGEANUM MEISN. SUBSP. CANESCENS (SOND.)
GERMISHUIZEN VAR. DISSECTUM
GERMISHUIZEN
450 0. DREGEANUM MEISN. SUBSP. CANESCENS (SOND.)
GERMISHUIZEN VAR. L I N E AR I FOL I UM
GERMISHUIZEN
470 0. DREGEANUM MEISN. SUBSP. CANESCENS (SOND.)
GERMISHUIZEN VAR. LOBOPHYLLUM
GERMISHUIZEN
490 0. DREGEANUM MEISN. SUBSP. CANESCENS (SOND.)
GERMISHUIZEN VAR. PILOSUM
GERMISHUIZEN
500 0. DREGEANUM MEISN. SUBSP. DREGEANUM
(-0. dregeanum Melsn. var. dregeanum) 4
(-0. dregel Melsn.) 1
(-0. natalense Schlechter) 1
550 0. DREGEANUM MEISN. SUBSP. LANCEOLATUM
GERMISHUIZEN
750 0. DREGEANUM MEISN. SUBSP. STREYI GERMISHUIZEN
800 0. DREGEANUM MEISN. SUBSP. SWAZICUM (BURTT
DAVY) GERMISHUIZEN
(■0. dregeanum Melsn. var. swazicum
(Burtt Davy) R.A. Grah.) 4
(■0. zeyherl Sond. var. swazicum Burtt
Davy) 4
0. dregeanum Melsn. var. canescens (Sond.) R.A.
Grah. - 0. DREGEANUM SUBSP.
CANESCENS VAR. CANESCENS
0. dregeanum Melsn. var. dregeanura - 0.
DREGEANUM SUBSP. DREGEANUM
0. dregeanum Melsn. var. pubescens (C.H. Wr . )
Burtt Davy - 0. DELAGOENSE
0. dregeanum Melsn. var. strlctum (C.H. Wr.)
R.A. Grah - 0. DELAGOENSE
0. dregeanum Melsn. var. swazicum (Burtt Davy)
R.A. Grah. - 0. DREGEANUM
SUBSP. SWAZICUM
0. dregel Melsn. - 0. DREGEANUM SUBSP.
DREGEANUM
0. natalense Schlechter ■ 0. DREGEANUM SUBSP.
DREGEANUM
0. zeyherl Sond. - 0. DREGEANUM SUBSP.
CANESCENS VAR. CANESCENS
0. zeyherl Sond. var. swazicum Burtt Davy - 0.
DREGEANUM SUBSP. SWAZICUM
CHENOPODI ACEAE
Contributed by B.J. Pienaar
2214000
2221000 -BETA L. *
1. TUTIN. 1964. FL. EUROP. 1: 91.
100 B. VULGARIS L. *
2229000 -ATRIPLEX L. Revision: M. No-bs (Cl).
1. HALL & CLEMENTS. 1923. PHYLOGENETIC
METHODS IN TAXONOMY 247-248.
1450 A. VESICARIA HEWARD EX BENTH. SUBSP.
APPENDICULATA (BENTH.) PARR-
SMITH
Contributed by B.J. Pienaar
2129000
2136000
150
-XIMENIA L.
X. AMERICANA L.
(-X. amerlcana L. var. microphylla Welw.
ex Oliv.) 1
( ■ X . rogersll Burtt Davy) 1
X. amerlcana L. var. microphylla Welw. ex Oliv.
2269000
1950
2650
3125
-SALSOLA L.
3. BOTSCH. 1978. BOT. ZHOURNAL 63,6
836.
4. BOTSCH. 1981. BOT. ZHOURNAL 66,7
1040.
5. BOTSCH. 1983. BOT. ZHOURNAL 68,9
1249.
S. CAUL I F LORA BOTSCH.
S. DENUDATA BOTSCH.
S. GARUBICA BOTSCH.
832-
1036-
1247-
Bothalia 19,2 (1989)
280
3350
372 5
3 7 50
37 7 5
4155
4350
4 750
5050
507 5
5550
55 7 5
56 50
63 50
6450
67 50
67 55
GEMMIPARA BOTSCH.
HOANIBICA BOTSCH.
HOTTENTOTICA BOTSCH.
HUABICA BOTSCH.
KLE I N FONTE I N I BOTSCH.
MARGINATA BOTSCH.
MIRABILIS BOTSCH.
OKAUKUEJENSIS BOTSCH.
OMARURUENSIS BOTSCH.
PROCERA BOTSCH.
PTILOPTERA BOTSCH.
ROB I NSON 1 1 BOTSCH.
SPENCER! BOTSCH.
SWAKOPMUNDI BOTSCH.
UGABICA BOTSCH.
UNJABICA BOTSCH.
AMARANTHACEAE Contributed by B.J. Pienaar
AIZOACEAE Contributed by B.J. Pienaar
2376000 -LIMEUM L.
1475 L. INDICUM STOCKS EX ANDERSON
2289000
2374000
MESEMBRYANTHEMACEAE
Contrlb. by B.J. Pienaar
. FIS SUM
FISSUM
A. DELAET 1 1
- A. DELAET I I
- A. TESTICULARE
DELAETII
A.. DELAETII
-ARGYRODERMA N.E. BR.
2. HARTMANN. 1977. MITT. INST. ALLG. BOT.
HAMBURG 15: 121-235.
A. amoenum Schwant. - A. PEARSONII
A. angus t 1 pe t al urn L. Bol. - A. CONGREGATUM
A. aureutn L. Bol. - A. DELAETII
A. auatrale L. Bol. - A. DELAETII
A. blandum L. Bol. - A. DELAETII
A. boreale L. Bol. - A. DELAETII
A. braunsll (Schwant.) Schwant. -
A. brevlpe8 (Schltr.) L. Bol. -
A. brevltubura L. Bol.
A. carlnatum L. Bol. p
A. carlnatum L. Bol. p
A. cltrlnum L. Bol. -
A. conclnnura Schwant.
A. CONGREGATUM L. BOL. %
(■A. ang us t 1 pe ta lum L. Bol.) 2
(-A. Jacobsenlanun Schwant.) 2
(-A. nortlerl L. Bol.) 2
(■A. peersll L. Bol.) 2
(■A. roolpanense L. Bol. p.p.) 2
A. CR ATE R I FORME (L. BOL.) N.E. BR.
(■A. pulvlnare L. Bol.) 2
(-A. subrotundum L. Bol.) 2
A. cuneat Ipetalum L. Bol. p.p. - A. DELAETII
A. cunea t 1 pe ta 1 um L. Bol. p.p. - A. PEARSONII
A. DELAETII MAASS
(■A. aureum L. Bol.) 2
(-A. auatrale L. Bol.) 2
(-A. blandum L. Bol.) 2
(■A. boreale L. Bol.) 2
(-A. brevltubum L. Bol.) 2
(«A. carlnatum L. Bol. p.p.) 2
(■A. cltrlnum L. Bol.) 2
(■A. conclnnum Schwant.) 2
(■A. c une a 1 1 pe t a 1 um L. Bol. p.p.) 2
( * A . denslpetalura L. Bol.) 2
(■A. forraoeum L. Bol.) 2
(■A. gregarlum L. Bol.) 2
(-A. latlfollum L. Bol.) 2
( ■ A . lesllel R. Br . ) 2
(■A. leucanthum L. Bol.) 2
( • A . longipes L. Bol.) 2
(■A. productum L. Bol.) 2
(■A. renlforme L. Bol.) 2
(-A. roolpanense L. Bol. p.p.) 2
( -A . roseuii Schwant. forma delaetll
Rowl . ) 2
(•A. roseum Schwant. forma roseurn) 2
(-A. schuldtii Schwant.) 2
(■A. speclosum L. Bol.) 2
(■A. splendens L. Bol.) 2
A. denslpetalum L. Bol. - A. DELAETII
A. dlgltlfollum (N.E. Br . ) Schwant. - A.
FISSUM
A. FISSUM (HAW.) L. BOL.
(■A. braunsll (Schwant.) Schwant.) 2
( ■ A . brevlpes (Schltr.) L. Bol.) 2
( • A . dlgltlfollum (N.E. Br . ) Schwant.) 2
(•A. hutchlnsonll L. Bol.) 2
(■A. lat Ipetalum L. Bol. var.
lat Ipetalum) 2
(-A. lat Ipetalum L. Bol. var. longltubun
L. Bol.) 2
( ■ A . lltorale L. Bol.) 2
(-A. orlentale L. Bol.) 2
A. formosura L. Bol. - A. DELAETII
A. framesll L. Bol. 1929 nec 1934 - A.
PEARSONII
A. FRAMES II L. BOL. SUBSP. FRAMES I I
( • A . framesll L. Bol. var. framesll) 2
(-A. framesll L. Bol. var. minus L.
Bol.) 2
A. framesll L. Bol. var. framesll ■ A.
FRAMES I I SUBSP. FRAMESII
A. framesll L. Bol. var. minus L. Bol. - A.
FRAMESII SUBSP. FRAMESII
A. gregarlum L. Bol. - A. DELAETII
A. hutchlnsonll L. Bol. - A. FISSUM
A. Jacobsen lan um Schwant. - A. CONGREGATUM
A. latlfollum L. Bol. - A. DELAETII
A. lat Ipetalum L. Bol. var. lat Ipetalum - A.
FISSUM
A. lat Ipetalum L. Bol. var. longltubum L. Bol.
- A. FISSUM
A. lesllel R. Br . - A. DELAETII
A. leucanthum L. Bol. • A. DELAETII
800
1000
1100
A. lltorale L. Bol. - A. FISSUM
A. longipes L. Bol. => A. DELAETII
A. luckoffll L. Bol. - A. PEARSONII
A. nortlerl L. Bol. - A. CONGREGATUM
A. orlentale L. Bol. - A. FISSUM
A. ovale L. Bol. - A. PEARSONII
A. PEARSONII (N.E. BR.) SCHWANT.
(■A. araoenura Schwant.) 2
(=-A. c une a 1 1 pe t a lum L. Bol. p.p.) 2
( ■ A . framesll L. Bol. 1 929 nec 1934) 2
( ■ A . luckoffll L. Bol.) 2
(-A. ovale L. Bol.) 2
(■A. schlechterl Schwant.) 2
A. peersll L. Bol. » A. CONGREGATUM
A. productum L. Bol. - A. DELAETII
A. pulvlnare L. Bol. - A. CRATERIF0RME
A. renlforme L. Bol. - A. DELAETII
A. roolpanense L. Bol. p.p. - A. CONGREGATUM
A. roolpanense L. Bol. p.p*. ■ A. DELAETII
A. roseum Schwant. forma delaetll Rowl. - A.
DELAETII
A. roseum Schwant. forma roseum » A. DELAETII
A. schlechterl Schwant. ■ A. PEARSONII
A. schuldtii Schwant. - A. DELAETII
A. speclosum L. Bol. - A. DELAETII
A. splendens L. Bol. - A. DELAETII
A. SUBALRUM (N.E. BR.) N.E. BR.
(-A. vllletll L. Bol . ) 2
A. subrotundum L. Bol. ■ A. CRATERIFORME
A. TESTICULARE (AIT.) N.E. BR.
(■A. carlnatum L. Bol. p.p.) 2
A. vllletll L. Bol. - A. SUBALBUM
5030 -C0N0PHYTUM N.E. BR.
8. HAMMER. 1988. ALOE 25,2: 34-35.
4250 C. BUYS IANUM A . R . MITCHELL & HAMMER
4300 C. CALCULUS (BERGER) N.E. BR. VAR. CALCULUS
4350 C. CALCULUS (BERGER) N.E. BR. VAR. KOMKANSICUM
(L. BOL.) RAWE
4400 C. CALCULUS (BERGER) N.E. BR. VAR. PROTUSUM L.
BOL.
5071 -MALEPH0RA N.E. BR.
1060 M. PURPURE0-CR0CEA (HAW.) JACOBSEN & SCHWANT.
2405073
445 5
-MESEMBRYANTHEMUM L.
M. SCHWANTES IA (L.) GLEN
-R00DIA N.E. BR.
1. N.E. BROWN. 1922 . F S A 2:
2405160
100
200
300
-TANQUANA H.E.K. HARTM. & LIEDE
2. HARTMANN & LIEDE. 1986. BOT. JB. 106:
433-485.
T. ARCHER I (L. BOL.) H.E.K. HARTM. & LIEDE
T. HILMARII (L. BOL.) H.E.K. HARTM. & LIEDE
T. PRISMATICA (MARLOTH) H.E.K. HARTM. & LIEDE
CARYOPHYLLACEAE (PART A) Contrlb. by B.J. Pienaar 2429000
2430000 -CERASTIUM L.
2. JALAS. 1964. FL . EUROP. 1: 142.
If. PRE HERBARIUM PRACTICE, FOLLOWING MOSCHL.
C. caespltosum Glllb. subsp. trlvlale (Link)
Hlltonen - C. F0NTANUM SUBSP.
TRIVI ALE
275 C. FONTANUM BAUMG. SUBSP. TRI VI ALE (LINK) JALAS
(■C. caespltosum Glllb. subsp. trivLale
(Link) Hlltonen) 2
(-C. holosteo Ides Fr.) 2
(-C. holosteoldes Fr . subsp. trlvlale
(Murb.) Mo sc hi) If
(■C. trlvlale Link) 2
(-C. vulgatura L.) 2
C. holosteoldes Fr . - C. FONTANUM SUBSP.
TRIVIALE
C. holosteoldes Fr.. subsp. trlvlale (Murb.)
Moschl ■ C. FONTANUM SUBSP.
TRIVIALE
C. trlvlale Link - C. FONTANUM SUBSP. TRIVIALE
C. vulgatum L. - C. FONTANUM SUBSP. TRIVIALE
ILLECEBRACEAE Contributed by B.J. Pienaar 2467000
2476000
200
300
400
500
600
700
-HERNIARIA L.
H. ERCKERTI I HERMANN SUBSP. ERCKERTI I VAR.
DEWET 1 1 HERMANN
H. ERCKERTII HERMANN SUBSP. ERCKERTI I VAR.
DINTERI CHAUDRI
H. ERCKERTII HERMANN SUBSP. ERCKERTII VAR.
ERCKERTII
H. ERCKERTII HERMANN SUBSP. PULVINATA CHAUDHR I
H. GRIMM 1 1 HERMANN
H. SCHLECHTER I HERMANN
LAURACEAE
Contributed by B.J. Pienaar
2782000
2801000 -Bellschmiedia Nees - DAHLGREN0DENDR0N
B. natalensls J.H. Ross ■ DAHLGREN0 DENDR0N
NATALENS IS
2813010 -DAHLGREN0DENDR0N J.J.M. V.D. MERWE & VAN WYK
1. VAN DER MERWE, VAN WYK & K0K. 1988. JL S.
AFR. BOT. 54,1: 80-87.
100 D. NATALENSIS (J.H. ROSS) J.J.M. V.D. MERWE &
VAN WYK
(■Bellschmiedia Nees) 1
PAPAVERACEAE Contributed by B.J. Pienaar 2833000
2853000 -PAPAVER L.
1. M0WAT & WALTERS. FL. EUROP. 1: 248-249.
150 P. ARGEM0NE L. *
Bothalia 19,2 (1989)
281
BRASSICACEAE Contributed by B.J. Pienaar 2863000
2875000 -HELIOPHILA L.
4. PIENAAR & NICHOLAS. 1988. BOTHALIA 18,2:
183-188.
1950 H. CORNELLS BERGIA PIENAAR & NICHOLAS
CRASSULACEAE Contributed by B.J. Pienaar 3161000
316AOOO -COTYLEDON L.
3237 C. ORBICULATA L. VAR. ORBICULATA
(-C. ramosissima Mill., non Haw.) 1
C. ramosissima Mill., non Haw. - C. ORBICULATA
VAR. ORBICULATA
3164010 -TYLECODON TOELKEN
2. VAN JAARSVELD. 1988. ALOE 25,2: 30-32.
50 T. BAYERI VAN JAARSVELD
3175000 -ADROMISCHUS LEM.
950 A. CRISTATUS (HAW.) LEM. VAR. CLAVIFOLIUS
(HAW.) TOELKEN
(■A. poe 1 In 1 1 z 1 anus Werderm.) 1
A. poe 1 1 n 1 1 z 1 anus Werderm. - A. CRISTATUS VAR.
CLAVIFOLIUS
ROSACEAE Contributed by B.J. Pienaar 3316000
3375000 -ALCHEMILLA L.
A 50 A. GALPINII HAUM. & BALLE
FABACEAE Contributed by G. Germlshulzen 3A36000
and B. D. Schr Ire
3AA3000
3AA3010
100
3AA6000
3 AA 9000
3653000
3657000
1650
2150
2650
32 50
5050
5130
6150
7 350
8 A 7 0
3662000
2050
A 2 50
A 7 00
-ALBIZIA DURAZZ.
A. dlstachya (Vent.) MacBride - PARASERIANTHES
LOPHANTHA SUBSP. LOPHANTHA
A. lophantha (Wllld.) Benth. - PARASERIANTHES
LOPHANTHA SUBSP. LOPHANTHA
-PARASERIANTHES NIELSEN
(Note change of spelling)
P. LOPHANTHA (WILLD.) NIELSEN SUBSP. LOPHANTHA
(•Acacia lophantha Wllld.) 1
(-Alblzia dlstachya (Vent.) MacBride) 1
(-Albizia lophantha (Wllld.) Benth.) 1
(-Mimosa dlstachya Vent.) 1
-ACACIA MILL.
A. lophantha Wllld. - PARASERIANTHES LOPHANTHA
SUBSP. LOPHANTHA
-MIMOSA L.
M. dlstachya Vent. - PARASERIANTHES LOPHANTHA
SUBSP. LOPHANTHA
-Borbonia L. - ASPALATHUS
B. le lan tha Phlll. - ASPALATHUS CRENATA
B. pinlfolia Marl. - ASPALATHUS LINEARIS
SUBSP. PINIFOLIA
-LOTONONIS (DC.) ECKL. & ZEYH.
1. HARVEY. 1862. FC 2: 60.
8. VAN WYK. 1988. BOTHALIA 18,1: 31-36.
9. VAN WYK. 1988. S. AFR. J. BOT. 54,2: 175-
181 .
10. VAN WYK. 1988. S. AFR. J. BOT. 54,6: 628.
L. CAERULESCENS (E. MEY.) B.-E. VAN WYK
(•Aspalathus caerulescens E. Mey.) 9
L. COMPTONI I B.-E. VAN WYK
L. DAHLGRENI I B.-E. VAN WYK
L. DISSITINODIS B.-E. VAN WYK
L. LENTICULA (E. MEY.) BENTH.
(-Crotalarla lentlcula E. Mey.) 9
L. LEUCOCLADA (SCHLTR . ) DUEMMER
(-Lebeckla leucoclada Schltr.) 1
L. MINIMA B.-E. VAN WYK
L. omahekensls Dlnter ex A. Schrelb. -
INDIGOFERA ARENOPHILA
L. PORRECTA (E. MEY.) BENTH.
L. SABULOSA SALTER
-LEBECKIA THUNB. Revision: A. Schreiber (M).
L. contamlnata Alt. f . non Wllld. - INDIGOFERA
F ILIFOLIA
L. contamlnata Ker-Gawl . non Thunb. -
INDIGOFERA F ILIFOLIA
L. leucoclada Schltr. - LOTONONIS LEUCOCLADA
L. nuda Sims - INDIGOFERA F ILIFOLIA
-ASPALATHUS L.
16. DAHLGREN. 1988. FS A 16,3: 1-A30.
A. acuminata Lara, subsp. magnlflora Dahlg. -
A. TULBAGHENSIS DAHLG.
A. adelphea Eckl. & Zeyh. - A. SUBTINGENS
A. AEMULA E. MEY.
( - A . serlcea Berg, subsp. aemula (E.
Mey . ) Dahlg . ) 16
(-Ononis splcata Thunb.) 1, 16
( -Paraspalathus aemula (E. Mey.) Presl)
1, 16
A. AMOENA (DAHLG.) DAHLG.
( -A . retrof lexa L. subsp. amoena Dahlg.)
10
A. argentea L. - A. CALEDONENSIS
A. bentharail Harv. - A. SPICATA
A. BREVICARPA (DAHLG.) DAHLG.
(-A. divarlcata Thunb. subsp. brevlcarpa
Dahlg.) 16
A. caerulescens E. Mey. - LOTONONIS
CAERULESCENS
A. CALEDONENSIS DAHLG.
( - A . argentea L.) 16
5000
5800
6350
7150
72 50
77 00
7800
8250
9050
9300
11600
11850
12600
12970
1 3010
1 3060
13160
132 10
1 3 230
1 3 2 AO
1 A1A0
1A350
1A500
1 A 5 7 0
1 A 6 00
(-Paraspalathus argentea (L.) Presl) 1,
16
A. CANDICANS AIT. F.
A. capltella Burch, ex Benth. - A. INOPS
A. CHORTOPHILA ECKL. b ZEYH.
(-A. frankenloldes DC. a. chortophlla
(Eckl. b Zeyh.) Harv.) 10
A. chortophlla Eckl. & Zeyh. subsp. congests
Dahlg. - A. CONGESTA
A. chortophlla Eckl. b Zeyh. subsp. kougaensls
Dahlg. - A. KOUGAENSIS
A. CLIFFORTIOIDES H. BOL.
( -A . splcata Thunb. subsp.
clif fort lo Ides (H. Bol. In
Schltr.) Dahlg.) 16
A. CONGESTA (DAHLG.) DAHLG.
( - A . chortophlla Eckl. b Zeyh. subsp.
congests Dahlg.) 16
A. CORDICARPA DAHLG.
A. CRENATA (L.) DAHLG.
( -A . leiantha (Phlll.) Dahlg.) 16
(-Borbonia leiantha Phlll.) 6, 14, 16
A. CUSPIDATA DAHLG.
A. cuspldata Dahlg. subsp. strlctlclada Dahlg.
- A. STR ICTICLADA
A. DENSIFOLIA BENTH.
A. divarlcata Thunb. subsp. brevlcarpa Dahlg. -
A. BREVICARPA
A. divarlcata Thunb. subsp. horlzontalls Dahlg.
- A. HOR IZONTAL IS
A. divarlcata Thunb. subsp. leptocoma (Eckl. b
Zeyh.) Dahlg. - A. LEPTOCOMA
A. EMPETR I FOLIA (DAHLG.) DAHLG.
(-A. retroflexa L. subsp. empetrl folia
Dahlg.) 16
A. erlclfolla L. subsp. puberula (Eckl. b
Zeyh.) Dahlg. - A. PUBERULA
A. ESTERHUYSENIAE DAHLG.
(-A. pumila Dahlg.) 16
A. frankenloldes DC. a. chortophlla (Eckl. b
Zeyh.) Harv. - A. CHORTOPHILA
A. gracill folia Dahlg. - A. JUNIPERINA SUBSP.
GRACILIFOLI A
A. HETEROPHYLLA L. F.
(-A. stachyera Eckl. b Zeyh.) 1
( -Paraspalathus hbterophylla ("Thunb.")
Presl) 1
A. heterophylla L. f. subsp. lagopus (Thunb.)
Dahlg. - A. LOTO IDES SUBSP.
LAGOPUS
A. heterophylla L. f. subsp. lotoldes (Thunb.)
Dahlg. - A. LOTOIDES SUBSP.
LOTO IDES
A. HORIZONTALIS (DAHLG.) DAHLG.
( -A . divarlcata Thunb. subsp.
horlzontalls Dahlg.) 16
A. INOPS ECKL. b ZEYH.
(-A. capltella Burch, ex Benth.) 1, 16
(-A. stenophylla Eckl. b Zeyh. subsp.
garclana Dahlg.) 16
A. ISOLATA (DAHLG.) DAHLG.
( -A . varians Eckl. b Zeyh. subsp.
lsolata Dahlg.) 16
A. JOUBERTIANA ECKL. b ZEYH.
A. Joubertiana Eckl. b Zeyh. subsp.
glabrlpetala Dahlg. - A.
SHAW I I SUBSP. GLABRIPETALA
A. Joubertiana Eckl. b Zeyh. subsp. longispica
Dahlg. - A. S HAW 1 1 SUBSP.
LONGISPICA
A. joubertiana Eckl. b Zeyh. subsp. shawil (L.
Bol.) Dahlg. - A. SHAWII
SUBSP. SHAWII
A. JUNIPERINA THUNB. SUBSP. GR AC ILIFOLIA
(DAHLG.) DAHLG.
(-A. gracillfolia Dahlg.) 16
A. JUNIPERINA THUNB. SUBSP. MONTICOLA DAHLG.
A. KATBERGENSIS (DAHLG.) DAHLG.
( - A . slmii H. Bol. subsp. katbergensls
Dahlg.) 16
A. KEEROMSBERGENS IS DAHLG.
A. KOUGAENSIS (GARAB. EX DAHLG.) OAHtG.
(-A. chortophlla Eckl. b Zeyh. subsp.
kougaensls Dahlg.) 16
A. lactea Thunb. subsp. adelphea (Eckl. b
Zeyh.) Dahlg. - A. SUBTINGENS
A. larlcifolia Lam. - A. UNIFLORA
A. LEBECKIO IDES DAHLG.
(-A. linearis (Burrn. f.) Dahlg. subsp.
latipetala Dahlg.) 16
A. leiantha (Phlll.) Dahlg. - A. CRENATA
A. LEPTOCOMA ECKL. b ZEYH.
(-A. divarlcata Thunb. subsp. leptocoma
(Eckl. b Zeyh.) Dahlg.) 16
A. leptophylla Eckl. A Zeyh. - A. UNIFLORA
A. LEUCOPHYLLA DAHLG.
A. leucophylla Dahlg. subsp. septentr ionalls
Dahlg. - A. PETERSONII
A. LINEARIFOLIA (BURM. F.) DC.
( - A . llnearlfolia DC. b. dlscreta Drege)
1, 16
( -A . rugosa Thunb. subsp. llnearlfolia
(Burn, f.) Dahlg.) 16
(-Trineurla llnearlfolia (DC.) Presl) 1,
16
A. llnearlfolia DC. b. dlscreta Drege - A.
LINEARIFOLIA
A. LINEARIS (BURM. F.) DAHLG.
( -A . linearis (Burm. f.) Dahlg. subsp.
pinlfolia (Marloth) Dahlg.) 16
(-Borbonia pinlfolia Marl.) 6, 15, 16
A. linearis (Burra, f.) Dahlg. subsp. latipetala
Dahlg. - A. LEBECKIOIDES
A. linearis (Burra, f.) Dahlg. subsp. pinlfolia
282
Bothalia 19,2 (1989)
15030
1 5050
16050
1 65 50
1 7050
1 7950
18850
20040
20800
21600
21700
21910
21920
21930
22000
22400
22900
23150
23350
24200
24350
24600
24800
(Marloth) Dahlg. - A. LINEARIS
A. L0T0 IDES THUNB. SUBSP. LAGOPUS (THUNB.)
DAHLG.
(-A. heterophylla L. f. subsp. lagopus
(Thunb.) Dahlg.) 16
(■Ononis lagopus Thunb.) 1, 16
A. LOTO IDES THUNB. SUBSP. LOTOIDES
(-A. heterophylla L. f. subsp. lotoldes
(Thunb.) Dahlg.) 16
(-A. procumbens E. Mey.) 1, 16
( -Paras palathus humlfusa Presl) 1, 16
( -Paras palathus lotoldes (Thunb.) Presl)
1 , 16
( -Paras palathus procumbens (E. Mey.)
Presl) 1, 16
A. NEGLECTA SALTER
( - A . splcata Thunb. subsp. neglecta
(Salter) Dahlg . ) 16
A. nervosa E. Mey. - A. SPICATA
A. OBTUSATA THUNB.
(-A. splnosa L. subsp. obtusata (Thunb.)
Dahlg.) 16
A. pachyloba Benth. subsp. succulentlfolla
Dahlg. - A. PACHYLOBA SUBSP.
V ILL IC AUL I S
A. PACHYLOBA BENTH. SUBSP. VILLICAULIS DAHLG.
(-A. pachyloba Benth. subsp.
succulentlfolla Dahlg.) 16
A. PETERS0NI I DAHLG.
( • A . leucophylla Dahlg. subsp.
septentrlonalls Dahlg.) 16
A. procumbens E. Mey. - A. LOTOIDES SUBSP.
LOTOIDES
A. PUBERULA (ECKL. & ZEYH.) DAHLG.
(-A. erlclfolia L. subsp. puberula
(Eckl. & Zeyh.) Dahlg.) 16
A. puralla Dahlg. - A. ESTERHUYSENIAE
A. remota Eckl. & Zeyh. - A. SPICATA
A. retroflexa L. subsp. amoena Dahlg. - A.
AM0ENA
A. RETROFLEXA L. SUBSP. ANGUSTI PETALA DAHLG.
A. retroflexa L. subsp. empetrlfolla Dahlg. -
A. EMPETRIFOLI A
A. RUGOSA THUNB.
A. rugosa Thunb. subsp. linearlfolla (DC.)
Dahlg. - A. LINEARIFOLIA
A. SECURIFOLI A ECKL. & ZEYH.
( -A . securifolla Eckl. & Zeyh. subsp.
crassa Dahlg.) 16
A. securifolla Eckl. 4 Zeyh. subsp. crassa
Dahlg. - A. S EC UR I F OL I A
A. SERICEA BERG.
( -Paraspalathus serlcea (Berg.) Presl) 1
A. serlcea Berg, subsp. aernula (E. Mey.) Dahlg.
- A. AEMULA
A. SHAW 1 1 L. BOL. SUBSP. GLABRIPETALA DAHLG.
(-A. Joubertlana Eckl. & Zeyh. subsp.
glabrlpetala Dahlg.) 16
A. SHAW 1 1 L. BOL. SUBSP. LONGISPICA (DAHLG.)
DAHLG.
( -A . Joubertlana Eckl. & Zeyh. subsp.
longispica Dahlg.) 16
A. SHAW I I L. BOL. SUBSP. SHAWII
(-A. Joubertlana Eckl. & Zeyh. subsp.
shavll (L. Bol.) Dahlg.) 16
A. SIMII H. BOL.
A. 8iall H. Bol. subsp. katbergensls Dahlg. -
A. KATBERGENSIS
A. s pha e r oc e pha la Schltr. - A. TRUNCATA
A. SPICATA THUNB.
( -A . benthaali Harv.) 5
( - A . nervosa E. Mey.) 5
(-A. remota Eckl. & Zeyh.) 5
A. splcata Thunb. subsp. c 1 1 f f or t lo lde s (H.
Bol. In Schltr.) Dahlg. - A.
CLIFFORTIOIDES
A. splcata Thunb. subsp. neglecta (Salter)
Dahlg. - A. NEGLECTA
A. splnosa L. subsp. obtusata (Thunb.) Dahlg. -
A. OBTUSATA
A. stachyera Eckl. & Zeyh. - A. HETEROPHYLLA
A. STENOPHYLLA ECKL. & ZEYR.
(■A. stenophylla Eckl. & Zeyh. subsp.
colorata Dahlg.) 16
A. stenophylla Eckl. & Zeyh. subsp. colorata
Dahlg. - A. STENOPHYLLA
A. stenophylla Eckl. & Zeyh. subsp. garclana
Dahlg. - A. INOPS
A. STRICTICLADA (DAHLG.) DAHLG.
( - A . cuspidate Dahlg. subsp.
strlctlclada Dahlg.) 16
A. SUBTINGENS ECKL. & ZEYH.
(-A. adelphea Eckl. & Zeyh.) 6, 16
(-A. lactea Thunb. subsp. adelphea
(Eckl. & Zeyh.) Dahlg.) 16
A. TRUNCATA ECKL. 6 ZEYH.
( -A . s phaerocepha la Schltr.) 1, 16
( - A . truncate Eckl. & Zeyh. subsp.
sphaerocephala (Schltr.)
Dahlg.) 16
A. truncate Eckl. & Zeyh. subsp. sphaerocephala
(Schltr.) Dahlg. - A. TRUNCATA
A. TULBAGHENSIS DAHLG.
(-A. acuminata Lara, subsp. magnlflora
Dahlg.) 16
A. UNIFLORA L.
( -A . larlclfolla Lam.) 10
(-A. leptophylla Eckl. & Zeyh.) 10
A. uniflora L. subsp. vl lldenovlana (Benth.)
Dahlg. - A. WI LLDENOWI AN A
BENTH.
A. VAR IANS ECKL. & ZEYH.
A. varlans Eckl. 6 Zeyh. subsp. Isolate Dahlg.
- A. ISOLATA
25550 A. WILLDENOWIANA BENTH.
(-A. uniflora L. subsp. wllldenovlana
( Benth . ) Dahlg . ) 16
3662050 -Paraspalathus Presl - ASPALATHUS
P. aemula (E. Mey.) Presl - ASPALATHUS AEMULA
P. argentea (L.) Presl - ASPALATHUS
CALEDONENS IS
P. heterophylla ("Thunb.") Presl - ASPALATHUS
HETEROPHYLLA
P. h um 1 f u sa Presl - ASPALATHUS LOTOIDES SUBSP.
LOTOIDES
P. lotoldes (Thunb.) Presl - ASPALATHUS
LOTOIDES SUBSP. LOTOIDES
P. procumbens (E. Mey.) Presl - ASPALATHUS
LOTOIDES SUBSP. LOTOIDES
P. serlcea (Berg.) Presl - ASPALATHUS SERICEA
3662100 -Trineurla Presl - ASPALATHUS
T. linearlfolla (DC.) Presl - ASPALATHUS
LINEARIFOLIA
3664000 -DICHILUS DC.
2. SCHUTTE & VAN WYK. 1988. S. SFR. J. BOT.
54,2: 182-186.
D. multlf lorus Burtt Davy - D. PILOSUS
400 D. PILOSUS CONRATH EX SCHINZ
(-D. multiflorus Burtt Davy) 2
(-D. pi 1 o 8 u 8 Conrath ex Schinz var.
multiflorus Burtt Davy) 2
( - D . p 1 1 o 8 us Kenslt) 2
D. pilosus Conrath ex Schinz var. multiflorus
Burtt Davy - D. PILOSUS
D. pilosus Kenslt - D. PILOSUS
450 D. REFLEXUS (N.E. BR.) A . L . SCHUTTE
3669000 -C ROTALAR I A L. Revision: R.M. Polhlll (K).
C. lenticula E. Mey. - L0T0N0NIS LENTICULA
3682000 -C YT IS US L.
C. psor aloldes L. - INDIGOFERA PSORALOIDES
3685000 -Ononis L. Southern African species moved to
ASPALATHUS, CROTALAR IA ,
INDIGOFERA, L0T0N0NIS
0. flliformis L. - INDIGOFERA SARMENTOSA
0. lagopus Thunb. - ASPALATHUS LOTOIDES SUBSP.
LAGOPUS
0. rnaur l tan lea L. - INDIGOFERA MAURITAN IC A
0. splcata Thunb. - ASPALATHUS AEMULA
3698000
-LOTUS L.
L. flliformis Berg. - INDIGOFERA SARMENTOSA
L. fruticosus Berg. - INDIGOFERA M AUR I TAN I C A
L. micro phyllus Hook. - INDIGOFERA GRACILIS
L. racemosus Poir. - INDIGOFERA MAURITANICA
3700000 -CYAMOPSIS DC.
100 C. SERRATA SCHINZ
(-Indigofera psammotropha H. Bol.) #
3702000
50
200
300
450
470
550
600
-INDIGOFERA L. Revision: B. Schrlre (PRE).
(Note all species are listed)
1. HARVEY. 1862. FC 2: 163-203.
2. GILLETT'. 1958. KEW BULL. ADD. SER. 1: 8.
3. SCHREIBER. 1970. FSWA 60: 38.
4. ROSS. 1972. FL. NATAL 200.
5. GILLETT, POLHILL & VERDCOURT. 1971. FTE A
3,1: 21 .
6. HILLIARD & BURTT. 1986. NOTES ROY. BOT.
GDN. EDINB. 43: 207-210.
7. JARVIE & STIRTON. 1986. BOTHALIA 16,2:
230.
8. JARVIE & STIRTON. 1987. BOTHALIA 17,1: 1-
6.
9. GERMISHUIZEN. 1987. 17,1: 33-34.
#. PRE HERBARIUM PRACTICE, FOLLOWING
SCHRIRE .
I. ACANTHOCLADA DINTER
(-1. longlspina Bak. f. ex Glllett) #
I. acanthorhachi s Dinter - LESSERTIA
ACANTHORHACHIS
I. accepta N.E. Br. - I. SETIFLORA
I. acutifolla Schinz - I. DISJUNCTA VAR.
DISJUNCTA
I. acutlsepala Conrath ex Bak. f. - I. ZEYHERI
I. ADENOCARPA E. MEY.
I. ADENOIDES BAK. F.
I. adonensls E. Mey. - I. POLIOTES
I. adscendens Eckl. & Zeyh. - I. HETEROPHYLLA
I. aeruginls Schweinf. - l. TR I GONE LLO I DE S
I. afflnls Harv. - I. BURKEANA
I. ALOPECUROIDES (BURM. F.) DC. VAR.
ALOPECUROIDES
(-1. corlacea Alt. var. alopecuro Ides
Harv.) #
I. ALOPECUROIDES (BURM. F.) DC. VAR. MINOR E.
MEY.
(-1. a lo pec ur o Id e 8 (Burn, f.) DC. var.
minor Eckl. & Zeyh.) #
(-1. corlacea Alt. var. minor (E. Mey.)
Harv.) #
(-1. maurltanlca (L.) Thunb. var. minor
E. Mey.) #
(-1. mlcrophylla Lara.) #
I. alopec uro Ides E. Mey. non DC. - I.
CANDOLLEANA
I. a lo pecuro lde s (Burra, f.) DC. var. minor
Eckl. & Zeyh. - I.
ALOPECUROIDES VAR. MINOR
I. alopecurus Schltr. - I. FOLIOSA
I. ALPINA ECKL. 6 ZEYH.
(-1. stlpularls Link.) #
I. ALTERNANS DC. VAR. ALTERNANS
Bothalia 19,2 (1989)
283
620
700
800
900
1000
1050
1200
1300
1400
1500
1600
1700
1900
2000
2 100
2200
2300
2400
2450
2600
2800
2900
3000
3100
3150
3200
3300
3350
3400
3450
3460
(■I. effusa E. Mey.) 2
(■I. enneaphylla Eckl. & Zeyh. non L.) It
I. ALTERNANS DC. VAR. MACRA BAK.
I. AMITINA N.E. BR.
I. AMOENA AIT.
I. araoena E. Mey. non Ait. - I. MEYER I ANA
I. ANA8IBENSIS A. SCHREIB .
I. angustata E. Mey. - I. STENOPHYLLA ECKL. 6
ZEYH .
I. angastlfolla Curtis non L. - I. VERRUCOSA
I. ANGUSTIFOLIA L. VAR. ANGUSTIFOLIA
I. angustifolla L. var . b r ac hy 8 1 ac hya DC. - I.
BRACHYSTACHYA
I. ANGUSTIFOLIA L. VAR. TENUIFOL I A (LAM.) HARV.
(-1. leptocaulls Eckl. & Zeyh.) #
(-1. 8trigosa Spreng.) #
I. angustifolla Thunb. non L. - I.
BRACHYSTACHYA
I. angus t iloba Bak. f. - I. TORULOSA VAR.
ANGUSTILOBA
I. anil L. - I. SUFFRUTICOSA
I. an t en nu 1 1 f e r a L. Bol. - I. ARENOPHILA
I. aphylla Breit. ex Link. - I. FILIFOLIA
THUNB .
I. AQUAE-NITENTIS BREM.
I. arenaria E. Mey. - I. EXIGUA
I. ARENOPHILA SCHINZ
(■I. an tennul 1 f e ra L. Bol.) 2
(■Lotononis otnahekensis Dinter ex
Schreiber) 3
I. ARGYRAEA ECKL . & ZEYH.
(■I. burchellii E. Mey. var. paucifolia
E. Mey.) #
(■I. nivea Willd. ex Spreng.) #
I. ARGYROIDES E. MEY.
(-1. engleri Bak. f.) 2, 3
(-1. s ax 1 co la Engl.) 2, 3
I. ARISTATA SPRENG.
I. arrecta Benth . ex Harv. non Hochst . ex A.
Rich. - I. CONFUSA
I. ARRECTA HOCHST. EX A. RICH.
I. arthrophylla Eckl. & Zeyh. - I. DENUDATA
I. ASTRAGALI NA DC.
I. ATRATA N.E. BR.
I. atrlnota N.E. Br. - I. REDUCTA
I. AURICOMA E. MEY.
(■I. a'uricoma E. Mey. var. cuneata Bak.
f.) 2, 3
I. auricoma E. Mey. var. cuneata Bak. f. ■ I.
AURICOMA
I. auricoma E. Mey. var. hololeuca (Benth. ex
Harv.) Gillett - I. HOLOLEUCA
I. BAINESII BAK.
(■I. varlabilis N.E. Br . p.p.) 2
I. bakerlana Viguier - I. DEMISSA
I. baukeana Vatke - I. SCHIMPERI VAR. BAUKEANA
I. BAUMIANA HARMS
I. BIFRONS E. MEY.
(-1. bifrons E. Mey. var. dlgitata E.
Mey . ) It
(-1. burchellii DC. non E. Mey.) It
I. bifrons E. Mey. var. digltata E. Mey. - I.
BIFRONS
I. bif rons E. Mey. var. trifollata E. Mey. -
I. CARDIOPHYLLA
I. BIJUGA VOGEL EX WALP.
I. bolusii N.E. Br . - I. SPICATA
I. BRACHYSTACHYA (DC.) E. MEY.
(■I. angustifolla L. var. br ac h y s t ac hy a
DC . ) #
(■I. angustifolla Thunb. non L.) It
I. brevifolia N.E. Br. - I. INYANGANA
I. BUCHANANI I 8URTT DAVY
I. burchellii DC. non E. Mey. - I. BIFRONS
I. burchellii E. Mey. var. raultifolia E. Mey. -
I. C0LLINA
I. burchellii E. Mey. var. paucifolia E. Mey. -
I. ARGYRAEA
I. BURKEANA BENTH. EX HARV.
( » I . affinis Harv.) 2, 3
(■I. gonioides Hochst. ex Bak. var.
damaren8is Bak. f.) 2, 3
I. cawa E. Mey. - I. HUMIFUSA
I. CANDICANS AIT.
I. candicans E. Mey. non Ait. - I.
HETEROPHYLLA
I. candicans Sieb. - I. FILIFORMIS
I. CANDIDISSIMA DINTER
I. CANDOLLEANA MEISN.
(-Anila mauritanica (L.) Kuntze var.
oligantha Kuntze) It
(■I. a lo pecuroid e s E. Mey. non DC.) #
(-1. coriacea Ait. var. hirta Harv.) It
(-1. mauritanica (L.) Thunb. var. erecta
Eckl. & Zeyh.) #
I. CAPI LLARIS THUNB.
I. CARDIOPHYLLA HARV.
(-1. bifrons E. Mey. var. trifollata E.
Mey.) #
I. carinata De Wild. - I. TRITA SUBSP.
SUBULATA
I. CECILII N.E. BR.
I. centrota Eckl. & Zeyh. ■ I. DENUDATA
I. CHARL I E R I A N A SCHINZ VAR. CHARLIERIANA
I. CHARLIERIANA SCHINZ VAR. LATA GILLETT
I. CHARLIERIANA SCHINZ VAR. SCABERRIMA (SCHINZ)
GILLETT
(■I. charlieriana Schinz var. sessllis
(Chiov.) Gillett) #
(■I. relaxata N.E. Br.) #
(-1. scaberrima Schinz) It
I. charlieriana Schinz var. sessllis (Chiov.)
Gillett - I. CHARLIERIANA VAR.
SCABERRIMA
3500
3550
3600
3700
3800
4000
4100
4110
4500
4600
4700
47 50
4780
4800
4900
5100
5200
5250
5300
5 500
55 50
5600
5700
5800
5900
6000
6100
6200
6 300
6600
6800
clnerascens Eckl. 4 Zeyh. - I. ZEYHERI
CIRCINNATA BENTH. EX HARV.
cognata N.E. Br. - I. VICIOIDES VAR.
ROGERS 1 1
COLLI NA ECKL. & ZEYH.
(•I. burchellii E. Mey. var. oultifolla
E. Mey.) #
C0LUTEA (BURM. F.) MERR.
(-Calega colutea Burn . f.) 2, 3
(-1. consangulnea Klotzsch) 2
(-1 . Junodli N.E. Br . ) 2
(■I. setlculosa Harv.) 2
(-1. vlscosa Lara.) 2
commiphoroides Chiov. - I. LUPATANA
COMMIXTA N.E. BR.
COMOSA N.E. BR.
(-1. malacoa tachys Benth. ex Harv. var.
semlnuda N.E. Br . ) #
compacta N.E. Br. - I. HILARIS
complanata Spreng. - I. PSORALOIDES
COMPLICATA ECKL. & ZEYH.
CONCAVA HARV.
CONFUSA PR A 1 N 6 BAK. P.
(■I. arrecta Benth. ex Harv. non Hochst.
ex A. Rich.) #
(■I. pretorlana Harms) #
consangulnea Klotzsch - I. COLUTEA
coriacea Ait. - I. MAURITANICA
coriacea Ait. var. alopecuroldes Harv. • I.
AL0PECUR0I DES VAR.
ALOPECUROI DES
coriacea Ait. var. cana Harv. - I.
MAURITANICA
coriacea Alt. var. hirta Harv. - I.
CANDOLLEANA
coriacea Ait. var. major E. Mey. - I.
MAURITANICA
coriacea Ait. var. minor (E. Mey.) Harv. -
I. ALOPECUROI DES VAR. MINOR
corniculata E. Mey. - I. TRISTIS
co ron i 1 lo ide s Jaub. & Spach. - I. TRITA
SUBSP. SCABRA
COSTATA GUILL. & PERR. SUBSP. MACRA (E.
MEY.) GILLETT
(•I. macra E. Mey.) 2, 3, 4
CREBRA N.E. BR.
CRYPTANTHA BENTH. EX HARV. VAR. CRYPTANTHA
CRYPTANTHA BENTH. EX HARV. VAR. 0CCIDENTALIS
BAK. F.
CUNEI FOLIA ECKL. & ZEYH. VAR. ANGUSTIFOLIA
HARV .
CUNEIPOLIA ECKL. & ZEYH. VAR. CUNEIFOLIA
(-1. florida E. Mey.) 1
CUNENENSIS TORRE
cylindrica DC. - I. FRUTESCENS
C YT I S 01 DES (L.) L.
(-1. lotoides auct. non Lam.) #
(■i. mucronata Lam.) #
(-Psoralea cytisoides L.) #
DALEOIDES BENTH. EX HARV. VAR. DALEOIDES
(-1. dodecaphylla Ficalho & Hiern) 2
DALEOIDES BENTH. EX HARV. VAR. GOSSWEILERI
BAK. F.
(-1. hololeuca Benth. ex Harv. var.
angolensis Bak. f.) 2
DAMARANA MERXM. & A. SCHREIB.
(■I. wilmaniae Bak. f. ex Gillett) 3
dealbata Harv. - I. SPINESCENS
DECLINATA E. MEY.
DECORA LINDL. *
deflexa Hochst. ex A. Rich. ■ I. PARVIFLORA
VAR. PARVIFLORA
DELAGOAENSIS BAK. F. EX GILLETT
DEMISSA TAUB.
(-1. bakerlana Viguier) 2
DENSA N.E. BR.
denudata L. f. var. dumosa (E. Mey.) Harv. -
I. DENUDATA
denudata L. f. var. luxurians Harv. - I.
DENUDATA
denudata L. f. var. s 1m p 1 ic i f o 1 ia Harv. -
I. DENUDATA
DILLWYNIOIDES BENTH. EX HARV.
DIMIDIATA VOGEL EX WALP.
d imorphophy 1 la Schinz ■ I. TRITA SUBSP.
SUBULATA
discolor E. Mey. - I. PROCUMBENS VAR.
PROCUMBENS
DISJUNCTA GILLETT VAR. DISJUNCTA
(■I. acutifolia Schinz) 2, 3
dlssimilis N.E. Br. - I. VICIOIDES VAR.
ROGERS 1 1
DISTICHA ECKL. & ZEYH.
(»I. enonensls E. Mey.) #
divaricate De Wild. - I. VICIOIDES VAR.
Bothalia 19,2 (1989)
284
VICIOIDES
I. dodecaphylla Ficalho & Hlern - I. DALEOIDES
VAR. DALEOIDES
6900 I. DOLICHOTHYRSA BAK. F.
7000 I. DREGEANA E. MEY.
(■I. gerrardiana Harv.)
I. durnosa E. Mey. - I. DENUDATA
I. echinata Willd. - I. NUMMULAR 1 1 FO L I A
I. effusa E. Mey. - I. ALTERNANS VAR.
ALTERNANS
7100 I. EGENS N.E. BR.
I. elLlptica E. Mey. - I. PAUCIFLORA'
I. endecaphylla Jacq. var. angustata Harv. -
I. SPICATA
I. engleri Bak. f. - I. ARGYROIDES
I. enneaphylla Eckl . & Zeyh. non L. - I.
ALTERNANS VAR. ALTERNANS
I. enonensis E. Mey. » I. DISTICHA
7300 l. ENORMIS N.E. BR.
I. erecta Eckl. & Zeyh. non Thunb. - I.
GRACILIS
I. erecta Thunb. - I. PORRECTA VAR. BICOLOR
7500 I. ERIOCARPA E. MEY.
I. erlocarpa E. Mey. var. wllliamsonli Harv. -
I. WILLI AMSONII
7600 I. EVANS IANA BURTT DAVY
(■I. pauxllia N.E. Br . ) #
7 700 I. EVANS 1 1 SCHLTR ,
7750 I. EXIGUA ECKL. & ZEYH.
(■I. arenarla E. Mey.) #
I. eyleslana auct. non Gillett - I. PROCUMBENS
I. falcata E. Mey. = I. SESSILIFOLIA
I. falcata E. Mey. var. glaberrima E. Mey. -
I. SESSILIFOLIA
I. falcata E. Mey. var. pubescens E. Mey. - I.
SESSILIFOLIA
7900 I. FASTIGIATA E. MEY.
( -I . faetlgiata E. Mey. var. angustata
Harv.) 2 , 4
(-1. rostrata Conrath) 2
I. fastlgiata E. Mey. var. angustata Harv. -
I. FASTIGIATA
I. ferruginea Schura. & Thonn. - I. HIRSUTA
VAR. HIRSUTA
8000 I. F ILICAULIS ECKL. & ZEYH.
(■I. subtlll8 E. Mey.) 1
I. fillfolia Ker-Gawl. - I. F ILIFOLIA
8100 I. FIL I FOLIA THUNB.
(-1. aphylla Breit. ex Link.) #
(-1. fillfolia Ker-Gawl.) t
( -I . Juncea DC . ) #
(■I. nuda (Sims) G. Don) it
(-Lebeckla contaminata Alt. f. non
Willd.) it
(-Lebeckla contaminata Ker-Gawl. non
Thunb . ) #
(-Lebeckla nuda Sims) #
8200 I. FILIFORMIS L. F.
(-1. candicans Sleb.) 1
(-1. flliformis L. f. var. adscendens
Eckl. & Zeyh.) it
(-1. flliformis L. f. var. planlfolia E.
Mey.) it
(-1. wynbergensls S. Moore) it
I. flliformis L. f. var. adscendens Eckl. &
Zeyh. - I. FILIFORMIS
I. flliformis L. f. var. planlfolia E. Mey. -
I. FILIFORMIS
8300 I. FILIPES BENTH. EX HARV.
8400 I. FLABELLATA HARV.
8500 I. FLAVICANS BAK.
I. fleckli Bak. f. - I. PECHUELtI
I. flexuoaa Eckl. & Zeyh. - I. DENUDATA
8700 I. FLORIBUNDA N.E. BR.
I. f lor Ida E. Mey. - I. CUNEIFOLIA VAR.
CUNEIFOLIA
8800 I. FOLIOS A E. MEY.
(-1. alopecurus Schltr.) #
(-1. vestlta Harv.) #
8900 I. FRONDOSA N.E. BR.
9000 I. FRUTESCENS L.
(-1. cylindrica DC.) it
9100 I. FULCRATA HARV.
I. fusca G. Don - I. HIRSUTA VAR. HIRSUTA
9250 I. GAIRDNERIAE HUTCH. EX BAK. F.
9300 I. GALPINII N.E. BR.
I. garkeana auct. non Vatke - I. HOMBLEI
I. geVardiana Wall, ex Bak. - I. HETERANTHA
I. gerrardiana Harv. - I. DREGEANA
9600 I. GIESSII A. SCHREIB.
9650 I. GIFBERGENS IS C.H. STIRTON & J.K. JARVIE
I. glabella Fourc. - I. VERRUCOSA
9700 I. GLAUCESCENS ECKL. & ZEYH.
9800 I. GLOMERATA E. MEY.
I. goniocarpa Bak. f. - I. LUPATANA
I. gonloldes Hochst. ex Bak. var. damarensis
Bak. f. - I. BURKEANA
9900 I. GRACILIS SPRENG.
(-1. erecta Eckl. & Zeyh. non Thunb.) #
(-1. setacea E. Mey.) #
(-Lotus raicr ophyl lus Hook.) #
10000 I. GRATA E. MEY.
I. grlquana Schltr. ex A. Zahlbr. - I.
LONGEBARBATA
10050 I. GRISOPHYLLA FOURC.
10200 I. GUERRANA TORRE
10300 I. GUTHRIEI H. BOL.
10400 I. HAMULOSA SCHLTR.
10450 I. HANTAMENSIS DIELS
I. he d ran oph y 11 a Eckl. & Zeyh. - I.
SESSILIFOLIA
10500 I. HEDYANTHA ECKL. & ZEYH.
(-1. secunda E. Mey.) #
I. he nd e c a ph y 1 1 a Jacq. - I. SPICATA
105 70
10600
10 7 0 0
10800
10900
11100
1 1 300
11350
11400
11550
11500
11600
11650
1 1700
11800
11900
11920
11950
12150
12200
12300
12400
12600
126 50
1 2700
12 7 50
12800
12900
13100
1 3200
1 3300
13600
1 3700
1 3800
I. HETERANTHA WALL. EX BRANDIS *
(-1. gerardiana Wall, ex Bak.) #
I. HETEROPHYLLA THUNB.
(-1. adscendens Eckl. & Zeyh.) #
(-1. candicans E. Mey. non Alt.) #
(-1. heterophylla Thunb. var.
tulbaghensls Bak. f.) #
I. heterophylla Thunb. var. montana Eckl. &
Zeyh. - I. SPINESCENS
I. heterophylla Thunb. var. tulbaghensls Bak.
f. - I. HETEROPHYLLA
I. HETEROTRICHA DC.
(-1 . rudis N.E. Br . ) 2, 3
I. heterotrlcha DC. var. ecklonil Harv. - I.
POLIOTES
I. heterotrlcha Eckl. & Zeyh. non DC. - I.
POLIOTES
I. HILARIS ECKL. & ZEYH.
(-1. compacta N.E. Br . ) #
(-1. hilarls Eckl. & Zeyh. var.
d rakensbergensis Bak. f.) #
(-1. hlrta E. Mey.) it
(-1. lelpzlglae Brem.) #
(-1. patula Bak.) 2
( -I. simllls N.E. Br . ) 2
(-1. wentzellana Harms) 2
I. hilarls Eckl. & Zeyh. var. drakensbergensls
Bak. f. - I. HILARIS
I. HIRSUTA L. VAR. HIRSUTA
(-1. ferruginea Schum. & Thonn.) 2
(-1. fusca G. Don) 2
I. hirsute L. var. polystachya Welw. ex Bak. -
I. LONGEBARBATA
I. hlrta E. Mey. - I. HILARIS
I. HISPIDA ECKL. & ZEYH.
(-1. lotoldes E. Mey.) #
(-1. rhodantha Fourc.) #
I. HOCHSTETTERI BAK. SUBSP. STREYANA ( MERXM . )
A. SCHREIB.
(-1. streyana Merxm.) 3
I. HOFMANNIANA SCHINZ
I. H0L0LEUCA BENTH. EX HARV.
(-1. aurlcoma E. Mey. var. hololeuca
(Benth. ex Harv.) Gillett) #
I. hololeuca Benth. ex Harv. var. angolensls
Bak. f. - I. DALEOIDES VAR.
G0SSWE ILERI
I. HOMBLEI BAK. F. & MARTIN
(-1. garkeana auct. non Vatke) #
I. HOLUBII N.E. BR.
I. HUMIFUSA ECKL. & ZEYH.
(-1. calva E. Mey.) #
I. HYBRIDA N.E. BR .
I. INCANA THUNB.
I. incana Thunb. var. ang us t Is t 1 pul a t a Bak. f.
- I. TOMENTOSA
I. INGRATA N.E. BR .
I. INHAMBANENSIS KL0TZSCH
(-1. polycarpa Benth. ex Harv.) 2, 4
I. intermedia Harv. - I. MEYERIANA
I. INYANGANA N.E. BR.
(-1. brevifolla N.E. Br . ) #
I. 1 0 N 1 1 J.K. JARVIE & C.H. STIRTON
I. Juncea DC. - I. FILIFOLIA
I. Junodil N.E. Br. - I. COLUTEA
I. krausslana Meisn . - I. DENUDATA
I. KROOKIl SCHLTR. EX A. ZAHLBR.
(-1. woodii H. Bol. var. parvifolia H.
Bol . ) #
I. LANGEBERGENSIS L. BOL.
1. laterltla Bertol. - I. N0TATA
I. LAXERACEMOSA BAK. F.
I. LEENDERTZIAE N.E. BR.
I. lelpzlglae Brem. - I. HILARIS
I. LEPIDA ECKL. & ZEYH.
I. LEPTOCARPA ECKL. & ZEYH.
I. leptocaul is Eckl. & Zeyh. - I. ANGUSTIFOLIA
VAR. TENUIFOLIA ,
I. leptophylla E. Mey. - I. VERRUCOSA
l. LIMOSA L. BOL.
I. LOBATA GILLETT
( -Microchar Is latifolia Benth. non
Mich.) it
I. LONGEBARBATA ENGL.
(-1. grlquana Schltr. ex A. Zahlbr.) 2
(-1. hlrsuta L. var. polystachya Welw.
ex Bak.) 2
(-1. schlechterl Bak. f.) 2, 4
I. LONGIPES N.E. BR.
(-Anlla tenuifolla (Lam.) Kuntze var.
fillfolia Kuntze) #
I. longisplna Bak. f. ex Gillett - I.
ACANTHOCLADA
I. lotoldes E. Mey. - I. HISPIDA
I. lotoldes auct. non Lam. - I. CYTIS0IDES
I. LUPATANA BAK. F.
(-1. coamlphoroides Chlov.) 2
(-1. goniocarpa Bak. f.) 2
I. LYALLI BAK. SUBSP. LYALLI
(-1. obermei jer lae Brem.) 2
I. LYDENBURGENSIS N.E. BR.
I. macra E. Mey. - I. COSTATA SUBSP. MACRA
I. malacostachys Benth. ex Harv. - I.
MELANADENIA
I. malacostachys Benth. ex Harv. var. macrura
Conrath ex Bak. f. - I.
MELANADENIA
I. malacostachys Benth. ex Harv. var. seminuda
N.E. Br. - I. C0M0SA
I. MAR IT IMA BAK.
I. MASONAE N.E. BR.
I. MAURITANICA (L.) THUNB.
(-1. corlacea Ait.) #
(■I. corlacea Alt. var. cana Harv.) #
Bothalia 19,2 (1989;
285
1 3900
14000
14050
14100
1 4200
1 4250
14300
14400
14500
14600
14700
14800
14900
1 4950
15100
1 5200
15300
15400
1 5500
15600
1 5700
15800
15900
16000
16200
1 6250
16300
16400
16450
16600
16660
16800
16850
1 6900
1 7000
17200
17350
1 7300
17400
(■I. corlacea Alt. var. major E. Mey.) #
(-Lotus fruticosus Berg.) #
(-Lotus raceraosus Poir.) 9
(-Ononis naur Itanlca L.) 9
I. raauritanlca (L.) Thunb. var. erecta Eckl. &
Zeyh. - I. CANOOLLEANA
I. raauritanlca (L.) Thunb. var. minor E. Mey. -
I. AL0PECUR0IDES VAR. MINOR
I. raearnsil Standley - I. SWAZIENSIS VAR.
SWAZIENSIS
I. MELANADENIA BENTH. EX HARV.
(-1. raalacostach-ys Benth. ex Harv.) If
(-1. malacoe t achya Benth. ex Harv. var.
macrura Conrath ex Bak. f.) #
I. MERXMUELLER I A. SCHREIB.
I. ME YER I ANA ECKL. & ZEYH.
(-1. araoena E. Mey. non Alt.) #
(-1. Intermedia Harv.) 9
I. MICRANTHA E. MEY.
I. MICROCARPA DESV.
I. mlcrophyl la Lam. - I. ALOPECUROIDES VAR.
MINOR
I. MIM0S0IDES BAK.
I. MISCHOCARPA SCHLTR .
I. MOLL IC0M A N.E. BR.
(-1. nelaonll N.E. Br.) 9
I. mollis E. Mey. - I. MOLLIS
I. MOLLIS ECKL. & ZEYH.
(-1. mollis E. Mey.) #
I. moniliformis Bak. f. - I. ORMOC AR POI D ES
I. MONOSTACHYA ECKL. & ZEYH.
(-1. oroboldes E. Mey.) #
I. mucronata Lam. - I. CYTISOIDES
I. MUNDIANA ECKL. & ZEYH.
I. nana Eckl. & Zeyh. - I. ZEYHERI
I. NATALENSIS H. BOL.
I. N E BROW N I AN A GILLETT
(“I. varlabllls N.E. Br . p.p.) 2
I. NEGLECTA N.E. BR.
I. nelsonll N.E. Br. - I. MOLLICOMA
I. nlgromontana Eckl. & Zeyh. - I. SPINESCENS
I. nltlda Salter - I. PSORALOIDES
I. nlvea Wllld. ex Spreng. -. I. ARGYRAEA
I. NOTATA N.E. BR.
(-1. laterltla Bertol.) #
I. nuda (Sims) O. Don - I. FILIFOLIA
I. NUDIC AULIS E. MEY.
I. NUMMULAR 1 1 FOLIA LIVERA EX ALSTON
( -Ac a n tho no t us echlnatus (Wllld.)
Benth.) 2
( -Hedysarum n urarn u 1 a r 1 1 f o 1 1 um L.) 2, 3
(-1. echlnata Wllld.) 2, 3
I. OBCORDATA ECKL. & ZEYH.
I. o be r me 1 J e r 1 a e Brem. - I. LYALLI SUBSP.
LYALLI
I. OBSCURA N.E. BR.
(»I. strlcta L. f. var. acuta Harv.) #
I. ollverl Schweinf. ex Engl. - I. SWAZIENSIS
VAR. SWAZIENSIS
I. ollverl Schweinf. ex Harms - I. SWAZIENSIS
VAR. SWAZIENSIS
I. ORMOCARPOIDES BAK.
(-1. moniliformis Bak. f.) 2
(-1. torulosa Bak.) 2
I. oroboldes E. Mey. - I. MONOSTACHYA
I. OVATA L. F.
(-1. sarmentosa L. f. var. latlfolla
Eckl. & Zeyh.) If
I. OVINA HARV.
I. OXALIDEA WELW. EX BAK.
(-1. supralevis N.E. Br . ) 2
I. OXYTROPIS BENTH. EX HARV.
I. PAPPEI FOURC.
I. parkerl Bak. - I. SPICATA
I. PARVIFLORA HEYNE EX WIGHT & ARN. VAR.
OCC I D6NTALIS GILLETT
I. PARVIFLORA HEYNE EX WIGHT & ARN. VAR.
PARVIFLORA
(-1. deflexa Hochst . ex A. Rich.) 2
I. patens Eckl. A Zeyh. - I. SESSILIFOLIA
I. patula Bak. - I. HILARIS
I. pau lflora E. Mey. non Eckl. & Zeyh. - I.
STRICTA
I. PAUCIFLORA ECKL. & ZEYH.
(-1. elliptic a E. Mey.) 9
I. pa ux 1 1 la N.E. Br. - I. EVANS IANA
I. PE ARS ONI I BAK. F.
I. PECHUELII KUNTZE
(-1. fleckii Bak. f.) 9
I. PENTAPHYLLA BURCH. EX HARV. NON MURR.
I. perplexa N.E. Br . - I. SWAZIENSIS VAR.
PERPLEXA
I. PLACIDA N.E. BR.
(-Anila rupestris (Eckl. & Zeyh.) Kuntze
var. glencoensis Kuntze) If
(-1. rufescens E. Mey.) 9
I. PLATYPODA E. MEY.
I. PODOPHYLLA BENTH. EX HARV.
I. POLIOTES ECKL. & ZEYH.
(-1. adonensls E. Mey.) #
(-1. heterotricha DC. var. ecklonii
Harv . ) #
(-1. heterotricha Eckl. & Zeyh. non DC.)
#
(-1. punctata Eckl. & Zeyh. non Thunb.)
#
(-1. rupestris Eckl. & Zeyh.) #
I. polycarpa Benth. ex Harv. - I.
INHAMBANENS IS
I. PONGOLANA N.E. BR.
I. PORRECTA ECKL. & ZEYH. VAR. BICOLOR HARV.
(-1. erecta Thunb.) #
I. PORRECTA ECKL. & ZEYH. VAR. PORRECTA
I. PRATICOLA BAK. F.
1 7 5 50
1 7600
17610
1 7 6 30
1 7 6 50
1 7800
1 7900
18000
1 8020
18050
18100
1 82 00
1 82 50
18300
1 8500
1 8600
18700
1 87 50
19000
19100
19150
19200
19300
19500
19700
19800
19850
19900
20000
I. pre tor lana Harms - I. CONPUSA
I. procurabens E. Mey. - 1. PROCUMBENS VAR.
CONCOLOR
I. PROCUMBENS L. VAR. CONCOLOR HARV.
(-1. procurabens E. Mey.) 9
I. procurabens L. var. discolor (E. Mey.) Harv.
- I. PROCUMBENS VAR.
PROCUMBENS
I. PROCUMBENS L. VAR. PROCUMBENS
(-1. discolor E. Mey.) 9
(-1. procurabens L. var. discolor (E.
Mey.) Harv.) 9
I. PROCUMBENS TORRE
(-1. eylestana auct. non Glllett) 9
I. psammotropha H. Bol. - CYAMOPSIS SERRATA
I. PSEUDOEVANSI I HILLIARD & BURTT
I. PSEUDO-INDIGOPERA ( MERXM . ) GILLETT
( -Ml cr ochar is galplnll N.E. Br . ) 2
( -M ic r och a r 1 s pseudo-l nd lgo f e ra Merxm.)
9
I. ps 1 locarpa Schltr. - I. SARMENTOSA
I. PSORALOIDES (L.) L.
(-Cytisus psoraloldes L.) #
(-1. coraplanata Spreng.) 9
(-1. nltlda Salter) 9
I. psoraloldes auct. non (L.) L. - I. RACEMOSA
I. punctata Eckl. 4 Zeyh. non Thunb. - I.
POLIOTES
I. punctata Thunb. - I. VERRUCOSA
I. PUNGENS E. MEY.
I. QUI NQUEFO LI A E. MEY.
I. RACEMOSA L.
(-1. psoraloldes auct. non (L.) L.) #
(-1. trifollata L.) 9
I. RADICIFERA CRONQUIST
I. RAUTANENI I BAK. F.
I. rechodes Eckl. & Zeyh. - I. DENUDATA
I. REDUCTA N.E. BR.
(-1. atrinota N.E. Br . ) #
I. REFLEXA E. MEY.
I. REHMANNII BAK. F.
I. relaxata N.E. Br . - T. CHARLIE R IANA VAR.
SCABERRIMA
I. retroflexa Baill. - I. TRITA SUBSP. SCABRA
I. rhodantha Fourc. - I. HISPIDA
I. RHYTIDOCARPA BENTH. EX HARV. SUBSP.
RHYTIDOCARPA
I. rlgescens E. Mey. var. lnermls E. Mey. - I.
DENUDATA
I. rlgescens E. Mey. var. splnosa E. Mey. - I.
DENUDATA
I. RIPAE N.E. BR.
I. rogersii R.E. Fr..- I. VICIOIDES VAR.
ROGERS 1 1
I. rostrata Conrath - I. FASTIGIATA
I. ROSTRATA H. BOL.
(-Anila zeyheri (Spreng.) Kuntze var.
macropnylla Kuntze) 9
I. RUBROGLANDULOSA GERMISHUIZEN
I. rudis N.E. Br. - I. HETEROTRICHA
I. rufescens E. Mey. - I. PLACIDA
I. rupestris Eckl. & Zeyh. - I. POLIOTES
I. SANGUINEA N.E. BR.
I. SARMENTOSA L. F.
(-1. psllocarpa Schltr.) 9
(-1. sarmentosa L. f. var. trifollata E.
Mey.) #
(-Lotus fillforml8 Berg.) 9
(■Ononis flliformis L.) 9
I. sarmentosa L. f. var. latlfolla Eckl. &
Zeyh. - I. OVATA
I. sarmentosa L. f. var. trifollata E. Mey. -
I. SARMENTOSA
I. saxlcola Engl. - I. ARGYROIDES
I. scaberrima Schlnz - I. CHARLI ER I ANA VAR.
SCABERRIMA
I. scabra Roth. - I. TRITA SUBSP. SCABRA
I. SCHIMPERI JAUB. & SPACH VAR. BAUKEANA
(VATKE) GILLETT
(-1. baukeana Vatke) 2, 3
I. SCHIMPERI JAUB. & SPACH VAR. SCHIMPERI
(-1. tettensls Klotzsch) 2, 3
I. SCHINZII N.E. BR.
I. schlechter 1 Bak. f. - I. LONGEBARBATA
I. secunda E. Mey. - I. HEDYANTHA
I. 8emlikien8is Robyns & Boutique - I.
VICIOIDES VAR. VICIOIDES
I. SESSILIFOLIA DC.
I. setacea E. Mey. - I. GRACILIS
I. seticulosa Harv. - I. COLUTEA
I. seticulosa Harv. var. luxurians H. Bol. -
I. SETOSA
I. SETIFLORA BAK.
(-1. accepts N.E. Br . ) 2
I. SETOSA N.E. BR.
(-1. seticulosa Harv. var. luxurians H.
Bol.) #
I. setulosa Bertol. - I. VERRUCOSA
I. SIEBERIANA SCHEELE
I. similis N.E. Br. - I. HILARIS
I. SORDIDA BENTH. EX HARV.
I. SPICATA FORSSK .
(-1. bolusii N.E. Br.) 2, 4
(-1. endec aphy 1 la Jacq. var. angustata
Harv.) 2, 4
(-1. hendecaphylla Jacq.) 2
(-1. parkerl Bak.) 2
286
Bothalia 19,2 (1989)
20100
20200
20400
20500
20700
20800
20850
20870
20900
21000
21100
21230
21250
21400
21450
2 1500
21550
21600
21700
2 1800
21900
2 1950
22000
22200
22 300
22 3 50
SPINESCENS E. MEY.
(■I. dealbata Harv.) #
(■I. heterophylla Thunb. var . nontana
Ec k 1 . & Ze yh . ) #
(-1. nigromontana Eckl . & Zeyh.) #
STENOPHYLLA ECKL. & ZEYH.
(DI. angustata E. Mey . ) #
(“I. zeyherl Sprang, ex Eckl. & Zeyh.
var. trlfoliolata Eckl. &
Ze yh . ) #
stlpularls Link. » I. ALPINA
streyana Merxa. - I. HOCHSTETTERI SUBSP.
STREYANA
STRICTA L. F.
(-1. pauciflora E. Mey. non Eckl. &
Zeyh.) 1
stricta L. f. var. acuta Harv. - I. OBSCURA
strict* L. f. var. peduncqlata Eckl. & Zeyh.
- I. ZEYHERI
strigosa Spreng. - I. ANGUSTIF0LIA VAR.
TENUIPOLIA
SUBC0RYMB0SA BAK. VAR. EYLESII BAK. F.
sublncana N.E. Br . - I. TRITA SUBSP.
SUBULATA
subtills E. Mey. - I. FILICAULIS
subulata Vahl ex Polr. - I. TRITA SUBSP.
SUBULATA
subulata Vahl ex Polr. var. scabra (Roth)
Melkle - I. TRITA SUBSP.
SCABRA
SUFFRUTICOSA MILL.
(-1 . anil L. ) #
SULCATA DC.
SUPERBA C.H. STIRTON
supralevls N.E. Br . » I. 0XALIDEA
SWAZIENSIS H. BOL. VAR. PERPLEXA (N.E. BR.)
GILLETT
(-1. perplexa N.E. Br.) 2
SWAZIENSIS H. BOL. VAR . ' SWAZ I ENSI S
(“I. aearnsll Standley) 2
(“I. ollverl Schvelnf. ex Engl.) 2
(»I. ollverl Schwelnf. ex Haris) 2
TEIXEIRAE TORRE
tenulcaulls Klotzsch » I. VICI0IDES V>?R.
VICIOIDES
TEN UISS IMA E. MEY.
t e t ragonoloba E. Mey. - I. TRITA SUBSP.
SUBULATA
tettensls Klotzsch - I. SCHIMPERI VAR.
SCHIMPERI
THESIOI DES J.K. JARVIE & C.H. STIRTON
TINCTORIA L. VAR. ARCUATA GILLETT
TOMENTOSA ECKL. & ZEYH.
(■I. lncana Thunb. var. angustlstlpulata
Bak. f.) #
torulosa Bak. - I. ORMOCA RPO I DE S
TORULOSA E. MEY. VAR. ANGUSTILOBA (BAK. F.)
GILLETT
(-1. angustlloba Bak. f.) 2
TORULOSA E. MEY. VAR. TORULOSA
transvaalensls Bak. f. - I. VICIOIDES VAR.
VICIOIDES
trifollata L. = I. RACEMOSA
TRIFOLIOIDES BAK. F.
TRIGONELLOIDES JAUB . & SPACH
(-1. aeruglnls Schwelnf.) 2
TRIQUETRA E. MEY.
TRISTIS E. MEY.
(-1. cornlculata E. Mey.)
TRISTOIDES N.E. BR.
(-1.
(-1.
(-1.
(-1.
(-1.
(-1.
(-1.
(-1
(-1.
(-1.
F. SUBSP. SCABRA (ROTH) DE KORT &
THI JSSE
cor o n 1 1 lo Id e 8 Jaub. & Spach.) 5
retrof lexa Balll.) 5
scabra Roth . ) 5
subulata Vahl ex Polr. var. scabra
(Roth) Melkle) it
trlta L. f. s u b 8 p . subulata (Vahl
ex Polr.) All var. scabra
(Roth) All) #
trlta L. f. var. scabra (Vahl ex
Polr.) All) #
F. SUBSP. SUBULATA (VAHL EX POIR.)
ALI
carlnata De Wild.) 2
d lior phophy 1 la Schlnz) 3
sublncana N.E. Br . ) #
subulata Vahl ex Polr.) 3
te t ragonoloba E. Mey.) #
trlta L. f. subsp. subulata (Vahl
ex Polr.) All var. subulata) #
trlta L. f. var. subulata (Vahl ex
Polr.) All) #
f. subsp. subulata (Vahl ex Polr.)
All var. scabra (Roth) All -
I. TRITA SUBSP. SCABRA
f. subsp. subulata (Vahl ex Polr.)
All var. subulata - I. TRITA
SUBSP. SUBULATA
trlta L. f. var. scabra (Vahl ex Polr.) All
- I. TRITA SUBSP. SCABRA'
trlta L. f. var. subulata (Vahl ex Polr.)
All - I. TRITA SUBSP. SUBULATA
varla E. Mey. - I. VICIOIDES VAR. R0GERSII
varlabll Is N.E. Br. p.p. - I. BAINESII
varlabllls N.E. Br . p.p. - I. NEBROWNI ANA
VELUTINA E. MEY.
VENUSTA ECKL. A ZEYH.
VERRUCOSA ECKL. & ZEYH.
(-1. angustl folia Curtis non L.) #
(■I. glabella Fourc . ) #
(-1. leptophylla E. Mey.) #
(-1. punctata Thunb.) #
(•I. setuloaa Bertol.) #
22450
22500
22800
23050
23000
23100
3702010
3702020
3703000
3715000
3718000
500
1200
2000
4000
4200
4400
(-1. zeyherl Spreng. ex Eckl. & Zeyh.
var. leptophylla (E. Mey.)
Harv.) #
I. vestlta Harv. - I. F0LI0SA
I. VICIOIDES JAUB. & SPACH VAR. ROGERSII (R.E.
FR.) GILLETT
(“I. cognata N.E. Br . ) 2
(-1. dissitnilis N.E. Br . ) #
(-1. rogersll R.E. Fr . ) 2
(-1. varla E. Mey.) #
I. VICIOIDES JAUB. & SPACH VAR. VICIOIDES
(•I. dlvarlcata De Wild.) 2
(“I. semllklensls Robyns & Boutique) 2
(-1. tenulcaulls Klotzsch) 2
(-1. transvaalensls Bak. f.) 2, 3
I. vlralnea E. Mey. - I. ZEYHERI
I. viscosa Lara. - I. COLUTEA
I. wentzellana Harms - I. HILARIS
I. WILL I AM SON 1 1 (HARV.) N.E. BR. *
(“I. erlocarpa E. Mey. var. wllllarasonli
Har v . ) 2 , 4
I. wllmanlae Bak, f. ex Glllett - I. DAMARANA
I. woodil H. Bo l . var. Intermedia H. Bol. - I.
WOO D 1 1 VAR. W00DII
I. W00DII H. BOL. VAR. LAXA H. BOL.
I. woodil H. Bol. var. parvlfolla H. Bol. - I.
KR00KII
I. WOO D 1 1 H. BOL. VAR. W00DII
( - I . woodil H. Bol. var. Intermedia H.
Bol.) #
I. wynbergensis S. Moore * I. FILIFORMIS
I. ZEYHERI SPRENG. EX ECKL. & ZEYH.
(■Anila zeyherl Kuntze var. normalis
Kun t z e ) #
(-1. acutlsepala Conrath ex Bak. f.) #
(■I. clnerascens Eckl. A Zeyh.) #
(■I. nana Eckl. A Zeyh.) #
(■I. stricta L. f. var. pedunculata
Eckl. A Zeyh.) #
(■I. vimlnea E. Mey.) #
I. zeyherl Spreng. ex Eckl. A Zeyh. var.
leptophylla (E. Mey.) Harv. -
I. VERRUCOSA
I. zeyherl Spreng. ex Eckl. A Zeyh. var.
trlfoliolata Eckl. A Zeyh. -
I. STENOPHYLLA
-Ac an thono t us Benth. - INDIG0FERA
A. echlnatus (Willd.) Benth. - INDIG0FERA
NUMMULARIIFOLIA
-Anila Kuntze - INDIG0FERA
A. mauritanlca (L.) Kuntze var. ollgantha
Kuntze - INDIG0FERA
CANDOLLEANA
A. rupestris (Eckl. A Zeyh.) Kuntze var.
glencoensis Kuntze -
INDIGOFERA PLACIDA
A. tenulfolia (Lam.) Kuntze var. flllfolla
Kuntze - INDIGOFERA L0NGIPES
A. zeyherl (Spreng.) Kuntze var. macrophylla
Kuntze - INDIGOFERA R0STRATA
A. zeyherl Kuntze var. normalis Kuntze -
INDIGOFERA ZEYHERI
-PS0RALEA L. Revision: C.H. Stirton (K).
P. cytisoides L. - INDIGOFERA CYTIS0IDES
-Galega L. Southern African species moved to
ASPALATHUS, CYCLOPIA, TEPHROSIA
G. colutea Burm. f. - INDIGOFERA COLUTEA
G. humills Thunb. - TEPHROSIA ANGULATA
G. pa 1 lens Ait. " TEPHROSIA CAPENSIS VAR.
CAPENSIS
-TEPHROSIA PERS. Revision: B.D. Schrire
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
(PRE) .
#. PRE HERBARIUM PRACTICE,, FOLLOWING
SCHRIRE .
araoena E. Mey. * T. KRAUSS I ANA
ANGULATA E. MEY.
(-Galega humills Thunb.) #
(-T. pallens (Ait.) Pers.) #
angu8tis8lma Engl. - T. L0NGIPES SUBSP.
LONGIPES
armltageana Chiov. - T. MACR0P0DA VAR.
MACR0P0DA
CAPENSIS (JACQ.) PERS. VAR. CAPENSIS
(-Galega pallens Ait.) it
(-T. ternatlfolia R.G.N. Young) #
damarensls Engl. - T. DREGEANA
dawsonli Bak. f. - T. LONGIPES SUBSP.
LONGIPES
dinteri Schlnz - T. DREGEANA
DREGEANA E. MEY.
( - T . damarensls Engl.) it
(-T. dinteri Schlnz) it
KRAUSSI ANA MEISN .
(-T.
linearis
LINEARIS
(-T.
LONGIPES
(-T.
(-T.
(-T.
(-T.
(-T.
amoena E. Mey.) it
(Willd.) Pers. subsp . discolor (E.
Mey.) Glllett) - T. LINEARIS
VAR. DISCOLOR
(WILLD.) PERS. VAR. DISCOLOR (E.
MEY.) BRUMMITT
linearis (Willd.) Pers. subsp.
discolor (E. Mey.) Glllett) it
MEISN. SUBSP. LONGIPES
angustlssima Engl.) 5
dawsonli Bak. f.) 5
longlpea Melan. subsp. longlpes
var. Icosisperma Brummltt) 5
lurlda Sond . var. druraraondii
Brumm 1 1 1 ) 5
lurlda Sond. var. llssocarpa
Bothalia 19,2 (1989)
287
A 7 00
67 50
Br umra i 1 1 ) 5
(-T. lur Ida Sond. var . lurida) 5
T. longipes Meian. subsp. long 1 pea var.
tcoalaperma Brummltt - T.
LONGIPES SUBSP. LONGIPES
T. lurida Sond. var. drumraondil Brummltt ■ T.
LONGIPES SUBSP. LONGIPES
T. lurida Sond. var. lissocarpa Brummltt - T.
LONGIPES SUBSP. LONGIPES
T. lurida Sond. var. lurida - T. LONGIPES
SUBSP. LONGIPES
T. MACR0P0DA (E. MEY.) HARV. VAR. MACR0P0DA
(-T. armitageana Chiov.) #
T. pallens (Alt.) Pers. - T. ANGULATA
T. PURPUREA -( L. ) PERS. SUBSP. LEPTOSTACHYA
(DC.) BRUMMITT VAR. PUBESCENS
BAK.
(-T. sparsiflora H.M. Forbes) #
T. sparsiflora H.M. Forbes - T. PURPUREA
SUBSP. LEPTOSTACHYA VAR.
PUBESCENS BAK.
T. ternatifolia R.G.N. Young - T. CAPENSIS
VAR. CAPENSIS
3746000 -Mlcrocharis Benth. - INDIGOFERA
M. galpinli N.E. Br. - INDIGOFERA PSEUDO-
INDIGOFERA
M. latifolia Benth non Mich. - INDIGOFERA
L0BATA
M. pseudo-ind igofe ra Merxm. - INDIGOFERA
PSEUD0-INDIG0FERA
( ■ D . caff rum Eckl. h Zeyh.) 2
( ■ D . scalpe DC.) 1, 2
(-D. 8 trangulaturo Wight & Arn.) 1, 2
(-Hedysarum repandum Vahl) l, 2
1200 D. SALICIPOLIUM (POIR.) DC. VAR. SALICIP0LIUA
( " D . grande E. Mey.) I, 2
("D. paleaceum GulIL. 6 Perr.) 1, 2
(-Hedysarum sallcifollum Poir.) 1, 2
D. scalpe DC. - D. REPANDUM
1300 D. SETIGERUM (E. MEY.) BENTH. EX HARV.
( - D . hirtura Gulll. h Perr.) 2
(-Nlcolsonia setlgera E. Mey.) 1, 2
D. spirale sensu Bak. - D. T0RTU0SUM
D. strangulatum Wight & Arn. - D. REPANDUM
1400 D. TORTUOSUM (SWARTZ) DC.
(-D. spirale sensu Bak.) I, 2
(-Hedysarum tortuosum Swartz) 1, 2
1500 0. VELUTINUM (WILLD.) DC.
( -D . laslocarpura (Beauv.) DC.) 2
(-Hedysarum laslocarpura Beauv.) 1, 2
(-Hedysarum velutlnum Wllld.) 1, 2
3807010 -Nlcolsonia Span. - DESMODIUM
N. barbata (L.) DC. var. argyrae (Welw. ex
Bak.) Schindl. - DESMODIUM
BARBATUM VAR. ARGYREUM
N. barbata (L.) DC. var. dlraorpt*a (Welw. ex
Bak.) Schindl. - DESMODIUM
BARBATUM VAR. DIM0RPHUM
N. caffra E. Mey. - DESMODIUM DREGEANUM
N. setigera E. Mey. - DESMODIUM SETIGERUM
3756000
100
-LESSERTIA DC. Revision: G.C. du
3. DINTER. 1932. FEDDE REP. 30:
L. ACANTHORHACHIS (DINTER) DINTER
(-Indigofera acanthorhachis
Pie s 8 i 8
202 .
Din t e r ) 3
3773000 -0RNITH0PUS L.
100 O. SATIVUS BROT. *
3778000 -Hedysarum L. Southern African species moved
to ALYSICARPUS, DESMODIUM,
ERIOSEMA, INDIGOFERA, WIBORGIA
H. canura J.F. Gmel. - DESMODIUM INCANUM
H. gangeticum L. - DESMODIUM GANGETICUM
H. incanura Swartz - DESMODIUM INCANUM
H. lasiocarpua Beauv. - DESMODIUM VELUTINUM
H. mac u 1 at um L. - DESMODIUM GANGETICUM
H. nummular iifolium L. - INDIGOFERA
NUMMULARIIFOLIA
H. racemosum Aubl. - DESMODIUM INCANUM
H. repandum Vahl - DESMODIUM REPANDUM
H. sallcifollum Poir. - DESMODIUM SALICIFOLIUM
VAR. SALICIFOLIUM
H. tortuosum Swartz - DESMODIUM TORTUOSUM
H. vaginale L. - ALYSICARPUS VAGINALIS VAR.
VAGINALIS
H. velutlnum Willd. - DESMODIUM VELUTINUM
3808000 -PSEUDARTHR IA WIGHT & ARN.
2. SCHRIRE. 1988. BOTHALIA 18,1: 11-24.
3810000 -ALYSICARPUS DESV.
2. SCHRIRE. 1988. BOTHALIA 18,1: 11-24.
A. glaber E. Mey. - A. RUGOSUS SUBSP.
PERENNIRUFUS
200 A. RUGOSUS (WILLD.) DC. SUBSP. PERENNIRUFUS J.
LEONARD
(-A. glaber E. Mey.) 2
(-A. violaceus (Forssk.) Schindl.) 2
( - A . wallichi Wight &« Arn.) 2
400 A. VAGINALIS (L.) DC. VAR. VAGINALIS
(-Hedysarum vaginale L.) 1, 2
A. violaceus (Forssk.) Schindl. - A. RUGOSUS
SUBSP. PERENNIRUFUS
A. wallichi Wight & Arn. - A. RUGOSUS SUBSP.
PERENNIRUFUS
3820000 -LESPEDEZA MICHX.
1. SCHRIRE. 1988. BOTHALIA 18,1: 11-24.
100 L. CUNEATA (DU MONT) G. DON *
(-L. sericea (Thunb.) Miq.) 1
L. sericea (Thunb.) Miq. - L. CUNEATA
3897000 -RHYNCHOSIA LOUR.
4750 R. NYASICA BAK.
3807000
300
400
600
650
750
1100
-DESMODIUM DESV.
2. SCHRIRE. 1988. BOTHALIA 18,1: 11-24.
D. barbatum (L.) Benth. subsp. dlraorphum (Welw.
ex Bak.) Laundon - D. BARBATUM
VAR. DIMORPHUM
D. BARBATUM (L.) BENTH. VAR. ARGYREUM (WELW. EX
BAK.) SCHUBERT
(-D. dimorphum Welw. ex Bak. var.
argyreum Welw. ex Bak.) 2
(-Nlcolsonia barbata (L.) DC. var.
argyrae (Welw. ex Bak.)
Schindl.) 2
D. BARBATUM (L.) BENTH. VAR. DIMORPHUM (WELW.
EX BAK.) SCHUBERT
(-D. barbatum (L.) Benth. subsp.
dimorphum (Welw. ex Bak.)
Laundon ) 1 , 2
(-D. dimorphum Welw. ex Bak.) 1, 2
(-Nlcolsonia barbata (L.) DC. var.
dimorpha (Welw. ex Bak.)
Schindl.) 1, 2
D. caffrum ( E .. Mey.) Druce - D. DREGEANUM
D. caffrum Eckl. & Zeyh. - D. REPANDUM
D. canura (J.F. Gmel.) Schinz & Thell. - D.
INCANUM
D. dimorphum Welw. ex Bak. - D. BARBATUM VAR.
DIMORPHUM
D. dimorphum Welw. ex Bak. var. argyreum Welw.
ex Bak. - D. BARBATUM VAR.
ARGYREUM
D. DREGEANUM BENTH.
( ■ D . caffrum (E. Mey.) Druce) 1, 2
(■Nlcolsonia caffra E. Mey.) 1, 2
D. GANGETICUM (L.) DC.
(-D. gangeticum (L.) DC. var. raaculatura
(L. ) Bak . ) l , 2
(-D. natalitum Sond.) 2
(-Hedysarum gangeticum L.) 1, 2
(-Hedysarum raaculatura L.) 1, 2
D. gangeticum (L.) DC. var. maculatum (L.) Bak.
D. GANGETICUM
D. grande E. Mey. - D. SALICIFOLIUM VAR.
SALICIFOLIUM
D. hirtum Gulll. & Perr. - D. SETIGERUM
D. INCANUM DC.
(-D. canum (J.F. Gmel.) Schinz & Thell.)
2
(-Hedysarum canura J.F. Gmel.) 2
(-Hedysarum incanum Swartz) 2
(-Hedysarum racemosum Aubl.) 2
D. laslocarpum (Beauv.) DC. - D. VELUTINUM
D. natalitum Sond. - D. GANGETICUM
D. paleaceum Gulll. & Perr. - D. SALICIFOLIUM
VAR. SALICIFOLIUM
D. REPANDUM (VAHL) DC.
3905000
350
450
1020
1350
1950
-VIGNA S A VI Revision: B.J. Pienaar (PRE).
#. PRE HERBARIUM PRACTICE, FOLLOWING
PIENAAJl .
V. decipiens Harv. - V. FRUTESCENS SUBSP.
FRUTESCENS VAR. FRUTESCENS
V. FRIESIORUM HARMS VAR. FRIESIORUM
V. FRUTESCENS A. RICH. SUBSP. FRUTESCENS VAR.
FRUTESCENS
( - V . decipiens Harv.)
(-V. longiloba Burtt Davy) 1, //
(-V. pongolensis Burtt Davy) 1, #
(-V. pseudotrlloba Harms) 1, #
V. huillensis Welw. ex Bak. - V. UNGUICULATA
SUBSP. DEKINDTIANA (HARMS.)
VERDC.
V. longiloba Burtt Davy - V. FRUTESCENS SUBSP.
FRUTESCENS VAR. FRUTESCENS
V. MONOPHYLLA TAUB .
V. pongolensis Burtt Davy - V. FRUTESCENS
SUBSP. FRUTESCENS VAR.
FRUTESCENS
V. pseudotrlloba Harms - V. FRUTESCENS SUBSP.
FRUTESCENS VAR. FRUTESCENS
V. PYMAEA R.E. FRIIS VAR. PYGMAEA
V. UNGUICULATA (L.) WALP. SUBSP. DEKINDTIANA
(HARMS) VERDC.
( - V . huillensis Welw. ex Bak.) 0
3910000 -DOLICHOS L.
3. HILLIARD & BURTT. 1988. NOTES R. BOT. GDN
EDINB. 45,1: 83-85.
150 D. ANGUST I SS IMUS E. MEY.
3910020 -MACROTYLOMA (WIGHT & ARN.) VERDC.
2. VERDCOURT. 1971. FTEA 4: 581-584.
500 M. UNIFLORUM (LAM.) VERDC. VAR. STENOCARPUM
.(BRENAN) VERDC.
GERANIACEAE Contributed by B.J. Pienaar 3924000
3926000
900
-SARCOCAULON (DC.) SWEET Revision: R.O.
Moffett (STE).
S. PATERSONII (DC.) G. DON
3928000
900
1 400
-PELARGONIUM L’HERIT.
20. NORDENSTAM. 1987. PL. SYST. EVOL. 155:
333-337.
21. VAN DER WALT & VAN ZYL. 1988. S. AFR. J.
BOT. 54,2: 145-171.
22. VAN DER WALT & VORSTER. 1988.
PELARGONIUMS OF SOUTHERN AFRICA
3: 1-149.
P. ANETHIFOLIUM (ECKL. & ZEYH.) STEUD.
P. ARIDUM R . A . DYER
288
Bothalia 19,2 (1989)
2 530
25 50
5000
5700
83 50
92 50
99 30
L 1350
11410
16030
(■P. dissectum (Eckl. & Zeyh.) Harv.) 22
P. CAESPITOSUM TURCZ. SUBSP. CAESPITOSUM
P. CAESPITOSUM TURCZ. SUBSP. CONCAVUM HUGO
P. DICHONDRIFOLIUM DC.
(■P. midd leton lanum Knath) 22
P. dissectum (Eckl. & Zeyh.) Harv. - P. ARIDUM
P. ELEGANS (ANDR.) WILLD.
(■P. ovale (Burn, f.) L'Herit. var.
ovat um Harv . ) 21
P. fumarloldes L'Herit. ex Harv. » P. MINIMUM
P. harveyanum Knuth - P. HYPOLEUCUM
P. HYPOLEUCUM TURCZ.
(-P. harveyanum Knuth) 22
P. LAXUM (SWEET) G. DON
P. ra idd le ton lanum Knuth ■ P. DICHONDRIFOLIUM
P. MINIMUM (CAV.) WILLD.
(-P. fumarloldes L'Herit. ex Harv.) 22
P. OVALE (BURM. F.) L'HERIT. SUBSP. HYALINUM
HUGO
P. ovale (Burra, f.) L'Herit. subsp. ovatum
Harv. - P. ELEGANS
P. OVALE (BURM. F.) L'HERIT. SUBSP.
VERONICIFOLIUM (ECKL. & ZEYH.)
HUGO
(-P. ver onlcae f ol lum (Eckl. & Zeyh.)
Steud.) 21
P. sldaefollura (Thunb.) Knuth - P. SIDOIDES
P. SIDOIDES DC.
(■P. sldaefollura (Thunb.) Knuth) 22
P. ve ro n ic ae f o 1 1 um (Eckl. & Zeyh.) Steud. - P.
OVALE SUBSP. VERONICIFOLIUM
RUTACEAE Contributed by M. Jordaan
and C . M . van Wyk
3986000
4037000 -AGATH0SMA WILLD. Revision: A. Bean (BOL).
#. PRE HERBARIUM PRACTICE, FOLLOWING BEAN.
11150 A. RUDOLPHII I. WILLIAMS
(■Acraadenia marlothll Duemmer) #
4040000 -ACMADENIA BART. & WENDL. F.
A. marlothll Duemmer - AGATH0SMA RUDOLPHII
4076000 - VE PR I S COMM. EX A. JUSS.
3. BOND & GOLDBLATT. 1-984. JL S. AFR. BOT.
13: 404.
150 V. LANCEOLATA (LAM.) G. DON
(■Toddalia lanceolata Lam.) 2, 3
(■V. undulata (Thunb.) Verdoorn & C.A.
Sm . ) 3
V. undulata (Thunb.) Verdoorn & C.A. Sm . - V.
LANCEOLATA
4077000 -TODDALIA JUSS.
T. lanceolata Lara. - VEPRIS LANCEOLATA
BURSERACEAE Contributed by M. Jordaan 4136000
4151000 -COMMIPHORA JACQ. Revision: J.J.A. van der
Walt (STE).
3. VAN DER WALT. 1986. FSA 18,3: 5-34.
MELIACEAE (PART A) Contributed by M. Jordaan 4155000
DICHAPETALACEAE Contributed by M. Jordaan 4283000
4285000 -TAPURA AUBL.
2. BRETELER. 1987. BELMONTIA 19: 56-60.
100 T. F ISCHERI ENGL.
(■T. flscherl Engl. var. pubescens
Verde. & Torre) 2
T. flscherl Engl. var. pubescens Verde. & Torre
- T. F ISCHERI
EUPHORBIACEAE
Contributed by M. Jordaan
4286000
-PSEUDOLACHNOSTYLIS PAX
2. RADCLIFFE-SMITH. 1987. FTEA EUPHORBIACEAE
1: 81.
P. deklndt 11 Pax - P. MAPROUNEIFOLI A
P. MAPROUNEIFOLIA PAX VAR. DEKINDTII (PAX)
RADCLIFFE-SM .
(-P. dekind til Pax) 2
P. MAPROUNEIFOLIA PAX VAR. MAPROUNEI POLIA
-PHYLLANTHUS L.
3. RADCLIFFE-SMITH. 1987. FTEA EUPHORBIACEAE
1 : 23.
#. PRE HERBARIUM PRACTICE, FOLLOWING
JORDAAN.
P. amapondena 1 s Sim - MARGARITARIA DISCOIDEA
VAR. FAGI FOLIA
P. ASPERULATUS HUTCH.
P. d iscoldeus (Balll.) Muell. Arg . -
MARGARITARIA DISCOIDEA VAR.
DISCOIDEA
P. flacourtloides Hutch. - MARGARITARIA
DISCOIDEA VAR. NITIDA
P. HETEROPHYLLUS E. MEY.
P. klrklanus Muell. Arg. - P. PINNATUS
P. PINNATUS (WIGHT) WEBSTER
(-P. kirkianus Muell. Arg.) 3
-MARGARITARIA L. F.
2. RADCLIFFE-SMITH. 1987. FTEA EUPHORBIACEAE
1: 67.
M. dlscoidea (Balll.) Webster subsp. discoldea
- M. DISCOIDEA VAR. DISCOIDEA
M. discoldea (Balll.) Webster subsp. nltlda
(Pax) Webster p.p. ■ M.
DISCOIDEA VAR. FAGIFOL IA
M. discoldea (Balll.) Webster subsp. nltlda
(Pax) Webster p.p. ■ M.
DISCOIDEA VAR. NITIDA
M. DISCOIDEA (BAILL.) WEBSTER VAR. DISCOIDEA
( -M . discoldea (Balll.) Webster subsp.
disco idea ) 2
(•Phyllanthus d iscoldeus (Balll.) Muell.
Arg.) 1
M. DISCOIDEA (BAILL.) WEBSTER VAR. FAGIFOLI A
(PAX) RADCLIFFE-SM.
( -M . discoldea (Balll.) Webster subsp.
nltlda (Pax) Webster p.p.) 2
( -Phyl lanthus amapondensls Sim) 1
M. DISCOIDEA (BAILL.) WEBSTER VAR. NITIDA (PAX)
RADCLIFFE-SM .
( “M . discoldea (Balll.) Webster subsp.
nltlda (Pax) Webster p.p.) 2
( -Phyllanthus flacourtloides Hutch.) 2
4155000 -CEDRELA P. BR.
2. WHITE & STYLES. 1986. FSA 18,3: 60.
4156000 -TOONA (ENDL.) M.J. ROEM.
2. WHITE & STYLES. 1986. FSA 18,3: 61.
MELIACEAE (PART B) Contributed by M. Jordaan 4159000
4171000 -TURRAEA L.
3. WHITE 6 STYLES. 1986. FSA 18: 45-47.
400 T. STREYI F. WHITE & STYLES
500 T. ZAMBESICA SPRAGUE & HUTCH.
MA LP IGH I ACEAE Contributed by M. Jordaan 4201000
4206000 -TRIASPIS BURCH.
250 T. HYPER ICO IDES (DC.) BURCH. SUBSP. NELSONII
( OL I V . ) IMMELMAN
( -T . thorncroftli Burtt Davy) 1
T. thorncroftli Burtt Davy - T. HYPERICOIDES
SUBSP. NELSONII
4219000 -S PHEDAMNOCARPUS PLANCH. EX BENTH. & HOOK. F.
3. DE VILLIERS & BOTHA. 1986. FSA 18: 66-69.
#. PRE HERBARIUM PRACTICE, FOLLOWING
JORDAAN .
600 S. TRANSVAALICUS (KUNTZE) BURTT DAVY
POLYGALACEAE Contributed by M. Jordaan 4273000
300
1550
24 50
6900
-POLYGALA L.
#*. PRE HERBARIUM PRACTICE, FOLLOWING
JORDAAN.
P. A LB I DA SCHINZ VAR. A LB I DA
P. EMPETRIFOLIA HOUTT.
(-P. teretlfolla Thunb.) ##
P. gagnebln lana Chod . - P. SPHENOPTERA
P. GRACILENTA BURTT DAVY
P. mural tloldes Eckl. & Zeyh. - MURALTlA
MURALTIOIDES
P. SPHENOPTERA FRESEN.
(■P. gagnebln lana Chod.) #
P. teretlfolla Thunb. - P. EMPETRIFOLIA
4278000 -MURALTlA JUSS.
2300 M. CHAMAEPITYS CHOD.
7400 M. MURALTIOIDES (ECKL. & ZEYH.) LEVYNS
(■Polygala muraltloldes Eckl. & Zeyh.) 1
4309000 -DRYPETES VAHL
2. RADCLIFFE-SMITH. 1987. FTEA EUPHORBIACEAE
• 1: 98.
#. PRE HERBARIUM PRACTICE, FOLLOWING
JORDAAN.
200 D. GERRARDI I HUTCH.
(■D. gerrardli Hutch, var. tomentosa
Radcllffe-Sm.) #
D. gerrardli Hutch, var. tomentosa Radcliffe-
Sm. - D. GERRARDI I
-ANTIDESMA L.
3. LEONARD. 1987. BULL. JARD. BOT. NAT.
BELG. 57: 453-454.
A. RUFESCENS TUL.
(■A. venosum auct. non E. Mey. ex Tul.)
A. venosum auct. non E. Mey. ex Tul
RUFESCENS
A.
4343000 -CLEISTANTHUS HOOK. F. EX PLANCH.
3. RADCLIFFE-SMITH. 1987. FTEA EUPHORBIACEAE
1: 133.
100 C. SCHLECHTERI (PAX) HUTCH. VAR. SCHLECHTER I
4345000 -BRI DELIA WILLD.
5. RADCLIFFE-SMITH. 1987. FTEA EUPHORBIACEAE
l: 125.
400
1100
-CROTON L.
4. RADCLIFFE-SMITH. 1987. FTEA EUPHORBIACEAE
1: 135-160.
C. MADANDENSIS S. MOORE
C. SYLVATICUS HOCHST.
-CAPERONIA ST. HIL.
3. RADCLIPPE-SMITH . 1987. FTEA EUPHORBIACEAE
1: 164.
-MICROCOCCA BENTH.
3. RADCLIFFE-SMITH. 1987. FTEA EUPHORBIACEAE
1: 260-264.
-ERYTHROCOCCA BENTH.
4. RADCLIPPE-SMITH. 1987. FTEA EUPHORBIACEAE
1: 271.
-ALCHORNEA SWARTZ
3. RADCLIFFE-SMITH. 1987. FTEA EUPHORBIACEAE
Bothalia 19,2 (1989)
289
800
2 550
1050
4416020
100
200
100
4498000
1: 251-260.
-AC A L Y PH \ L .
3. RADCLIFFE-SMITH. 1 98 7 . FTE A EUPHOR B I ACF.AE
1: 185-213.
A. FRUTICOSA FOKSSK. VAR. FRUT1COSA
A. senensls Klotzsch - A. VILLICAULIS
A. VI LLICAULIS A. RICK.
(■A. senensls Klotzsch) 3
-TRAGI A L.
3. RADCLIFFE-SMITK. 1987. KEW BULL. 42,2:
396-397 .
T. d urbane ns 1 s Kuntze ■ T. GLABRATA VAR.
GLABRATA
T. GLABRATA (MUELL. ARG . ) PAX & HOFFM. VAR.
GLABRATA
( ■ T . durbanensls Kuntze) 3
T. natalensls Sond. - TRAGIELLA NATALENSIS
T. PRIONOIDES RADCLIFFE-SM .
-TRAGIELLA PAX & K . HOFFM.
1. RADCLIFPE-SMITH. 1987. FTEA E UP KOR B I AC E A E
1: 320.
T. NATALENSIS (SOND.) PAX & K. HOFFM.
(aSphaerostyll9 natalensls (Sond.)
Croizat) 1
(■Tragla natalensls Sond.) 1
-S phae r o 6 t y 1 1 s Baill. Southern African taxa
moved to TRAGIELLA
S. natalensls (Sond.) Croizat - TRAGIELLA
NATALENSIS
-DALECHAMPI A L.
3. RADCLIFFE-SMITH. 1987. FTEA EUPKORBIACEAE
1: 287.
D. CAPENSIS SPRENG. F.
(-D. klrkll Praln) 3
D. klrkll Praln - D. CAPENSIS
D. SCANDENS L. VAR. CORDOFANA (WEBB) MUELL.
ARG .
-ALEURITES FORST.
A. ford 1 1 Hem s 1 . * - VERNICIA FORDII
A. raontana (Lour.) Wilson * - VERNICIA MONTANA
-VERNICIA LOUR.
1. RADCLIFFE-SMITH. 1987. FTEA EUPHORB I AC EA £
1 : 181 .
V. FORDII ( HEMS L . ) AIRY SHAW *
(■Aleurites fordll Herasl.) 1
V. MONTANA LOUR. *
(■Aleurites raontana (Lour.) Wilson) 1
-JATROPHA L.
2. RADCLIFFE-SMITH. 198 7 . FTEA E UP HOR B I ACE A E
1: 343-365.
J. raessinica E . A . Bruce = J. SPICATA
J. SPICATA PAX
(*J. raessinica E.A. Bruce) 2
-MANIHOT MILL.
2. RADCLIFFE-SMITH. 1987. FTEA E UP HO R B l AC E A £
1: 367.
M . ESCULENTA CRANTZ *
(■M. utilisslma Pohl *) 2
M. utilisslma Pohl * = IM . ESCULENTA *
-CLUTIA L. Revision: D. Koutnlk (BOL) & G.L.
Webs ter ( DAV) .
4. RADCLIFFE-SMITH. 1987. FTEA EUPKORBIACEAE
1: 332-339.
-SUREGADA ROXB. EX ROTTL.
2. RADCLIFFE-SMITH. 1987. FTEA EUPKORBIACEAE
1: 376-377.
-SAPIUM P.BR.
4. RADCLIFFE-SMITK. 1987. FTEA EUPKORBIACEAE
1: 390.
S. ELLIPTIC UM (KRAUSS) PAX
ANACARDIACEAE
Contributed by M. Jordaan
BOT.
52: 369-371.
BOT. 52,4:
BOT. 54,5:
BOT. 54,6:
369-
501-
5 34-
1150
2150
6150
10200
102 26
19850
239 50
27150
-EUPHORBIA L.
7. LEACH. 1986. S. AFR
8. LEACH. 1986. S. AFR
371 .
9. LEACH. 1986. S. AFR
503 .
10. LEACH. 1986. S. AFR
540.
PRE HERBARIUM PRACTICE, FOLLOWING
LEISTNER & JORDAAN.
E. ANOPLIA STAPF
E. BAYER I LEACH
E. CYATHOPHORA J. MURR. *
(-Poinsettia cyathophora (J. Murr.)
Bartl. *) //#
E. GENICULATA ORTEG. *
(■Poinsettia genlculata (Orteg.)
Klotzsch & Garcke) //// *
6. GENTILIS N.E. BR. SUBSP. GENTILIS
E. GENTILIS N.E. BR. SUBSP. TANQUANA LEACH
E. PEDEMONTANA LEACH
E. SILIC IICOLA DINTER
E. VAALPUTS I ANA LEACH
-POINSETTIA R. GRAH.
n. PRE HERBARIUM PRACTICE, FOLLOWING
LEISTNER & JORDA'AN.
P. cyathophora (J. Murr.) Bartl. * ■ EUPHORBIA
CYATHOPHORA
P. genlculata (Orteg.) Small * ■ EUPHORBIA
GENICULATA
4563000
350
100
600
900
3800
-LANNEA A. RICH.
L. SCHIMPERI (HOCHST. EX A. RICH.) ENCL. VAR.
SCHIMPERI
-RHUS L.
4. MOFFETT. 1988. S. APR. J. BOT. 54,2: 172.
R. ACOCKSII MOPPETT
R. BOLUS I I SOND. EX ENGL.
R. CILIATA LICHT. EX SCHULT.
R. LAEVIGATA L.
(-R. laevigata L. var. atomarla (Jacq.)
R . Pe rnande a ) #
(-R. laevigata L. var. latlfolla
(Schonl.) R. Pernandes) #
(-R. laevigata L. var. vllloaa (L. f.)
R. Fernandes) #
R. laevigata L. var. atomarla (Jacq.) R.
Fernandes - R. LAEVIGATA
R. laevigata L. var. latlfolla (Schonl. )R.
Pernandes - R. LAEVIGATA
R. laevigata L. var. vlllosa (L. f.) R.
Pernandes - R. LAEVIGATA
R. longl9plna sensu Schonl. p.p. non Eckl. A
Zeyh - R. PTEROTA
R. PTEROTA PRESL
(-R. long lsplna sensu Schonl. p.p. non
Eckl. & Zeyh) 9
CELASTRACEAE
Contributed by M. Jordaan
4618000
100
4627000
150
4630000
-Celastrus L. Southern African taxa placed In
MAYTENUS and PUTTERLICKIA
C. cordatus E. Mey. ex Sond. - MAYTENUS
CORDATA
C. procurabens L. f. - MAYTENUS PROCUMBENS
-MAYTENUS MOLINA
#. PRE HERBARIUM PRACTICE, FOLLOWING
JORDAAN.
M. CORDATA (E. MEY. EX SOND.) LOES.
(■Celastrus cordatus E. Mey. ex Sond.) #
( -Gymnos por la cordata (E. Mey. ex Sond.)
Sim) #
M. PROCUMBENS (L. F.) LOES.
(■Celastrus procurabens L. f.) 1
( -Gymno 8 por 1 a procurabens (L. f.) Loes.)
1
-EMPLEUR I DIUM SOND.
1. GOLDBLATT, TOBE ET AL. 1985. ANN.
MISSOURI BOT. GDN. 72: 167-
183.
E. JUNIPERINUM SOND.
-Gymnospor la (Wight & Arn.) Hook. f. ■ MAYTENUS
G. cordata (E. Mey. ex Sond.) Sim - MAYTENUS
CORDATA
G. procurabens (L. f.) Loes. - MAYTENUS
PROCUMBENS
-PUTTERLICKIA ENDL. Revision: A . E . van Wyk
(PRU) .
1. DAVIDSON. 1927. BOTHALIA 2: 336-338.
3. VAN WYK & MOSTERT. 1987. S. AFR. J. BOT.
53,4: 267-270.
P. RETROSPINOSA VAN WYK & MOSTERT
-PTEROCELASTRUS MEISN.
1. DAVISON. 1927. BOTHALIA 2: 321-326.
SAPINDACEAE
1927. BOTHALIA 2:
Contributed by M. Jordaan
326-336.
4723000
4726000 -CARDIOSPERMUM L.
200 C. GRANDIFLORUM SWARTZ
300 C. HALICACABUM L. VAR. HALICACABUM
350 C. HALICACABUM L. VAR. MICROCARPUM (KUNTH)
BLUME
( -C . mlcrocarpum Kunth) 1
C. raicrocarpum Kunth ■ C. HALICACABUM VAR.
MICROCARPUM
MELIANTHACEAE Contributed by M. Jordaan
4853000 -BERSAMA FRESEN. Revision: P.D.F.
TILIACEAE Contributed by M. Jordaan
4975000
-TRIUMFETTA L.
1. WILD. 1984 . F S A 2 1 , 1 : 20-28.
MALVACEAE
Contributed by M. Jordaan
4853000
Kok (PRU).
4937000
4980000
5020000
100
-LAVATERA L.
L. ASSURGENTIFLORA KELLOGG *
(Garden escape collected in the Cape.
2916 (PORT NOLLOTH): Port Nolloth (-BD)
Zletsman 821)
-CIENFUEGOSIA CAV.
C. anoraala (Wawra ex Wawra & Peyr.) Guerke -
GOSSYPIUM ANOMALUM
C. trlphylla (Harv.) K. Schum. ■ GOSSYPIUM
TRIPHYLLUM
-GOSSYPIUM L.
3. VOLLESEN. 1987. KEW BULL. 42,2: 337-349.
G. ANOMALUM WAWRA EX WAWRA & PEYR. SUBSP.
ANOMALUM
290
Bothalia 19,2 (1989)
( -Cienf uegosia anomala (Wawra ex Wawra &
Pey r . ) Guerke ) 2
G. anomalum Wawra ex Wawra & Peyr. subsp .
triphyllum (Harv. ) Roberty -
G. TRIPHYLLUM
200 G. HERBACEUM L. SUBSP. APRICANUM (WATT)
VOLLESEN
( -G . herbaceum L. var . africanura (Watt)
Hutch . & Ghose ) 3
(-G. ob t us t f o 1 ium Roxb. ex G. Don var.
afrlcanun Watt) 1
(-G. transvaalenae Watt) 3
G. herbaceura L. var. afrlcanun (watt) Hutch. &
Ghose - G. HERBACEUM SUBSP.
AFRICANUM
G. obtu8l follura Roxb. ex G. Don var. afrlcanun
Watt - G. HERBACEUM SUBSP.
AFRICANUM
G. transvaalense Watt - G. HERBACEUM SUBSP.
AFRICANUM
( -Cienf uegosia trlphylla (Harv.) K.
Schum . ) 2
(-G. anomalum Wawra ex Wawra & Peyr.
subsp. triphyllum (Harv.)
Roberty) 3
STERCULIACEAE Contributed by K. Imraelman 5044000
5056000
8 5 50
1 3300
15100
1 7 700
2 3200
-HERMANNIA L. Revision: B. de Winter (PRE).
H. abyssinica (Harv.) K. Schum. - H.
QUARTI NI ANA SUBSP. STELLULATA
H. adenotricha K. Schum. - H. QUART IN IANA
SUBSP. STELLULATA
H. ERODIO IDES (BURCH. EX DC.) KUNTZE
(»H. resedifolia (Burch.) R.A. Dyer) #
(-Mahernia resedaefolia Burch.) #
H. ex 1 1 1 8 Bur 1 1 Davy - H. QUARTINI ANA SUBSP.
STELLULATA
H. geraini flora Dinter & Engl. - H. QUARTINIANA
SUBSP. STELLULATA
H. HETEROPHYLLA (CAV.) THUNB.
(-H. huralf usa Hochr . ) #
(-H. humili9 Thunb.') #
H. humifusa Hochr. - H. HETEROPHYLLA
H. humilis Thunb. - H. HETEROPHYLLA
H. JAC0BEIF0LIA (TURCZ.,) R-A- OYER
H. MARGINATA (TURCZ.) PILLANS
H. QUARTINIANA A. RICH. SUBSP. STELLULATA (K.
SCHUM.) DE WINTER
(-H. abyssinica (Harv.) K. Schum.) 1
(-H. adenotricha K. Schum.) #
( - H . exilis Burtt Davy) #
( -H . geminiflora Dinter & Engl.) 1
(-H. schinzii K. Schum.) 1
(-H. sinuata Burtt Davy) #
(-H. stellulata (Harv.) K. Schum.) #
(-Mahernia abyssinica Hochst. ex Harv.)
1, #
(-Mahernia stellulata Harv.) #
H. resedifolia (Burch.) R.A. Dyer - H.
ERODIO IDES
H. schinzii K. Schum. - H. QUARTINIANA SUBSP.
STELLULATA
H. sinuata Burtt Davy - H. QUARTINIANA SUBSP.
STELLULATA
H. stellulata (Harv.) K. Schum. - H.
QUARTINIANA SUBSP. STELLULATA
5056010 -Mahernia Schum. - HERMANNIA
M. abyssinica Hochst. ex Harv. - HERMANNIA
QUARTINIANA SUBSP. STELLULATA
M. resedaefolia Burch. - HERMANNIA ER0DI0IDES
M. stellulata Harv. - HERMANNIA QUARTINIANA
SUBSP. STELLULATA
BEG0NIACEAE Contributed by C.M. van Wyk 5396000
5486000 -NESAEA COMM. EX JUSS.
100 N. ANAGA LL0 IDES KOEHNE
(-N. loandenals auct. non (Welw. ex
Hiern) Koehne) #
500 N. DINTERI KOEHNE
(-N. transvaallca Fernandes) #
N. f lorlbunda Sond. - N. RADICANS VAR.
FLORIBUNDA
N. kuntzei Koehne - N. SCHINZII
N. loandenals auct. non (Welw. ex Hiern) Koehne
- N. ANAGALLOIDES
1100 N. 0ND0NGANA KOEHNE
(-N. ondonga subsp. ondonga var.
evansiana (A. Fernandes &
Diniz) A. Fernandes) #
N. ondonga subsp. ondonga var. evansiana (A.
Fernandes & Diniz) A. Fernandes
- N. ONDONGA
1190 N. RADICANS GUILL. A PERR. VAR. FLORIBUNDA
(SOND.) A. FERNANDES
( -N . florlbunda Sond.) #
1195 N. RADICANS GUILL. & PERR. VAR. RADICANS
N. rivulare (Wood & Evans) Koehne - N.
SCHINZII
1800 N. SARCOPHYLLA (WELW. EX HIERN) KOEHNE
1900 N. SCHINZII KOEHNE
( -N . kuntzei Koehne) #
(■N. rivulare (Wood & Evans) Koehne) #
N. transvaallca Fernandes - N. DINTERI
RHYNCHOCALYCACEAE Contributed by K. Immelman 5495000
5495000 -RHYNCHOCALYX OLIV.
2. TOBE & RAVEN. 1984. ANN. MISSOURI B0T.
GDN 41: 836-843.
COMBRETACEAE Contributed by C.M. van Wyk 5536000
5538000 -COMBRETUM LOEFL. Revision: E.F. Hennessy (U.
D-W. ) .
600 C. CELASTROI DES WELW. EX LAWS.
ONAGRACEAE Contributed by C.M. van Wyk
5819000 -GAURA L.
200 G. LINDHEIMERI ENGELM. & GRAY *
ARALI ACEAE Contributed by C.M. van Wyk
5872010 -SEEMANNARALIA VIG.
100 S. GERRARDII (SEEMANN) HARMS
APIACEAE Contributed by C.M. van Wyk
5918000 -SANICULA L.
100 S. ELATA BUCH.-HAM. EX D. DON
5970000 -CONI UM L.
100 C. CHAER0PHYLL0IDES (THUNB.) SOND.
5791000
5839000
5893000
5994000 -BUPLEURUM L.
B. difforme L. - ANGINON DIFF0RMIS
5994010 -ANGINON RAF.
1. BURTT. 1988. NOTES R. BOT. GDN EDINB.
45,1: 90-91.
100 A. DIFFORME (L.) B.L. BURTT
(-Bupleurum difforme L.) 1
( -Rhy t icarpus difforrais (L.) Briq.) I
200 A. RUGOSUM (THUNB.) RAF.
( -Rhyt icar pus rugosus (Thunb.) Sond.) 1
300 A. SWELLENDAMENSIS (ECKL. & ZEYH.) B.L. BURTT
( -Rhy t icar pus ecklonls Sond.) 1
( - Rhy t i c a r pus swe l 1 e nd amen s l s (Eckl. &
Zeyh . ) Briq.) 1
5397000 -BEGONIA L.
1300 B. S UTHERLANDI I HOOK. F.
THYMELAEACEAE Contributed by C.M. van Wyk 5429000
320
2100
2700
5950
-GNIDIA L.
G. CAFFRA ( MEISN . ) GILG
G. GYMNOSTACHYA (C.A. MEY.) GILG
G. hoepfneriana Gilg - G. KRAUSSIANA VAR.
KRA US S I ANA
G. KRAUSSIANA MEISN. VAR. KRAUSSIANA
(»G. hoepfneriana Gilg.) 14
( -La s l o s 1 phon hoe p f ne r i a n us Vatke ex
Gilg) 5, 14
G. SIMILIS C.H. WR.
5996000 -Rhyticarpus Sond. - ANGINON
R. difformis (L.) Briq. - ANGINON DIFFORME
R. ecklonis Sond. - ANGINON SWELLENDAMENSIS
R. rugosus Sond. - ANGINON RUGOSUM
R. s we 1 1 e nd arae n s i s (Eckl. & Zeyh.) Briq. -
ANGINON SWELLENDAMENSIS
6013000 -DEVERRA DC.
300 D. DENUDATA (VIV.) PFISTERER 6 POOL. SUBSP.
APHYLLA (CHAM. & SCHLECHTD.)
PFISTERER & P0DL.
6046010 -Thunbe rg 1 e 1 la Wolff - ITASINA
T. filiformis ( Ko so-Po 1 Jansky ) Wolff - ITASINA
FILIF0LIA
5435010 -Lasiosiphon Fresen. - GNIDIA
L. hoe p f ne r 1 an u s Vatke ex Gilg - GNIDIA
KRAUSSIANA VAR. KRAUSSIANA
LYTHRACEAE Contributed by K. Immelman 5473000
6046020 -ITASINA RAF.
1. BURTT. 1988. NOTES R. BOT. GDN EDINB.
45,1: 93-94.
100 I. FILIFOLIA (THUNB.) RAF.
(-Thunberg iel la filiformis (Koso-
Poljansky) Wolff) 1
5474000 -AMMANNIA L.
#. PRE HERBARIUM PRACTICE, FOLLOWING
IMMELMAN.
200 A. BACCIFERA L.
(-A. wormskloldii Fisch. & C.A. Mey.) #
(-A. wormskloldii Fisch. & C.A. Mey.
var. alata Koehne) #
400 A. SENEGALENSIS LAM. EX POIR.
A. wormskloldii Fisch. & C.A. Mey. - A.
BACCIFERA
A. wormskloldia Fisch. & C.A. Mey. var. alata
Koehne - A. BACCIFERA
6054000 -DIPL0L0PHI UM TURCZ.
6062000 -FOENICULUM MILL.
F. VULGARE MILL. *
6069000 -He t er o p t i 1 i s E. Mey. ex Meisn. - DASISPERMUM
H. suf frut lcosa (Berg.) Leute - DASISPERMUM
SUFFRUTICOSUM
6069010 -Cnldium Cusson - DASISPERMUM
C. s uf f r u t 1 c o s um Cham. & Schlechtd. -
DASISPERMUM SUFFRUTICOSUM
5476000 -LYTHRUM L.
2. S0NDER. 1894. PC 2: 516.
6069020
-DASISPERMUM RAF.
Bothalia 19,2 (1989)
291
1. BURTT. 1988. NOTES R. BOX. GDN EDINB.
45,1: 92-93.
100 D. SUFFRUTICOSUM (BERG.) B.L. BURTT
(■Cnidlura 8uf f rut icosara Cham. &
Sc h lech td . ) 1
( -He te ropt 11 is suffruticosa (Berg.)
Le u t e ) 1
( -Aua t roo lmusops dispar (N.E. Br.) A.
Meeuee ) 1
200 V. MARGINATA (N.E. BR.) AUBREV.
(■Au8troilau8op8 marglnata (N.E. Br.) A.
Meeuee) 1
LOGANIACEAE Contributed by M. Jordaan 6447000
6078000 -ANNES0RRHIZA CHAM. 6 SCHLECHTD.
7. BURTT. 1988. NOTES R. BOT. GDN EDINB.
45,1: 92 .
A. gummifera (L.) Jackson - A. INEBRIANS
650 A. INEBRIANS (THUNB.) WIJNANDS
( “ A . guramlfera (L.) Jackson) 6
(-Glia gummifera (L.) Sond . ) 6
6078010 -Glia Sond. - ANNES0RRHIZA
G. gummifera (L.) Sond. - ANNESORRHIZA
INEBRIANS
6116000 -PEUCEDANUM L.
3. BENTHAM & HOOKER. 1867. GEN 1,3: 902.
5. DRUDE. 1898. PFLANZENFAM . 3,8: 237.
7. PHILLIPS. 1917. ANN. S. AFR. MUS. 16:
108 .
14. BURTT. 1988. NOTES R. BOT. GDN EDINB.
45,1: 94.
3100 P. UPINGTONIAE (SCHINZ) DRUDE
(-Lefebvrea uplngtoniae Schinz) 5*
6116020 -Lefebvrea A. Rich. - PEUCEDANUM
L. uplngtoniae Schinz - PEUCEDANUM UPINGTONIAE
ERICACEAE Contributed by C.M. van Wyk 6179000
6237000
1 940
1 9780
20650
21530
21570
2 1860
21940
40320
44740
50350
53140
59070
64840
-ERICA L.
28. OLIVER. 1 987 . S. AFR. J. BOT. 53,6: 455-
458 .
E. accoramoda t a Klotz9ch ex Benth. var.
ebracteata H. Bol. - E.
LASC IVA
E. ALTIPHILA E.G.H. OLIVER
(-Philippia altlcola E.G.H. Oliver) 28
(-Philippia trl9tis H. Bol.) 28
E. DRACOMONTANA E.G.H, OLIVER
(-Philippia drakensbergensis E.G.H.
Oliver) 28
E. ELSIEANA (E.G.H. OLIVER) E.G.H. OLIVER
(-Philippia elsieana E.G.H. Oliver) 28
E. ESTER I ANA E.G.H. OLIVER SUBSP. ESTERIANA
(-Philippia esterhuyseniae E.G.H. Oliver
subsp. esterhuyseniae) 28
E. ESTERIANA E.G.H. OLIVER SUBSP.
SWARTBERGENSIS (E.G.H. OLIVER)
E.G.H. OLIVER
(-Philippia esterhuyseniae E.G.H. Oliver
sub9p. 8 wa r t be r g en a i s E.G.H.
Oliver) 28
E. EVANS 1 1 (N.E. BR.) E.G.H. OLIVER
(-Philippia evansli N.E. Br.) 28
E. EXLEEANA E.G.H. OLIVER
(-Philippia leeana Klotzsch) 28
(-E. accommodata Klotzsch var.
ebracteata H. Bol.) 28
(-Philippia stokoei L. Guthrie) 28
E. MADIDA E.G.H. OLIVER
(-Philippia irrorata E.G.H. Oliver) 28
E. NOTHOLEEANA (E.G.H. OLIVER) E.G.H. OLIVER
(-Philippia notholeeana E.G.H. Oliver)
28
(-Philippia pallida L. Guthrie) 28
E. PETRICOLA E.G.H. OLIVER
(-Philippia petrophila E.G.H. Oliver) 28
E. PROCA VI ANA (E.G.H. OLIVER) E.G.H. OLIVER
(-Philippia procaviana E.G.H. Oliver) 28
E. SIMII (S. MOORE) E.G.H. OLIVER
(-Philippia sirnii S. Moore) 28
E. TRISTIS BARTL.
(-Philippia absinthoides (Thunb.) E.G.H.
Oliver) 28
(-Philippia chamis8onis Klotzsch) 28
6240000 -Philippia Klotzsch - ERICA
P. absinthoides (Thunb.) E.G.H. Oliver - ERICA
TRISTIS
P. altlcola E.G.H. Oliver - ERICA ALTIPHILA
P. chamlssonis Klotzsch » ERICA TRISTIS
P. drakensbergensls E.G.H. Oliver - ERICA
DRACOMONTANA
P. elsieana E.G.H. Oliver - ERICA ELSIEANA
P. esterhuyseniae E.G.H. Oliver subsp.
esterhuyseniae - ERICA
ESTERIANA SUBSP. ESTERIANA
P. esterhuyseniae E.G.H. Oliver subsp.
swar tbe rgens is E.G.H. Oliver -
ERICA ESTERIANA SUBSP.
SWARTBERGENSIS
P. evansli N.E. Br . - ERICA EVANSII
P. irrorata E.G.H. Oliver - ERICA MADIDA
P. leeana Klotzsch - ERICA EXLEEANA
P. notholeeana E.G.H. Oliver - ERICA
NOTHOLEEANA
P. pallida L. Guthrie - ERICA PELTATA
P. petrophila E.G.H. Oliver - ERICA PETRICOLA
P. procaviana E.G.H. Oliver - ERICA PROCAVIANA
P. slmll S. Moore ■ ERICA SIMII
P. stokoei L. Guthrie - ERICA ACCOMMODATA VAR.
EBRACTEATA
P. trlstis H. Bol. - ERICA CAESPITOSA
SAPOTACEAE Contributed by
Wyk 6353000
6386020 -VITELLARIOPSIS ( BAILL . ) DUBARD
100 V. DISPAR (N.E. BR.) AUBREV.
6460000 -STRYCHNOS L.
S. lnnocua Del. subsp. dysophylla (Benth.)
Verdoorn - S. MADAGASCAR I ENS IS
S. lnnocua Del. subsp. gerrardll (N.E. Br . )
Verdoorn - S. MADAGASCAR I ENS I S
400 S. MADAGASCAR I ENS IS POIR.
(-S. lnnocua Del. subsp. dysophylla
(Benth.) Verdoorn) 2
(-S. lnnocua Del. subsp. gerrardll (N.E.
Br . ) Verdoorn) 2
APOCYNACEAE Contributed by C.M. van Wyk 6549000
6680000 -ADENIUM ROEM. 6 SCHULT.
(-A. obesura (Forssk. ) Roem. & Schult.
var. multlflorum (Klotzsch)
Codd) 2
A. obeauni (Forssk.) Roem. & Schult. var.
multlflorum (Klotzsch) Codd -
A. MULTIFLORUm
6688000 -STROPHANTHUS DC.
2 BEENTJE. 1982. BELMONTIA 13: 17-163.
PER I PLOCAC EAE Contributed by C.M. van Wyk 6729000
6740000 -CRYPTOLEPIS R. BR.
150 C. DELAGOENSIS SCHLTR .
6747000 -RAPH IONACME HARV. H.J.T Venter ( BLF-U ) .
8. VENTER 6 VERHOEVEN. 1988. S. AFR. J. BOT.
54,6: 603-606.
9. VENTER & VERHOEVEN. 1988. S. APR. J. BOT.
54,6: 607-610.
840 R. LOBULATA VENTER & VERHOEVEN
870 R. LUCENS VENTER & VERHOEVEN
A SC LE P I ADAC E AE Contributed by M. Jordaan 6752000
and C.M. van Wyk
6787010
400
1400
2400
2900
3000
-PACHYCARPUS E. MEY.
3. SMITH. 1988. JL . S. AFR. BOT. 54,5: 399-
439 .
(-P. inconetana N.E. Br . ) 3
(-P. valldus (Schltr.) N.E. Br . ) 3
P. CAMPANULATUS (HARV;) N.E. BR. VAR.
SUTHERLANDI I N.E. BR.
(-P. gerrardll (Harv.) N.E. Br . ) 3
P. gerrardll (Harv.) N.E. Br. - P.
CAMPANULATUS VAR. SUTHERLANDI I
P. grandif lorus (L. f.) E. Mey. var.
chrysanthus N.E. Br. - P.
GRANDIFLORUS VAR. GRANDIFLORUS
P. grandif lorus (L. f.) E. Mey. var.
e 1 a toe a r 1 na t us N.E. Br . - P.
GRANDIFLORUS VAR. GRANDIFLORUS
elatocarlnatus N.E. Br . ) 3
P. GRANDIFLORUS (L. F.) E. MEY. VAR. TOMENTOSUS
(SCHLTR.) N.E. BR.
P. inconstans N.E. Br . - P. ASPERIFOLIUS
P. insignis (Schltr.) N.E. Br . - P.
TRANS VAALENSIS
P. RIGIDUS E. MEY.
P. rigidu8 E. Mey. var. tridens E. Mey. - P.
RIGIDUS
P. STENOGLOSSUS (E. MEY.) N.E. BR.
P. TRANSVAALENSIS (SCHLTR.) N.E. BR.
(-P. insignis (Schltr.) N.E. Br.) 3
P. valldus (Schltr.) N.E. Br . - P.
ASPERIFOLIUS
6789000 -STENOSTELMA SCHLTR.
(Note change in genus number)
6791QOO
1150
3350
4400
-ASCLEPIAS L. Revision: A. Nicholas (PRE).
5. HILLIARD & BURTT. 1986. NOTES R. BOT. GDN
EDINB. 43,2: 192-193.
6. NICHOLAS. 1987. BOTHALIA 17,1: 17-23.
7. NICHOLAS. 1987. BOTHALIA 17,1: 29-43.
A. COMPRESSIDENS (N.E. BR.) A. NICHOLAS
( - A . navicularis (E. Mey.) Schltr. var.
coopressldens N.E. Br . ) 6
A. GORDON-GRAYAE A. NICHOLAS
A. NAVICULARIS (E. MEY.) SCHLTR.
A. navicularis (E. Mey.) Schltr. var.
compressidens N.E. Br . - A.
COMPRESSIDENS
6870000 -BRACHYSTELMA R. BR.
3. HILLIARD & BURTT. 1988. NOTES R. BOT. GDN
EDINB. 45,1: 77.
4. BALKWILL & BALKWILL. 1988. JL S. AFR.
BOT. 54,1: 60-62.
1 77 5 B. DYER I K. BALKWILL & M.-J. BALKWILL
6874000 -C EROPEGI A L.
C. flanaganii (Schltr.) Huber var. alexandrlna
Huber - RIOCREUXIA ALEXANDRINA
C. flanaganii (Schltr.) Huber var. fallax Huber
- RIOCREUXIA WOODII
C. flanaganii (Schltr.) Huber var. flanaganii -
RIOCREUXIA FLANAGANII
292
Bothalia 19,2 (1989)
6875000
150
250
320
700
-RIOCREUXIA DECNE.
2. DYER. 1983. CEROPEGIA, BRACHYSTELMA &
RIOCREUXIA IN S. AFR. 233.
R. ALEXANDRINA (HUBER) R . A . DYER
(-Ceropegia flanaganil (Schltr.) Huber
var. alexandrina Huber) 1, 2
(-R. flanaganil Schltr. subsp.
alexandrina (Huber) R.A. Dyer)
2
R. BURCHE LLI I K. SCHUM.
(-R. flanaganil Schltr. subsp. segregata
R.A. Dyer) 2
R. flanaganil Schltr. subsp. alexandrina
(Huber) R.A. Dyer - R.
ALEXANDRINA
R. FLANAGAN 1 1 SCHLTR.
R. flanaganil Schltr. subsp. segregata R.A.
Dyer - R. BURC HE LLI I
R. flanaganil Schltr. subsp. voodii (N.E. Br . )
R.A. Dyer - R. WOODII
R. WOODII N.E. BR.
(-Ceropegia flanaganil (Schltr.) Huber
var. fallax Huber) 1, 2
( -R . flanaganil Schltr. subsp. woodii
(N.E. Br.) R.A. Dyer) 2
6885070
200
-ORBEOPSIS LEACH
0. CAUDATA (N.E
Revision: L.C. Leach ( NBG ) .
BR.) LEACH SUBSP. RHODES IACA
(LEACH) LEACH
6885100 -TROMOTRICHE HAW. Revision: L.C. Leach (NBG).
2. LEACH. 1984. JL. S. AFR. BOT. 50,4: 549-
562.
BORAGINACEAE Contributed by W.G. Welman 7038000
7043000
100
150
200
1. BAKER & WRIGHT. 1905. FTA 4,2: 25.
2. FRIEDRICH-HOLZHAMMER. 1967. FSWA 119: 2-
3.
E. AMOENA KLOTZSCH
E. hottentotlca Burch. - E. RIGIDA
E. OBTUSIFOLIA HOCHST. EX DC. •
E. RIGIDA (THUNB.) DRUCE
(-Capraria rigida Thunb.) 2
( - E . hottentotlca Burch.) 2
H. macro phylla Thunb. - DIASCIA MACROPHYLLA
H. MONTANA L. F.
(-H. pachyceras Diels) 3
(-H. racemosa (Houtt.) Merrill) 3
H. nana Diels - H. CENTRODES
H. pachyceras Diels - H. MONTANA
H. racemosa (Houtt.) Merrill - H. MONTANA
H. SABULOSA L. F.
H. unilabiata Thunb. - DIASCIA UNI LAB I ATA
7521000
60
83
88
96
97
105
no
112
113
1 15
117
203
210
220
PH YLLOPODIUM BENTH.
4. HILLIARD. 1989.
45,3:
P. ANOMALUM HILLIARD
NOTES R. BOT. GDN EDINB.
481-491 .
P .
P.
P.
P .
P.
P .
P.
P.
P.
P .
P.
P .
P .
P.
P .
P.
P.
P .
P.
(-Polycarena plantaginea auct.) 4
CAESPITOSUM HILLIARD
calvum Hiern - MELANOS PERMUM TRANS VAALE NS E
COLLINUM (HIERN) HILLIARD *
(-Polycarena collina Hiern) 4
(-Polycarena pubescens Benth. p.p.) 4
DOLOMITICUM HILLIARD
ELEGANS ( CHOIS Y ) HILLIARD
(-P. 1 Inear If o lium H. Bol.) 4
(-Polycarena linearifolia (H. Bol.)
Levyns) 4
(-Selago elegans Choisy) 4
glutlnosum Schltr. - TRIEENEA GLUTINOSA
krebsianum Benth. - GLEKIA KREBS IANA
linearif ol lure H. Bol. - P. ELEGANS
LUPULIFORME (THELL.) HILLIARD
(-Polycarena lupuliformis Thell.) 4
MAX I I (HIERN) HILLIARD
(-Polycarena maxii Hiern) 4
MICRANTHUM (SCHLTR.) HILLIARD
(-Polycarena rariflora Benth. var.
micrantha Schltr.) 4
MIMETES HILLIARD
NAMAENSE (THELL.) HILLIARD
(-Polycarena namaensis Thell.) 4
PUBIFLORUM HILLIARD
rupestre Hiern - MELANOS PERMUM RUPESTRE
RUSTII (ROLFE) HILLIARD
(-Selago rustii Rolfe) 4
schlechteri Hiern - TRIEENEA SCHLECHTERI
TWEEDENSE HILLIARD
VISC IDISSIMUM HILLIARD
VERBENACEAE
Contributed by W.G. Welman
7192000 -HOLMSKIOLDIA RETZ.
H. speciosa Hutch. & Corbishley - KAROMIA
S PEC I OS A
H. tettensis auett. non (Klotzsch) Vatke -
KAROMIA SPECIOSA
7192010 -KAROMIA P. DOP
1. FERNANDES. 1985. GARCIA DE ORTA 7(1-2):
33-46.
100 K. SPECIOSA (HUTCH. & CORBISHLEY) R. FERNANDES
(-Holmskioldia speciosa Hutch. &
Corbishley) 1
(-Holmskioldia tettensis auett. non
(Klotzsch) Vatke) 1
LAMIACEAE Contributed by W.G. Welman 7210000
7359000 -SYNCOLOSTEMON E. MEY. EX BENTH.
2. CODD. 1988. BOTHALIA 18,1: 92-93.
875 S. RAMULOSUS E. MEY. EX BENTH.
7366000 -OCIMUM L.
0. fissilabrum Briq. - BECIUM FILAMENTOSUM
0. knyanum Vatke - BECIUM FILAMENTOSUM
0. rautanenii Briq. - BECIUM FILAMENTOSUM
D. s tenog loss urn Briq. - BECIUM FILAMENTOSUM
7366010 -BECIUM LINDL.
2. SEBALD. 1988. STUTTGARTER BEITR . NATURK.
SER. A, 419: 51.
250 B. FILAMENTOSUM (FORSSK.) CHIOV.
( » B . knyanum (Vatke) N.E. Br . ex Broun &
Massey) 2
( - B . obovatum (E. Mey. ex Benth.) N.E.
Br. var. knyanum (Vatke)
Cufod.) 1, 2
( -Oc 1 mum fissilabrum Briq.) 1, 2
(-Ocimum knyanum Vatke) 1, 2
( -Oc i mum rautanenii Briq.) I, 2
(-Ocimum stenoglossum Briq.) 1, 2
B. knyanum (Vatke) N.E. Br. ex Broun & Massey -
B. FILAMENTOSUM
B. obovatum (E. Mey. ex Benth.) N.E. Br . var.
knyanum (Vatke) Cufod. - B.
FILAMENTOSUM
SCROPHULARIACEAE (PART A)
Contributed by
W.G. Welman
7471000 -DIASCIA LINK & OTTO Revision: O.M. Hilliard &
B.L. Bur 1 1 (E); K. Steiner
( NBG)
6. HILLIARD & BURTT. 1988. NOTES R. BOT. GDN
EDINB. 45,1: 87-88.
3750 D. TRANSKEIANA HILLIARD & BURTT
7472000 -HEMIMERIS L. P.
1. HIERN. 1904. FC 4,2: 164.
2. DIELS. 1910. BOT. JB. 44: 121.
3. GRANT. 1938. ANN. MISS. BOT. CARD. 25:
435-453.
100 H. CENTRODES HIERN
( -H . nana Diels) 3
H. elegans Hiern - DIASCIA CAPSULARIS
200 H. GRACILIS SCHLTR.
100
-GLEKIA HILLIARD
1. HILLIARD. 1989.
45,3:
G. KREBSIANA (BENTH.)
( -Phyl lopod ium
NOTES R. BOT. GDN EDINB.
481-491.
HILLIARD
krebsianum Benth.) 1
7522000
1050
1350
1550
1775
1930
1935
2 350
3175
-POLYCARENA BENTH.
4. HILLIARD. 1989. NOTES R. BOT. GDN EDINB.
45,3: 481-491.
P. AEMULANS HILLIARD
P. BATTENI ANA HILLIARD
P. calva (Hiern) Levyns - MELANOS PERMUM
TRANSVAALENSE
P. collina Hiern - PHYLLOPODIUM COLLINUM
P. COMPTONI I HILLIARD
P. discolor Schinz - MELANOS PERMUM FOLIOSUM
P. EXIGUA HILLIARD
P. foliosa Benth. - MELANOSPERMUM FOLIOSUM
P. FORMOSA HILLIARD
P. glutinosa (Schltr.) Levyns - TRIEENEA
GLUTINOSA
P. GRACILIS HILLIARD
P. LILACINA HILLIARD VAR. LILACINA
P. LILACINA HILLIARD VAR. DIFFICILIS HILLIARD
P. linearifolia (H. Bol.) Levyns -
PHYLLOPODIUM ELEGANS
P. lupuliformis Thell. - PHYLLOPODIUM
LUPULIFORME
P. maxii Hiern - PHYLLOPODIUM MAXII
P. namaensis Thell. - PHYLLOPODIUM NAMAENSE
P. NARDOUWENSIS HILLIARD’
P. plantaginea auct. - PHYLLOPODIUM ANOMALUM
P. pubescens Benth. p.p. - PHYLLOPODIUM
COLLINUM
P. rariflora Benth. var. micrantha Schltr. -
PHYLLOPODIUM MICRANTHUM
P. rupestris (Hiern) Levyns - MELANOSPERMUM
RUPESTRE
P. schlechteri (Hiern) Levyns - TRIEENEA
SCHLECHTERI
P. SUBTILIS HILLIARD
P. transvaalensis Hiern - MELANOSPERMUM
TRANSVAALENSE
7522010
100
200
300
400
500
600
-MELANOSPERMUM HILLIARD
1. HILLIARD. 1989. NOTES R. BOT. GDN EDINB.
45,3: 481-491.
M. FOLIOSUM (BENTH.) HILLIARD
(-Polycarena discolor Schinz) l
(-Polycarena foliosa Benth.) 1
M. ITALAE HILLIARD
M. RUDOLF 1 1 HILLIARD
M. RUPESTRE (HIERN) HILLIARD
( -Phyllopodlum rupestre Hiern) 1
(-Polycarena rupestris (Hiern) Levyns) l
M. SWAZICUM HILLIARD
M. TRANSVAALENSE (HIERN) HILLIARD
(-Phyllopodlum calvum Hiern) 1
(-Polycarena calva (Hiern) Levyns) 1
(-Polycarena transvaalensis Hiern) 1
7522020
100
200
300
-TRIEENEA HILLIARD
1. HILLIARD. 1989.
45,3:
T. ELSIAE HILLIARD
T. FRIGIDA HILLIARD
T. GLUTINOSA (SCHLTR.
NOTES R. BOT.
481-491.
) HILLIARD
GDN EDINB.
Bothalia 19,2 (1989)
293
( -Phyl lopod lum glutlnosum Schltr.) 1
(-Polycarena glutlnosa (Schltr.) Levyna)
1
400 T. LANCILOBA HILLIARD
500 T. LAS IOCEPHALA HILLIARD
600 T. LAXIFL0RA HILLIARD
700 T. LONGIPEDICELLATA HILLIARD
800 T. SCHLECHTERI (HIERN) HILLIARD
( -Phyl lo pod 1 um schlechterl Hiern) 1
(-Polycarena schlechterl (Hiern) Levyna)
1
900 T. TAYL0RII HILLIARD
7 562000 -LINDERNIA ALLION I
5. PHILCOX. 1987. BOL. SOC. BROT. 60: 267-
270.
50 L. CONFERTA (HIERN) PHILCOX
(-Ilyaanthes conferta Hiern) 5
600 L. PULCHELLA (SKAN) PHILCOX
(-Ilyaanthes pulchella Skan) 5
700 L. WILMSII (ENGL. & DIELS) PHILCOX
(-Ilyaanthes muddil Hiern) 5
(-Ilyaanthes wllmali Engl. & Diels) 5
7564000 -ILYSANTHES RAF.
I. conferta Hiern - LINDERNIA CONFERTA
I. muddil Hiern - LINDERNIA WILMSII
I. pulchella Skan - LINDERNIA PULCHELLA
I. wllmali Engl. & Diels - LINDERNIA WILMSII
SELAGINACEAE
7568000
rlbuted by W.G. Welman
300
350
-SELAG0 L.
S. elegans Cholsy - PHYLL0P0DIUM ELEGANS
S. rustii Rolfe - PHYLLOPODIUM RUSTII
-MICRODON CHOISY
1. ROLFE. 1901. FC 5,1: 175-177.
2. LEVYNS . 1948. JL S. APR. BOT. 14: 85
M. CAPITATUS (BERG.) LEVYNS
( -M . ovatus (L.) Cholsy) 2
(-Selago capitata Berg.) 2
M. CYLINDRICUS E. MEY.
(-Selago splcata Link) 1
M. LINEARIS CHOISY
M. LUCIDUS (VENT.) CHOISY
(-Selago 1 uc Ida Vent.) I
M. ORBICULARIS CHOISY
M. ovatus (L.) Cholsy - M.. CAPITATUS
SCROPHULARIACEAE (PART B)
Contributed by
W.G. Welman
100
350
-Gerardla L. - CYCNIUM
G. tubulosa L. f. - CYCNIUM TUBULOSUM SUBSP.
TUBULOSUM
-SOPUBIA BUCH.-HAM. EX D. DON
S. fas t lg lata Hiern - S. KARAGUENSIS VAR.
KARAGUENSIS
S. KARAGUENSIS OLIV. VAR. KARAGUENSIS
( - S . fast lglata Hiern) 2
-CYCNIUM BENTH. EMEND. ENGL.
3. HANSEN. 1978. DANSK BOT. ARKIV 32,3: 1-
72.
C. ADONENSE E. MEY. EX BENTH. SUBSP. ADONENSE
C. TUBULOSUM (L. F.) ENGL. SUBSP. TUBULOSUM
ACANTHACEAE
8007000
Contributed by W.G. Weln
7906000
-ASYSTAS I A BLUME
A. natalensis C.B. Cl. - SALPINCTIUM NATALENSE
A. stenoslphon C.B. Cl. - SALPINCTIUM
STENOSIPHON
8007010 -SALPINCTIUM T.J. EDWARDS
1. EDWARDS 4 GETLIFFE NORRIS. 1989. S. AFR.
J. BOT. 55,1: 6-10.
100 S. HIRSUTUM T.J. EDWARDS
200 S. NATALENSE (C.B. CL.) T.J. EDWARDS
(-Asystasia natalensis C.B. Cl.) 1
300 S. STENOSIPHON (C.B. CL.) T.J. EDWARDS
(-Asystasia stenoslphon C.B. Cl.) 1
RUBIACEAE Contributed by W.G. Welman 8119000
8471000 -DIODIA GRONOV. EX L.
2. HILLIARD & BURTT. 1988. NOTES R. BOT. GDN
EDINB. 45,1: 86.
ASTERACEAE
itributed by W.G. Weln
8729000
100
150
-ETHULIA L . F.
1. GILBERT & JEFFREY. 1988. KEW BULL. 43,2:
165-193.
E. CONYZOIDES L. F. SUBSP. CONYZOIDES
E. CONYZOIDES L. F. SUBSP. KRAUSSII (WALP.) M.
1100
1550
1800
1850
2100
2300
2400
2500
3000
8816030
1 50
850
9065000
V. CINEREA (L.) LESS. VAR. CINEREA
( -Cony za clnerea L.) 13
V. COLORATA (WILLD.) DRAKE SUBSP. COLORATA
V. GALPINII KLATT
( -Pegole 1 1 la tenella DC.) 15
V. GLABRA (STEETZ) VATKE VAR. GLABRA
(-Linzla glabra Steetz var. glabra) 8
V. GLABRA (STEETZ) VATKE VAR. LAXA (STEETZ)
BRENAN
(-Linzla glabra Steetz var. laxa Steetz)
13
V. INHACENSIS G.V. POPE
V. kraussll Sch. Blp. ex Walp. - V.
OL IGOCE PHALA
V. leptolepls Bak. - V. ADOENSIS VAR.
KOTSCHYANA
V. ME IOSTEPHANA C. JEFFREY
V. MYRIANTHA HOOK. F.
( -V . ampla 0. Hoffra.) 13
(-V. podocoma Sch. Blp. ex Vatke) 13
( - V . s 1 1 pulacea Klatt) 13
V. NATALENSIS SCH. BIP. EX WALP.
V. NESTOR S. MOORE
V. OLIGOCEPHALA (DC.) SCH. BIP. EX WALP.
( -V . kraussll Sch. Blp. ex Walp.) 13
V. podocoma Sch. Blp. ex Vatke - V. MYRIANTHA
V. stenolepls Ollv. - V. ANTHELM INTICA
V. sti pulacea Klatt - V. MYRIANTHA
V. wood 1 1 0. Hoffm. - V. ADOENSIS VAR.
KOTSCHYANA
-Linzla Sch. Blp. - VERNONIA
L. glabra Steetz var. glabra - VERNONIA GLABRA
VAR. GLABRA
L. glabra Steetz var. laxa Steetz - VERNONIA
GLABRA VAR. LAXA
-ADENOSTEMMA J.R. & G. FORST.
2. KING & ROBINSON. 1987. MONOGRAPHS SYST.
BOT. 22: 477.
A. dregei DC. - A. VISCOSUM
A. natalense DC. - A. VISCOSUM
A. perrottetii DC. - A. VISCOSUM
A. VISCOSUM J.R. FORST. & G. FORST.
( - A . dregei DC.) 2
( - A . natalense DC.) 2
( - A . perrottetii DC.) 2
-Eupatorium L. - AGERATINA, CAMPULOCLI NI UM ,
CHROMOLAENA, PEGOLETTIA,
PTERONIA, STOMATANTHES
E. macrocephalum Less. - C AM PULOCLINI UM
MACROCEPHALUM
E. rugosum Houtt. - AGERATINA ALT I S S IMA
-AGERATINA SPACH
A. ALTISSIMA (L.) R.M. KING & H. ROBINSON *
(-Eupatorium rugosum Houtt.) 1
-CAMPULOCLINI UM DC.
1. KING 6 ROBINSON. 1987. MONOGRAPHS SYST.
BOT. 22: 519.
C. MACROCEPHALUM (LESS.) DC. *
(-Eupatorium macrocephalum Less.) 1
-CONYZA LESS.
C. c In
L.
VERNONIA CINEREA VAR. CINEREA
-ANAXETON GAERTN.
2. NORDENSTAM. 1988. S. AFR. J. BOT. 54,6:
631-632 .
A. LUNDGRENI I B. NORD.
-Iphiona Cass. Southern African species moved
to ANISOTHRIX, ANTI PHI ONA ,
PEGOLETTIA
I. dentata (H. Bol.) H. Bo 1 . - ANISOTHRIX
KUNTZEI
I. Integra Compton - ANISOTHRIX INTEGRA
-ANISOTHRIX 0. HOFFM.
1. ANDERBERG. 1988. BOT. JAHRB. SYST. 109,3:
363-372.
A. INTEGRA (COMPTON) A. ANDERB.
(-Iphiona Integra Compton) 1
A. KUNTZEI 0. HOFFM.
(-Iphiona dentata (H. Bol.) H. Bol.) 1
(-Pegolettla dentata H. Bol.) 1
-PEGOLETTIA CASS.
1. ANDERBERG. 1986. CLADISTICS 2,2: 158-186.
P. acuminata DC. - P. RETROPRACTA
P. dentata H. Bol. - ANISOTHRIX KUNTZEI
P. RETROPRACTA (THUNB.) KIES
(-P. acuminata DC.) 1
P. tenella DC. - VERNONIA GALPINII
-MATRICARIA L.
cut iloba (DC.) Harv. - MYXOPAPPUS
ACUTILOBUS
fricana Berg. - ONCOSIPHON AFRICANUM
lb Ida (DC.) Fenzl ex Harv. - FOVEOLINA
ALBIDA
Harv
FOVEOLINA
jtoma (DC.) Fenzl
DICHOTOMA
rata DC. - ONCOSIPHON GLABRATUM
L f era (Thunb.) Fe^zl ex Harv. -
ONCOSIPHON PILULIFERUM
11 f lor a (Thunb.) Fenzl ex Harv. -
ONCOSIPHON GRANDIFLORUM
jtlfolia S. Moore - FOVEOLINA ALBIDA
a (Thunb.) DC. - ONCOSIPHON AFRICANUM
lifera (L.f.) Druce - ONCOSIPHON
294
Bothalia 19,2 (1989)
PILULIFERUM
M. sabulosa Wolley-Dod - ONCOSIPHON SABULOSUM
M. schlnz lana Thell. - FOVEOLINA SCHINZIANA
M. achlechteri H. Bol. ex Schltr. - ONCOSIPHON
SCHLEC HTER I
M. tenella DC. - FOVEOLINA TENELLA
934 1070 -Tanacetum Harv. - ATH ANAS I A , COTULA, FOVEOLINA,
HIPPIA, MYXOPAPPUS, PENTZIA,
ONCOSIPHON, STCHISTOSTEPHIUM
T. acutilobum DC. - MYXOPAPPUS ACUTILOBUS
T. al bid urn DC. - FOVEOLINA ALB I DA
T. grandl f lorura Thunb. - ONCOSIPHON
GRANDIPLORUM
T. suf f rut Icoaum L. - ONCOSIPHON S UFFRUT ICOSUM
9351000
-COTULA L.
C. globifera Thunb.
C. ptlullfera L. f.
C. t anac etlfolla L.
ONCOSIPHON PILULIFERUM
ONCOSIPHON PILULIFERUM
ONCOSIPHON SUFFRUTICOSUM
(-Cotula pllul If era L. f.) 1
(-Matricaria globifera (Thunb.) Fenzl ex
Harv.) 1
(-Matricaria pllullfera (L. f.) Druce) 1
( -Pentzla globifera (Thunb.) Hutch.) 1
( -Pentzla pllullfera (L. f.) Fourc.) 1
600 0. SABULOSUM (WOLLEY DOD) KALLERS JO
(-Matricaria sabulosa Wolley Dod) 1
(-Pentzla sabulosa (Wolley Dod) Hutch.)
1
700 0. SC HLECHTER I (H. BOL.) KALLERS JO
(-Matricaria achlechteri H. Bol. ex
Schltr.) 1
800 0. SUFFRUTICOSUM (L.) KALLERSJO
(-Cotula tanacetlfolla L.) 1
(-Pentzla suffruticosa (L.) Hutch, ex
Merxm . ) 1
(-Pentzla tanacetlfolla (L.) Hutch.) 1
(-Tanacetum auf f r ut lcosum L.) 1
9366000
1825
-PENTZIA THUNB. Revlalon: K. Bremer & S.
Kallerajo (S)
9. KALLERSJO. 1988. BOT. J. LINN. SOC. 96:
299-322 .
P. acutlloba (DC.) Hutch. - MYXOPAPPUS
ACUTILOBUS
P. alblda (DC.) Hutch. - FOVEOLINA ALB I DA
P. annua DC. - FOVEOLINA ALB I DA
P. dlchotoma DC. - FOVEOLINA DICHOTOMA
P. eenll S. Moore - RENNERA EENII
P. galplnil Hutch. - MYXOPAPPUS HEREROENSIS
P. globifera (Thunb.) Hutch. - ONCOSIPHON
PILULIFERUM
P. grandiflora (Thunb.) Hutch. - ONCOSIPHON
GRANDIPLORUM
P. hereroenala 0. Hoffn. - MYXOPAPPUS
HEREROENS IS
P. Intermedia Hutch. - ONCOSIPHON INTERMEDIUM
P. LANATA HUTCH.
P. 1 ax a Brem. & Oberrn. - RENNERA LAXA
P. membranacea Hutch. - FOVEOLINA ALB I DIFORMI S
P. pllullfera (L. f.) Fourc. - ONCOSIPHON
PILULIFERUM
P. sabulosa (Wolley-Dod) Hutch. - ONCOSIPHON
SABULOSUM
P. achlnzlana (Thell.) Merxm. & Eberle -
FOVEOLINA SCHINZIANA
P. suffruticosa (L.) Hutch, ex Merxm. -
ONCOSIPHON SUFFRUTICOSUM
P. tanacetlfolla (L.) Hutch. - ONCOSIPHON
SUFFRUTICOSUM
9366010 -RENNERA MERXM.
1. KALLERSJO. 1988. BOT. J. LINN. SOC. 96:
299-322.
25 R. EENII (S. MOORE) KALLERSJO
(-Pentzla eenll S. Moore) 1
50 R. LAXA (BREM. f, OBERM.) KALLERSJO
(-Pentzla laxa Brem & Oberm.) 1
100 R. LIMNOPHILA MERXM.
9366030 -MYXOPAPPUS KALLERSJO
1. KALLERSJO. 1988. BOT. J. LINN. SOC. 96:
299-322.
100 M. ACUTILOBUS (DC.) KALLERSJO
(-Matricaria acutlloba (DC.) Harv.) 1
(-Pentzla acutlloba (DC.) Hutch.) 1
(-Tanacetum acutilobum DC.) 1
200 M. HEREROENSIS (0. HOFFM.) KALLERSJO
(-Pentzla galplnil Hutch.) 1
(-Pentzla hereroenala 0. Hoffm.) 1
9366040
100
200
300
400
500
-FOVEOLINA KALLERSJO
1. KALLERSJO. 1988. BOT. J. LINN. SOC. 96:
299-322.
F. ALB I DA (DC.) KALLERSJO
(-Matricaria alblda (DC.) Fenzl ex
Harv.) 1
(-Matricaria hir8utlfolla S. Moore) 1
(-Pentzla alblda (DC.) Hutch.) 1
(-Pentzla annua DC.) 1
(-Tanacetum albldum DC.) 1
F. ALB IDIFORMIS (THELL.) KALLERSJO
(-Matricaria albldlformla Thell.) 1
(-Pentzla membranacea Hutch.) 1
F. DICHOTOMA (DC.) KALLERSJO
(-Matricaria dlchotoma (DC.) Fenzl ex
Harv.) 1
(-Pentzla dlchotoma DC.) 1
F. SCHINZIANA (THELL.) KALLERSJO
(-Matricaria achlnzlana Thell.) 1
(-Pentzla achlnzlana (Thell.) Merxm. &
Eberle*) 1
F. TENELLA (DC.) KALLERSJO
(-Matricaria tenella DC.) 1
9406000 -CINERARIA L.
10. HILLIARD. 1989. NOTES R. BOT. GDN EDINB.
45,2: 185-187.
125 C. ALBOMONTANA HILLIARD
3275 C. VAGANS HILLIARD
9366020
100
200
300
400
500
-ONCOSIPHON KALLERSJO
1. KALLERSJO. 1988. BOT. J. LINN. SOC. 96:
299-322 .
0. AFRICANUM (BERG.) KALLERSJO
(-Matricaria afrlcana Berg.) 1
(-Matricaria capenala L.) 1
(-Matricaria hlrta (Thunb.) DC.) 1
0. GLABRATUM (THUNB.) KALLERSJO
(-Matricaria glabrata (Thunb.) DC.) 1
0. GRANDIPLORUM (THUNB.) KALLERSJO
(-Matricaria grandiflora (Thunb.) Fenz
ex Harv.) 1
(-Pentzla grandiflora (Thunb.) Hutch.)
(-Tanacetum grandlflorum Thunb.) 1
0. INTERMEDIUM (HUTCH.) KALLERSJO
(-Pentzla Intermedia Hutch.) 1
0. PILULIFERUM (L. F.) KALLERSJO
(-Cotula globifera Thunb.) 1
9420000 -OTHONNA L. Revlalon: B. Nordenatam (S)
5925 0. OVALIFOLIA HUTCH.
9438000 -BERKHEYA EHRH . Revlalon: S. Roeasler (M)
7. HILLIARD. 1989. NOTES R. BOT. GDN EDINB.
45,2: 183-185.
4925 B. LEUCAUGETA HILLIARD
9500000
150
175
2 00
300
-OLDENBURG I A LESS.
1. BOND. 1987. S. AFR. J. BOT.
500.
0. arbuacula DC. - 0. GRANDIS
0. GRANDIS (THUNB.) BAILL .
(-Arnica grandla Thunb.) 1
(-0. arbuacula DC.) 1
0. INTERMEDIA BOND
0. PAPIONUM DC.
0. PARADOXA LESS.
53,6:
Each contributor is acknowledged at the beginning of
the groups for which he/she is responsible. Although staff
of the Botanical Research Institute have final responsibility
for maintaining Taxon-PRECIS, we acknowledge with
gratitude the co-operation of other botanists in reporting
changes. Mrs J. Mulvenna spent hours slaving over a hot
terminal to add all the changes to PRECIS, thereby also
helping immeasurably in catching and eliminating incon-
sistencies.
REFERENCES
ARNOLD, T.H. in press. Computerization of the curatorial and service
functions of the National Herbarium, Pretoria. Transvaal Muse-
um Monograph 9.
CROSBY, M.R. & MAGILL, R.E. 1981. A dictionary of mosses.
Missouri Botanical Garden, St Louis.
DYER, R.A. 1975. The genera of southern African flowering plants. Vol.
1. Dicotyledons. Botanical Research Institute, Pretoria.
DYER, R.A. 1976. The genera of southern African flowering plants. Vol.
2. Monocotyledons. Botanical Research Institute, Pretoria.
GIBBS RUSSELL,' G.E., WELMAN, W.G., GERMISHUIZEN, G.,
RETIEF, E., PIENAAR, B.J., REID, C., FISH, L., VAN ROOY,
J., VAN WYK, C.M., KALAKE, E. & STAFF 1988. New taxa,
new records and name changes for southern African plants.
Bothalia 18: 293-304.
GROLLE, R. 1983. Nomina generica hepaticarum: references, types and
synonymies. Acta Botanica Fennica 121: 1—62.
MORRIS, J.W. & GLEN, H.F. 1978. PRECIS, the National Herbarium
of South Africa (PRE) Computerised Information System. Taxon
27: 449-462.
SCHELPE, E.A.C.L.E. & ANTHONY, N.C. 1986. Pteridophyta. InO.A.
Leistner, Flora of southern Africa. Botanical Research Institute,
Pretoria.
Bothalia 19.2: 295-312 (1989)
ANNUAL REPORT OF THE BOTANICAL RESEARCH INSTITUTE
1988/1989
1st April 1988— 31st March 1989
CONTENTS
Introduction 295
Reports of divisions 296
Staff list 307
Publications by the staff 310
INTRODUCTION
It can be reported with satisfaction that the threat of total
dismemberment of the Botanical Research Institute, which
would have been the result of the implementation of the
Commission for Administration’s Report of February 1988,
has been averted. The efforts needed to forestall what
would have been a disastrous event for botany in South
Africa have taken an inescapable toll of day to day activities
and the delays in the process of the amalgamation of the
Botanical Research Institute and the National Botanic
Gardens, in addition, have had a strongly negative
influence on staff morale.
It is regretted that the BRI has had to sacrifice eight posts
to the newly established Pasture Research Centre of the
Department of Agriculture and Water Supply, but it is
gratifying that the relinquishing of these posts was not
accompanied by the loss of any* of the botanical functions
for which the BRI is responsible. The botanical mandate
of the BRI has therefore been transferred to the new
organization, as a whole. The new national botanical
organization will produce major benefits for all concerned,
if managed efficiently and in the national interest. In this
context it is important to emphasize that the new
organization will have a wider scope than the BRI,
including not only research but also relevant aspects of
horticulture and conservation. In addition, the education
of the public towards the realization of their responsibility
and role in maintaining a stable plant environment, has
an equal priority. The object of the new organization will
therefore be to function as the primary promoter of
botanical extension, research and services in the State
Sector. This implies that its development programmes will
have to be geared towards satisfying the primary needs of
the state in the fields mentioned.
The research output of the Institute, in spite of the
generally unsettling effect of the amalgamation, remained
at a high level, the total of publications by staff being 85
papers of different kinds.
The journals published by the Institute appeared on
schedule. The fiftieth volume of The Flowering Plants of
Africa underwent a facelift and appeared with an attractive
illustrated cover and now includes an editorial. Two parts
of Bothalia, the house journal of the Institute, were
published. In the Flora of southern Africa the appearance
of the monographic account of the legume genus
Aspalathus (Papilionoideae) by the late Rolf Dahlgren set
a standard unlikely to be easily surpassed.
The third edition of Acocks’s Veld Types of South Africa
appeared during the year and contains a new selection of
veld photographs of higher quality. This re-issue, like its
forerunners, should prove to be a popular addition to our
publications.
Computers are playing an increasing role in the
organization. The use of DELTA, the international
computerized system of preparing manuscripts on
taxonomic subjects, is gaining momentum in the Institute
and is likely to revolutionize the process of writing of flora
handbooks. The procurement of this system for the BRI
again places it ahead of many other organizations.
Generated by the computerized PRECIS data system a list
of nearly 750 new names and name changes has appeared
in the journal Bothalia. This annual contribution, compiled
from information gathered in scanning over 120 journals,
updates the List of species of southern African plants (edn
2) announced in the 1987/88 Annual Report, thus extending
the usefulness of this standard work. Keeping abreast of
name changes remains a problem of considerable
magnitude, which is made manageable by the unique
service provided to users by the BRI.
The response to the demonstration of the computeriza-
tion of the curatorial and service functions of the National
Herbarium, at the AETFAT Congress in Hamburg in 1988,
showed that in this respect also, the Institute has maintained
its leading position. The establishment of
a database on indigenous southern African food plants
promoted during the AETFAT Congress, was also
favourably received.
Two major plans to improve accommodation for staff
and to ensure the safety of the collections were finalized
during the year under review. Firstly the installation of
an air-conditioning and a gas-flooding system to protect
the irreplaceable collections of the National Herbarium
against fire was successfully completed. In addition, the
expansion of accommodation in the basement of the main
building in Pretoria, by enclosing the areas under the
western wing, was completed and greatly improved the
296
Bothalia 19,2 (1989)
facilities for decontaminating plant materials before these
are brought into the building.
In general the Institute had a good year and it is to be
hoped that the large number of new initiatives which are
being developed will receive added stimulus once the
unsettling effects of the process of amalgamation have
abated. We look forward to a bright future.
HERBARIUM DIVISION
For the third successive year, under-staffing and a heavy
workload have placed staff of the Institute’s four herbaria
under considerable pressure, affecting all major functions,
namely, curation, research and herbarium services. This
situation should improve considerably in the following
year, with the introduction of a handling fee for plant
identifications and through the selective control of
accessions into the National Herbarium.
National Herbarium, Pretoria (PRE)
Curation
Scientific staff continued to scan over 120 scientific jour-
nals as well as books for taxonomic and nomenclatural
changes covering the Flora of southern Africa region.
These contained 172 articles of direct relevance to southern
Africa. A total of 744 changes was recorded affecting about
3% of the flora. These included: 412 new names (of which
about 293 were new taxa), 205 names reduced to
synonymy and 44 orthographic corrections.
An extension of the known distribution area was
recorded for 45 taxa, mostly in terms of provincial
boundaries (Transvaal 1 new taxon, OFS 3, Natal 4, Cape
18, Transkei 5, Botswana 1 and Swaziland 3) with 10 new
records for southern Africa.
Computerization
The in-house herbarium computer system is functioning
well and continues to be an important aid, resulting in both
improved efficiency and better management. New
applications developed on the Burroughs B28 system
include: merging outstanding specimen loan data with a
standard loan expiry letter and the use of the computer
to selectively control the accessioning of specimens into
the National Herbarium.
Accommodation
The major disruptive changes that the National
Herbarium and Mary Gunn Library underwent, with the
installation of fire protection (halon flooding) and air
conditioning, came to an end towards the middle of the
year, much to the relief of all herbarium staff. This was
followed immediately, however, by three further additions
to the Herbarium, namely, the installation of three working
bays on the south side of each herbarium wing, provision
of a special room for decontaminating and drying
specimens and a new SEM room and various other rooms
adjacent to the Moss Herbarium. All these alterations are
now complete and the herbarium is, at last, functioning
normally again.
Visitors
In addition to numerous local visitors from various
universities, Government institutes, nature conservation
departments etc., together with members of the general
public, the herbarium was also consulted by officers and
personnel from Lesotho, Botswana, Swaziland, Bophuthat-
swana, Venda and Transkei.
A number of overseas botanists visited the Institute and
Herbarium. These included: Prof. D. Wiens (Utah, USA);
Dr C.L. Calvin (Oregon, USA); Miss S. Liede (Hamburg,
W Germany); Prof, and Mrs D. & U. Miiller-Doblies (Ber-
lin, W Germany); Dr R.N. Lester (Birmingham,
England); Prof. J.R. Harborne (Reading, England); Mr
and Mrs la Croix (Darling, England); Dr F. Bauman
(Amsterdam, Netherlands); Prof. R.M. Schuster
(Massachusetts, USA); Dr S.P. Doolan (Oxford, England);
Dr D.J. Lehmiller (Texas, USA); Dr M. Hale (Washington,
USA); Miss S. Andrews (Kew, England); Dr P. Perrino
(Italy); Dr C. Laghetti (Italy); Dr D.A. Preston (Indiana,
USA).
Collecting expeditions
Due to financial and other constraints, few collecting
expeditions were undertaken this year. Those that were
undertaken included trips to Namaqualand (Riccia and
general collecting); northern Natal and eastern Transvaal
( Vigna and Polygonaceae); eastern OFS (general
collecting) and SW Cape ( Pentameris ).
Herbarium services
Plant identifications numbering 211 batches and 13 229
specimens were undertaken for officers of this Institute,
various state departments, provincial administrations,
universities and neighbouring States. The major users of
the service were: staff of the BRI 26% of the material
identified; Universities 20%; nature conservation 16%;
museums 16%; botanic gardens 10%; private collectors
5%; agriculture 3%; forestry 2% and others 2%.
Identifications for 233 visitors numbered 1 152. Of the
specimens received as gifts or part of exchange agreements
1 620 had to be given names or renamed. Seventy-one
specimens were identified for Onderstepoort involving
stock poisonings. Enquiries received by telephone totalled
978. New accessions to the herbarium numbered 22 886
for southern Africa and ± 6 166 for tropical Africa and
other areas.
Specimen loans sent to local and overseas herbaria
numbered 85 (comprising 11 046 specimens) with 101
existing loans returned in full and 31 loans returned in part
(8 167 specimens were received back in total). The total
number of outstanding loans is 308 (35 425 specimens).
Some 10930 specimens were despatched as part of
exchange agreements and 5 953 were received by PRE.
Specimens received as gifts totalled 1 519 while 2 Oil
specimens were dispatched as gifts in return.
Research and related activities
Contributions to the lichen flora (F. Brusse). Six new
species were described and ten new combinations made
Bothalia 19,2 (1989)
in the genus Parmelia. Two new species were also
described for Moronea and Fuscidea, one in each genus.
Eight new lichen records were made for southern Africa.
Ricciaceae (Hepaticae) (S.M. Perold). Three papers were
published during the year on: Riccia nigrella DC. and R.
campbelliana Howe (both new records for southern Africa)
and two new species R. argenteolimbata and R. alboma-
ta. Two papers are in press involving R. montana and R.
alboporosa (new species) and R. albolimbata and the sta-
tus of R. albosquamata. Four more papers have been sub-
mitted for publication. Final drafts of 30 Riccia species
have been written up for the revision of the family. Five
new species in the section Pilifer are still being
investigated. Distribution maps for 35 species are now
complete. The artwork for 32 species is also complete.
Contributions to the moss flora (J. van Rooy) . The genera
Rhachithecium, Rhabdoweisia and Orthotrichum have
been completed for the 3rd fascicle of the flora. Three
undescribed species and two new records for Africa have
been identified in the family Orthotrichaceae. Work on
the illustrations and distribution maps for this family is
nearing completion. Work on the family Polytrichaceae
for the 4th fascicle has commenced.
Revision of Carex (Cyperaceae) (C. Reid). Seventeen taxa
have been delineated. Two formerly described species
cannot be upheld and one requires further investigation.
One putative hybrid has been identified. All loan
specimens have been identified and their qualitative and
quantitative characters are currently being measured.
Loans from three institutions are awaited. Typification is
75 % complete with mainly the cosmopolitan taxa still out-
standing.
Polygonaceae (G. Germishuizen). A revision of the genus
Oxygonum has now finally been completed and written
up in the Flora of southern Africa format. Work on the
genera Fagopyrum, Emex and Rumex is still in progress.
Revision o/Vigna (Fabaceae) (B.J. Pienaar). This revision
is now approaching completion. Thirteen species are
recognised for which the descriptive work is largely
complete.
Revision of Convolvulaceae (W.G. Welman). Ninety-one
descriptions of the Convolvulaceae in Prof. A.D.J.
Meeuse’s manuscript were shortened to fit the Flora of
southern Africa format.
Transvaal Wild Flowers (Vol. 2) ( G . Germishuizen).
Descriptions were completed for 60 species taking the total
to 150. The artist, Mrs Anita Fabian, has completed
illustrations for 350 taxa which means that this component
of Volume 2 is 75 % complete.
Flora of the Transvaal (E. Retief and P. Herman). This
flora is to closely resemble, with some additions, the The
flora of Natal by J.H. Ross (1972). To date, data sheets have
been completed for 73 families, 182 genera and 861 taxa.
This is 11,8% of the total number of taxa for the region.
Research support
Scanning electron microscope (S.M. Perold). Some 2 974
SEM micrographs were prepared for the following
officers: Dr R. Ellis (grass leaf surfaces); Mr N. Barker
(grass caryopses); Dr H. Glen ( Aloe leaf surfaces); Mr
J. van Rooy (moss leaves and capsules); Mrs S. Perold
(Riccia thalli and spores); Mr G. Germishuizen
(Oxygonum)\ Miss C. Reid (Carex caryopses); Mr F.
Brusse (lichen thalli); Mr D. McDonald (Pelargonium
pollen); Dr K.L. Immelman (Justicia pollen); Dr B. de
Winter (Gorteria) and Miss E. Retief (Cyphostemma
leaves). Towards the end of 1988 the SEM unit was moved
to its new premises in the basement next to the Cryptogam
Herbarium.
Expansion of collections from poorly represented areas
(various contributors). North-western Cape — holdings for
the sixteen 1/4° grids visited, increased by 71% from 132
to 447 collections; eastern Transvaal — although most of
the twenty-one 1/4° grids visited were fairly well represent-
ed in PRE, two were not and increased by 43% from 74
to 126 collections.
Expansion of the fruit, seed and spirit collections (E. Retief
and G. Germishuizen). The fruit and seed collections
remained unchanged at 4 567 and 4 134 respectively. The
spirit collection was expanded from 3 726 to 3 801 col-
lections.
Publications
Seventeen articles appeared in local (15) as well as
overseas (2) journals. A further 31 articles are in press
awaiting publication.
Natal Herbarium, Durban (NH)
No staff changes occurred during the year. Miss R.
Williams continued her good work as Curatrix of the
Herbarium closely assisted by Mrs H.E. Noble and the
rest of the herbarium staff.
Mr A.M. Ngwenya attended the 1988 International
Diploma Course in Herbarium Techniques. This was a
three month course which was held at the Royal Botanic
Gardens, Kew (Figure 1).
A total of 3 101 specimens were identified, 412 visitors
were attended to, 838 specimens were sent out on loan
and accessions to the herbarium numbered 2 261.
One thousand and fifty-three Dicotyledon specimens
from the Cape have been sent to the National Herbarium
and 434 to the Stellenbosch Herbarium.
Albany Museum Herbarium, Grahamstown (GRA)
The post of Herbarium Assistant remained unfilled from
November 1987 to October 1988 when Mrs Louise Verwy
was appointed. During the eleven intervening months the
herbarium staff, and in particular Mrs E. Brink, Curatrix
of the herbarium, were under considerable pressure trying
to deal with the workload due to the vacant post.
298
Bothalia 19,2 (1989)
FIGURE 1. — Mr Alfred Ngwenya of
the Natal Herbarium receives
his International Diploma in
Herbarium Techniques from
Dr G.T. Prance, Director of the
Royal Botanic Gardens, Kew,
4 November 1988.
Dr Amy Jacot Guillarmod is established in her part-time
post as Curator of the Pocock Marine Algae. Mr Neil
Abrahams, who has given voluntary assistance in the
herbarium for the past 21 years, has become an integral
part of the ‘family’ and is sorely missed whenever he leaves
town for commitments elsewhere.
The location of this herbarium within a large Museum
is one of the factors which gives a strong bias towards the
public service performed by this Unit. It is also clear that
Grahamstown’s chief ‘export’, education, plays a rote in
the community’s expectations of the Museum. Action was
taken in August to limit the access of visitors to the
herbarium, to give staff more time to perform curatorial
tasks. Public access to the herbarium is now restricted to
between 08h00 and 12h00 on week-days. Although visitors
adapted quickly to this change, the actual number of
visitors during the year increased slightly to 751.
Other information services also increased for the year
with plant identifications numbering 2 440, telephone
queries 600 and correspondence 238. Display in the
herbarium totalled 39, many of these made possible by
material brought in by members of the public. Material
displayed in the Museum entrance decreased slightly to
six, compared to nine displays last year.
Many of the identifications done were of plant fragments
either for the State Veterinarian (seven visits) or the
Museum’s archaeologist (five visits).
The numbers of specimens mounted (1 507) and
accessioned (2 279) differ very little from last year. A back-
log, however, does exist that needs attention. Nine loans
to other institutions were dealt with, four (393 sheets) were
sent out and five (202) were refiled. The herbarium staff
continued to assist Mrs G.D. Court who is busy with the
Namaqualand Flora. Eighteen loans (2 616 sheets) were
either received or returned on her behalf to other herbaria.
The Grahamstown Nature Reserve was visited 27 times
during this year. The Nature Reserve Caretaker is just able
to keep the alien infestation at its present level and, alone,
is unable to really make any inroads into it. It is much
to his credit that the Reserve still shows up clearly against
the surrounding weed-infested slope as a protected area.
Government Herbarium, Stellenbosch (STE)
The herbarium remains in the temporary accommoda-
tion provided by the University in the old Carnegie
Library. The renovations to the Natural Sciences building
are progressing well and the herbarium should move back
during the latter half of 1989.
During the year 2 496 specimens were identified for the
staff, various government and provincial bodies, and for
private firms and individuals. This is less than in previous
years and has provided time for staff to do meaningful
curatorial work and research. Telephone enquiries during
the year numbered 211 with 272 visitors requiring
information. Accessions to the herbarium were 2 240
specimens. The recording of identifications and listing of
loans is now working well in a microcomputer-based
system.
Mr Oliver continued his work on minor genera of the
Ericaceae concentrating on an extremely variable complex
of four genera with 65 species. He delivered a paper on
his generic concepts in the capsular genera at a Heather
Symposium held by the Linnean Society of London.
Thereafter he worked on the collections at the Royal
Botanic Gardens, Kew, and the British Museum. Mrs
Beyers has begun work on the thymelaeaceous genera
Lachnaea and Cryptadenia. Mrs Fellingham has been
investigating some problem groups in the genus Cliffortia
(Rosaceae).
Collecting trips were mainly confined to local ones in
search of research material. However, Mrs Fellingham
Bothalia 19,2 (1989)
299
spent four days collecting in an undercollected grid area
near Sutherland.
FLORA RESEARCH DIVISION
Dr G.E. Gibbs Russell who has led this division since
November 1984 is presently working as Database
Co-ordinator of the Flora of North America project at the
Missouri Botanical Garden. Dr O.A. Leistner has been
temporarily put in charge of the division.
Flora of southern Africa (FSA)
With the aid of the computer-based Register of Southern
African Plant Taxonomic Projects, research work of the
co-operators and potential contributors to this huge project
has been co-ordinated. Although the Department no longer
finances research contracts on this project, taxonomists
outside the Institute have continued to support the work.
Minor purchases of art work and manuscripts could be
made from the divisional budget. The seventh meeting of
the FSA working group was held during the Congress of
the South African Association of Botanists (SAAB) in
Pretoria during January 1989. The Association was
requested to support the creation of a fund to finance work
undertaken for the FSA. News on the FSA was publicised
in Forum Botanicum , the newsletter of the Association.
Vol. 16,3,6 of the FSA was published. It deals with the
genus Aspalathus of the family Fabaceae and comprises
278 species, including A. linearis the Rooibos tea. The
work written by the late Prof. Rolf Dahlgren of the
University of Copenhagen, Denmark, is illustrated by
146 figures and 123 distribution maps.
Bryophyta: a total of 15 families, comprising 41 genera
and 65 species, have been revised and written up in
FSA format by Dr R.E. Magill of the Missouri
Botanical Garden and Mr J. van Rooy. Five species have
been illustrated by Miss G.C. Condy.
Vol. 2: Poaceae — Ehrharteae. Dr G.E. Gibbs Russell
continued her revision of the genus Ehrharta in co-
operation with Dr R.P. Ellis. The Dura group of this
genus was published in Bothalia, the Ramosa group
was submitted for publication and morphological data
for the Capensis group were recorded in the DELTA
system for computerizing plant descriptions.
The computerized manual for the grasses of southern
Africa, designed as a precursor to Vol. 2 of the FSA,
is nearing completion. Generic descriptions were drawn
up by Dr G.E. Gibbs Russell from the data base of
world grasses maintained at the Australian National
University, Canberra. The other members of the group
working on this manual are Mrs L. Fish, Dr H.M. An-
derson, Miss M. Koekemoer, Mr N. Barker and Mrs
W.J.G. Roux. Some 700 of the 970 taxa to be dealt with
in the work have been written up and illustrations were
completed for about 180 genera.
Vol. 5: Liliaceae — Aloinae. The manuscript of Aloe by
Dr H.F. Glen and Mr D.S. Hardy has been edited and
is almost ready for publication. The possibility of
having this genus published by the private sector in a
format going beyond that of the normal Flora volumes
is being investigated. The manuscript of Kniphofia by
Dr L.E. Codd has been edited but the material on
Gasteria has been temporarily withdrawn by the author.
Liliaceae — Asparagoideae. DrK.L. Immelman who took
over the incomplete manuscript of Protasparagus from
Mrs A. A. Mauve (Obermeyer) has nearly completed
the work. She checked all specimens in the National
Herbarium and drew up new distribution maps for all
the accepted 69 species in the genus.
Vol. 8: Orchidaceae. Except for the revision of the genus
Disperis, which is now being undertaken by Mr J.
Manning, the manuscript of this family is complete and
has been submitted to the editor.
Vol. 9: the manuscripts by Dr K.L. Immelman on the
families Salicaceae, Fagaceae, Urticaceae and
Piperaceae are awaiting publication, pending
completion of work on Myricaceae and Moraceae.
Vol. 11: Mesembryanthemaceae. DrH.E.K. Hartmann of
the University of Hamburg, W Germany and her
students have revised several more genera in the
Leipoldtiinae including Cephalophyllum and
Pleiospilos.
Vol. 12: Portulacaceae. Dr H.F. Glen and Mr D.S. Hardy
have started on a revision of this family for the FSA.
A prototype character list for use with DELTA was
generated and tested using the data in H.R. Toelken’s
revision of Talinum. A list of names, types, literature
sources and other useful information was drawn up.
Vol. 16: Fabaceae. Mr B.D. Schrire who is preparing an
account of the tribe Indigoferae has drawn up a checklist
of Indigofera comprising information on almost 300
taxa. A synopsis of the species of the genus is well
advanced. Some 40 to 50 new taxa need to be described
in the genus. About 140 name changes in Indigofera
were entered into the update of Taxon-PRECIS.
Vol. 23: Lythraceae, Lecythidaceae and Rhizophoraceae.
Except for one uncertain taxon, all taxonomic and
nomenclatural decisions have been taken for the
revision of the Lythraceae. The treatment of the one
species of the Lecythidaceae has been completed. In
Rhizophoraceae manuscripts of the three genera with
a single species each in the region are largely complete.
The fourth genus, Cassipourea, is considered to
comprise four species and three subspecies.
Vol. 24: Prof. E.F. Hennessy at the University of Durban-
Westville has continued her work on Combretum and
Prof. A.E. van Wyk of the University of Pretoria his
work on Myrtaceae and Melostomataceae.
Vol. 25: Ericaceae. Mr E.G.H. Oliver has concentrated
on four of the ‘minor genera’ in the family and has sunk
the southern African members of Philippia under Erica.
This brings the number of species in this genus, the
largest genus of flowering plants on the subcontinent,
to about 660.
Vol. 26: Convolvulaceae: Miss W.G. Welman has adapted
most of the work by Prof. A.D.J. Meeuse to Flora
300
Bothalia 19,2 (1989)
format. Descriptions of ten species not included in
Meeuse’s revision of 1958 have also been completed.
Vol. 30: Pedaliaceae — Gesneriaceae. Prof. H.-D.
Ihlenfeldt of the University of Hamburg, West Germany
and his students have progressed with their work on
Pedaliaceae, Martyniaceae and Orobanchaceae.
Vol. 30: Acanthaceae — Justiciae. The revision of the
genus Justicia by Dr K.L. Immelman, for which she was
awarded a Ph.D. , has been adapted to FSA format and edit-
ed. This genus and the other large genus Monechma still
await the completion of a few minor genera by research-
ers outside the Institute.
Vol. 33: Asteraceae — Inuleae. Miss M. Koekemoer has
started on a revision of Disparago. A preliminary analysis
has shown that ten taxa can be distinguished of which three
are undescribed.
Pretoria Flora
Dr O.A. Leistner completed the family Asteraceae. The
last group dealt with was group 8 comprising 36 species
of Helichrysum and related genera. All families have now
been written up. Some updating of existing manuscript
remains to be done as well as completing the introductory
chapter.
Namaqualand Flora
The contract awarded by the Department to Dr C.
Boucher of the University of Stellenbosch to produce an
identification manual for the flora of Namaqualand was
renewed. After some initial technical problems good
progress has been made with this project.
Palaeoflora of southern Africa
Dr J.M. Anderson and Dr H.M. Anderson have
completed Vol. 2 dealing with the Gymnosperms
(excluding Dicroidium) of the Molteno Formation and the
work has gone to press. Camera-ready copy of 567 pages
has been provided comprising 331 photographic plates
(with a total of 3 386 individual photos), 1 107 line
drawings, 154 tables and 117 maps. The work deals with
111 species belonging to 23 genera. Vol. 3, which will cover
the lower plant groups, including ferns, is in preparation.
Liaison Officer, Kew
Mr B. Schrire ended his term of office and was
succeeded by Mr A. Nicholas. They dealt with 178 queries
and requests from both South Africa and overseas. Mr
Schrire worked out the nomenclature of the estimated 300
species of Indigofera in southern Africa after a study tour
of 15 European herbaria. He is writing up this large genus
with the aid of the DELTA programme. The research work
of Mr Nicholas was mainly focussed on the section
Trichocodon of the asclepiadaceous genus Pachycarpus.
He also delivered a seminar on the Asclepiadaceae to the
School of Botany, Trinity College, Dublin. A selection of
tree paintings from the Botanical Research Institute
exhibited at a Royal Horticultural Society show in London
won a Grenfell Silver-gilt medal.
DATA SUBDIVISION
The subdivision continued to co-ordinate the computer
requirements of the Institute. The Institute’s present
computer systems include two large data storage systems
on the B7900 mainframe of the Department of Agriculture
(PRECIS and PHYTOTAB), SABINET links to IBM
mainframe for the library, as well as numerous smaller
PC systems, including a new IBM compatible PC for the
vegetation ecology section, which replaces the old Hewlett
Packard 9845B. The Herbarium mini system, a Unisys
B26, is also linked via a modem to the B7900 mainframe.
PRECIS, at present under the jurisdiction of the
Herbarium Division, is managed by Dr H.F. Glen and Mr
N.P. Barker. PRECIS has continued to grow, with label
information on Approximately 18 000 specimens added
over the past year. The present number of specimens
catered for by the data base totals 661 000.
No fewer than 82 requests tor information were
received, the majority being from universities (25
requests). Other requests received came from 12 other
government departments, nine individuals, eight other
botanical gardens and five from overseas. There were also
23 requests from staff of the Institute and Units.
Taxon-PRECIS continues to be updated regularly with
new and corrected names, the annual list of which appears
in this edition of Bothalia. The maintenance of this
information, a task co-ordinated by Mrs B.C. de Wet and
Mrs J.M. Mulvenna, is assisted by the herbarium mini
computer which has now been programmed to record all
changes in the PRECIS format. This allows the easy
updating of PRECIS. It is hoped that a similar system will
be programmed in the near future, allowing the label
information for specimen labels typed on the Herbarium
mini system to be formatted and entered directly into
PRECIS-compatible files.
Forty-eight data sets obtained from theses and journals
are presently on the PHYTOTAB data base. Data files
required for the synthesis of these matrices are being
loaded by Mrs W. Jones. Mr R.H. Westfall and Mrs B.C.
de Wet have refined (PHYTO 00 — key construction) and
developed (PHYTO 39 — environmental analysis)
programs for the data base.
Smaller systems continuing on the B7900 include the
Garden Records system, developed by Mrs B.C. de Wet
and maintained by Mrs K.P. Clarke, and the PHOTOS data
base maintained by Miss A.P. Backer. The latter system
holds information on photographic vegetation records for
the Vegetation Ecology Division.
STRUCTURE AND FUNCTION DIVISION
Under the leadership of Dr J.J. Spies and in spite of
limited manpower, this division has had a productive year.
A total of thirteen articles were published in scientific
journals and ten papers, as well as four poster papers, were
read at congresses (of these, one paper and one poster were
presented overseas). Mesdames H. du Plessis and E.J.L.
Saayman shared the prize for the best poster at the
Congress of the South African Genetic Society.
Bothalia 19,2 (1989)
Cytogenetics
During August 1988 Dr J.J. Spies undertook a study tour
to the USA and Canada where he read a paper and
presented a poster at the XVIth International Genetic
Congress in Toronto. A visit to Dr G. Davidse (Missouri
Botanical Garden, St Louis, Missouri) made it possible
to complete an article on the genus Ehrharta and to make
good progress with two other articles. This co-operation
is the result of Dr Davidse’s visit as research associate to
the BRI in 1987.
Work on the grass tribe Arundineae has progressed well
during the past year: three basic chromosome numbers,
i.e. 6, 7 and 13, were found and it has been determined
that large-scale evolution of the karyotype occurs in this
group. This could indicate how the tribe has evolved
and how an unfavourable environment may influence
evolutionary development.
Mrs H. du Plessis studied the cytogenetic aspects of the
important pasture grass, Heteropogon contortus : the
species has a basic chromosome number of 10, both
aneuploidy and polyploidy occur and paracentric
inversions are found in some specimens. These phenomena
explain the tremendous morphological variation in the
species and show that any effort to improve the plant should
be preceded by a cytogenetic study.
Mrs E.J.L. Saayman established that the C3 form of
Alloteropsis semialata is diploid and the C4 form is
hexaploid. Further study is needed to determine whether
the two forms are merely mimics, whether the C4 form
is caused by an additional genome or whether pleiotropism
and/or multifactorial heredity have caused the evolutionary
change.
Mrs E. van der Merwe’s cytogenetic investigation of
certain Digitaria eriantha strains selected for breeding
better strains, indicates that some strains exhibit abnormal
meiotic chromosome behaviour. The associated decline
in fertility renders these strains unsuitable for use in
breeding programmes. The available commercial strains
contain different polyploidy levels. Because diploid plants
are usually more fertile than polyploid plants, natural
selection against polyploidy occurs. The occurrence of
sexual diploid plants entails that seed has to be increased
in isolation in order to diminish variation within strains.
Here one has to guard constantly against inbreeding.
A study of the tribe Ehrharteae showed that the tribe
has a secondary basic chromosome number of x = 12.
Polyploidy occurs and the highest polyploidy level is
decaploid. B-chromosomes occur in some specimens.
Comparative anatomy
Dr R.P. Ellis continued to work with Dr H.P. Linder
of the University of Cape Town on taxonomic studies of
the genus Pentaschistis . A revision of the genus has been
completed. The names of 67 species were typified and 16
new taxa described. The study revealed that, in addition
to spikelet morphology, leaf anatomy is very valuable in
the delimitation of species. An extensive anatomical atlas
of the genus is being compiled, comprising detailed studies
of the glands and the general biology of the genus. They
301
will serve as a basis for further research on the delimitation
of generic groupings in the Arundineae.
Mary Gunn Library
The librarians, Mrs E. Potgieter and B.F. Lategan, have
had a busy year. The demand for library services has
increased remarkably. This fact is illustrated by the
following figures, with those of the previous year indicated
in brackets: books and journals loaned 2 186 (1 750),
interlibrary loans handled 2 139 (2 123), enquiries 3 188
(2 629), photocopies 36 236 (28 360), books and journals
bound 370 (270), books purchased 96 (224) and journals
received 417 (411). Additional manpower is urgently needed
to assist with the interlibrary loans and the associated
photocopying. Apart from the tasks mentioned above, the
entire journal collection was rearranged on new shelves
installed to accommodate the anticipated expansion in the
next decade.
Photography
Mrs A.J. Romanovski has once again experienced a very
busy year as indicated by the following figures: 1 569
photographs were taken, 785 films were developed and
19 078 prints were made, of which 6 994 were for
publication purposes.
VEGETATION ECOLOGY DIVISION
The functions of the Vegetation Ecology Division under
Dr J.C. Scheepers are to study the vegetation of South
Africa and its ecological relations. This work involves
three main aspects: the identification, description,
classification and mapping of the various kinds of
vegetation; study of the ecological relationships of different
kinds of vegetation — with one another and with the
environment — and of the various processes and
mechanisms that determine the behaviour of plant
communities; and the application of such ecological
knowledge to the management and utilization of
vegetational resources (Figure 2).
Members of the Vegetation Ecology Division had the
pleasure and privilege of meeting Professors Sandro and
Erika Pignatti of Italy, during a Workshop on the
phytosociological processing of large data sets and also
during subsequent field excursions during January, 1989.
Transvaal bushveld and forest studies
Mr R.H. Westfall is studying the vegetation ecology of
the Sour Bushveld of the Waterberg area of the central
Transvaal. He has sampled 63 stands representing about
14 different vegetation types. A new method of illustrating
vegetation structure using layer diagrams generated by a
portable computer for each stand was developed. This
method emphasizes the field layer cover as a proportion
of the total layer cover. In only 12 % of the stands sampled
was the field layer cover greater than 50%. The method
could prove very effective in identifying areas with a high
erosion potential.
The vegetation ecology of the Upper Limpopo River
Valley in the western Transvaal bushveld is being
investigated by Mr J.M. van Staden. Particular attention
302
FIGURE 2. — A fence line on the Grootfontein Experimental Farm near
Middelburg, Cape, showing the contrasting effects of winter
(Feb. —Aug.) and summer(Aug.-Feb.) grazing on a Grassy
Shrubland.
is being paid to bush encroachment and its effects on the
grass cover. In this way, base-line data will also be obtained
for monitoring the dynamics of veld condition and trend.
The work of Mr G.B. Deall on the Sabie transect has
been completed. A trilogy of definitive publications is in
press. A number of other texts is in preparation as
unpublished reports and articles for publication.
Coastal studies
With the finalizing of the report on the conservation
priorities in the tract between the Mozambique border and
Sodwana, Dr P.J. Weisser has completed his coverage of
the vegetation of the coastal dunes of KwaZulu. It is
intended to publish this report in due course. Other data
gathered during the subproject are to be worked up and
published as a series of research papers.
The effects of the 1988 floods on the wetlands of the
lower 5 km of the Orange River have been investigated
by Mr M.G. O’Callaghan. In this area, 320 ha of wetland
vegetation was destroyed, owing mainly to the deposition
of sand as bedload. Approximately 50 ha of potential
wetland area was added. To clarify and authenticate
herbarium and other records, over 500 Sarcocornia and
Salicornia specimens from various herbaria were
identified, necessitating 12 000 anatomical sections. It was
found that 2% had been identified as the wrong genus and
30% had been identified as the wrong species. Of the
Bothalia 19,2 (1989)
latter, 9,4% were hybrids, whereas 8,4% were previously
indeterminate.
Cape fynbos studies
The ecological study by Mr H.C. Taylor of Mountain
Fynbos in the Cederberg is well advanced. Field work has
been completed for the northern sector. Phytosociological
classification of the data by means of the PHYTOTAB
program package has been carried to an advanced stage.
Apart from two distinct thicket communities, six major
fynbos groupings have become apparent, reflecting the
major habitats within the study area. Minor refractory
groups are at present receiving closer attention.
Mr DJ. McDonald is making good progress on his
ecological study of the fynbos and other vegetation of the
Langeberg. Field sampling of the first transect through the
Boosmansbos Wilderness Area is complete. Analysis of
the data is well advanced and three vegetation types have
been identified: Wet Fynbos, Mesic-Arid Mountain
Fynbos and Afromontane Forest. Soil samples from
selected releves have been analysed and related to the
different vegetation types. Sampling of a second transect
at Swellendam is well advanced with 41 releves completed.
Grassland studies
Field sampling of the grassland vegetation of the
Amersfoort area of the Eastern Transvaal Highveld has
been completed by Miss B.J. Turner. She experienced
much difficulty in the data processing and the
interpretation of the results to arrive at a practically useful
classification. The data have been analysed and several
communities have been distinguished. The report on this
work is progressing well and is nearing completion.
Mr P.J.J. Breytenbach has made good progress on his
study of the vegetation in the Grootvlei area of the south-
central Transvaal Highveld. He has nearly completed the
field sampling. Preliminary data processing has yielded
very promising results. This study is intended to provide
a link between the eastern and the western sectors of the
Grassland Biome.
Karoo studies
The reconnaissance study of the Karoo and other
vegetation in the Graaff-Reinet and Middelburg areas of
the eastern Karoo Region has been completed by Mr A. R.
Palmer. Eight manuscripts have been prepared for
publication dealing with the syntaxonomy, synecology,
plant-soil relationships, phytogeography and mapping of
vegetation. The syntaxa are classified into five classes, nine
orders and seventeen communities. The classes
distinguished are Grasslands, Karoo Shrublands, Karoo
Dwarf Shrublands, Subtropical Transitional Thicket and
Riparian Thicket (Figure 3). The distribution of syntaxa
corresponds to the steep precipitation gradients
encountered in the study area.
Mr J.F. van Blerk has commenced a reconnaissance
study of the Succulent Karoo vegetation in the Knersvlakte
vicinity of the Van Rhynsdorp-Nuwerus area. This is a
daunting task still in the early stages of preparation of field
work.
Bothalia 19,2 (1989)
303
Central technical support services
During the year, 583 references from 1987 and 1988 have
been encoded and loaded onto ATMS by Miss A.P. Backer.
Several literature searches were undertaken for
researchers.
Further progress has been made on the ecological data
bank by Mrs W. Jones. There are 45 data sets available
on the computer of which 78% of the species names and
49% of the community names are available. The first
retrieval program, viz. PHYTO 200, designed to retrieve
species associated in a releve with a particular species,
is running successfully. One enquiry from outside South
Africa has already been successfully answered with the
aid of this program.
EXPERIMENTAL ECOLOGY DIVISION
The objective of experimental ecology is to provide a
predictive understanding of processes at the ecophysiologi-
cal level of the individual as well as at the population,
community and ecosystem levels. To provide focus within
this wide hierarchical range of systems, the Experimental
Ecology Division, under Dr M.C. Rutherford, has
concentrated on problem-related research of critical
processes of stressed systems. This has concerned specific
problems of the more arid, drought-stressed ecosystems,
the alien invasive plant-stressed ecosystems as well as the
special stresses that develop in the Fynbos Biome through
substrate disturbance.
Various levels of active research liaison continue with
the Botany Department of the University of Cape Town,
with other researchers within the co-operative Fynbos and
Karoo Biome Programmes, and with officers of the Winter
Rainfall and Karoo Regions of the Department of
Agriculture and Water Supply. In this regard. Dr
Rutherford is now co-supervisor, of two Ph.D. degree
projects and one M.Sc. degree project whereas Mr G.W.
Davis is the convenor of the highly successful Working
Group on Commercial Wildflower Resources of the
Fynbos Biome Project under the auspices of the
Foundation for Research Development. This working
group is stimulating effective dialogue between
conservation managers (chiefly from forestry and nature
conservation bodies), biological researchers (from both
universities and state institutions) and commercial
wildflower producers to promote far-sighted planning for
utilization of fynbos veld as a renewable resource.
Fynbos reproductive ecology
Dr C.F. Musil has found that the post-fire reproductive
success, measured in terms of seedling/parent ratios, of
lowland fynbos species with small seeds growing in sand
plain communities infested by alien Acacia saligna was
not significantly different to that in surrounding natural
vegetation. Indigenous species with large seeds exhibited
a significant reduction in post-fire reproductive success
in the alien-infested communities. Efficiency of post-fire
vegetative reproduction among indigenous species in alien-
infested communities was similar to that in surrounding
natural vegetation. Season of burn (autumn or spring) had
no significant effect on total, indigenous seedling
population densities. However, higher densities of
sprouters were found after autumn than after spring burns,
which may be related to increased mortalities among
sprouters during the dry summer period.
Fynbos transformation studies
An experimental plant community at a mountain fynbos
study site, which had been subjected to experimental
disturbance by tilling, was shown by Mr G.W. Davis and
Mr A.R Flynn to retain altered compositional characteris-
tics for at least three years following disturbance. Relative
to an adjacent undisturbed control, it maintained: poorer
species richness; a lower index of diversity (Shannon-
Wiener index); and a higher index of dominance (Simpson
index). Germination in seeds of the dominant shrub species
( Leucadendron xanthoconus ) was shown to be affected by
ambient conditions known to be altered by tillage.
Germination success under controlled environment
conditions was inversely related to water stress, depth of
seed burial, and an ambient daytime temperature greater
or less than an optimum of approximately 15 °C.
FIGURE 3. — Researchers from the
Karoo Biome Project surveying
Dwarf Shrubland near Prince
Albert.
304
Ecophysiology in alien invasive plant-stressed ecosystems
Dr Rutherford’s and Mr J. de W. Bosenberg’s simulation
of Acacia cyclops canopy radiation attenuation by
replicated field shade units has started to identify shade-
sensitive species in the second year of treatment. It appears
that the negative reaction of species, such as
Anthospermum spathulatum, to A. cyclops might be
attributable to light limitation of obligate heliophytic
species. However, several other amensalistic species,
relative to A. cyclops, have not yet been adversely affected
by one-and-a-half years of strongly reduced light levels,
and allelopathic tests are consequently being initiated.
In a comparative study of photosynthetic responses of
A. cyclops and of an indigenous colonizer of similar
habitats, Leucospermum rodolentum. Dr Rutherford, Mr
G.F. Midgley and Mr Bosenberg, have related carbon
uptake patterns to photosynthetically active radiation, leaf
temperature, vapour pressure deficit, stomatal conductance
and xylem pressure potentials. Although the two seedling
populations had similar light compensation points, at high
light intensities L. rodolentum seedlings had much higher
photosynthetic rates and were even less light saturated
when compared with the seedlings of A. cyclops. These
findings are being related to the patterns of establishment
of these species in disturbed habitats.
Nutrient and water relations in Karoo
Mr G.F. Midgley has determined that, in an arid region,
success of certain deciduous shrubs relative to evergreens
may be determined primarily by soil nitrogen status, with
soil water availability a secondary factor. This is possibly
explained by deciduous forms having a higher nutrient
requirement than evergreens. In a field-simulated drought,
succulent plant forms were more negatively affected than
wer# non-succulents. These results suggest that rainfall
frequency and soil nutrient status are important
determinants of vegetation composition in semi-arid parts
of the western Karoo, and that leaf habit and water storage
characteristics may determine the distribution and success
of perennial plant forms in these areas.
Plant population response ecology in Karoo
Mr L.W. Powrie has been constructing a computer
database of important responses to major events in the
autecology of karroid plant populations, using both the
formal, scientific and the largely untapped informal,
undocumented information sectors. To date he has
collected data from 21 agricultural workers in the Karoo
for creating the database. There are data on some 226
worksheets, from which 61 partial species descriptions
have been entered into the computer database. A list of
some 1 250 botanical and common names has been
compiled from several sources. The unusual rains, floods
and large locust outbreaks during 1988, have provided
further opportunities for observing responses of Karoo
plants.
PLANT EXPLORATION DIVISION
The division, under Mr M.J. Wells continued to
concentrate on problem plants and food plants, but the loss
of research staff resulted in greater emphasis being placed
Bothalia 19,2 (1989)
on the scientific information and garden utilization
services.
Conservation of germ plasm
With the field work on Mr T.H. Arnold’s investigation
of tribal crop plants having been completed, no more seed
samples were forthcoming from this source. Seed
previously collected was distributed: 10 samples of Vigna
unguiculata and six of Voandzeia subterranea to France.
Mrs H. Joffe collected 84 seed and 23 vegetative samples
for distribution to local nurseries, in order to encourage
cultivation of indigenous plants.
Indigenous food plants
Mr A. A. Balsinhas abstracted information from 60
publications, bringing the references consulted for the
national food plants data bank to a total of 284. The newly
consulted references contributed information about 2 354
species and resulted in the addition of 96 new names to
the list of southern African food plants which now includes
1 793 species.
Miss S.E. Chadwick prepared a manuscript on Tylosema
esculentum, the 10th of 14 priority food plants of the veld
that are being studied. Photographs and diagnostic
descriptions are required before these texts are ready for
publication. She also prepared the text for a plate in The
Flowering Plants of Africa on another plant with edible
seeds — Citrullus ecirrhosus.
Primitive crop plants of African origin
Mrs L.D. Jacobs completed the preparation of material
resulting from the previous year’s field trips. This included
56 tannin tests and 250 colour tests on Sorghum seed, and
the completion of 215 data sheets on Pennisetum. She also
dissected and prepared slides of spikelets from 61
collections of Sorghum and Pennisetum.
Data was prepared for a paper to be entitled A survey
of frequency of and preference for primitive crops’ in five
ethnic regions (KwaZulu, KwaNdebele, Bophuthatswana,
Capri vi and Kavango). This will complement a survey,
previously completed, on other ethnic regions. In addition
maps were prepared of the occurrence (and preference
ratings) for all regions of the following crops: Sorghum,
Pennisetum, Zea, Citrullus, Cucurbita, Lagenaria,
Voandzeia, Vigna, Arachis, Phaseolus and Cajanus.
Catalogue of problem plants
More information about listed and additional species was
filed by Mr Wells for possible future editions of the
catalogue. The existing catalogue, despite its incomplete-
ness, continues to be invaluable in answering queries
regarding weeds and invaders.
Plant invaders
A series of 14 radio talks on the introduction of invader
plants was prepared and recorded by Mr Wells. Mrs
D.M.C. Fourie also recorded radio talks on our publication
‘Declared weeds and invader plants’. Although Miss L.
Henderson (of the Plant Protection Research Institute) is
Bothalia 19,2 (1989)
305
no longer on our staff we continue to assist her with her
plant invader surveys, where possible, and Mrs H. Joffe
accompanied her on a field trip to the eastern Cape.
Garden utilization
Mrs H. Joffe continued to increase the level of garden
utilization. She made 430 seed collections of which 150
were to meet requests from researchers, 91 were for the
herbarium’s carpological collection, 82 for the spirit
collection, and 23 were channelled back to the nursery
for propagative purposes. She also made 380 collections
of flowering or vegetative material, of which 57 were to
meet requests from local or overseas researchers, 202 were
added to the herbarium spirit collection and 42 were
channelled back to the nursery for propagative purposes.
Eleven collections weye made for the artist to figure.
Colour slides taken, annotated and filed totalled 450. Some
130 herbarium specimens were made, mainly as vouchers
for plants that were photographed or from which seed or
other collections were made.
Scientific information services
Mrs D.M.C. Fourie handled 286 written and 143
telephonic requests for material and information, and dealt
with 32 visitors and four groups. She also catalogued and
filed 442 articles and letters in our special information
files, in preparation for future enquiries.
Among the requests dealt with were advice on new
stamp issues and texts for stamps illustrating indigenous
plants.
A layman 's guide to botanical publications
The draft list was circulated to publishers and to
botanical organizations for comment and suggested
additions. Response was good and about 100 references
were added, bringing the total to 455. The list can now
be finalized and the references classified.
A survey of cultivated plants in South Africa
Ms Susyn Andrews, horticultural botanist from Kew,
was assisted in making contacts with local horticulturalists.
Her survey, when completed, will be of great assistance
in identifying and naming cultivated plants. She has
already shown that many names in use in the trade are
incorrect or out of date.
History of the Botanical Research Institute
Mrs Fourie, who has been gathering historical material
from Miss M.D. Gunn and others, completed the history
of the Institute up until the end of Dr Pole Evans’ tenure
(1939). A talk entitled The early years’ was given to staff
members and a few invited guests. She has been fortunate
in being able to assemble a fascinating photographic
record. Work will now begin on the next period, and
ultimately the completed work will be published.
Liaison services
In the absence of a liaison officer Mrs Fourie accepted
visits of a few groups that she felt merited special
assistance, e.g. black school teachers and overseas
conference delegates. Mr Wells also organized visits from
delegates to the congress of the South African Association
of Botanists. Mrs Joffe assisted a few garden tours, but
many requests for tours of the Institute and garden had
to be turned down.
PRETORIA NATIONAL BOTANICAL GARDEN
Under the direction of the curator, Mr D.H. Dry, the
garden staff were able to continue with new developments,
despite increased garden maintenance needs that resulted
from heavy rains. Coping with aggressive weed growth
was a particular problem.
A field trip to the northern Transvaal, undertaken by
Mr D.S. Hardy and Mr D.J.F. Strydom added valuable live
and herbarium material, as well as seeds to the existing
collections. Mr N.F. van Zyl accompanied by Dr D.
Lehmiller, an American Crinum enthusiast, managed to
collect live material of all Namibian Crinum species bar
one elusive species. Mrs K.P. Clarke, the garden records
officer, recorded 647 new accessions to the garden, of
which about 100 are classified as rare or endangered
species.
Messrs Strydom and N. A. Klapwijk sealed three of the
four earth dams in the general section of the garden with
bentonite clay. The pond in front of the main building was
also sealed and the fountain made operative. The area
around the entrance gate to the main building was
landscaped by using boulders as the main feature. The
fountain and the boulder ‘koppie’ greatly improve the
approaches to the building.
In the Coastal Forest Biome near the Silverton gate, two
new plant beds were made, while additional plantings were
made to other existing beds in the Biome. Mr L.C.
Steenkamp, who recently turned 73, and his team of
workers paved 480 m of new nature trails.
The old extractor fan and obsolete wet wall of the orchid
house were renovated by Messrs Klapwijk and Van Zyl
and installed in the smaller ornamental glasshouse where
they function very well.
Mr Hardy was sponsored to attend the International
Symposium on Bulbous & Cormous Plants, arranged by
the American Plant Life Society at the Irvine campus of
the University of California. He also delivered eight talks
on South African succulents at various venues in the USA.
Mr Van Zyl resigned from the Institute at the end of
February 1989: unfortunately staff with experience in our
garden seem to be very sought after!
BIOSYSTEMATICS DIVISION
This division, under the direction of Dr O. A. Leistner,
has devoted itself largely to the editing and typesetting of
the publications of the Institute. Mrs E. du Plessis assisted
with editing The Flowering Plants of Africa and Flora of
Southern Africa and undertook most of the translation
work for the Institute. Mrs B.A. Momberg assisted with
editing Bothalia and Memoirs of the Botanical Survey of
South Africa. An increased amount of typesetting was done
in-house by Mrs S.S. Brink.
306
Bothalia 19,2 (1989)
Bothalia
Numbers 1 and 2 of Vol. 18 and the index to Vol. 17
were published.
Flora of southern Africa (FSA)
Vol. 16,3,6, dealing with the 278 species of Aspalathus
of the Fabaceae was published.
The Flowering Plants of Africa/Die Blomplante van Afrika
As mentioned in the introduction a new green cover was
designed for this journal with a reduced print of one of
the plates in the journal on the front. Beginning with
volume 50,1 which was published during the year, each
volume will consist of 2 numbers with one number
containing 20 plates published per year.
Palaeoflora of southern Africa
Vol. 2 of the Molteno formation has gone to press.
Memoirs of the Botanical Survey of South Africa
No. 57, the third edition of John Acocks’s (Figure 4)
classical work Veld types of South Africa (Figure 5) was
published. Plant names have been updated and minor
adjustments and corrections made in the text. The text was
set at the Institute. Many new and more relevant photos
were chosen to illustrate veld types.
SEMINARS
The seminar committee, under the chairmanship of Dr
J.J. Spies organised the following lectures:
1988.05.20
Mr J. van Blerk (BRI): Die verwerking van Acocks se
ongepubliseerde velddata met behulp van moderne
rekenaarmetodes.
1988.05.26
Dr D.J.B. Killick (BRI): Linnaean typification.
1988.05.26
Mr T. Arnold (BRI): Computerization of curatorial and
plant identification functions of the National Herbarium.
1988.08.25
Prof. D. Wiens (Department of Biology, University of
Utah, Salt Lake City, Utah 84112, USA): Reproductive
failure in plants: a genetic hypothesis for extinction.
FIGURE 4.— John Phillip Harison Acocks (1911-1979).
1988.09.15
Dr C.L. Calvin (Department of Biology, Portland State
University, Portland, OR 97207, USA): A quantative
approach to studying endophytic system development in
mistletoes.
1988.11.18
Mrs H. du Plessis (BRI): Die gebruik van sitogenetika in
die taksonomie van Tribolium.
1988.12.01
Mrs D. Fourie (BRI): The history of the BRI: the early
years.
1989.02.17
Dr H. Anderson (BRI): The palaeoflora of South Africa
— past, present and future.
FIGURE 5.— The 3rd edition of
Acocks’s Veld types of South
Africa was published at the be-
ginning of 1989. Persons large-
ly involved in the production of
this work were from left to
right: Dr O.A. Leistner (edi-
tor); Mrs B.A. Momberg (as-
sistant editor); Mrs S.S. Brink
(typesetter) and Dr J.C.
Scheepers (photographs).
Bothalia 19,2 (1989)
BOTANICAL RESEARCH INSTITUTE
307
Scientific, Technical and Administrative Staff
(31st March 1989)
Director
B. de Winter, M.Sc., D.Sc. (Taxonomy of Poaceae, es-
pecially Eragrostis, and of Hermannia: plant geography)
Deputy Director
D.J.B. Killick, M.Sc., Ph.D., F.L.S. (General taxonomy;
nomenclature; mountain ecology and editing)
ADMINISTRATION DIVISION
Chief Provisioning Ad-
ministration Clerk ....
Senior Provisioning Ad-
ministration Clerks ...
Personal Secretary to
Director
Senior Registration
Clerk
Registration Clerk
Accounting Clerk
Receptionist
Typists
J.T.C. Snyman
(Head of Division)
Mrs E.S. Smit
Mrs A.E. Engelbrecht
Mrs M.M. Loots
Mrs I.J.H. Joubert
Mrs R.W.R. Koehne
G. Herman
Mrs A.E. van der Merwe
Mrs S.S. Brink
Mrs E.L. Bunton*
Mrs S.M. Thiart
HERBARIUM DIVISION
Wing B (Piperaceae— Oxalidaceae)
Senior Agricultural
Researcher
Senior Agricultural
Researcher
Chief Agricultural
Research Technician ..
Administrative Assistant
III
G. Germishuizen, M.Sc.
(Taxonomy of Polygon-
aceae; plant identifications)
Miss E. Retief, M.Sc.
(Plant identifications)
Mrs B.J. Pienaar, B.Sc. Hons
(Taxonomy of Vigna\ plant
identifications)
C. Letsoalo
Wing C (Linaceae— Asclepiadaceae)
Senior Agricultural
Researcher
Senior Agricultural
Technician
Assistant Agricultural
Research Technician ..
Agricultural Research
Assistant
Herbarium Assistant ....
Mrs C.M. van Wyk, M.Sc.
(Plant identifications)
Mrs M. Jordaan, B.Sc.
(Plant identifications)
N.P. Barker, B.Sc. Hons.
(Taxonomy of Pentameris
and Pseudopentameris)
A. A. Balsinhas**
(Plant identifications)
Mrs J.L.M. Grobler*
Officer-in-Charge T.H. Arnold, M.Sc.
Acting Miss W.G. Welman, M.Sc.
NATIONAL HERBARIUM, PRETORIA (PRE)
Assistant Director T.H. Arnold, M.Sc.
(Curator; taxonomy of
Ficinia)
Herbarium Assistant .... Mrs C.J. van Niekerk
Wing A (Pteridophytes- Monocotyledons)
Senior Agricultural
Researcher
Chief Agricultural
Research Technician ..
Herbarium Assistant ....
Administrative Assistant
in
Miss C. Reid, B.Sc. Hons
(Taxonomy of Carex\ plant
identifications)
Mrs L. Fish, B.Sc.
(Plant identifications)
Mrs S. Burger
S. Makgakga
* Half-day
**Part-time assistance
— Asteraceae)
Wing D (Convolvulaceae
Senior Agricultural
Researcher
Senior Agricultural
Researcher
Principal Agricultural
Research Technician ..
Administrative Assistant
III
Cryptogamic Herbarium
Agricultural Researcher
Assistant Agricultural
Researcher
Chief Agricultural
Research Technician ..
Herbarium Assistant ....
Miss W.G. Welman, M.Sc.
(Plant identifications)
P.P.J. Herman, M.Sc.
(Plant identifications)
Mrs M.J.A.W. Crosby* B.Sc.
(Plant identifications)
F.A. Brusse, M.Sc.
(Lichens)
J. van Rooy, B.Sc. Hons
(Musci)
Mrs S.M. Perold* B.Sc.
(S.E.M. technician;
taxonomy of Ricciaceae)
Mrs L. Filter*
J. Phahla
308
Bothalia 19,2 (1989)
Herbarium services
Senior Agricultural
Technician
Herbarium Assistants ...
TVpist
Administrative Assistant
III
Mrs M. Jordaan, B.Sc.
(Controlling Officer)
Mrs M. Dednam*
(Plant identification services)
Mrs M.Z. Heymann*
(Loans and exchanges)
Miss M. Francis
Mrs M. Cloete*
G. Lephaka
(Preparation and packaging)
NATAL HERBARIUM, DURBAN (NH)
Assistant Agricultural
Researcher
Provisioning Admini-
strative Clerk
Administrative Assis-
tants III
Administrative Assis-
tants II
Miss R. Williams, B.Sc. Hons
(Curator; plant identifica-
tions)
Mrs H.E. Noble*
A. M. Ngwenya
T.B. Sikhakhane
B. M. Mbonambi
S.B. Nzimande (Gardener)
GOVERNMENT HERBARIUM, GRAHAMSTOWN (GRA)
Senior Agricultural
Researcher
Herbarium Assistant ....
Administrative Assis-
tants III
Administrative Assis-
tant I
Mrs E. Brink, B.Sc.
(Curator; plant identifica-
tions)
Mrs L.M. Verwey
A.D. Booi
R. Klaas (Grahamstown
Nature Reserve)
J. Zenzile
GOVERNMENT HERBARIUM, STELLENBOSCH (STE)
Senior Agricultural
Researcher
Assistant Agricultural
Researcher
Agricultural Research
Technician
Administrative Assis-
tants m
E.G.H. Oliver, M.Sc.
(Curator; taxonomy of Erica-
ceae)
Mrs J.B.P. Beyers, B.Sc. Hons
(Plant identifications)
Mrs A.C. Fellingham, B.Sc.
(Plant identifications)
Mrs J. Leith
Miss E. van Wyk
VEGETATION ECOLOGY DIVISION
Assistant Director J.C. Scheepers, M.Sc., D.Sc.
(Vegetation ecology, espe-
* Half-day
Senior Agricultural
Researchers
Agricultural Researcher
Assistant Agricultural
Researchers
Principal Agricultural
Research Technicians .
dally of forest/woodland/
grassland relationships;
conservation and land-use
planning; phytogeography)
D.J. McDonald, M.Sc.
(Mountain fynbos ecology
and phytosociology; Braun-
Blanquet approach and
techniques)
A.R. Palmer, Ph.D.
(Karoo ecology; remote sen-
sing; nature conservation;
vegetation mapping)
H.C. Taylor, M.Sc.
(Mountain fynbos and forest
ecology; Braun-Blanquet ap-
proach and techniques; con-
servation)
P.J. Weisser, Ph.D.
(Forest ecology; air-photo in-
terpretation and mapping;
reedswamp ecology; Zululand
coast dune vegetation; con-
servation)
R.H. Westfall, M.Sc.
(Ecology and phytosociology
of Transvaal bushveld; ecol-
ogical data and literature
storage, retrieval and proces-
sing; syntaxonomic nomen-
clature)
M.G. O’Callaghan, M.Sc.
(Estuarine ecology and phyto-
sociology; land-use planning
and management; nature con-
servation)
P.J.J. Breytenbach, B.Sc. Hons
(Grassland ecology; pasture
science; nature conservation)
Miss B.J. Turner, B.Sc. Hons
(Grassland ecology; pasture
science; nature conservation)
J.F. van Blerk, B.Sc. Hons.
(Succulent Karoo ecology;
ecological literature; pasture
science; photography)
J.M. van Staden, B.Sc. Hons.
(Bushveld ecology; pasture
science; nature conservation;
photography)
Miss A.P. Backer, B.Sc.
(Ecological data processing
and presentation; ecological
literature; nature conservation;
air-photo interpretation and
cartography; photography)
M.D. Panagos, N. Dipl. Agric.
(Bot.Res.)(Computer science;
data processing; sampling
and monitoring vegetation
and environment)
Mrs J. Schaap, H.P.E.D.
(Draughtsmanship and carto-
Bothalia 19,2 (1989)
309
graphy; artwork, layout and
design)
Agricultural Research
Technicians Mrs W. Jones, B.Sc.
(Computer science; ecological
data processing and presen-
tation; remote sensing; air-
photo interpretation and
cartography)
W.J. Myburgh, B.Sc.
(Grassland ecology; pasture
science; nature conservation)
C.M. van Ginkel, N. Dipl.
(Nat. Cons.) (Karoo ecology;
nature conservation; photo-
graphy; remote sensing)
R Zeier, B.Sc.
(Ecological data processing
and presentation; ecological
field methods; ecological
literature; Succulent Karoo
ecology)
Agricultural Research
Assistant Miss H.M.Moolman, TELE. D.
(Technical, editorial and
administrative support func-
tions)
Senior Provisioning
Administration Clerk .. Mrs E.W. Lewis
Agricultural Research
Assistant D.M. de Witt
(Laboratory, field and cura-
torial assistance)
Genera] Assistant S.V. Dolo
PLANT STRUCTURE AND FUNCTION DIVISION
Officer-in-Charge J.J. Spies, M.Sc., Ph.D.
CYTOGENETICS
Assistant Director
Agricultural Researcher
Assistant Agricultural
Researcher
Agricultural Research
Technician
Agricultural Research
Assistant
J.J. Spies, M.Sc., Ph.D.
(Cytogenetics of grasses)
Mrs E. van der Merwe, M.Sc.
(Cytogenetics of Digitaria)*
Mrs H. du Plessis, B.Sc. Hons
(Cytogenetics of grasses)
Mrs E. J. L. Saayman, B.Sc. Hons
(Cytogenetics of grasses)
Miss C.C. Steyn
(Microtechnique)
EXPERIMENTAL ECOLOGY DIVISION
Assistant Director M.C. Rutherford, M.Sc., Ph.D.,
Dipl. Datamet. (Primary pro-
duction ecology of terrestrial
ecosystems; experimental
ecological studies in strand-
veld, fynbos and Karoo)
Senior Agricultural
Researcher C.F. Musil, M.Sc., Ph.D.
(Reproductive ecophysiology
in fynbos)
Agricultural Researchers
G.W. Davis, M.Sc.
(Transformations of fynbos
ecosystems by the wild flower
picking industry)
G.F. Midgley, B.Sc. Hons
(Plant stress ecology in Karoo
ecosystems)
L.W. Powrie, M.Sc.
(Plant population response
ecology in Karoo)
Senior Research
Technicians A.P Flynn, B.Sc.
(Fynbos ecology; plant com-
munity development)
J. de W. Bosenberg,B.Sc. Hons
(Fynbos and Karoo ecology;
monitoring effects of alien
plants on strandveld and fyn-
bos)
* Grassland Research Centre
** Library Services, Department of National Education
t Half-day
COMPARATIVE PLANT ANATOMY
Specialist Scientist R.P. Ellis, M.Sc., D.Sc.
(Anatomy of grasses)
Agricultural Research
Assistant Mrs A.G. Botha
(Microtechnique)
PHOTOGRAPHIC SERVICES
Photographer Mrs A.J. Romanovski
MARY GUNN LIBRARY
Senior Librarian . Mrs E. Potgieter, B.Bibl.**
Library Assistant Mrs B.F. Lategant**
PRETORIA NATIONAL BOTANICAL GARDEN
Chief Agricultural
Research Technician .. D.H. Dry, NTC Dip. (Hort.)
(Curator; special interest
Ocotea bullata and Xerophyta
retinervis)
Chief Agricultural
Research Technicians . D.S. Hardy
(Nursery supervision, succu-
lents and orchids)
D.J.F. Strydom, NTC Dip.
(Hort.), Dip. Rec. PA.
(Supervision northern section
of the garden)
Agricultural Research
Technicians N.A. Klapwijk, NDH
(Supervision southern section
of garden)
310
Bothalia 19,2 (1989)
N.F. van Zyl, NDH
(Propagation for main plant-
ings)
Agricultural Research
Assistant Mrs K.P. Clarke
(Garden records)
Farm Foremen L.C. Steenkamp
(Supervision of labour)
G.R. Lubbe
(Workshop and stores)
PLANT EXPLORATION DIVISION
Assistant Director
Chief Agricultural
Research Technician ..
Senior Agriculture
Research Technician ..
Agricultural Research
Technician
M.J. Wells, M.Sc.
(Weeds research, botanical
horticulture, fynbos utiliza-
tion and conservation)
Mrs D.M.C. Fourie* B.Sc.
(Scientific information ser-
vice)
Mrs H. Joffe* B.Sc.
(Garden utilization)
A. A. Balsinhas**
(Indigenous food plant data
bank)
FLORA RESEARCH DIVISION
O.A. Leistner, M.Sc., D.Sc.,
F.L.S.
H.F. Glen, M.Sc., Ph.D.
(Taxonomy of Aloe)
J.M. Anderson, M.Sc., Ph.D.
(Palaeobotany, plant geogra-
phy)
* Half-day
** Part-time assistance
*** Biometry & Datametrics
Assistant Director
Senior Agricultural
Researcher
Assistant Specialist
Scientist
Agricultural Researchers
Assistant Agricultural
Researcher
Agricultural Research
Technician
Graphic Artist
Agricultural Research
Assistant
Miss K.L. Immelman, M.Sc.,
Ph.D. (Taxonomy, especially
Acanthaceae, Lythraceae,
Urticaceae)
B.D. Schrire, M.Sc.
(Taxonomy of Fabaceae,
electronic data processing)
H . M . Anderson, M . Sc. , Ph. D.
(Palaeobotany)
Miss M. Koekemoer, B.Sc.
Hons (Grass species mono-
graphs)
Miss G.C. Condy, M.A.
Mrs W.J.G. Roux*
(Plant distributions, specimen
administration)
DATA SUBDIVISION
Data Officer
Agricultural Research
Technician
Data typist
Agricultural Research
Assistants
T.H. Arnold, M.Sc.
Mrs B.C. de Wet, B.Sc., B.A.,
H.D.L.S.***
(Garden records, program-
ming for PHYTOTAB and
taxon-PRECIS)
Mrs J.M. Mulvenna
(Data input)
Mrs E.B. Evenwel
(Quality control for PRECIS)
Mrs B.J. Harris
(New specimen encoder for
PRECIS)
BIOSYSTEMATICS DIVISION
Assitant Director
Senior Liaison Officer . .
Senior Agricultural
Research Technician ..
O.A. Leistner, M.Sc., D.Sc.,
F.L.S. (Editing)
Mrs E. duPlessis, B.Sc. Hons,
S.E.D. (Editing and trans-
lating)
Mrs B.A. Momberg* B.Sc.
(Editing)
PUBLICATIONS BY THE STAFF
(1988.04.01-1989.03.31)
BOUCHER, C. & SHEPHERD, PA. 1988. Plant communities of the
Pella site. In M.L. Jarman, A description of the Fynbos Biome
Project intensive study site at Pella : 38 —76. Occasional Report
No. 33. Foundation for Research Development, CSIR, Pretoria.
BRUSSE, F.A. 1988a. Five new species of Parmelia (Parmeliaceae,
lichenized Ascomycetes) from southern Africa, with new
combinations and notes, and new lichen records. Mycotaxon 31:
533-555.
BRUSSE, F.A. 1988b. Lithoglypha , a new lichen genus from Clarens
Sandstone. Bothalia 18: 89—90.
BRUSSE, F.A. 1988c. A new species of Porpidia from the Drakens-
berg. Bothalia 18: 93-94.
BRUSSE, F.A. 1988d. Schizodiscus , a new porpidioid lichen genus from
the Drakensberg. Bothalia 18: 94-96.
BRUSSE, F.A. 1989. Two new species of Parmelia (Parmeliaceae,
Lichenes), further new combinations and notes and additional
new lichen records from southern Africa. Mycotaxon 34:
399-406.
CODD, L.E. 1988a. Resuscitation of Syncolostemon ramulosus E. Mey.
ex Benth. Bothalia 18: 92—93.
CODD, L.E. 1988b. Obituary: Robert Allen Dyer (1900-1987). Botha-
lia 18: 131-133.
COWLING, R.M., CAMPBELL, B.M., MUSTART, P, McDONALD,
D.J., JARMAN, M.L. & MOLL, E.J. 1988. Vegetation
classification in a floristically complex area: the Agulhas Plain.
South African Journal of Botany 54: 290—300.
Bothalia 19,2 (1989)
311
DAVIDSE, G. 1988. A revision of Prionanihium (Poaceae: Arundineae).
Bothalia 18: 143—153.
DEALL, G.B. 1988. Review: Planting a Bible garden, by F. Nigel Hepper
Bothalia 18: 135.
DRY, D.H. 1988a. Gerhard groet as lektor. Parks Administration Apr : 14
DRY, D.H. 1988b Danie Dry se. Parks Administration Apr 14—16.
DRY, D.H. 1988c. Redaksioneel: Hoer produktiwiteit. SAVKO NT Nuus
July: £.
DRY, D.H. 1988d. Ervaring deur deelnemende bestuur. SAVKO NT Nuus
July: 20.
DRY, D.H. 1988e. Editorial: Co-operative education: how well are we
coping? SASCE NT News July: 2.
DRY, D.H. 1989a. Redaksioneel: SAVKO NT Nuus ontgroei homselP.
SAVKO Nuus 1:2.
DRY, D.H. 1989b. Redaksioneel: Nasionale simposium. SAVKO Nuus 1:2.
DRY, D.H. 1989c. Editorial: Is a new system of education viable? SASCE
News 1: 2.
DU PLESSIS, E. 1988. Review: Dictionary of southern African place
names, by P.E. Raper. Bothalia 18: 135.
DU PLESSIS, H. & SPIES, J.J. 1988. Chromosome studies on African
plants. 8. Bothalia 18: 119—122.
ELLIS, R.P. 1988a. Leaf anatomy of the South African Danthonieae
(Poaceae). XVI. The genus Urochlaena. Bothalia 18: 101—104.
ELLIS, R.P. 1988b. Leaf anatomy of the South African Danthonieae
(Poaceae). XVII. The genus Chaetobromus. Bothalia 18:
195-209.
ELLIS, R.P. 1988c. Leaf anatomy and systematics of the genus Panicum
(Poaceae: Panicoideae) in southern Africa. Monographs in
Systematic Botany 25: 129—156.
ELLIS, R.P. 1988d. A review of comparative leaf blade anatomy in the
systematics of the Poaceae: the past twenty-five years. In T.R.
Soderstrom et al.. Grass systematics and evolution: 3—10.
Smithsonian Institution Press, Washington D.C.
GERMISHUIZEN, G. 1988a. Oxvgonum dregeanum complex. Bothalia
18: 173-181.
GERMISHUIZEN, G. 1988b. Reinstatement of Oxygonum acetosella.
Bothalia 18: 187.
GERMISHUIZEN, G. 1988c. Adenolobus pechuelii subsp. pechuelii.
The Flowering Plants of Africa 50: t. 1979.
GERMISHUIZEN, G. & GLEN, H.F. 1988. Bauhinia natalensis. The
Flowering Plants of Africa 50: t. 1980.
GIBBS RUSSELL, G.E. 1988. Distribution of subfamilies and tribes
of Poaceae in southern Africa. Monographs in Systematic Bota-
ny 25: 555-566.
GIBBS RUSSELL, G.E. 1989. The PRECIS computer system and its
applications for the Flora of southern Africa. In N. Morin,
Floristics for the 21st Century: the report. American Institute of
Biological Sciences, Washington DC.
GIBBS RUSSELL, G.E. & ELLIS, R.P. 1988. Taxonomy and leaf
anatomy of the genus Ehrharta (Poaceae) in southern Africa: the
Dura group. Bothalia 18: 165—172.
GIBBS RUSSELL, G.E. & SPIES, J.J. 1988. Variation in important South
African pasture grasses: I. Morphological and geographic
variation. Journal of the Grassland Society of Southern Africa
5: 15-21.
GIBBS RUSSELL, G.E., WELMAN, W.G., GERMISHUIZEN, G.,
RETIEF, E., PIENAAR, B.J., REID, C., FISH, L., VAN ROOY,
J.. VAN WYK, C.M. & KALAKE, E. 1988. New taxa, new
records and name changes for southern African plans. Bothalia
18: 293-304.
GLEN, H.F. 1988a. Astridia velutina. The Flowering Plants of Africa
50: t. 1970.
GLEN, H.F. 1988b. Aloe plicatilis. The Flowering Plants of Africa 50:
t. 1972.
GLEN, H.F. & HARDY, D.S. 1988a. The identity of Aloe penduliflora
Bak. Kew Bulletin 43: 523—529.
GLEN, H.F. & HARDY, D.S. 1988b. Aloe capitata var. capitata. The
Flowering Plants of Africa 50: t. 1973.
GLEN, H.F. & HARDY, D.S. 1988c. Nepenthes madagascariensis. The
Flowering Plants of Africa 50: t. 1974.
HARDY, D.S. 1988a. Stapelia clavicorona. Aloe 25: 36.
HARDY, D.S. 1988b. Pterodiscus aurantiacus , a case of mistaken iden-
tity. Aloe 25: 2d.
HARDY, D.S. 1988c. Aloe haworthioides. The Flowering Plants of Africa
50: t. 1971.
IMMELMAN, K.L. Justicia betonica. The Flowering Plants of Africa
50: t. 1969.
KILLICK, D.J.B. 1988a. Review: The botany of the southern Natal
Drakensberg, by O.M. Hilliard & B.L. Burtt. Bothalia 18:
135-136.
KILLICK, D.J.B. 1988b. Review: The botany of the Commelins, by DO.
Wijnands. Bothalia 18: 330-332.
KILLICK, D.J.B. 1988c. Adansonia digitata L. The Flowering Plants
of Africa 50: t. 1967.
KILLICK, D.J.B. 1988d. Streptocarpus hilsenbergii R. Br. var.
hilsenbergii. The Flowering Plants of Africa 50: t. 1975.
KILLICK, D.J.B. & DU PLESSIS, E. 1988. History of The Flowering
Plants of Africa and an index to volumes 1 to 49. The Flowering
Plants of Africa. Index to Vols 1-49, pp. 44.
KILLICK, D.J.B. & OSBORNE, R. 1988. The National Herbarium.
Encephalartos 16: 26—21.
LEISTNER, O.A. 1988a. Podocarpus latifolius. The Flowering Plants
of Africa 50: t. 1965.
LEISTNER, O.A. 1988b . Podocarpus elongatus. The Flowering Plants
of Africa 50: t. 1966.
LEISTNER, O.A. 1988c. Translation of obituary: Herman Merxmiiller
(1920-1988), by Jiirke Grau. Bothalia 18: 325-327.
LEISTNER, O.A. 1988d. Review: Flora Zambesiaca. Vol. 8, Part 1,
by E. Launert. South African Journal of Botany 54 : 307.
MACDONALD, I.A.W., CLARK, D.L. & TAYLOR, H.C. 1989. The
history and effects of alien plant control in the Cape of Good
Hope Nature Reserve, 1941—1987. South African Journal of Botany
55: 56-75.
McDONALD, Of. 1988. A synopsis of the plant communities of
Swartboschkloof, Jonkershoek, Cape Province. Bothalia 18:
233-260.
McDONALD, D.J. & MORLEY, M. 1988. a checklist of the flowering
plants and ferns of Swartboschkloof, Jonkershoek, Cape Province.
Bothalia 18: 261-270.
O'CALLAGHAN, M.G. 1988. Soutmoerasse van groot belang. In Weskus
Nasionale Park — Supplement to Die Burger , 19th Aug. 1988.
OLIVER, E.G.H. 1988a. Studies in the Ericoideae (Ericaceae). VI. The
generic relationship between Erica and Philippia in southern
Africa. Bothalia 18: 1-10.
OLIVER, E.G.H. 1988b. Coilostigma zeyherianum — a correction.
Bothalia 18: 185.
OLIVER, E.G.H. 1988c. Aristea biflora. The Flowering Plants of Africa
50: t. 1978.
OLIVER, E.G.H. 1989. Bot. Soc's active octogenarians: congratulations
Dr. Codd. Veld & Flora 74: 130.
PALMER, A.R. 1988a. Nature reserves and hiking trails. In R.A. Lubke,
F.W. Gess & M.N. Bruton, A field guide to the Eastern Cape
Coast: 429-438. Wildlife Society of Southern Africa, Gra-
hamstown.
PALMER, A.R. 1988b. Ecoclass: landscape classification using an expert
system. Expert 88. Proceedings of a symposium. CSIR, Pretoria.
PALMER, A.R., CROOK, B.J.S. & LUBKE, R.A. 1988. Aspects of
the vegetation and soil relationships in the Andries Vosloo Kudu
Reserve, Cape Province. South African Journal of Botany 54:
309-314.
PANAGOS, M .D. 1988. The role of the technician in botanical research.
SASCE NT News July: 12-13.
PEROLD, S.M. & VOLK, O H. 1988a. Studies in the genus Riccia
(Marchantiales) from southern Africa. 8. R. campbelliana
(subgenus Riccia), newly recorded for the region. Bothalia 18:
37-42.
PEROLD, S.M. & VOLK, O H. 1988b. Studies in the genus Riccia
(Marchantiales) from southern Africa. 9. R. nigrella and the status
of R. capensis. Bothalia 18: 43-49.
PIENAAR, B.J. & NICHOLAS, A. Heliophila comellsbergia, a new
species from the Richtersveld. Bothalia 18: 183-185.
REID, C. 1988. Review: Flora of Australia (Vol. 45), Hydatellaceae to
Liliaceae, by Alexander S. George. Bothalia 18: 330.
RETIEF, E. & NICHOLAS, A. 1988. The cigarette beetle Lasioderma
serricome (F.) (Coleoptera: Anobiidae): a serious herbarium pest.
Bothalia 18 : 97—99.
RUSSELL, S. & VAN ROOY, J. 1988. The Bryoflora of Namibia.
Monographs in Systematic Botany 25: 453—460.
SCHRIRE, B.D. 1988. A synopsis of the tribe Desmodieae (Fabaceae)
in southern Africa. Bothalia 18: 11—24.
SODERSTROM, T.R. & ELLIS, R.P. 1988a. The woody bamboos
(Poaceae: Bambuseae) of Sri Lanka: a morphological anatomical
study. Smithsonian Contributions to Botany 72: 1—75.
SODERSTROM, T.R. & ELLIS, R.P. 1988b. The position of bamboo
genera and allies in a system of grass classification. In T.R.
Soderstrom et al.. Grass systematics and evolution: 225—238.
Smithsonian Institution Press, Washington D.C.
SPIES, J.J. & DU PLESSIS, H. 1988. Chromosome studies on African
plants. 6. Bothalia 18: 111—114.
312
Bothalia 19,2 (1989)
SPIES, J.J. & GIBBS RUSSELL, G.E. 1988. Variation in important South
African pasture grasses: II. Cytogenetic and reproductive
variation. Journal of the Grassland Society of southern Africa
5: 22-25.
SPIES, J.J. & VOGES, S.P. 1988. Chromosome studies on African plants.
7. Bothalia 18: 114-119.
TAYLOR, H.C. 1988. Review: The ecology and management of biological
invasions in southern Africa, by I.A.W. MacDonald, F.J. Kruger
& A. A. Ferrar. Bothalia 18: 329—330.
VAN WYK, B-E., NOVELLIE, P.A. & VAN WYK, C.M. 1988. Flora
of the Zuurberg National Park. 1. Characterization of major
vegetation units. Bothalia 18: 211-220.
VAN WYK, B-E., VAN WYK, C.M. & NOVELLIE, P.A. 1988. Flora
of the Zuurberg National Park. 2. An annotated checklist of ferns
and seed plants. Bothalia 18: 221—232.
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. argenteolimbata and R. albornata. Bothalia 18: 155 —
163.
WEISSER, P.J., BACKER, A.P. & VAN EEDEN, S. 1988. Aerial
photographs as a long-term data source for vegetation studies.
In I.A.W. MacDonald & R.J.M. Crawford, Long-term data series
relating to southern Africa ’s renewable natural resources. South
African National Scientific Programmes Report No. 157:
251-252. CSIR, Pretoria.
WELLS, M.J. 1988. Smodingium argutum. The Flowering Plants of Africa
50: t. 1968.
WESTFALL, R.H. & DE WET, B.C. 1988. New programs for
preliminary sequencing of species and releves in phytosociological
data sets. Bothalia 18: 122-123.
WESTFALL, R.H. & PANAGOS, M.D. 1988. The plant number
scale — an improved method of cover estimation using variable-
sized belt transects. Bothalia 18: 289-291.
Bothalia 19,2: 313-318 (1989)
OBITUARY
INEZ CLARE VERDOORN (1896-1989)
Friends and colleagues were saddened to hear of the
death on the 2nd April 1989 of Inez Clare Verdoom (Figure
1), the doyenne of South African botany.
She was born in Pretoria on the 15th June 1896. Her
father was a surveyor’s draughtsman and her mother was
the sister of the noted Afrikaans writer, poet, lawyer and
naturalist, Eugene Marais.
During her early school years Inez was troubled by ill-
health. She later attended the Pretoria High School for
Girls and the Loreto Convent where she matriculated in
1916. In her last year at school she wrote an essay which
was to prove crucial to her career and to have a marked
influence on botanical science in South Africa. After a
school visit to the Division of Botany, which at the time
was headed by the dynamic Dr Illtyd Pole Evans, she
penned her impressions of the visit and these were included
in the school’s letter of thanks to the Division. When the
post of herbarium assistant fell vacant, Pole Evans was
determined to acquire the services of Inez Verdoorn. Af-
ter matriculating she had started work in the war office
of Transport and Remounts, but thinking this might not
be a sufficiently permanent position, she transferred to
the Controller and Auditor General’s office. When she was
informed of the vacant herbarium post, she was unsure
as to whether she should make the change, as she was hap-
py in her work in spite of finding it rather dull. But Pole
Evans, who had decided that Inez Verdoorn should join
his Division, used his influence to obtain her transfer to
his staff.
She started work at the Division on the 1st April 1919.
As an herbarium assistant she was responsible for the
pressing, drying and mounting of hundreds of specimens
(Figure 2). However, Inez’s inherent curiosity did not allow
her to settle happily into this mundane work. One day she
asked Miss Sydney Stent, the curator of the herbarium,
why two plants that looked so similar should have two
completely different names. The plants in question were
a species of Chrysanthemum and one of Pentzia. She was
instructed to take Volume 3 of Flora capensis and find out
for herself. With a primitive microscope Inez studied the
dissected flowers and in her own words ‘I can’t tell you
what it did to me — it was so wonderful’. She had found
the path she was to follow for the rest of her life. And
it was perhaps during these early formative years in the
herbarium that the qualities which were to make her great
became evident. Her sense of humour stood her in good
stead as she was teased endlessly by her more learned col-
leagues Sydney Stent and Zoe Findley. But the atmosphere
in the herbarium was one of real happiness together with
complete dedication to the work and Inez must have played
a large part in the cameraderie that developed amongst
the members of staff. She had a genuine love for and in-
terest in people from all walks of life, was a devout Chris-
tian, and when in the years to come she was to be honoured
in many ways, her abiding humility was to prove one of
her greatest attributes.
In 1920 Inez tried to obtain a higher qualification and
enrolled as a part-time student at the Transvaal University
College. Here she coped well with botany and zoology
but found chemistry a problem. It was difficult to manage
the extramural work and ill-health was again causing
problems. Pole Evans advised her to forget about university
and concentrate on her herbarium work. It was therefore
left to her to provide her own botanical education. By 1925
the young botanist had achieved such a degree of
proficiency that she was considered a suitable successor
FIGURE I. — Inez Clare Verdoom
(1896—1989), still at work in
1978.
314
Bothalia 19,2 (1989)
FIGURE 2. — Inez Verdoorn and Jim
Howlett in ‘New Herbarium’,
± 1923.
to the first South African Liaison Officer at the Royal
Botanic Gardens, Kew. Her work at this mecca of the
botanical world was to serve as a further inspiration to
her research.
On her return to South Africa in 1927 she completed
one of her important early works: a revision of the genus
Crotalaria in south and south-east tropical Africa. Her
first scientific publication had been ‘The genus Fagara
which appeared in 1919. In total she published more than
300 articles and papers (Appendix). Amongst the more
important of these were revisions of genera which, apart
from those already mentioned, included Plinthus, Crinum,
Waltheria, Cola, Melhania and Hermannia subgenus
Hermannia. Families revised for the Flora of southern
Africa were Oleaceae, Salvadoraceae, Loganiaceae,
Gentianaceae, Zamiaceae and Welwitschiaceae.
Having been responsible for her own botanical education,
Inez was always aware of the need to teach laymen about
the wonders of the plant world. A large number of her
publications were prepared with this communication link
in mind. Amongst these were An introduction to botany
and to a few Transvaal flowers (1934), Edible wild fruits
of Transvaal (1938), articles for a number of primary
school books, as well as a large number of articles in
popular journals. She also contributed extensively to
botanical articles in the twelve volumes of the Standard
encyclopedia of South Africa (1970—1972). She was the sin-
gle most important contributor of text in the Flowering
Plants of Africa series and provided a formidable total of
159 descriptions to accompany the botanical illustrations.
Inez Verdoorn is considered to be the main instigator and
stimulus behind the Flora of southern Africa project
(Figure 3). Her last major generic revision, that of
Hermannia subgenus Hermannia, was published in 1980.
With the publication of a few minor papers up to 1984 she
achieved a time span of 65 years between her first and
final papers. This is a record of scientific productivity
which will not easily be matched. In recognition of her
achievements the following publications were dedicated
to her: Volume 28 of Flowering Plants of Africa, Volume
40 of the Journal of South African Botany and Eve Palmer’s
Afield guide to the trees of southern Africa.
In 1944 Inez became one of only two employees in the
Department of Agriculture to be promoted to a professional
post without the prescribed academic qualifications. She
was placed in charge of the National Herbarium with the
rank of Senior Professional Officer, a post she held until
her retirement in 1951. This bold step by bureaucracy was
vindicated when her achievements in the field of botany
were acknowledged with the award of an honorary Ph.D.
from the University of Natal in 1967.
As a member of various scientific societies Inez met
researchers in many fields of biological science. She was
always grateful to have learned so much from and been
in touch with so many of the noted scientists of the early
period of her work. People such as Pijper (Pathology),
Jansen and Smit (Entomology), Theiler (Veterinary
Science), Broom and Dart (Palaeontology) and Bigalke
(Zoology) were to have a profound influence on the young
girl who avidly listened to all the papers presented at
scientific meetings. Inez was to offer her services to two
of these scientific societies in the years to come. She was
President of the South African Biological Society in 1956
and President of Section B of the South African
Association for the Advancement of Science in 1963—1964.
She was also a founder member of the South African
Association of Botanists as well as a member of the South
African Association of University Women, the Association
for the Taxonomic Study of the Flora of Tropical Africa
(AETFAT) and the South African Council of English
Education. She was honoured by some of these associa-
tions with the presentation of a number of awards. She
received the Senior Captain Scott Medal from the South
African Biological Society in 1952 and the South African
Association for the Advancement of Science presented her
with their Certificate of Merit in 1977 and their Silver
Medal in 1980. In 1981 she received the Senior Medal for
Botany from the South African Association of Botanists.
Bothalia 19,2 (1989)
315
Inez had a deep love of the veld and, in spite of
difficulties with transport in the early days, she managed
to make frequent field trips. Through the years she
personally collected more than 4 000 botanical specimens
and she joined forces with other collectors such as Codd
and Dyer in expeditions which were to greatly increase
the holdings of the National Herbarium. She was
responsible for describing numerous new species. One of
the highlights of her career was the description of a new
Encephalartos species in 1945 which she named eugene-
maraisii in honour of her uncle who had been the first
person to inform her about the cycad in the Waterberg.
She herself was commemorated by Dr Percy Phillips, her
one-time Chief, who named the genus Inezia after her.
A number of species named ‘verdoorniae’ also
acknowledge her contribution to botany. She was honoured
in the entomological world when a beetle found on the
cycad she had named after her uncle was named
Apinotropis verdoornae. This new genus of beetle was
found at the same time that she collected the cycad in 1944.
Following her retirement in 1951 Inez worked in a
temporary capacity until 1968 after which she continued
with her botanical research for a further 12 years. In her
late eighties she would still board the bus which
transported her from central Pretoria to the Botanical
Research Institute which had become her second home.
In all weathers she walked the long path through the garden
to and from the herbarium and the bus stop.
The esteem in which Inez Verdoorn was held by her
colleagues is illustrated in a statement made by Dr R. Allen
Dyer, her close friend and Chief, when he proposed that
the University of Natal award her with an honorary
doctorate. He wrote: ‘I received a number of scientific
honours over the years but I have no hesitation in saying
that much of what I achieved in my profession was due
to her stimulating personality and example’. Throughout
her career she had certainly set an example to both young
and old botanists. She herself emphasized the fact that she
had been inspired by three men: Dr Illtyd Pole Evans, Sir
Arnold Theiler and General Jan Christiaan Smuts. She felt
that these men epitomised the qualities of hard work,
integrity, a love of nature and the environment, a love of
humanity and a national pride. The example they set was
one that she followed diligently. Her qualities are well
summed up in the motivation read at the presentation of
her doctorate in 1967: ‘by her dedication to the ideals of
science, by her modest and unobtrusive dignity and her
resolution in defending those ideals, by the outstanding
quality of her work, and her kindliness and courtesy as
a botanical guide, philosopher and friend to presidents,
prime ministers, professors, students and scholars she has
become South Africa’s most widely respected botanist’.
If Inez had ever made a plea for a long and fruitful life
it could not have been more aptly put than in a verse by
the botanical artist Cythna Letty, an associate and lifelong
friend:
God grant me many years
That I may roam
Unhindered on the Great Karroo, to know
Each plant in flower
Each bush in bloom
She was indeed granted those years and used them to the
full.
;
FIGURE 3. — Publication in 1963 of Volume 26 of the Flora of southern Africa , the first volume in the series. From left. Dr R.A. Dyer,
Dr L.E. Codd, I.C. Verdoorn and Dr B. de Winter.
316
Bothalia 19,2 (1989)
PUBLICATIONS OF I.C. VERDOORN
THE FLOWERING PLANTS OF (SOUTH) AFRICA
VERDOORN, I.C. 1940a. Lobelia rosulala. Vol. 20: t. 790.
—1940b. Gladiolus varius var. elatus. Vol. 20: t. 791.
—1940c. Echinops amplexicaulis. Vol. 20: t. 796.
— 1940d. Thunbergia gibsonii. Vol. 20: t. 800.
—1941a. Thunbergia erecta. Vol. 21: t. 801.
—1941b. Streptocarpus grandis. Vol. 21: t. 803.
—1941c. Streptocarpus reynoldsii, sp. nov. Vol. 21: t. 804.
— 1941d. Sansevieria desertii. Vol. 21: t. 806.
— 1941e. Stapelia mcloughlinii , sp. nov. Vol. 21: t. 812.
— 1941f. Streptocarpus johannis, sp. nov. Vol. 21: t. 813.
— 1941g. Haworthia umbomboensis , sp. nov. Vol. 21: t. 818.
— 1941h. Verbascum temacha , sp. nov. Vol. 21: t. 834.
— 1941i. Dierama reynoldsii sp. nov. Vol. 21: t. 836.
—1943a. Aloe ortholopha. Vol. 23: t. 882.
—1943b. Cryptostephanus vansonii, sp. nov. Vol. 23: t. 885.
—1943c. Crinum buphanoides. Vol. 23: t. 887.
—1943d. Dichrostachys glomerata. Vol. 23: t. 894.
— 1943e. Hypericum sonderi. Vol. 23: t. 897.
— 1943f. Nautochilus labiatus. Vol. 23: t. 901.
— 1943g. Striga elegans. Vol. 23: t. 907.
— 1943h. Aloe reitzii. Vol. 23: t. 911.
— 1943i. Stapelia meintjiesii , sp. nov. Vol. 23: t. 917.
—1944a. Staavia glutinosa. Vol. 24: t. 927.
—1944b. Bulbinella setosa. Vol. 24: t. 939.
—1944c. Eucomis pole-evansii . Vol. 24: t. 953.
— 1944d. Eucomis humilis. Vol. 24: t. 954.
— 1944e. Eucomis vandermerwei, sp. nov. Vol. 14: t. 955.
—1946a. Halleria lucida. Vol. 25: t. 961.
—1946b. Kalanchoe crundallei, sp. nov. Vol. 25: t. 967.
—1946c. Cissus unquiformifolius . Vol. 25: t. 972.
— 1946d. Combretum microphyllum. Vol. 25: t. 978.
— 1946e. Haworthia pallida. Vol. 25: t. 989.
— 1946f. Barleria obtusa. Vol. 25: t. 998.
—1947a. Encephalartos villosus. Vol. 26: t. 1001, 1002.
—1947b. Albuca transvaalensis . Vol. 26: t. 1009.
—1947c. Aloe trichosantha. Vol. 26: t. 1014.
— 1947d. Aloe debrana. Vol. 26: t. 1016.
— 1947e. Bulbine tortifolia , sp. nov. Vol. 26: t. 1019.
— 1947f. Hypoxis rigidula. Vol. 26: t. 1021.
— 1947g. Clematopsis kirkii. Vol. 26: t. 1026.
— 1947h. Psoralea pinnata var. latifolia. Vol. 26: t. 1029.
—1948. Bulbine stenophylla , sp. nov. Vol. 27: t. 1044.
—1949a. Plectranthus ciliatus. Vol. 27: t. 1051.
—1949b. Encephalartos ngoyanus, sp. nov. Vol. 27: t. 1054.
-1949c. Eulophia complanata , nom. nov. Vol. 27: t. 1056.
-1949d. Hypoxis nitida , sp. nov. Vol. 27: t. 1058.
— 1949e. Catophractes alexandri. Vol. 27: t. 1060.
-1949f. Encephalartos lebomboensis, sp. nov. Vol. 27: t. 1079.
—1950a. Ximenia cajfra. Vol. 28: t. 1081.
—1950b. Caralluma camosa. Vol. 28: t. 1085.
—1951a. Crotalaria recta. Vol. 28: t. 1104.
—1951b. Celsia brevipedicellata. Vol. 28: t. 1105.
—1951c. Oncoba spinosa. Vol. 28: t. 1111.
— 1951d. Sansevieria dooneri. Vol. 28: t. 1114.
—1952a. Eulophia watkinsonii. Vol. 29: t. 1127.
—1952b. Lissochilus platypetalus. Vol. 29: t. 1128.
—1952c. Polystachya golungensis. Vol. 29: t. 1131.
— 1952d. Eulophia hians. Vol. 29: t. 1132.
— 1952e. Habenaria aberrans. Vol. 29: t. 1133.
— 1952f. Ochna pulchra. Vol. 29. t. 1139.
—1953a. Dombeya rotundifolia . Vol. 29: t. 1143.
—1953b. Crinum graminicola, sp. nov. Vol. 29: t. 1155.
—1953c. Crinum bulbispermum. Vol. 29: t. 1159.
—1954a. Erica drakensbergensis . Vol. 30: t. 1161.
—1954b. Eulophia cucullata. Vol. 30: t. 1171.
—1954c. Disa nervosa. Vol. 30: t. 1173.
— 1954d. Disa patula var. transvaalensis. Vol. 30: t. 1174.
— 1954e. Acampe pachyglossa. Vol. 30: t. 1175.
—1955a. Menodora africana. Vol. 30: t. 1187.
—1955b. Galtonia viridiflora, sp. nov. Vol. 30: t. 1188.
—1955c. Polygala virgata. Vol. 30: t. 1192.
— 1955d. Hexalobus glabrescens. Vol. 30: t. 1195.
—1956a. Erica holtii. Vol. 31: t. 1202.
—1956b. Celtis africana. Vol. 31: t. 1210.
—1956c. Acacia karroo. Vol. 31: t. 1220.
— 1956d. Schrebera argyrotricha. Vol. 31: t. 1228.
— 1956e. Eulophia clitellifera. Vol. 31: t. 1235.
—1957a. Strychnos usambarensis. Vol. 32: t. 1242.
—1957b. Amorphophallus abyssinicus. Vol. 32: t. 1251.
—1957c. Polystachya tayloriana. Vol. 32: t. 1256.
— 1957d. Polygala myrtifolia. Vol. 32: t. 1259.
—1958a. Salacia rehmannii. Vol. 32: t. 1270.
—1958b. Jasminum multipartitum. Vol. 32: t. 1272.
-1959a. Watsonia densiflora. Vol. 33: t. 1293.
—1959b. Watsonia densiflora, colour form. Vol. 33: t. 1294.
—1959c. Bauhinia esculenta. Vol. 33: t. 1311.
— 1959d. Nervilia grandiflora. Vol. 33: t. 1312.
— 1959e. Cyrtorchis praetermissa. Vol. 33: t. 1313.
— 1959f. Albuca aperta, sp. nov. Vol. 33: t. 1314.
— 1959g. Tritonia nelsonii. Yol. 33: t. 1315.
—1961a. Aloe secundiflora. Vol. 34: t. 1341.
—1961b. Aloe monotropa, sp. nov. Vol. 34: t. 1342.
—1961c. Podranea ricasoliana. Vol. 34: t. 1347.
— 1961d. Podranea brycei. Vol. 34: t. 1348.
— 1961e. Bulbine abyssinica. Vol. 34: t. 1350.
— 1961f. Crinum moorei. Vol. 34: t. 1351.
— 1961g. Aloe tenuior. Vol. 34: t. 1352.
—1962a. Gladiolus pole-evansii, sp. nov. Vol. 35: t. 1373.
—1962b. Crinum delagoense. Vol. 35: t. 1389.
—1962c. Aloe jucunda, Vol. 35: t. 1390.
— 1962d. Aloe soutpansbergensis. Vol. 35: t. 1391.
-1963a. Aloe bakeri. Vol. 36: t. 1401.
-1963b. Aloe bellatula. Vol. 36: t. 1402.
-1963c. Habenaria comuta. Vol. 36: t. 1404.
—1963d. Bonatea antennifera. Vol. 36: t. 1405.
— 1963e. Aloe isaloensis. Vol. 36: t. 1419.
—1964a. Crinum baumii. Vol. 36: t. 1432.
—1964b. Crinum variabile. Vol. 36: t. 1433.
—1964c. Peltophorum africanum. Vol. 36: t. 1434.
—1965a. Nymania capensis. Vol. 37: t. 1454.
—1965b. Crinum campanulatum . Vol. 37: t. 1455.
—1966a. Aloe albiflora. Vol. 37: t. 1466.
—1966b. Aloe arenicola. Vol. 37: t. 1467.
-1966c. Thespesia acutiloba. Vol. 37: t. 1468.
— 1966d. Holothrix orthoceras. Vol. 37: t. 1469.
— 1966e. Hibiscus calyphyllus. Vol. 37: t. 1470.
—1967a. Dombeya pulchra. Vol. 38: t. 1489.
—1967b. Aloe inermis. Vol. 38: t. 1516.
—1967c. Aloe rauhii. Vol. 38: t. 1517.
— 1967d. Eulophia hereroensis. Vol. 38: t. 1518.
-1968a. Sphenostylis marginata. Vol. 39: t. 1521.
—1968b. Aloe dhalensis. Vol. 39: t. 1522.
—1968c. Crinum paludosum, sp. nov. Vol. 39: t. 1523.
—1969a. Lonchocarpus nelsii. Vol. 39: t. 1543.
—1969b. Exacum affine. Vol. 40: t. 1576.
-1969c. Crinum minimum. Vol. 40: t. 1577.
-1970a. Aloe capitata. Vol. 40: t. 1595.
—1970b. Aloe viridiflora. Vol. 40: t. 1598.
—1970c. Hermannia althaeifolia. Vol. 41: t. 1603.
— 1970d. Hermannia angularis. Vol. 41: t. 1604.
— 1970e. Hermannia comosa. Vol. 41: t.1605.
— 1970f. Hermannia diversistipula var. graciliflora, var. nov. Vol. 41:
t. 1620.
—1972a. Hermannia prismatocarpa . Vol. 42: t. 1628.
—1972b. Crinum carolo-schmidtii. Vol. 42: t. 1629.
—1972c. Crinum eucrophyllum. Vol. 42: t. 1642.
— 1972d. Crinum rautanenianum. Vol. 42: t. 1643.
-1972e. Aloe gariepensis. Vol. 42: t. 1654.
—1973. Crinum crassicaule. Vol. 43: t. 1676.
—1974a. Haemanthus magnificus. Vol. 43: t. 1681.
—1974b. Hermannia concinnifolia, sp. nov. Vol. 43: t. 1691.
—1976. Hermannia confusa. Vol. 43: t. 1718.
—1977a. Aloe namibensis. Vol. 44: t. 1730.
—1977b. Aloe dewinteri. Vol. 44: t. 1752.
—1977c. Aloe asperifolia. Vol. 44: t. 1753.
—1979. Aloe corallina. Vol. 44: t. 1788.
-1983a. Crinum firmifolium. Vol. 47: t. 1874.
—1983b. Crinum asiaticum. Vol. 47: t. 1875.
—1984. Dombeya macrantha. Vol. 48: t. 1896.
BALLY, P R O. & VERDOORN, I.C. 1956. Aloe peckii, sp. nov. Vol.
31: t. 1214.
CHRISTIAN, H.B. ex VERDOORN, I.C. 1951a. Aloe munchii, sp. nov.
Bothalia 19,2 (1989)
317
Vol. 28: t. 1091.
CHRISTIAN, H.B. ex VERDOORN, I.C. 1951b. Aloe mcloughlinii, sp.
nov. Vol. 28: t. 1112.
CHRISTIAN, H.B. & VERDOORN, I.C. 1952. Aloe nyeriensis, sp. nov
Vol. 29: t. 1126.
HARDY, D. & VERDOORN, I.C. 1970. Aloe ballii. Vol. 40: t. 1589.
VERDOORN, I.C. & CHRISTIAN, H.B. 1940. Aloe marsabitensis, sp.
nov. Vol. 20: t. 798.
—1941. Aloe mzimbana , sp. nov. Vol. 21: t. 838.
-1946. Aloe torrei , sp. nov. Vol. 25: t. 987.
—1947. Aloe magnidentata, sp. nov. Vol. 26: t. 1015.
VERDOORN, I.C. & HARDY, D. 1965. Aloe prinslooi. Vol. 37: t. 1453.
VERDOORN, I.C. & LEIGHTON, F. 1947. Omithogalum leptophyllum.
Vol. 26: t. 1038.
BOTHALIA
VERDOORN, I.C. 1928. A revision of the Crotalarias of south and south-
east tropical Africa. Vol. 2: 371—420.
—1939. Three species of Strychnos with, 1-seeded fruits. Vol. 3: 583—588.
—1941. The genus Plinthus. Vol. 4: 177—179.
—1951a. South African species of Acacia with glandular glutinous pods.
Vol. 6: 153-160.
—1951b. Commiphora neglecta, sp. nov. (Burseraceae). Vol. 6: 214-215.
—1951c. Salsola rabieana , sp. nov. (Chenopodiaceae). Vol. 6: 218—219.
— 1951d. Encephalartos humilis, sp. nov. (Cycadaceae). Vol. 6: 220-221.
— 1951e. Ochna glauca, sp. nov. (Ochnaceae). Vol. 6: 232—233.
—1954a. The nomenclature of the Cape Acacia. Vol. 6: 409-413.
—1954b. Airiplex suberecta, sp. nov. (Chenopodiaceae). Vol. 6: 418—421.
—1954c. Encephalartos eximus, sp. nov. (Cycadaceae). Vol. 6: 426—428.
—1956. The Oleaceae of southern Africa. Vol. 6: 549—639.
—1958. Notes on the Loganiaceae, Salvadoraceae and Oleaceae in South
Africa. Vol. 7: 11-15.
—1961. Chironia stokoei, sp. nov. (Gentianaceae). Vol. 7: 458—459.
—1961. Notes on Gentianaceae. Vol. 7: 459—463.
—1962. Salacia wardii , sp. nov. (Hippocrateaceae). Vol. 8: 114-115.
—1964a. Melhania Integra , sp. nov. (Sterculiaceae). Vol. 8: 177—178.
—1964b. Melhania poly gama, sp. nov. (Sterculiaceae). Vol. 8: 178-179.
—1966a. Contemporary botanists and botanical collectors. Frederick
Ziervogel van der Merwe. Vol. 8, Supplement No. 1: 59-60.
—1966b. A new species of Dombeya and a new variety of Dombeya rotun-
difolia (Sterculiaceae). Vol. 9: 143-145.
—1969a. A new species of Crinum, C. foetidum (Amaryllidaceae). Vol.
10: 56-58.
-1969b. A new species of Hermannia, El. umbratica (Sterculiaceae).
Vol. 10: 76-77.
—1969c. New names in Hermannia. H. antonii and H. repetenda (Ster-
culiaceae). Vol. 10 : 77—79.
— 1969d. New combinations in the genus Hermannia. H. burchellii and
H. cuneifolia var. glabrescens (Sterculiaceae). Vol. 10: 79-81.
— 1969e. A new species of Corchorus, C. sulcatus (Tiliaceae). Vol. 10:
81-82.
—1972a. A new species of Hermannia, H. muirii Pillans (Sterculiaceae).
Vol. 10: 571-572.
—1972b. Two new varieties of Hermannia filifolia (Sterculiaceae). Vol.
10 : 572 - 573.
—1973. The genus Crinum in southern Africa (Amaryllidaceae). Vol.
11: 27-52.
—1974. A new species of Hermannia, H. helicoidea (Sterculiaceae). Vol.
11: 288.
—1975a. A new species of Hermannia, H. litoralis (Sterculiaceae). Vol.
11: 518.
—1975b. Typification of Hermannia rugosa (Sterculiaceae). Vol. 11: 519.
—1980. Revision of Hermannia subgenus Hermannia in southern Africa
(Sterculiaceae). Vol. 13: 1—63.
—1981a. Revision of Melhania in southern Africa (Sterc liaceae). Vol.
13 : 263 -273.
—1981b. The genus Waltheria in southern Africa (Sterculiaceae). Vol.
13 : 275 -276.
—1981c. The genus Cola in southern Africa (Sterculiaceae). Vol. 13:
277-279.
BRUECKNER, A. & VERDOORN, I.C. 1954. Airiplex erosa, sp. nov.
(Chenopodiaceae). Vol. 6: 416—418.
CHRISTIAN, H.B. & VERDOORN, I.C. 1954. Aloe babatiensis, sp.
nov. (Liliaceae). Vol. 6: 440 -442.
DYER, R.A. & VERDOORN, I.C. 1951. Encephalartos natalensis, sp.
nov. Vol. 6: 205-211.
OBERMEYER, A. A., SCHWEICKERDT, H.G. & VERDOORN, I.C.
1937. An enumeration of plants collected in the northern Transvaal.
Vol. 3: 223-258.
STANDARD ENCYCLOPAEDIA OF SOUTHERN AFRICA
VERDOORN, I.C. 1970a. Acacia. Vol. 1: 10—11.
—1970b. Ana-tree. Anaboom. Witapiesdoring. (Acacia albida). Vol. 1:
345-346.
-1970c. Apiesdoring. (Acacia galpinii). Vol. 1: 488.
— 1970d. Bokbaai-vygie. Sandvygie. Dorothea-flower. (Dorotheanthus bel-
lidiformis and some other species of Dorotheanthus). Vol 2:
399-400.
—1971a. Camel-thorn. Kameeldoring. (Acacia giraffae). Vol. 3: 6—7.
-1971b. Cassias. Vol. 3: 122.
—1971c. Caterpillar bean. (Zomia capensis). Vol. 3: 126.
-1971d. Citrus family. (Rutaceae). Vol. 3: 243.
— 1971e. Clovers. ( Trifolium spp.). Vol. 3: 274.
— 1971f. Dubbeltjie. Dubbeltjiedoring. Duiwelsdis. Duiweltjies. Duiwel-
tjiesdoring. (Emex australis; Tribulus spp. and Dicerocaryum
zanguebaricum) . Vol. 4: 95—96.
— 1971g. Enkeldoring. Engelse doring. Oudoring. ( Acacia robusta). Vol.
4: 368.
— 1971h. Forest fever-tree. Koorsboom. Cabbage-tree. Tobacco tree. (An-
thocleista grandiflora = A. zambesiaca). Vol. 4: 480—481.
-1972a. Fruits, wild. Vol. 5 : 79 - 84.
-1972b. Ganna family. (Chenopodiaceae). Vol. 5: 117.
—1972c. Geelhaak. Yellow-thorn. (Acacia Senegal). Vol. 5: 133.
— 1972d. Gemsbok bean. Gemsbokboontjie. Braaiboontjies. (Bauhinia
esculenta). Vol. 5: 135—136.
— 1972e. Gentian family. (Gentianaceae). Vol. 5: 149-150.
— 1972f. Gloxinia and Streptocarpus family. (Gesneriaceae). Vol. 5: 210.
— 1972g. Hibiscus family. (Malvaceae). Vol. 5: 518—519.
— 1972h. Huilbos. African wattle. (Peltophorum africanum). Vol. 5: 652.
— 1972i. Ironwood. White ironwood. (Vepris undulata = V. lanceolata).
Vol. 6: 149.
— 1972j. Kaffirboom. Natal Kaffirboom. ( Erythrina humeana = E.
humei). Vol. 6: 264.
—1972k. Kaffir orange family. (Loganiaceae). Kaffir orange. Swart
klapperboom. Mkwakwa. ( Strychnos madagascariensis =
Strychnos innocua subsp. dysophylla). Vol. 6: 266—267.
—19721. Kanniedood. Corkwood. Cork-tree. Kurkhout. Kurkboom. (Com-
miphora pyracanthoides subsp. glandulosa = C. glandulosa).
Vol. 6: 292.
—1972m. Legume family. (Leguminosae). Vol. 6 : 575 —576.
— 1972n. Lemon-thorn, lowveld. ( Fagara humilis ). Vol. 6: 583.
— 1972o. Lion’s-eye. (Wormskioldia longepedunculata) . Vol. 6: 651.
— 1972p. Lucky-bean. Love-bean. Crab-eye. Minnie-minnie. (Abrus
precatorius) . Vol. 7: 57.
— 1972q. Melianthus family. (Melianthaceae). Vol. 7: 324.
— 1972r. Monkey orange. Klapper. Botterklapper. (Strychnos pungens).
Vol. 7: 514-515.
—1973a. Paperbark-thorn. Verveldoring. Kurk-kameeldoring. (Acacia
sieberana var. woodii = A. woodii). Vol. 8: 441.
—1973b. Pear, wild. Wild plum. Blombos. Dikbas. Drolpeer. (Dombeya
rotundifolia) . Vol. 8: 484.
—1973c. Quiver-tree. Kokerboom. ( Aloe dichotoma). Vol. 9 : 207.
—1973d. Rattle-bush. Klapperbossie. Styfsiektebos(sie). ( Crotalaria
burkeana). Vol. 9: 250.
— 1973e. Sagewood family. (Loganiaceae). Vol. 9: 458.
— 1973 f . Sekelbos. Sickle-bush. (Dicrostachys cinerea). Vol. 9 : 575.
—1974a. Sterculia family. (Sterculiaceae). Vol. 10: 276.
—1974b. Suurklapper. ( Strychnos cocculoides). Vol. 10: 363.
-1974c. Swarthaak. (Blou, Bruin)haakdoring. Hakiesdoring. Wag-n-
bietjie. ( Acacia mellifera subsp. detinens). Vol. 10: 369.
— 1974d. Sweet-pea, wild. ( Sphenostylis angustifolia) . Vol. 10: 385.
— 1974e. Tamarisk. Abiquasgeelhout. ( Tamarix usneoides = T. austro-
africana). Vol. 10: 404—405.
MISCELLANEOUS PUBLICATIONS
VERDOORN, I.C. 1919. The genus Fagara, as represented in the South
African herbaria. Journal of Botany 57: 201—205.
-1920. The order Primulines, as represented in the Transvaal. South
African Journal of Science 16: 365—375.
-1922. Notes on Aponogeton distachyos. South African Journal of Natural
History 3,2: 17-19.
—1924a. The flowering parasitic plants. South African Journal of Natural
History 4: 221-228.
—1924b. The kameel-doorn. Journal of Agriculture 8: 414-416.
—1924c. Melasma. Journal of Agriculture 9: 448—450.
— 1924d. Phyllanthus cedrelifolius, sp. nov. Kew Bulletin 1924: 259.
—1926a. Revision of the African Toddalieae. Kew Bulletin 1926: 389.
318
Bothalia 19,2 (1989)
—1926b. Review: A book of South African flowers, by Barclay, Bolus
and Steer. Kew Bulletin 1926: 432.
—1927. Stapelia tapscottii, sp. nov. Kew Bulletin 1927: 357.
-1929. Notes on the vegetation of the Fountains Valley, Pretoria. South
African Journal of Science 29: 190—194.
—1933. The baobab. South African Journal of Science 30: 255—257.
-1934. An introduction to botany and a few veld flowers.
—1935a. Toddaliopsis bremekampii , sp. nov. Kew Bulletin 1935: 204.
—1935b. Ficus smutsii, sp. nov. Kew Bulletin 1935: 205.
-1935c. The African resurrection plant. South African Garden and
Country Life 25: 21.
—1937a. Notes on the flora of southern Africa: Syncolostemon eriocepha-
lus , sp. nov. Kew Bulletin 1937: 447.
—1938. Edible wild fruits of Transvaal . Department of Agriculture and
Forestry, Bulletin 185.
—1939a. Eetbare veldvrugte van Transvaal. Departement van Landbou
en Bosbou, Bulletin 185.
—1939b. Notes on indigenous plants and cut flowers. South African
Horticultural Journal 1,3: 20-21.
—1939c. Chance meetings with Amaryllids in four corners of the South
African veld. Herbertia 6: 111—112.
—1940a. Indigenous plants as cut flowers, continued. South African
Horticultural Journal 2,2: 13—14.
—1940b. Bauhinia galpinii N.E. Br. South African Horticultural Journal
2,3: 12.
-1940c. Spring in the Union Buildings wild flower garden. South African
Horticultural Journal 3,1: 13.
-1940d. A chance letter among dried plants. South African Horticultural
Journal 3,1: 19—20.
—1941a. Hasie. Our Animal Magazine 1,5.
—1941b. Tambotie, the African jumping bean. Our Animal Magazine 1,7.
—1941c. Desmonema fragosum , sp. nov., from the northern Transvaal,
Journal of South African Botany 7: 209—211.
—1942. Inleiding tot plantkunde en tot enige Transvaalse veldblomme.
—1945a. On the genus Aloe: preparation of herbarium material at Pretoria.
Chronica Botanica 9: 150—151.
—1945b. A new species of Encephalartos from the Waterberg, E. eugene-
maraisii. Journal of South African Botany 11: 1—3.
-1947a. Rika en die mossies. Lentedae, Hfst. 19, Std 1.
—1947b. Die diederikkie. Lentedae , Hfst. 27, Std 2.
—1947c. Die vriendelike boom en die verkleurmannetjie. Lentedae , Hfst.
28, Std 2.
— 1947d. Vroeg in die more. Lentedae , Hfst. 29, Std 2.
— 1947e. Vroeg in die more. Lentedae , Hfst. 30, Std 2.
— 1947f. Nuus van die blinkblaar-wag:n-bietjie. Lentedae , Hfst. 13, Std 3.
— 1947g. Blinkblaar-wag:n-bietjie se boodskap. Lentedae , Hfst. 14, Std 3.
— 1947h. Die liguster vertel. Lentedae, Hfst. 24, Std 4.
-1947i. In veld en kloof, lentedae, Hfst. 31, Std 4.
— 1947j. Die wederopstanding plant. Lentedae, Hfst. 32, Std 4.
—1947k. Springboontjies. Lentedae , Hfst. 6, Std 5.
—1949. Salix species. One of our nature’s fodder trees. Farming in South
Africa 24: 253-254.
—1951a. Encephalartos natalensis. The cycad of the valley of a thousand
hills. South African Journal of Science 48: 162.
—1951b. A cycad tour of the eastern Cape. African Wild Life 5: 153—159.
—1954. The Transvaal Ericaceae. Journal of South African Botany 20:
91-115.
—1955. Reflections on old Pretoria. Public Servant, November.
—1956a. Cycads in the Transvaal Province. Trees in South Africa 8,2: 3—9.
—1956b. Crinum macowanii. Journal of South African Botany 22: 79—83.
—1957a. The baobab or kremetartboom. African Wild Life 11: 95—99.
—1957b. An assessment of voluntary associations that advance the study
of biology in southern Africa. Presidential address. South African
Biological Pamphlet 19: 6—13.
-1962. Encephalartos species in Natal. Trees in South Africa 14: 50—55.
—1963a. Encephalartos species in Natal. Reprinted in Aloe 1,1.
—1963b. Oleaceae. Flora of southern Africa 26: 100—132.
—1963c. Salvadoraceae. Flora of southern Africa 26: 132—134.
—1963d. Loganiaceae. Flora of southern Africa 26: 134-171.
— 1963e. Our veld flowers. Crinums for the garden, Cape heaths. Farming
in South Africa 39,8: 50—52.
— 1963f. Living fossils. Farming in South Africa 39,8: 56.
— 1963g. A new species of Salsola with notes on its affinities, S. henriciae.
Journal of South African Botany 29: 5—9.
—1964a. Orchids as I see them. Transvaal Orchid Society Bulletin 10.
—1964b. Notes on an interesting South African orchid. Transvaal Orchid
Society’ Bulletin 11.
—1964c. Cycads in the Transvaal Province. Reprinted in Aloe 2,1: 9—14.
— 1964d. A cycad tour of the eastern Cape. Reprinted in Aloe 2,2: 9-14.
-1965. Plant taxonomy in South Africa. Presidential address. South
African Journal of Science 61: 43—50.
—1966a. A definition and name for an eastern Cape Crinum, C.
macowanii subsp. confusum, subsp. nov. South African Journal
of Botany 32: 67-71.
-1966b. Welwitschiaceae. Flora of southern Africa 1: 48-51.
-1966c. Identity of John Forbes’s Crinum. Wild Life 20: 197-203.
—1967a. Onze inheemse borne, Adansonia. Nieuws uit Zuid-Afrika 7.
—1967b. Onze inheemse planten, Encephalartos spp. Nieuws uit Zuid-
Afrika 7.
-1967c. Onze inheemse planten, Crinum. Nieuws uit Zuid-Afrika 7.
-1969. Review of Leeuwenberg’s Strychnos in Africa. South African
Journal of Science 65: 326.
-1972a. Foreword to Aloes of the South African Veld, by Bornman &
Hardy.
—1972b. The genus Hermannia. Veld & Flora 2,3: 43—44.
—1974. Our cycads in perspective. Aloe 12,3: 101—106.
-1983a. Diary of an expedition — 1937, Part Four. Aloe 20,1: 28-29.
-1983b. Diary of an expedition. Part Six (should read Five). Aloe 20,1:
40-41.
DYER, R.A. & VERDOORN, I.C. 1966. Zamiaceae. Flora of southern
Africa, 1: 3—34.
—1968—69. Encephalartos manikensis and its allies. Kirkia 7: 147-158.
STEYN, D., VAN DER WALT, S.J. & VERDOORN, I.C. 1948. The
seed of some species of Encephalartos. A report on their toxicity.
South African Medical Journal 22 : 758 —760.
VERDOORN, I.C. & COLLETT, D.G. 1974. Fibre yielding wild stock
rose. Farming in South Africa 22: 499—501.
VERDOORN, I.C. & MARAIS, W. 1963. Gentianaceae. Flora of
southern Africa 26: 171—243.
D.M.C. FOURIE
Bothalia 19,2: 319-320 (1989)
Book Reviews
TREES AND SHRUBS OF THE WITWATERSRAND, MAGALIES-
BERG AND PILANESBERG by J. VAN GOGH & J.M. ANDERSON
1988. Struik Publishers, Oswald Pirow St, Foreshore, Cape Town 8001.
Pp. 112. Size 280 x 215 mm. ISBN 0 86977 719 X. Price: hard cover
R34.95 + GST.
This book has an unusual arrangement in that there is a minimum of
text, most information being presented pictorially. Authors’ notes and
an explanation of how to use the book are followed by descriptive pages,
each with two to four species, alternating with facing, detailed, water-
colour illustration pages showing the species described opposite.
Descriptive pages show for each species, a habit drawing, a scale giv-
ing the approximate height of a mature specimen in its favoured habitat
for the study area, very brief notes mainly on bark and foliage, a draw-
ing indicating habitat and a distribution map.
The habitat drawing is purely topographical, no account being taken
of soil types. It is in effect an oblique aerial view from the east of a
stylised portion of Magaliesberg range and surrounds, with the particular
species marked thereon in green. A key drawing in the explanatory notes
defines the various habitats.
On the distribution maps the occurrence of a species is indicated by
a green area of varying size depending on the extent of the presence.
With the aid of a transparent overlay, on which a particular point has
been marked, positioned successively on each of the maps, it should
theoretically be possible to ascertain the various species which occur
at the selected point. However, the maps are very small and minor
occurrences of a species may be very difficult to discern, particularly
in artificial light, and in the case of minute green areas situated next
to rivers and dams which are shown in blue. A slight re-arrangement
would have made it possible to increase the size of these maps by a not
insignificant 15%, without altering the overall page layout.
Draughtsmanship of the coloured illustrations is, in general, reasonably
good but the painting has, in most cases, a slap-dash or hurried appearance
which is surprising as this work was not done in the field. The lack of
variation in the greens has been criticised by various indigenous tree
enthusiasts and a colour expert. This could be due to a low standard
of the colour Rpoto-prints from which the paintings were done, or to
poor colour reproduction, or both.
The book title is decidedly ill-chosen. ‘Trees and shrubs of the
Witwatersrand’ appears in large print followed by an unobtrusive
‘Magaliesberg and Pilanesberg’ in small print — as though it was an
afterthought. The first portion is identical to the title of the Tree Society
work, first published in 1964 and now well into its third edition. This
must inevitably lead to confusion. In addition, it should have been the
trees and shrubs of the Pilanesberg and Magaliesberg, in that order, that
required any emphasis, as the flora of those areas is likely to be with
us a good deal longer than that of the Witwatersrand, which is rapidly
vanishing under tar and concrete. Incidentally, both authorship and
publisher of the Tree Society publication were incorrectly attributed in
the list of reference works.
The authors are of course to be commended for the time and effort
expended. It is no mean accomplishment to put together a work of this
sort. They have provided a valuable record of the species which occur,
or did once occur, in an area which, taken overall, is surely the most
severely threatened in the Transvaal. They have also furnished us with
a handy identification guide having considerable utility in the field of
conservation.
J.D. CARR
FLORA OF AUSTRALIA (VOL. 19), Myrtaceae— Eucalyptus, An-
gophora by G.M. CHIPPENDALE. 1988. Australian Government Pub-
lishing Service , G.P.O. Box 84, Canberra, A.C.T. 2601. Pp. 542, line
drawings, eight colour plates, maps. Price: soft cover edn, Cat. No.
88 0515 5, US $44,95; case bound edn. Cat. No. 87 0157 2, US $59,95.
This welcome addition to the Flora of Australia covers the two closely
related genera Eucalyptus and Angophora. The former has become a
familiar part of the South African scene since its first introduction 150
years ago and plays a major role in the economy; the latter, while less
commonplace, has secured a small niche for itself in the realm of ar-
boriculture.
To botanists and foresters in South Africa, particularly, the volume
under review represents a refreshingly pragmatic approach to the subjects
of eucalypt classification and identification. In a number of important
respects it departs significantly from the several monographic treatments
that have appeared since the mid-nineteenth century.
A concise account of Eucalyptus L’Herit., embracing its chief mor-
phological features, distribution and uses, is followed by a brief review
ot additional floral and other characters which, although of taxonomic
significance, are not mentioned either in the foregoing or in the succeeding
descriptions of individual species (which go to make up the great bulk
of the volume). The discussion centres chiefly round stamens and anthers,
the operculum, ovules and the cotyledons, and reference is made to the
use of these organs by previous authors to separate taxa and groups of
taxa within the genus. Bentham (1867) in Flora australiensis, followed
by Von Mueller (1879—1884) in Eucalyptographia , based his series largely
on staminal characters. Blakely (1934) in A key to the eucalypts also used
stamens, particularly anthers, as a basis for his classification of species
into sections. The result was not altogether satisfactory, for when other
characters were taken into account, his classification was seen to be
‘artificial’. Moreover Blakely’s key was found unworkable in practice by
South African forest botanists. Maiden (1929-1931) in A critical review
of the genus Eucalyptus (Vol. VIII, Parts 3-5) arranged the majority
of taxa known at the time according to the shape of the cotyledons. Carr
& Carr (1959) in Developmental morphology of the floral organs in
Eucalyptus, showed that the genus could be divided into two groups,
the first comprising species with one operculum ‘Monocalyptus’ and the
second those with two ‘Symphyomyrtus’.
In the present work a wide range of readily observable features,
including habit, bark, phyllotaxy, leaf morphology, inflorescence, flower
buds and fruits, is substituted for a single character to separate groups
and individual taxa. A dichotomous key in two stages is provided, the
first stage differentiating 20 groups of taxa and the second breaking these
down to the constituent 5D species, subspecies and varieties. The groups
are again ‘artificial’ in as much as each comprises an assemblage of taxa
without close phyletic affinities. Geographical criteria are incorporated
in the second stage of the key, but these are ancillary and seldom if ever
critical when deciding which of two alternatives to opt for. A notable
contribution in its own right, the key is simple to use and represents a
signal advance on previous devices of the kind (Card Sorting Key not
excepted).
On taxonomic grounds the genus is divided into 92 series. These do
not feature in the key, neither is a synopsis given of the systematic
relationships perceived to exist within the genus. The series circumscrip-
tions, are to some extent based on an assessment of those proposed by
Pryor & Johnson (1971) in A classification of the eucalypts, but never-
theless deviate from them substantially.
Descriptions of individual species need to be read in conjunction with
those of the series because characters common throughout a given series
are not repeated. However, because the number of species in a series
never exceeds 21, this does not require a great deal of manual dexterity
or mental agility. The buds and fruits of almost all taxa are illustrated
by effective line drawings, which would seem to have been prepared
specially for this volume and not merely reproduced from Maiden’s
Critical review. Notes appended to the species descriptions draw attention
to distinguishing features and could contribute much to the accuracy and
speed of identifications.
The text has been purged of a great many redundant or obscure epithets
long bandied about in the botanical and forestry literatures. These have
been relegated to the categories of ‘presumed hybrids’ and ‘doubtful
names’.
Angophora, comprising seven species, is a straightforward revision
of the genus along the lines adopted for Eucalyptus.
320
Bothalia 19,2 (1989)
In the succeeding pages, distribution maps of all the taxa dealt with
in the text are given in the same order as that in which the descriptions
appear. Although small in scale, they provide a fair indication of the
geographical range of each taxon.
Latin diagnoses of new taxa, hitherto unpublished combinations and
lectotypifications are all given in an Appendix. This is followed by a
Supplementary glossary, a list of Abbreviations and contractions , an Index
to genera and species and a map of Australia showing Botanical regions.
The author points out that recent years have witnessed an upsurge of
research on the genus Eucalyptus. A number of important papers have
been published and others are in preparation. These investigations will
doubtless lead to refinements in the classification of the eucalypts as time
passes. A review even now in progress is expected to result in many
changes being proposed at and above the rank of species. This
development was foreshadowed by Johnson (1975) in Problems of species
and genera in Eucalyptus (Myrtaceae), when he discussed the desirability
of splitting off several distinct genera. A decision in this regard would
depend on the wideness of polyphylesis in the genus relative to that in
other myrtaceous genera and also on ‘practical convenience’. Appropos
the latter consideration, Johnson poses the question: ‘Could the users
of botanical names tolerate formal division of Eucalyptus into 9 or 10
genera?' He continues: ‘Only about 100 species would be in Eucalyptus
sensu stricto . . The reaction of South African forestry practitioners,
if not botanists, to such a development would be one of dismay. In the
opinion of the reviewer, a move of the kind would seriously damage the
credibility of botanical names as aids to communication and the transfer
of information. Stability and simplicity are essential if botanical
nomenclature is to serve the interests of other disciplines.
South African foresters, and probably the majority of botanists, will
be well content with the conservative approach adopted by the author
of this volume with respect to taxonomic issues. They would assuredly
wish the work to remain the standard reference on the subject for many
years to come.
(
R.J. POYNTON*
* South African Forestry Research Institute, RO. Box 727, Pretoria 0001
THE GARDENER’S LABYRINTH by THOMAS HILL, edited by
RICHARD MABEY. 1988. Oxford University Press, Walton St, Oxford
OX2 6DP, England. Pp. 221, 40 colour plates, 110 black and white
reproductions, soft cover, stitched. Price: R47.70 + GST.
This delightful production is an augmented version of the first popular
gardening book in the English l&nguage, which was first published in
1577. It proved very popular and ran to six editions over a period of 75
years. The publication under review is based on the 1652 edition, but
excludes Hill’s astrologically based notes on medicinal uses
of specific plants. It retains, however, a table which serves as a precis
of the content of these notes.
Mabey's introduction is a scholarly and entertaining review of Hill’s
text, as seen in its social, botanical, horticultural, literary and historical
perspectives. No apology is needed for quoting from it: ‘As a study of
plants it is overshadowed by the massive herbals of the Elizabethan
botanists Turner and Gerard. Even as a garden encyclopaedia it cannot
compare with John Parkinson’s Paradisi in Sole . . . But from its very
first pages, The labyrinth ’s descriptions of how to test soil between the
fingers, build a rose arch, mulch a vegetable bed, leave you in no doubt
that this is a pioneering work ... It is the enthusiastic, coaxing, down-
to-earth voice of the incipient garden columnist ... Some of his advice
is remarkably precocious ... yet running through the book is a ... to
most modern minds, patently impractical line of advice which springs
from the old tradition of natural magic . . . these passages . . . give an insight
into this crucial moment in history, when the old beliefs were colliding
with the burgeoning mood of reason and practicality’.
The Gardemer 's labyrinth contains 37 chapters, each with one to two
pages of text, copiously illustrated. The chapter headings deal with good
garden practice, as varied as:
Of plainer instructions for the chusing of a battle ground, with other
matter necessary.
To have a wall of Rosemary of a great height quickly, and
Of helps against Scorpions, Todes, Garden Mice, Weasels, and all other
beasts, wasting herbs and fruits.
They are followed by a second part which details the cultivation,
qualities, and uses of more than 50 different herbs, vegetables and flowers.
Both parts benefit greatly from Richard Mabey’s second contribution,
an excellent glossary, identifying plants by their common names, and
explaining the terms used. I discovered to my sorrow that a garden was
not recognized as a ‘battle ground’ in the modern sense, but that ‘battle’
in this context means ‘fertile’. I would suggest that for this glossary alone
the Gardener 's labyrinth is worthy of a place on most library shelves.
But, undoubtedly, the illustrations also add a great deal. They include
contemporary paintings, drawings and engravings, as well as embroidery,
skilfully chosen to portray the techniques, beliefs, moods, landscapes,
cultivars and bounty of those times. The colour reproductions are
generally good, but a number of the black and white prints suffer from
very uneven register. This failing aside, the Gardener's labyrinth is
extremely rewarding, both in text and illustration. It invokes humility
and wonderment that so much was known and thought about, so long
ago, and that some of it has been forgotten. An educational treat.
M.J. WELLS
321
INDEX -
Acacia
dealbata Link, 237
mearnsii De Wild., 237
Acanthaceae, 151, 209
Actiniopteris semiflabellata Pichi-Sermolli , 171
Adhatoda
leptantha (Nees) Nees, 209
tubulosa Nees, 209
Africa
central, 167
southern, 9, 17, 125, 151, 157, 161
ailment, 111
air pores, 9
alien, 237
Amauropelta oppositiformis (C. Chr.) Holtt., 173
Amphibolia L. Bol. ex Toelken & Jessop, 179
maritima L. Bol. ex Toelken & Jessop, 182
analysis, 183
anatomy, 9, 45, 134
culm, 189
leaf, 41, 189, 217
Anderson, J.M. Review: The Banksia Atlas, Australian Flora and Fauna
Series No. 8, ed. by Anne Taylor and Stephen Hopper. 1988,
142
aneuploidy, 125
Annual report of the Botanical Research Institute 1988/1989, 295
Anthemideae (Asteraceae), 27, 29
Arecaceae, 213
Amell, S., 17
Arriaga, Mirta O. , 45
Arundinella
berteroniana (Schult.) Hitchc. & Chase, 45
hispida (Willd.) Kuntze, 45
nepalensis Trin. , 45
Arundinelleae (Poaceae), 45
Arundineae (Poaceae), 217
Arundinoideae (Poaceae), 134
Asplenium
buettneri Hieron. ex Brause, 171
parablastophorum Braithewaite, 173
sebungweense J. E. Burrows, 173
uhligii Hieron., 173
Asteraceae, 27, 29
Asthenatherum mossamedense (Rendle) Conert, 41
Aulojusticia Lindau, 209
linifolia Lindau, 210
Backer, A.P. , 91
Barker, N.P. The caryopsis surface of Pentameris and Pseudopentameris
(Arundinoideae, Poaceae) revisited, 134
Barker, N.P., 275
Bilderdykia Dumort. , 175
Britz, P.J., 268, 270
Brusse, F. A new species of Inezia (Anthemideae) from the north-eastern
Transvaal (Asteraceae), 27
Brusse, F. A new Phymaspermum (Anthemideae) species from dolomite
areas of the Wolkberg (Asteraceae), 29
Brusse, F. A new species of Fuscidea (Lichenes) from the Cape Fold
Mountains (Fuscideaceae), 35
Brusse, F. A new species of Maronea (Lichenes) from the Drakensberg
(Fuscideaceae), 36
Bryophyta, 276
Burrows, J.E. New taxa, combinations and records of Pteridophyta from
southern and central Africa, 167
C4, 41, 45
calcium salts, 9
camping kitchen, 273
Capensis, 189
Cape Province, 1, 7, 35
north-western, 7
Carr, J.D. Review: Trees and shrubs of the Witwatersrand, Magalies-
berg and Pilanesberg, by J. van Gogh & J.M. Anderson, 319
Carr, J.D. & Retief, E. A new species of Combretum from Natal
(Combretaceae), 38
caryopsis, 134
central Africa, 167
Centropodia mossamedensis (Rendle) T.A. Cope, 41
checklist, 91, 183
Eastern Transvaal Escarpment, 92
Pteridophytes of Transkei, 186
Chromolaena odorata (L.) R.M. King & H. Robinson, 237
chromosome numbers, 125
-INDEKS
chromosome studies, 125
climate, 53
coastal communities, 237
comb, nov., 171, 209
Combretaceae, 38
Combretum mkuzense Carr & Retief, 38
Crotalarieae (Fabaceae), 1, 32
Ctenopteris rigescens (Bory ex Willd.) J. Sm., 171
culm, 45
culm anatomy, 189
cytology, 125
Danthonieae (Poaceae), 41, 217
Davidse, G., 125
Deall, G.B. & Backer, A.P. The vegetation ecology of the Eastern
Transvaal Escarpment in the Sabie area. 3. Annotated checklist, 91
Deall, G.B., Scheepers, J.C. & Schutz, C.J. The vegetation ecology of
the Eastern Transvaal Escarpment in the Sabie area. 1. Physical
environment, 53
Deall, G.B., Theron, G.K. & Westfall, R.H. The vegetation ecology
of the Eastern Transvaal Escarpment in the Sabie area. 2.
Floristic classification, 69
Deall, G.B. & Westfall, R.H. Improving the resolution of floristic/habitat
pattern correlations on phytosociological tables, 263
Dehn, M., 179
De Wet, B.C., Gibbs Russell, G.E., Germishuizen, G., Schrire, B.D.,
Jordaan, M., Pienaar, B.J., Welman, W.G., Reid, C., Van Wyk,
C.M., Fish, L., Immelman, K.L., Van Rooy, J., Glen, H.F. &
Barker, N.P. New taxa, new records and name changes for
southern African plants, 275
Dicotyledoneae, 97, 279
distinctive cells, 45
distribution, 9
double bundle sheath, 45
Drakensberg, 36
drier-transporters, 269
Dryopteris
dracomontana Schelpe & N.C. Anthony, 183
oppositiformis C. Chr., 173
tsaratananensis C. Chr., 173
Eastern Transvaal Escarpment, 53, 69, 91
ecology, 53, 69, 91. 183, 263
vegetation, 53, 69, 91
Ehrharta Thunb. , 125, 189
Ramosa group, 189
aphylla Schrad., 191
var. fasciculata Stapf, 191
var. filiformis Nees, 193
digyna Thunb., 190
filiformis Mez, 193
ramosa (Thunb.) Swartz, 190
subsp. aphylla (Schrad.) Gibbs Russell, 191
subsp. ramosa, 191
var. aphylla (Schrad.) Gluckmann ex Adamson, 191
rehmannii Stapf 192
subsp. filiformis (Stapf) Gibbs Russell, 193
subsp. rehmannii, 193
subsp. subspicata ( Stapf) Gibbs Russell, 194
var. filiformis Stapf, 193
subspicata Stapf, 194
Ehrharteae (Poaceae), 125
Ellis, R.P. Leaf anatomy of the South African Danthonieae (Poaceae).
XVIII. Centropodia mossamedensis, 41
Ellis, R.P. , 45, 189
Ellis, R.P. Leaf anatomy of the South African Danthonieae (Poaceae).
XIX. The Genus Prionanthium, 217
endemic, 9
environment, 53
ethnobotany, 225
Fabaceae, 1, 7, 32
field-data capture, 267
field drier, 270
Fish, L., 277
floristic classification, 69
floristic/habitat correlation, 263
forest, 237
Fourie, D.M.C. Obituary: Inez Clare Verdoom (1896—1989), 313
fruits, 175
Fuscidea hottentotta Brusse, 35
Fuscideaceae (Lichenes), 35, 36
fynbos, 189
Fynbos Biome, 189
322
Gendarussa leptantha Nees, 209
geology, 53
Germishuizen, G. Oxygonum altissimum, a new species from central
Somalia (Polygonaceae), 210
Germishuizen, G., 281—287
Germishuizen, G., Kok, P.D.F. & Robbertse, P.J. Polygonum hydropiper
in southern Africa (Polygonaceae), 211
Germishuizen, G., Robbertse, P.J. & Kok, P.D.F. The genera Polygonum
and Bilderdykia (Polygonaceae) in southern Africa: morphology
and taxonomic value of the ocrea and fruit, 175
Gibbs Russell, G.E., 275
Gibbs Russell, G.E. & Ellis, R.P. Taxonomy and leaf anatomy of the
genus Ehrharta (Poaceae) in southern Africa: the Ramosa
group, 189
glands, linear, 217
Glen, H.F., 275
Grammitis
flabelliformis sensu Morton, 171
rigescens (Bory ex Willd.) J.E. Burrows, 171
grassland, 237
guide for authors to Bothalia, 145
Gymnospermae, 93
Hartmann, H.E.K. & Dehn, M. A re-examination of the genus
Arnphibolia (Mesembryanthemaceae), 179
Henderson, L. Invasive alien woody plants of Natal and the north-eastern
Orange Free State, 237
herbarium labels, 267
Hutchings, Anne. A survey and analysis of traditional medicinal plants
as used by the Zulu, Xhosa and Sotho, 111
Hutchings, A., 183
Hutchings, A. Observations on plant usage in Xhosa and Zulu medicine,
225
Hymenophyllum
ciliatum Swartz var. splendidum (V.d. Bosch) C. Chr. , 171
plumieri Hooker & Grev., 171
splendidum V.d. Bosch, 169
Immelman, K.L. Studies in the southern African species of Justicia and
Siphonoglossa (Acanthaceae): palynology, 151
Immelman, K.L. Siphonoglossa and Aulojusticia in southern Africa
(Acanthaceae), 209
Immelman, K.L., 290
Inezia speciosa Brusse, 27
invasive plants (woody), 237
Johnson, C.T. & Hutchings, A. A contribution to the pteridophyte flora
of Transkei, 183
Jordaan, M.. 288-290, 291
Justicia L. , 151
leptantha (Nees) Lindau, 209
pulegioides subsp. late-ovata C.B. CL, 209
tubulosa (Nees) T. Anders., 209
key, 175
Bilderdykia sp., 175
Ehrharta subspp., 191. 193
Polygonum spp., 175
Siphonoglossa spp. & subspp., 209
Killick, D.J.B. Two nomenclatural problems involving Article 63, 133
Killick, D.J.B. Review: Southern African botanical literature 1600—1988
SABLIT, compiled by A.S. Kerkham, 141
Kok, P.D.F., 175, 211
Kranz, 41, 45
KwaZulu, 237
Lantana camara L. , 237
leaf anatomy, 41, 189, 217
lectotypification, 179
Lichenes, 35, 36
linear glands, 217
Lotononis (DC.) Eckl. & Zeyh., 1, 7
sect. Polylobium (Eckl. & Zeyh.) Benth., 1
sect. Telina (E. Mey.) Benth. , 1
azureoides B-E. van Wyk, 1
elongata (Thunb.) D. Dietr., 32
gracilifolia B-E. van Wyk, 3
involucrata group, 7
lamprifolia B-E. van Wyk, 4
racemiflora B-E. van Wyk, 1
Lygodium kerstenii Kuhn, 183
map cabinet, 272
Marchantiales, 9, 17, 157, 161
Maronea afroalpina Brusse, 36
Marsilea farinosa Launert subsp. arrecta J.E. Burrows, 169
medicinal plants. 111
medicine, 225
meiosis, 125
Meliaceae. 31
Melica ramosa Thunb., 190
Mesembryanthemaceae, 179
Mohria caffrorum (L.) Desv. var. ferruginea J.E. & S.M. Burrows, 168
Monocotyledoneae, 93, 277
morphology, 175
Mtunzini, Zululand, 213
NADP-me, 45
name changes, 275
Natal, 38, 237
new records, 31, 157, 167, 183, 275
new species, 1, 3, 4 , 7, 9, 12 , 27, 29, 35, 36, 38, 157, 167, 173, 209, 210
new taxa, 1, 3, 4 , 7, 9, 12 , 27, 29, 35, 36, 38, 167, 168, 209, 210, 275
nomenclature, 133
north-eastern Orange Free State, 237
north-western Cape, 7
obituary, 137, 313
ocrea, 175
Oetosis ensiformis (Swartz) Greene, 171
Ophioglossum
convexum J.E. Burrows, 167
rubellum Welw. ex A. Braun, 168
thomasii Clausen, 167
Orange Free State, north-eastern. 237
Oxygonum altissimum Germishuizen, 210
palynology, 151
Panagos, M.D. Review: Combretaceae in southern Africa, by J.D. Carr.
1988, 141
Panagos, M.D., 268, 272 , 273
Panagos, M.D., Britz, P.J. & Westfall, R.H. Plant collecting apparatus
for taxonomic and ecological studies. 5. A gas drier for field
drying of plant specimens, 270
Panagos, M.D. & Westfall, R.H. Plant collecting apparatus for
taxonomic and ecological studies. 1. A lightweight plastic plant
press for on-site specimen pressing, 266
Panagos, M.D. & Westfall, R.H. Plant collecting apparatus for
taxonomic and ecological studies. 4. Drier-transporters for plant
presses, 269
Panicoideae (Poaceae), 45
Peckham, G.D. & Van Jaarsveld, F.A. New botanical perspectives on
the origin of the Raphia palms at Mtunzini (Arecaceae), 213
Pentameris Beauv., 134
Pentaschistis triseta (Thunb.) Stapf 217
Perold, S.M. Studies in the genus Riccia (Marchantiales) from southern
Africa. 11. Riccia montana and R. alboporosa, a further two new
white-scaled species of the group ‘Squamatae’, 9
Perold, S.M. Studies in the genus Riccia (Marchantiales) from southern
Africa. 12. Riccia albolimbata and the status of R. albosquamata,
white-scaled species originally described by Arnell, 17
Perold, S.M. Studies in the genus Riccia (Marchantiales) from southern
Africa. 13. A new species, R. hantamensis , in section Pilifer and
a new record for R alatospora, 157
Perold. S.M. Studies in the genus Riccia (Marchantiales) from southern
Africa. 14. R. concava section Pilifer, 161
Persicaria hydropiper (L.) Spach, 211
pharmacognosy, 225
Phymaspermum argenteum Brusse, 29
physiography, 53
phytosociological classification, 263
phytosociology, 69
PH YTOTABr 69
Pienaar, B.J., 279 -281, 287
Pilifer, section (Riccia), 157, 161
plant collecting apparatus, 266 - 274
plant press, 266
Poaceae, 41, 45, 125, 134, 189, 217
Polygonaceae, 175, 210, 211
Polygonum L. , 175
hydropiper L. , 211
polyploidy, 125
Polypodium rigescens Bory ex Willd., 171
Poynton, R.J. Review: Flora of Australia (Vol. 19), Myrtaceae —
Eucalyptus, Angophora, by G.M. Chippendale, 319
practitioners, 225
Prionanthium
dentatum (L. f. ) Henr. , 217
ecklonii (Nees) Stapf, 217
pholiuroides Stapf, 217
Pseudopentameris Conert, 134
Psidium guajava L. , 237
Pteridophyta, 92, 167, 183, 276
publications
by I.C. Verdoorn, 316
by Staff of BRI, 310
Ramosa eroup (Ehrharta, Poaceae), 189
323
Raphia, 213
rare, 9
Reid, C., 276 -278
reticulation, 9
Retief, E., 38
Riccia
subgen. Riccia, 9
sect. Pilifer Volk, 157, 161
sect. Riccia, 9
group Squamatae, 9
alatospora Volk & Perold, 159
albolimbata S. Amell, 17
alboporosa Perold, 12
aibosquamata 5. Arnell, 22
concava Bisch. , 163
hantamensis Perold, 157
montana Perold, 9
Robbertse, P.J. , 175, 211
Rubus spp. , 237
Saayman, E.J.L., 125
Sabie, Transvaal, 53, 69, 91
Salix babylonica L. , 237
Sanchez, Evangelina, Arriaga, Mirta O. & Ellis, Roger P. Kranz
distinctive cells in the culm of Arundinella (Arundinelleae;
Panicoideae; Poaceae), 45
savanna, 237
scales, 9
white, 17
Scheepers, J.C., 53
Schrire, B.D. Obituary: Rudolf Georg Strey (1907-1988), 137
Schrire, B.D., 281—287
Schutz, C.J., 53
Siphonoglossa Oerst. , 151
leptantha (Nees) Immelman, 209
subsp. late-ovata (C.B. Cl.) Immelman, 209
subsp. leptantha, 209
linifolia (Lindau) C.B. Cl., 210
nkandlaensis Immelman, 209
nummularia S. Moore, 209
tubulosa (Nees) Benth. ex Lindau, 209
soil, 53
Solanum mauritianum Scop., 237
Somalia, central, 210
Sotho, 111
southern Africa, 9, 17, 125, 151, 157, 161, 167, 189, 209, 211, 275
Sphaerocionium splendidum (V.d. Bosch) Copeland, 171
Spies, J.J. , Saayman, E.J.L., Voges, S.P. & Davidse, G. Chromosome
studies on African plants. 9. Chromosome numbers in Ehrharta
(Poaceae: Ehrharteae), 125
sp. nov., 1, 3, 4 , 7, 9, 12 , 27, 29, 35, 36, 38, 157, 167, 173, 209, 210
spore ornamentation, 9
Squamatae group (Riccia), 9
staff list of BRI, 307
stat. nov., 191, 193
Strey, Rudolf Georg (1907-1988), 137
subsp. nov., 169
survey, 237
taxonomy, 1, 7, 9, 17, 27, 29, 31, 32 , 35, 36, 38, 151, 157, 161, 167, 175,
179, 183, 189, 209, 217, 263
Thelypteris
oppositiformis (C. Chr.) Ching, 173
strigosa sensu Schelpe, 173
tsaralananensis (C. Chr.) Ching, 173
Theron, G.K., 69
Transkei, 183
Transvaal
Eastern Escarpment, 53, 69, 91
north-eastern, 27
Sabie, 53, 69, 91
Wolkberg, 29
Turraea pulchella (Harms) Pennington, 31
Van Jaarsveld, F.A., 213
Van Rooy, J., 276
Van Staden, J.M., 273
Van Wyk, B-E. Studies in the genus Lotononis (Crotalarieae, Fabaceae).
2. Three new species of the section Telina from the Cape Province,
1
Van Wyk, B-E. Studies in the genus Lotononis (Crotalarieae, Fabaceae).
5. A new species of the L. involucrata group (section Polylobium)
from the north-western Cape Province, 7
Van Wyk, B-E. The identity of Lotononis elongata (Crotalarieae,
Fabaceae), 32
Van Wyk, C.M., 290, 291
var. nov., 168
vegetation ecology, 53, 69, 91
Verdoorn, Inez Clare (1896-1989), 313
Vittaria
ensiformis Swartz, 171
plantaginea Bory, 171
Voges, S.P, 125
Wells, M.J. Review: The gardener’s labyrinth, by Thomas Hill, ed.
Richard Mabey, 320
Welman, W.G., 292-294
Westfall, R.H., 69, 263, 266, 269, 270
Westfall, R.H. Plant collecting apparatus for taxonomic and ecological
studies. 2. Coldat: a field-data capture program for collector’s
data and herbarium labels, 267
Westfall, R.H., Britz, P.J. & Panagos, M.D. Plant collecting apparatus
for taxonomic and ecological studies. 3. A new top-loading plant
press for off-site specimen pressing, 268
Westfall, R.H. & Panagos, M.D. Plant collecting apparatus for taxonomic
and ecological studies. 6. A transportable map cabinet for
vehicle and office use, 272
Westfall, R.H., Panagos, M.D. & Van Staden, J.M. Plant collecting
apparatus for taxonomic and ecological studies. 7. A transportable
camping kitchen for vehicle use, 273
white scales, 17
Williams, R. Turraea pulchella rediscovered (Meliaceae), 31
winter rainfall area, 125
Wolkberg, Transvaal, 29
Xhosa, 111. 225
Xiphopteris
flabelliformis sensu Schelpe, 171
rigescens (Bory ex Willd.) Alston, 171
Zulu, 111, 225
Zululand, 213
(
'
' Si-
X