Bothalia
’N TYDSKRIF VIR PLANTKUNDIGE NAVORSING
A JOURNAL OF BOTANICAL RESEARCH
Vol. 20,1
May/Mei 1990
PUBLICATIONS OF THE NATIONAL BOTANICAL INSTITUTE, PRETORIA
PUBLIKASIES VAN DIE NASIONALE BOTANIESE INSTITUUT, PRETORIA
Obtainable from the National Botanical Institute, Private Bag X101,
Pretoria 0001, Republic of South Africa. A current price list of all avail-
able publications will be issued on request.
BOTHALIA
Bothalia is named in honour of General Louis Botha, first Premier
and Minister of Agriculture of the Union of South Africa. This
house journal of the National Botanical Institute, Pretoria, is devoted to
the furtherance of botanical science. The main fields covered are
taxonomy, ecology, anatomy and cytology. Two parts of the journal
and an index to contents, authors and subjects are published
annually.
Verkrygbaar van die Nasionale Botaniese Instituut, Privaatsak X101,
Pretoria 0001, Republiek van Suid-Afrika. ’n Geldige lys van alle beskik-
bare publikasies kan aangevra word.
Bothalia is vemoem ter ere van Generaal Louis Botha, eerste Eerste
Minister en Minister van Landbou van die Unie van Suid-Afrika. Hierdie
lyfblad van die Nasionale Botaniese Instituut, Pretoria, is gewy aan die
bevordering van die wetenskap van plantkunde. Die hoofgebiede wat gedek
word, is taksonomie, ekologie, anatomie en sitologie. Twee dele van die
tydskrif en ’n indeks van die inhoud, outeurs en onderwerpe verskyn
jaarliks.
MEMOIRS OF THE BOTANICAL SURVEY OF SOUTH AFRICA
MEMOIRS VAN DIE BOTANIESE OPNAME VAN SUID-AFRIKA
The memoirs are individual treatises usually of an ecological nature, ’n Reeks van losstaande omvattende verhandelings oor vemaamlik
but sometimes dealing with taxonomy or economic botany. ekologiese, maar soms ook taksonomiese of plantekonomiese onderwerpe.
THE FLOWERING PLANTS OF AFRICA / DIE BLOMPLANTE VAN AFRIKA
This serial presents colour plates of African plants with accompanying
text. The plates are prepared mainly by the artists at the National Botanical
Institute. Many well known botanical artists have contributed to the series,
such as Cythna Letty (over 700 plates), Kathleen Lansdell, Stella Gower,
Betty Connell, Peter Bally and Fay Anderson. The Editor is pleased to
receive living plants of general interest or of economic value for
illustration.
From Vol. 50, one part of twenty plates is published annually. A volume
consists of two parts. The publication is available in English and Afrikaans.
Hierdie reeks bied kleurplate van Afrikaanse plante met bygaande teks.
Die skilderye word meestal deur die kunstenaars van die Nationale
Botaniese Instituut voorberei. Talle bekende botaniese kunstenaars het
tot die reeks bygedra, soos Cythna Letty (meer as 700 plate), Kathleen
Lansdell, Stella Gower, Betty Connell, Peter Bally en Fay Anderson.
Die Redakteur verwelkom lewende plante van algemene belang of
ekonomiese waarde vir afbeelding.
Vanaf Vol. 50 word een deel, bestaande uit twintig plate, jaarliks
gepubliseer. ’n Volume bestaan uit twee dele. Die publikasie is beskikbaar
in Afrikaans en Engels.
FLORA OF SOUTHERN AFRICA / FLORA VAN SUIDELIKE AFRIKA
A taxonomic treatise on the flora of the Republic of South Africa,
Ciskei, Transkei, Lesotho, Swaziland, Bophuthatswana, South West
Africa/ Namibia, Botswana and Venda. The FSA contains descriptions
of families, genera, species, infraspecific taxa, keys to genera and
species, synonymy, literature and limited specimen citations, as well as
taxonomic and ecological notes. Also available in the FSA series are the
following:
’n Taksonomiese verhandeling oor die flora van die Republiek van Suid-
Afrika, Ciskei, Transkei, Lesotho, Swaziland, Bophuthatswana, SWA/
Namibie, Botswana en Venda. Die FSA bevat beskrywings van families,
genusse, spesies, infraspesifieke taksons, sleutels tot genusse en spesies,
sinonimie, literatuur, verwysings na enkele eksemplare, asook beknopte
taksonomiese en ekologiese aantekeninge. Ook beskikbaar in die FSA-
reeks is die volgende:
The genera of southern African flowering plants, by/deur R.A. Dyer, Vol. 1 Dicotyledons (1975); Vol. 2
Monocotyledons (1976).
Keys to families and index to the genera of southern African flowering plants, by/deur R.A. Dyer (1977).
Plant exploration of southern Africa by Mary Gunn & L.E. Codd. Obtainable ffom/Beskikbaar van: A. A. Balkema
Marketing, Box/Posbus 317, Claremont 7735, RSA.
PALAEOFLORA OF SOUTHERN AFRICA / PALAEOFLORA VAN SUIDELIKE AFRIKA
A palaeoflora on a pattern comparable to that of the Flora of
southern Africa. Much of the information is presented in the form
of tables and photographic plates depicting fossil populations. Now
available:
’n Palaeoflora met ’n uitleg vergelykbaar met did van die Flora van suide-
like Afrika. Baie van die inligting word aangebied in die vorm van tabelle
en fotografiese plate waarop fossiele populasies afgebeeld word. Reeds
beskikbaar:
Molteno Formation (Triassic) Vol. 1. Introduction. Dicroidium , by/deur J.M. & H.M. Anderson.
Molteno Formation (Triassic) Vol. 2. Gymnosperms (excluding Dicroidium ), by/deur J.M. & H.M. Anderson.
Prodromus of South African Megafloras. Devonian to Lower Cretaceous, by/deur J.M. & H.M. Anderson.
Obtainable ffom/Beskikbaar van: A. A. Balkema Marketing, Box/Posbus 317, Claremont 7735, RSA.
BOTHALIA
’N TYDSKRIF VIR PLANTKUNDIGE NAVORSING
A JOURNAL OF BOTANICAL RESEARCH
Volume 20,1
Editor/Redakteur: O.A. Leistner
Assisted by B.A. Momberg
Editorial Board/Redaksieraad
D.F. Cutler
B. de Winter
P.H. Raven
J.R Rourke
M.J. Werger
Royal Botanic Gardens, Kew, UK
Botanical Research Institute, Pretoria, RSA
Missouri Botanical Garden, St Louis, USA
National Botanic Gardens, Kirstenbosch, RSA
University of Utrecht, Utrecht, Netherlands
Editorial Committee/Redaksiekomitee
B. de Winter
D.J.B. Killick
O.A. Leistner
B.A. Momberg
M.C. Rutherford
J.C Scheepers
ISSN 0006 8241
Issued by the National Botanic Gardens/Botanical Research Institute, Private Bag X101, Pretoria 0001,
South Africa
Uitgegee deur die Nasionale Botaniese Tuine/Navorsingsinstituut vir Plantkunde, Privaatsak X101
Pretoria 0001, Suid-Afrika
1990
CONTENTS — EMHOUD
Volume 20,1
1. Studies in the genus Lotononis (Crotalarieae, Fabaceae). 9. Four new species of the L. pentaphylla group,
section Lipozygis. B-E. VAN WYK 1
2. Studies in the genus Lotononis (Crotalarieae, Fabaceae). 12. Four new species of the L. falcata group,
section Leptis. B-E. VAN WYK 9
3. Studies in the genus Lotononis (Crotalarieae, Fabaceae). 13. Two new species and notes on the occur-
rence of cleistogamy in the section Leptis. B-E. VAN WYK 17
4. Studies in the genus Riccia (Marchantiales) from southern Africa. 15. R. hirsuta and R. tomentosa,
sp. nov., two distinct species previously treated as one. O.H. VOLK and S.M. PEROLD .... 23
5. Studies in the genus Riccia (Marchantiales) from southern Africa. 16. R. albomarginata and R. simii,
sp. nov. S.M. PEROLD 31
6. New species of Erica (Ericaceae) from the Cape Province. E.G.H. OLIVER 41
7. Studies in the southern African species of Justicia and Siphonoglossa (Acanthaceae): seeds. K.L.
IMMELMAN 49
8. Studies in the southern African species of Justicia and Siphonoglossa (Acanthaceae): indumentum.
K.L. IMMELMAN 61
9. Notes on African plants:
Asclepiadaceae. Corona lobe variation and the generic position of Asclepias macra. A. NICHOLAS
and D.J. GOYDER 87
Asteraceae. A new species of Pterothrix (Gnaphalieae) from the northern Cape. F. BRUSSE ... 67
Fabaceae. Studies in the genus Lotononis (Crotalarieae. 10. L. esterhuyseniana, a new species
from the south-western Cape. B-E. VAN WYK 70
Fabaceae. Studies in the genus Lotononis (Crotalarieae). 11. A new species of the section
Leobordea from north-western Namibia. B-E. VAN WYK 73
Fabaceae. Studies in the genus Lotononis (Crotalarieae). 14. Three new species of the sections
Telina and Polylobium. B-E. VAN WYK 75
Liliaceae/Asphodelaceae. The correct author citations of Aloe bowiea and A. myriacantha
(Alooideae). G.F. SMITH 80
Poaceae. Two new species of Stipagrostis (Aristideae) from the dune-Namib Desert, Namibia.
B. DE WINTER 82
10. Vegetative morphology and interfire survival strategies in the Cape Fynbos grasses. H.P. LINDER and
R.P. ELLIS 91
11. The ecology of the False Bay estuarine environments. Cape, South Africa. 1. The coastal vegetation.
M. O’CALLAGHAN , 105
12. The ecology of the False Bay estuarine environments, Cape, South Africa. 2. Changes during the last
fifty years. M. O’CALLAGHAN 113
13. Miscellaneous note:
Notes on the distribution and habitat of Aloe bowiea (Liliaceae/Asphodelaceae: Alooideae) — an
endangered and little known species from the eastern Cape. G.F. SMITH and A.E. VAN
WYK 123
14. Obituary: Mary Davidson Gunn (1899—1989). D.M.C. FOURIE 127
15. Book reviews 131
16. Guide for authors to Bothalia 133
Digitized by the Internet Archive
in 2016
https://archive.org/details/bothaliavolume2020unse
Bothalia 20,1: 1-7 (1990)
Studies in the genus Lotononis (Crotalarieae, Fabaceae). 9. Four new
species of the L. pentaphylla group, section Lipozygis
B-E. VAN WYK*
Keywords: Fabaceae, Lotononis section Lipozygis , sectional limits, new species
ABSTRACT
L. pentaphylla (E. Mey.) Benth. and related species were previously shown to be very different from the rest of the section
Lipozygis (E. Mey.) Benth. of Lotononis (DC.) Eckl. & Zeyh. These species are all annuals and can easily be distinguished
by their capitate inflorescences, sessile flowers and indehiscent, wind-dispersed pods. Four new species of this group are
described, namely L. oligocephala B-E. van Wyk, L. globulosa B-E. van Wyk, L. laticeps B-E. van Wyk and L. longicephala
B-E. van Wyk.
UITTREKSEL
Daar is voorheen aangetoon dat L. pentaphylla (E. Mey.) Benth. en verwante spesies baie verskillend is van die res van
die seksie Lipozygis (E. Mey.) Benth. van Lotononis (DC.) Eckl. & Zeyh. Hierdie spesies is almal jaarplante en kan maklik
onderskei word aan hul hofievormige bloeiwyses, sittende blomme en nie-oopspringende, windverspreide peule. Vier nuwe
spesies van hierdie groep word beskryf, naamlik L. oligocephala B-E. van Wyk, L. globulosa B-E. van Wyk, L. laticeps
B-E. van Wyk en L. longicephala B-E. van Wyk.
INTRODUCTION
Lotononis pentaphylla (E. Mey.) Benth. , L. polycephala
(E. Mey.) Benth., L. anthylloides Harv., L. bolusii
Diimmer and L. rosea Diimmer differ from all other
annual species of Lotononis (DC.) Eckl. & Zeyh. in their
densely capitate inflorescences, sessile flowers and
indehiscent, wind-dispersed pods (Van Wyk 1989). These
five species were previously included by Bentham (1843),
Harvey (1862) and Diimmer (1913) in the section Lipozygis
(E. Mey.) Benth., but their annual habit and unusual
morphology have apparently been overlooked. Four new
species that clearly belong to this group are described
below.
Wind-dispersal is not uncommon in the tribe
Crotalarieae but it is usually accomplished by winged
pods, as in the genus Wiborgia Thunb. Personal observa-
tions have shown that wind-dispersal also occurs in
Lotononis benthamiana Diimmer and in Lebeckia
melilotoides Dahlgren. In these two species, the highly
persistent corolla acts as a wing to facilitate dispersal. L.
pentaphylla and its allies however, show an unusual and
interesting mode of seed dispersal. The tiny pods are few-
seeded, totally indehiscent and are completely enclosed
by a densely hirsute, much-inflated calyx. When the seeds
have matured, the calyx with its enclosed pod is shed from
the peduncle. At this stage, the petals are crumpled up and
withered, but the total absence of a pedicel and the hairy,
inflated calyx result in a very effective dispersal by wind.
Only a slight wind is necessary to move the pod (diaspore)
over long distances by a rolling rather than floating action.
Morphological features associated with this dispersal
mechanism are unique within the genus Lotononis and
therefore valuable as diagnostic characters. The total
* Department of Botany, Rand Afrikaans University, P. O. Box 524,
Johannesburg 2000.
MS. received: 1989.04.12.
absence of a pedicel is the most obvious one, and perhaps
also the most useful.
1. Lotononis oligocephala B-E. van Wyk, sp. nov. ,
L. polycephalae (E. Mey.) Benth. valde affrnis, sed foliis
minoribus 5-foliolatis (in L. polycephala 3-foliolatis),
inflorescentiis floribus bracteisque minoribus, lobis calycis
brevioribus latioribusque, atque corolla omnino glabra (in
L. polycephala dense pubescente) differt.
TYPE. — Cape Province, 2918 (Gamoep): Areb, ± 27
miles [43,2 km] NE of Springbok, 25.07.1972, Van der
Westhuizen 276 (PRE, holo.; K, MO, iso.).
Prostrate annual up to 0,8 m wide. Branches sparsely
leafy, densely silky. Leaves invariably 5-foliolate,
(6— )10 — 18( — 24) mm long; petiole ± as long as the
terminal leaflet or longer; leaflets relatively small, broadly
obovate, (3 — )5 — 8(— 10) mm long, (2— )3— 5(— 7) mm
wide, base cuneate, apex truncate to emarginate, densely
silky on both surfaces. Stipules single at each node,
lanceolate to broadly ovate, 3-4 mm long, 2-3 mm wide,
densely silky on both surfaces. Inflorescences in terminal
heads, the heads somewhat flattened, 4— 8-flowered;
peduncle variable in length, 5—50 mm long; bracts large,
broadly obovate, ± 4 X ± 4 mm; bracteoles absent.
Flowers relatively small, 12—14 mm long, yellow; pedicel
absent. Calyx subequally lobed but with the lower lobe
slightly narrower than the upper four lobes, the sinuses
of equal depth; lobes broadly triangular, acute. Standard
oblong, as long as the keel; claw 4-5 mm long; lamina
± 8 mm long, 5—6 mm wide, without lobes or callosities,
glabrous but with a few hairs dorsally along the middle.
Wing petals oblong, ± as long as the keel, glabrous;
auricle small, ± 1 mm long; apex obtuse; sculpturing in
4—5 rows of intercostal lunae, fading into transcostal
lamellae towards the auricle. Keel petals obovate-oblong,
only slightly auriculate; claw 5 mm long; lamina 9 mm
long, 5 mm wide, glabrous; apex rounded. Androecium
2
Bothalia 20,1 (1990)
FIGURE 1. — Lotononis oligocephala.
A, flowering twig; B1 & B2,
leaves and stipules: 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; G1
& G2, pistils: Gl, from young
flower, G2 from older flower;
HI & H2, bracts; II, 12 & 13,
anthers: II, dorsifixed anther,
12, carinal anther, 13, long
basifixed anther. All from Van
der Westhuizen 276. Scales in
mm.
long and narrow; anthers dimorphic; basifixed anthers
oblong, slightly longer than the small ovoid dorsifixed
anthers; carinal anther similar to dorsifixed anthers.
Gynoecium sessile; pistil very small, ovoid-oblong,
pubescent, with 5—12 ovules; style long and slender. Pods
and seeds unknown (Figure 1).
L. oligocephala is closely related to L. polycephala (E.
Mey.) Benth. but can easily be distinguished by the much
smaller and 5-foliolate leaves (3-foliolate in L. polycepha-
la), the smaller inflorescences and flowers, the smaller
bracts, the shorter and wider calyx lobes, and the corolla,
which is totally glabrous (densely pubescent in L. poly-
cephala). This species is known only from the type collec-
tion, which is from northern Namaqualand (Figure 2).
CAPE. — 2918 (Gamoep): Areb, ± 27 miles [43,2 km] NE of Springbok
(—AC), Van der Westhuizen 276 (PRE, holo.; K, MO, iso.).
2. L. globulosa B-E. van Wyk, sp. nov., L. pentaphyl-
lae (E. Mey.) Benth. et L. bolusii Diimmer similis, sed
bracteis maximis late ovatis, floribus paulo maioribus.
vexillo suborbiculari (non oblongo) et foliis semper
3-foliolatis (folia quidem nonnulla 5-foliolata in L.
pentaphylla et L. bolusii). Praesertim similis L. laticipi
B-E. van Wyk, sed ab illo specie inflorescentiis globosis
(non discoideis), bracteis maioribus, vestitura densius
hirsuta, vexillo longiore, lobis calycis latioribus, atque
lobis calycis duobus superioribus latioribus quam inferiori-
bus (superioribus inferioribus aequantibus in L. laticipe
differt).
TYPE. — Cape Province, 3320 (Montagu): 29,5 km
from Touws River to Laingsburg, near Tweedside,
13.10.1986, B-E. van Wyk 2210 (PRE, holo.).
Prostrate annual up to 0,4 m wide. Branches sparsely
leafy, densely to sparsely hirsute. Leaves invariably
3-foliolate, (5 — )8 — 16( — 32) mm long; petiole as long or
usually longer than the terminal leaflet; leaflets compara-
tively small, oblanceolate to obovate, (3 — )5 — 10( — 14) x
(1 — )3 — 4( — 6) mm, base cuneate, apex rounded or rarely
emarginate, abaxial surface sparsely hirsute, adaxial
Bothalia 20,1 (1990)
3
surface glabrous. Stipules single at each node, lanceolate
to oblong, 3-6 x ± 1 mm. Inflorescences in terminal
heads, the heads globose, 8-20-flowered; peduncle
variable in length, usually short, 5-25 mm long; bracts
large, very broadly ovate, (4— )7— 10 x (4— )7— 10 mm;
bracteoles absent. Flowers relatively small, 9-10 mm long,
yellow; pedicel absent. Calyx subequally lobed but with
the two upper lobes slightly wider than the lower lobes,
the sinuses of ± equal depth; lobes narrowly triangular,
acute. Standard suborbicular, as long as the keel or slightly
shorter; claw ± 4 mm long; lamina 5—6 x 5—8 mm,
without lobes or callosities, pubescent over most of the
abaxial surface. Wing petals oblong, + as long as the keel,
pubescent along the apex; auricle small, ± 0,5 mm long;
apex obtuse; sculpturing in 4—5 rows of intercostal lunae,
fading into transcostal lamellae towards the auricle. Keel
petals obovate, only slightly auriculate; claw ± 4 mm
long; lamina 4-6 x 3—4 mm, pubescent over most of
the surface; apex rounded. Androecium long and narrow;
anthers dimorphic; basifixed anthers oblong, slightly
longer than the small ovoid dorsifixed anthers; carinal
anther similar to dorsifixed anthers. Gynoecium sessile;
pistil very small, ovoid-oblong, pubescent, with 5—9
ovules; style long and slender. Pods very small, ovoid,
+ 4 x 2,5 mm, much inflated laterally, totally
indehiscent, enclosed by the persistent and much-inflated
calyx; upper suture minutely verrucose, 2— 3-seeded.
Seeds suborbicular, ±1,5 mm in diameter, testa minutely
but densely tuberculate (Figure 3).
L. globulosa is similar to L. pentaphylla (E. Mey.)
Benth. and L. bolusii Diimmer but differs in the very large,
broadly ovate bracts, the slightly larger flowers, the
suborbicular (not oblong) standard petal and in the
consistently 3-foliolate leaves (always at least some leaves
5-foliolate in L. pentaphylla and L. bolusii). It is
particularly similar to L. laticeps B-E. van Wyk, but differs
from this species in the globose (not discoid) inflores-
cences, the larger bracts, the more hirsute vestiture, the
longer standard petal, the wider calyx lobes and the two
upper calyx lobes, which are wider than the lower lobes
(upper lobes as wide as the lower lobes in L. laticeps).
These differences are clearly shown in Figures 3 & 4.
This species is known from a limited area in the south-
western Cape (Figure 2), where it is perhaps more
common than the very poor herbarium record would
suggest.
CAPE. — 3319 (Worcester): Ceres Division, Gydouw (-AB), Leipoldt
3123 (BOL, K); Ceres District, Laken Vlei (— BC), Compton 12074
(NBG), Levyns 1053 (BOL, SAM). 3320 (Montagu): 29,5 km from Touws
River to Laingsburg, near Tweedside (—AD), B-E. van Wyk 2210 (PRE,
holo.), 2211 (JRAU).
3. L. laticeps B-E. van Wyk, sp. nov., L. globulosae
B-E. van Wyk valde affinis, sed capitulis discoideis (non
globosis), bracteis minoribus, vestitura sparsiori breviori,
vexillo breve carina valde breviori (vexillum carinam in
L. globulosa speciebusque affinibus aequans) differt. A
L. globulosa calyce minori sub-pariter lobato (superiori-
bus inferioribus haud latioribus), lobis angustioribus etiam
differt. A L. pentaphylla, L. bolusii, L. roseaque etiam
folios semper 3-foliolatis, bracteis valde maioribus late
ovatis (non linearibus nec lanceolatis) atque vexillo
suborbiculari (non oblongo) etiam differt.
TYPE. — Cape Province, 3219 (Wuppertal): Ceres
District, Stompiesvlei, Swartruggens (sandy stony plateau,
3500 ft.), 19.11.1961, Esterhuysen 29334 (BOL, holo.; C,
K, M, MO, S, iso.).
Prostrate annual up to 0,3 m wide. Branches sparsely
leafy, minutely hirsute. Leaves invariably 3-foliolate,
(6 — )12 — 15( — 17) mm long; petiole ± as long as the
terminal leaflet or slightly longer; leaflets relatively small,
oblanceolate to obovate, (3 — )5 — 8(— 10) x (1,5 — )3 — 4( — 5)
mm, base cuneate, apex rounded to truncate, sparsely
hirsute on both surfaces. Stipules single at each node,
lanceolate to oblong, 3-5 mm x ± 1 mm. Inflorescences
in terminal heads, the heads discoid (wider than long),
8— 20-flowered; peduncle variable in length, 3—28 mm
long; bracts large, broadly ovate, 4—5 x 4—5 mm;
bracteoles absent. Flowers relatively small, 9—10 mm long,
yellow; sessile. Calyx subequally lobed, the sinuses of ±
equal depth; lobes narrowly linear, acute. Standard
suborbicular, much shorter than the keel; claw 1-2 mm
long; lamina + 3 x + 4 mm, without lobes or callosities,
pubescent over most of the abaxial surface. Wing petals
oblong, shorter than the keel, pubescent along the apex;
auricle small, ± 0,5 mm long; apex obtuse; sculpturing
in 4—5 rows of intercostal lunae, fading into transcostal
lamellae towards the auricle. Keel petals oblong, only
slightly auriculate; claw + 2,5 mm long; lamina + 6 x
± 3 mm, pubescent over most of the surface; apex
rounded. Androecium long and narrow; anthers dimorphic;
basifixed anthers oblong, slightly longer than the small
ovoid dorsifixed anthers; carinal anther similar to dorsi-
fixed anthers. Gynoecium sessile; pistil very small, ovoid-
oblong, pubescent, with 2—4 ovules; style long and
slender. Pods and seeds unknown (Figure 4).
FIGURE 2.— The known geographical distribution of Lotononis
oligocephala. A; L. globulosa, A; L. laticeps, ■; and L.
longicephala, #.
4
Bothalia 20,1 (1990)
FIGURE 'S.—Lotononis globulosa. A1 & A2, inflorescences: Al, young inflorescence, A2, mature inflorescence showing the globose shape
and large bracts; Bl, B2 & B3, leaves and stipules: B1 & B2, adaxial view, B3, 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; HI, mature fruit (dispersal unit or
diaspore), showing the persistent inflated calyx which totally encloses the pod; H2 & H3, pods with the calyx removed: H2, lateral
view, H3, top view; II, 12 & 13, anthers: II, long basifixed anther, 12, carinal anther, 13, dorsifixed anther; J1 & J2, flowers in lateral
view; K, bract; L, seed in lateral view. All from Van Wyk 2210 except J1 from Van Wyk 2211, 12 from Leipoldt 3123. Scales in mm.
Bothalia 20,1 (1990)
5
FIGURE 4. — Lotononis laticeps. A,
flowering twig; Bl, B2 & B3,
leaves and stipules: Bl, abaxial
view, B2 & B3, adaxial 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; HI & H2, bracts; II, 12
& 13, anthers: II, basifixed
anther, 12, carinal anther, 13,
dorsifixed anther; J, flower in
lateral view showing the short
standard petal. All from
Esterhuysen 29334. Scales in
mm.
This poorly known species has so far been recorded only
from a single locality in the Ceres District (Figure 2). It
is very closely related to L. globulosa, but can easily be
distinguished by the short standard petal. Other diagnostic
characters (see Figures 3 & 4) are given under L.
globulosa.
CAPE.— 3219 (Wuppertal): Ceres District, Stompiesvlei, Swartruggens
(—DC), Esterhuysen 29334 (BOL, holo.; C, K, M, MO, S, iso.).
4. L. longicephala B-E. van Wyk, sp. nov., distincta
sine affinitatibus manifestis. Similis est L. pentaphyllae
(E. Mey.) Benth. et L. bolusii Diimmer, sed ab illis
speciebusque omnibus affinibus foliis semper 3-foliolatis,
capitulis valde minoribus oblongis (non globosis nec
discoideis), floribus leguminibusque valde minoribus
differt. A L. globulosa B-E. van Wyk et L. laticipe B-E.
van Wyk etiam bracteis linearibus inconspicuis (non
magnis ovatis) differt.
TYPE. — Cape Province, 3319 (Worcester): flats east
of Prince Alfred’s Hamlet, 10.10.1974, Oliver 5063 (PRE,
holo.; K, MO, STE, iso.).
Prostrate annual, 0,5— 0,8 m wide. Branches sparsely
leafy, sparsely pubescent. Leaves invariably 3-foliolate,
(5— )10 — 14( — 22) mm long; petiole ± as long as the
terminal leaflet; leaflets relatively small, oblanceolate to
obovate, (3— )5— 9(— 12) x (1,5 — )3 — 4(— 6) mm, base
cuneate, apex truncate to emarginate, abaxial surface
sparsely pubescent, adaxial surface glabrous. Stipules
single at each node, lanceolate, 2— 6 X ± 1 mm. Inflores-
cences in terminal or subterminal heads, the heads globose
when young, oblong when fully developed, (4—)
12— 52-flowered; peduncle variable in length, 5-24 mm
long; bracts small, linear or narrowly lanceolate, 2—3 mm
long, up to 0,5 mm wide; bracteoles absent. Flowers very
small, ± 6 mm long, yellow; pedicel absent. Calyx
subequally lobed, the sinuses of ± equal depth; lobes
6
Bothalia 20,1 (1990)
FIGURE 5 .—Lotononis longicephala . A, flowering twig, showing the elongated (spicate) inflorescences; B1 & B2, leaves and stipules: Bl, abaxial
view, B2, adaxial view; C, calyx opened out with the upper lobes to the left (vestiture not shown); D1 & D2, standard petals: Dl, abaxial
view, D2, adaxial view; El & E2, wing petals; FI & F2, keel petals (note sculpturing on F2); Gl, pistil; G2, young pod; HI, mature
fruit (dispersal unit or diaspore), showing the persistent inflated calyx which totally encloses the pod; H2 & H3, pods with the calyx removed:
H2, lateral view, H3, top view; II, 12 & 13, anthers: II, long basifixed anther, 12, carinal anther, 13, dorsifixed anther; J, flower in lateral
view; K, bract; L, seeds in lateral view, showing the sparsely tuberculate surface. All from Van Wyk 2200 except C, Dl, E2, F2, G2 &
K from Esterhuysen 29299. Scales in mm.
Bothalia 20,1 (1990)
7
narrowly triangular, acute. Standard suborbicular, as long
as the keel; claw ± 1,5 mm long; lamina ± 2,5 x ±
4 mm, without lobes or callosities, abaxially pubescent
over most of the basal part. Wing petals oblong, almost
as long as the keel, glabrous except for a few hairs on the
auricle and near the attachment of the claw; auricle small;
apex obtuse; sculpturing in 4-5 rows of intercostal lunae,
fading into transcostal lamellae towards the auricle. Keel
petals oblong, only slightly auriculate; claw ± 2 mm long;
lamina ± 3 x ± 1,5 mm, pubescent at least towards the
apex; apex rounded. Androecium long and narrow; anthers
dimorphic; basifixed anthers broadly oblong, much larger
than the small ovoid dorsifixed anthers; carinal anther
similar to dorsifixed anthers. Gynoecium sessile; pistil very
small, ovoid-oblong, pubescent, with 3-4 ovules; style
long and slender. Pods very small, ovoid, ± 2,5 x ± 1,5
mm, much inflated laterally, totally indehiscent, enclosed
by the persistent and much-inflated calyx; upper suture
minutely verrucose, 2— 3-seeded. Seeds suborbicular, ±
1,2 mm in diameter, testa sparsely tuberculate (Figure 5).
L. longicephala is a distinct species with no obvious
affinities. It is similar to L. pentaphylla and L. bolusii,
but differs from these and all related species in the
consistently 3-foliolate leaves, the much smaller and
oblong (not globose or discoid) heads, the much smaller
flowers and the much smaller pods. It differs from L.
globulosa and L. laticeps also in the inconspicuous, linear
bracts (Figure 5). L. longicephala is known only from the
vicinity of Touws River in the south-western Cape (Figure
2).
CAPE. — 3219 (Wuppertal): Ceres District, E foot of Schurweberg
(next to Bokkeveld Tafelberg) (—CD), Esterhuysen 20631 (BOL); near
the base of Schurweberg Peak (—CD), Esterhuysen 29299 (BOL, C, K,
S). 3319 (Worcester): flats east of Prince Alfred’s Hamlet (—AD), Oliver
5063 (PRE, holo.; K, MO, STE, iso.); Verkeerdevlei, 64,5 km from
Ceres to Touws River (-BD), B-E. van Wyk 2241 (BOL, C, GRA, JRAU,
K, MO, NBG, PRE, SAAS, STE). 3320 (Montagu): 29,5 km from Touws
River to Laingsburg, near Tweedside (—AD), B-E. van Wyk 2200 (JRAU),
2202 (BOL), 2203 (GRA), 2204 (K), 2205 (MO), 2206 (NBG), 2207
(PRE), 2208 (S), 2209 (NH, SAAS, STE).
ACKNOWLEDGEMENTS
I thank the Directors and staff of the cited herbaria for
the loan of specimens. Dr H. F. Glen (Botanical Research
Institute, Pretoria) kindly translated the diagnoses. The
taxonomic study of Lotononis is a registered 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. The London
Journal of Botany 2: 594—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.
HARVEY, W.H. 1862. Leguminosae. In W.H. Harvey & O.W. Sonder,
Flora capensis 2: 47—66. Hodges & Smith, Dublin.
VAN WYK, B-E. 1989. Studies in the genus Lotononis (Crotalarieae,
Fabaceae). VIII. A new species of the L. corymbosa group and
notes on the taxonomy of the section Upozygis. South Afiican
Journal of Botany 55: 528—532.
Bothalia 20,1: 9-16 (1990)
Studies in the genus Lotononis (Crotalarieae, Fabaceae). 12. Four new
species of the L. falcata group, section Leptis
B-E. VAN WYK*
Keywords: Fabaceae, Lotononis section Leptis, new species, southern Africa
ABSTRACT
The taxonomic position of Lotononis falcata (E. Mey.) Benth. and related species is briefly discussed. These species were
previously included in the section Leptis (Eckl. & Zeyh.) Benth. but new information indicates an affinity with the section
Oxydium Benth. Four new species of this group are described: L. fruticoides B-E. van Wyk, L. pachycarpa Dinter ex
B-E. van Wyk, L. lineanfolia B-E. van Wyk and L. schreiberi B-E. van Wyk. The known geographical distribution and
diagnostic characters of the new species are given.
UITTREKSEL
Die taksonomiese posisie van Lotononis falcata (E. Mey.) Benth. en verwante spesies word kortliks bespreek. Hierdie
spesies is voorheen ingesluit in die seksie Leptis (Eckl. & Zeyh.) Benth. maar nuwe inligting dui op ’n verwantskap met
die seksie Oxydium Benth. Vier nuwe spesies van hierdie groep word beskryf: L. fruticoides B-E. van Wyk, L. pachycarpa
Dinter ex B-E. van Wyk, L. linearifolia B-E. van Wyk en L. schreiberi B-E. van Wyk. Die bekende geografiese verspreiding
en diagnostiese kenmerke van die nuwe spesies word aangegee.
INTRODUCTION
A study of Lotononis falcata (E. Mey.) Benth. and its
allies [previously included in the section Leptis (Eckl. &
Zeyh.) Benth.] has shown the presence of four undescribed
species. The purpose of this paper is to describe the new
species and to show that the L. falcata group is better
placed in the section Oxydium Benth. than in Leptis.
The original concept and limits of the genus Leptis Eckl.
& Zeyh. were considerably broadened by Bentham (1843)
when he included it as a section within Lotononis (DC.)
Eckl. & Zeyh. As presently circumscribed (Diimmer
1913), Leptis is undoubtedly an artificial group. It appears
to have been used as a convenient position for species that
did not seem to fit comfortably elsewhere.
Similarities and differences between the section
Oxydium and various groups of the section Leptis are given
in Table 1. The L. laxa and L. falcata groups clearly have
more in common with Oxydium than with Leptis sensu
stricto. This is indicated by chemical similarities as well
as the subequally lobed calyx, the basally dilated standard
petal, the strongly dimorphic anthers and the keel petals,
which are often beaked. In the L. falcata group however,
the keel petals are usually not distinctly beaked but all the
other characters are typical for Oxydium. The panduri-
form shape of the standard petal (see Figures 1, 3-5) is
a useful diagnostic character for the L. falcata group and
the annual habit also distinguishes this group from L. laxa
and related species.
Lotononis fruticoides B-E. van Wyk, sp. nov., L.
falcatae valde affinis sed habitu valde robustiore (planta
annua fruticiformis 0,3 -0,6 m alta), pedunculis longis (L.
falcata pedunculis ± absentibus), inflorescentiis
* Department of Botany, Rand Afrikaans University, R O. Box 524,
Johannesburg 2000.
MS. received 1989.06.06.
plerumque 3:floris (plerumque 1- vel rarius 2-floris in L.
falcata), foliolis angustioribus acutis, leguminibus valde
brevioribus, distributione magis orientali differt.
TYPE. — Cape Province, 3224 (Graaff-Reinet):
Naudesberg Pass, 39 km from Graaff-Reinet, 31.08.1986,
Van Wyk 2020 (PRE, holo.; K, MO, NBG, SAAS, iso.).
Robust annual up to 0,6 m high and wide. Branches
divaricate, rigid, sparsely leafy, glabrescent. All mature
parts (except the corolla) sparsely and minutely strigillose.
Leaves trifoliolate, (6 — )12— 25( — 36) mm long; petiole ±
half as long as the terminal leaflet; leaflets very narrowly
elliptic to linear, (4-)10-20(-30) x (0,5 — )1 — 2 mm,
sparsely and inconspicuously strigillose on both surfaces.
Stipules absent or when very rarely present then single
at each node, linear, 1 mm long. Inflorescences leaf-
opposed, distinctly pedunculate, umbellately (1 — )3( — 5)-
flowered; peduncle short, (3 — )5 — 10( — 27) mm long; bracts
minute, linear, ± 0,5 mm long; bracteoles absent. Flow-
ers 8-10 mm long, yellow; pedicel short, 1-1,5 mm long.
Calyx subequally lobed but with the lower lobe narrower
than the upper four, the sinuses of equal depth; lobes nar-
rowly triangular, acute. Standard panduriform, as long as
the keel; claw much dilated at the base, 2,5-3 x 1—1,5
mm at the base, with a small central callosity; lamina 5—7
X 4-5 mm, without lobes or callosities, glabrous but with
a few minute hairs dorsally along the middle, often with
a reddish brown line along the midrib. Wing petals ob-
long, slightly shorter than the keel, folded into a long shal-
low pocket near the base, glabrous; apex rounded;
sculpturing indistinct or absent. Keel petals half oblong-
elliptic, auriculate, glabrous; apex acute but not beaked.
Androecium with the anthers markedly dimorphic;
basifixed anthers linear, more than 2x longer than the
small ovoid dorsifixed anthers; carinal anther intermediate
in size and shape. Gynoecium subsessile; pistil ovoid-
oblong, sparsely pubescent; style only slightly curved.
Pods subsessile, shortly oblong, 10-14 x 2,5— 3,5 mm,
10
Bothalia 20,1 (1990)
FIGURE 1. — Lotononis fruticoides. A, inflorescence and flower; B, leaf in adaxial view (note absence of stipules); Cl & C2, calyces opened
out with the upper lobes to the left; D1 & D2, standard petals, showing the panduriform shape, dilated claw and central callosity; E, wing
petal (note absence of sculpturing); F, keel petal; G, pistil; HI & H2, pods: HI, in lateral view, H2, in top view, II, 12 & 13, anthers.
II, basifixed anther, 12, carinal anther, 13, dorsifixed anther; J, seed, showing the tuberculate surface. All from Van Wyk 2020 except C2
& D1 from Schutte 220. Scales in mm.
Bothalia 20,1 (1990)
11
TABLE 1. — Summary of similarities and differences between Lotononis section Oxydium and various groups of the section Leptis
compressed, glabrous, apex somewhat cuspidate, upper
suture ± smooth, ± 6-seeded, the seeds cn 1,5-2 mm
long funicles. Seeds subtri angular, ± 1,5 mm in diameter;
testa pale yellow, often with dark purple marks, densely
but minutely tuberculate (Figure 1).
L. fruticoides is closely related to L. falcata, but differs
in the much more robust habit (a shrub-like annual of
0,3— 0,6 m high), the long peduncles (peduncles ± absent
in L. falcata), the predominantly 3-flowered inflorescences
(usually 1- or rarely 2-flowered in L. falcata), the narrower
and acute leaflets, the much shorter pods and in the more
eastern distribution. A form of L. falcata from the
Calvinia-Sutherland area is rather similar to L. fruticoides
and may be confused with it, but in this form the leaflets
are broadly obovate to oblanceolate and the habit totally
different. L. fruticoides has been recorded only from the
south-eastern regions of the Cape Province, where it
appears to be very common (Figure 2). It is highly
cyanogenic (Van Wyk 1989), and stock losses reported
from the Beaufort West area (Burtt Davy 1912) may be due
to this species rather than Dichilus gracilis Eckl. & Zeyh. ,
with which it is often confused. The chromosome number
of L. fruticoides (2n = 18) has been reported by Van Wyk
& Schutte (1988) and the presence of pyrrolizidine
alkaloids by Van Wyk & Verdoom (1989).
CAPE. — 3125 (Steynsburg): Middelburg division, Bangor Farm
(—AC), Bolus s.n. sub BOL 14075 (BOL). 3222 (Beaufort West): ± 80
km west of Beaufort West, back of the mountain on Farm Ezels Korn,
adjoining the Farm Layton (— AB), Shearing 1272 (JRAU); Karoo National
Park, rocky plateau just above Wagenpad Dam (-AD), Bengis 474 (PRE);
Molteno Pass near Beaufort West (-BA), Van Wyk 2137 (JRAU, NBG,
PRE); start of Molteno Pass (— BC), Schutte 219 (GRA, JRAU, K, MO,
NBG, PRE), 220 (BOL, GRA, JRAU, PRE, SAAS, STE). 3223 (Riet-
bron): Courlands Kloof, Nelspoort (-AA), Pearson 832 (NBG). 3224
(Graaff-Reinet): Naudesberg Pass, 39 km from Graaff-Reinet (-BA),
Van Wyk 2020 (PRE, holo.; JRAU, K, MO, NBG, iso.), 2021 (GRA,
JRAU, PRE, S, SAAS), 2022 (JRAU, NBG,
PRE), 2023 (BOL, BLFU, JRAU, STE), 2024 (JRAU, KMG, PRE,
STE), 2025 (JRAU, M, NH, WIND), 2026 (JRAU); C. M. van Wyk
s.n. (JRAU).
L. pachycarpa Dinter ex B-E. van Wyk, sp. nov.
L. pachycarpa Dinter in sched. Amphinomia curtii (Harms) Schreiber
sensu Schreiber: 286 (1957), pro parte. L. leptoloba auct. non H. Bol.:
Schreiber: 82 (1970).
L. linearifoliae valde affinis, sed foliolis oblanceolatis
vel obovatis (non linearibus), stipulis valde maioribus,
pube patentiore, leguminibus ovoideis turgidibus. A L.
curtii forma magnitudineque florium, vexilla panduri-
formi, pedicellis longioribus atque forma leguminum
differt. Etiam cum L. leptoloba confusa est, sed species
altera pedicellos brevissimos in pedunculis longis
(pedicellus longus, pedunculo ± deficiente in L. pachy-
carpa) et flores omnino dissimiles magnitudine et forma
habet. L. leptoloba habet alas valde longiores, vexillam
valde maiorem omnino forma dissimilem, legumina
oblongos (non ovoideos), et distributionem geographicam
dissimilem.
TYPE. — Namibia, 2615 (Liideritz): Halenberg,
29.08.1929, Dinter 6648 (PRE, holo.; BM, BOL, K, M,
SAM, STE, iso.).
FIGURE 2. — The known geographical distribution of Lotononis
fruticoides, •; L. pachycarpa, A; L. linearifolia, A; and L.
schreiberi, ■.
12
Bothalia 20,1 (1990)
FIGURE 3. — Lotononis pachycarpa.
A1 & A2, inflorescences and
flowers (note the long
pedicels); B1 & B2, leaves in
adaxial view; C, calyx opened
out with the upper lobes to the
left; D, standard petal; E, wing
petal; F, keel petal; G, pistil;
HI, H2, H3 & H4, pods: HI
& H2, in lateral view, H3, in
top view, H4, after dehiscence
as viewed from inside (note
verrucose upper suture and
long funicles); II, 12 & 13,
anthers: II, basifixed anther,
12, carinal anther, 13,
dorsifixed anther; J, seed,
showing the tuberculate
surface. A1 from Dinter 4070;
A2, B2, HI, H2, H3 & H4
from Dinter 6648; C, D, E, F,
G, D, 12 & D from MerxmUller
& Giess 28505; B1 & J from
Giess, Volk & Bleissner 5462 .
Scales in mm.
Small annual up to 0,1 m high and up to 0,3 m wide.
Branches divaricate, slender, sparsely leafy. All mature
parts (except the corolla) densely but minutely silky-
strigillose. Leaves trifoliolate, (6-)8-16(-26) mm long;
petiole long or longer than the terminal leaflet; leaflets
oblanceolate to obovate, (2 — )4 — 8( — 14) x (1 — )2 — 3 mm,
sparsely silky-strigillose on both surfaces. Stipules single
at each node, small, lanceolate, 1 — 3( — 4) mm long.
Inflorescences leaf-opposed, invariably single-flowered;
peduncle ± absent, up to 0,5 mm long; bracts very small,
linear, up to 1,5 mm long; bracteoles absent. Flowers
small, 4,5—7 mm long, yellow; pedicel long and slender,
as long or longer than the calyx, up to 10 mm long. Calyx
subequally lobed but with the lower lobe a little narrower
than the upper four and the lateral sinuses slightly
shallower than the upper and lower ones; lobes triangular,
acute. Standard broadly panduriform, as long as the keel;
claw very short, slightly dilated at the base, 1—2 x ±
1 mm, without callosities; lamina 4-5 x 3—4 mm,
without lobes or callosities, glabrous but with a few hairs
dorsally along the middle. Wing petals oblong, slightly
shorter than the keel; apex rounded; sculpturing in
± 3 rows of intercostal lunae fading into a few trans-
costal lamellae towards the auricle. Keel petals
subtriangular, auriculate, glabrous; apex acute but not
beaked. Androecium with the anthers markedly dimorphic;
basifixed anthers oblong, more than 3x longer than
the small ovoid dorsifixed anthers; carinal anther slightly
larger than the dorsifixed anthers. Gynoecium sub-
sessile; pistil ovoid-oblong, pubescent; style only slightly
curved. Pods subsessile, small, ovoid, 6—10 x 3—4 mm,
much inflated laterally, minutely silky-strigillose,
apex obtuse, upper suture minutely but distinctly
verrucose, 8— 15-seeded, the seeds on up to 2 mm long
funicles. Seeds suborbicular, ± 1,5 mm in diameter;
testa brown, densely but minutely tuberculate (Figure
3).
L. pachycarpa is closely related to L. linearifolia, but
differs in the oblanceolate (not linear) leaflets, the much
Bothalia 20,1 (1990)
13
larger stipules, the more spreading vestiture and the ovoid,
turgid pods (Figure 3). From L. curtii it differs in the shape
and size of the flowers, the panduriform standard petal,
the longer pedicels and also in the shape of the pods. It
has also been confused with L. leptoloba (Schreiber 1970)
but the inflorescence structure is only superficially similar
(a long pedicel with the peduncle ± absent in L.
pachycarpa-, a short pedicel on a long peduncle in L.
leptoloba). The flowers are also totally different in size
and structure. L. leptoloba has much longer wing petals,
a much laiger and differently shaped standard petal, oblong
(not ovoid) pods, and a different geographical distribution.
L. pachycarpa occurs in southern Namibia (Figure 2),
whereas L. leptoloba has been recorded only from the
western and north-western parts of the Cape Province.
NAMIBIA. — 2615 (Liideritz): Halenberg (-DA), Dinter 4070 (BOL,
PRE, SAM), 6648 (PRE, holo.; BM, BOL, K, M, SAM, STE, iso.).
2616 (Aus): 12 miles [19,2 km] west of Aus on the way to Liideritz (-CA),
Giess, Volk & Bleissner 5462 (PRE, M, MO, WIND). 2716 (Witputz):
9 km N of Rosh Pinah (—DC), Merxmiiller & Giess 28505 (M, WIND).
2816 (Oranjemund): Namib, west of Obibberge, 14 km S of Obib Wasser
(—BA), Merxmiiller & Giess 32363 (M).
L. linearifolia B-E. van Wyk, sp. nov., L. pachy-
carpae similis, sed foliolis gracilibus linearibus, stipulis
inconspicuis, pube breviori plus arete appressa ac
leguminibus oblongo-linearibus tantum parum inflatis
differt. Etiam similis est L. falcatae, sed ab ilia specie-
busque affmibus pedicello longo gracili (calyce longiore)
ac foliolis anguste linearibus, plus dense sericeo-strigillosis
differt.
TYPE. — Cape Province, 2822 (Glen Lyon): Hay 0.303
[see map in Wilman (1946)], 09.07.1936, Acocks 506 (PRE,
holo.; BOL, KMG, PRE, iso.).
Very small annual up to 0,1 m high and 0,25 m wide.
Branches procumbent, slender, sparsely leafy. All mature
parts (except the corolla) densely but minutely strigillose.
Leaves trifoliolate, slender, (12— )18— 28(— 42) mm long;
petiole usually very much longer than the terminal leaflet;
leaflets very narrowly oblanceolate or usually linear,
(4 — )6 — 14( — 18) x (0,5 — )1 — 1,2( — 2) mm, densely but
minutely strigillose on both surfaces. Stipules single at
each node, inconspicuous, linear, up to 1 mm long. In-
florescences leaf-opposed, invariably single-flowered;
peduncle ± absent, up to 0,5 mm long; bracts very small,
linear, up to 1 mm long; bracteoles absent. Flowers small,
4,5—7 mm long, yellow; pedicel long and slender, as long
or much longer than the calyx, up to 8 mm long. Calyx
subequally lobed but with the lower lobe a little narrower
than the upper four and the lateral sinuses slightly
shallower than the upper and lower ones; lobes triangular,
acute. Standard broadly panduriform, as long as the keel;
claw short, slightly dilated at the base, ± 1,5 x ± 1 mm,
with a central callosity; lamina ±4,5 x ±3 mm, without
lobes or callosities, glabrous but with a few hairs dorsally
along the middle. Wing petals oblong, slightly shorter than
the keel; apex rounded; sculpturing in ± 3 rows of inter-
costal lunae fading into a few transcostal lamellae towards
the auricle. Keel petals shortly oblong, auriculate,
glabrous; apex acute but not beaked. Androecium with the
anthers markedly dimorphic; basifixed anthers oblong,
more than 3x longer than the small ovoid dorsifixed
anthers; carinal anther slightly larger than the dorsifixed
anthers. Gynoecium subsessile; pistil oblong, pubescent;
style curved upwards. Pods subsessile, oblong to linear,
(8 — )10 — 14 x 2,5-3 mm, not inflated laterally, minutely
strigillose, apex obtuse, upper suture distinctly and evenly
verrucose, 10— 15 -seeded, the seeds on up to 1,5 mm long
funicles. Seeds suborbicular, ± 1,2 mm in diameter; testa
brown, densely but minutely tuberculate (Figure 4).
A distinct species, similar to L. pachycarpa but differs
in the slender, linear leaflets, the inconspicuous stipules,
the shorter and more closely appressed vestiture and the
oblong-linear and only slightly inflated pods. It is also
similar to L. falcata, but differs from this and related
species in the long slender pedicel (much longer than the
calyx) and the linear, densely silky-strigillose leaflets
(Figure 4).
L. linearifolia was listed as an unidentified species by
Wilman (1946: 52) and the two specimens from Namibia
were cited by Schreiber (1970: 82) as perhaps belonging
to L. leptoloba sensu Schreiber (= L. pachycarpa). It is
highly cyanogenic (Van Wyk 1989) and, according to notes
on the Kinges specimen in M, is said to have caused sheep
losses. The only known localities (southern Namibia and
the northern Cape) are given in Figure 2, but the species
is probably not as rare as the scanty herbarium record
would suggest.
NAMIBIA. — 2616 (Aus): Liideritz District, Farm Weissenbom
(-AB), Kinges 2433 (M); Namib plain between Neisip and Eureka
(—AD), Merxmiiller & Giess 2876 (M). 2822 (Glen Lyon): Hay 0.303
[near the Orange River W of Groblershoop, see map in Wilman (1946)]
(-CD), Acocks 506 (PRE, holo.; BOL, KMG, PRE, iso.).
L. schreiberi B-E. van Wyk, sp. nov. , species distincta
sine affinitatibus manifestis. L. leptolobae superficialiter
similis, sed stipulis perpusillis inconspicuis, pedunculo
subnullo, vexillo panduriformi (non late cordato) conspicue
striato pubescenti, apice acuto (non obtuso) basi dilatato,
antheris plus valde dimorphis, leguminibus breviter
ellipticis (non oblongis) cum ±12-15 seminibus (non ±
30 seminibus) differt. Etiam L. falcatae speciebusque
affmibus similis, sed ab illis speciebusque omnibus
sectionis Oxydii alis carinisque pubescentibus differt.
TYPE. — Namibia, 2113 (Cape Cross): Cape Cross,
about 0,5 km from the coast, 29.04.1965, Giess 8707 (PRE,
holo.; K, M, WIND, iso.).
Small annual up to 0,1 m high and 0,3 m wide. Branches
prostrate, divaricate, often densely leafy. All mature
parts densely to sparsely pubescent. Leaves trifoliolate,
variable in size, (7— )10— 20(— 28) mm long; petiole as long
or often much longer than the terminal leaflet; leaflets
oblanceolate to obovate, (3 — )5 — 10( — 14) x (2— )3— 5(— 7)
mm; vestiture very variable, densely to sparsely pubescent
on both surfaces, adaxial surface glabrescent and
often totally glabrous at maturity. Stipules single at each
node, minute and inconspicuous, linear, up to 2 mm
long. Inflorescences leaf-opposed, single-flowered, very
rarely 2-flowered; peduncle ± absent, up to 0,5 mm long;
bracts very small and inconspicuous, linear, up to 1 mm
long; bracteoles absent. Flowers 9-11 mm long, yellow;
14
Bothalia 20,1 (1990)
FIGURE 4. — Lotononis linearifolia.
A, inflorescence and flower
(note the long pedicel); B1 &
B2, leaves in adaxial view; C,
calyx opened out with the
upper lobes to the left; D,
standard petal, showing the
panduriform shape of the
lamina, dilated claw and
central callosity; E, wing petal;
F, keel petal; G, pistil; HI, H2
& H3, pods (note verrucose
upper suture): HI & H2, in
lateral view, H3, in top view;
II, 12 & 13, anthers: II,
basifixed anther, 12, carinal an-
ther, 13, dorsifixed anther; J,
seed, showing the tuberculate
surface. All from Acocks 506
except B1 from Merxmuller &
Giess 2876 and H2, H3 & J
from Kinges 2433. Scales in
mm.
pedicel short, 2—3 mm long. Calyx subequally lobed;
lobes long, very narrowly triangular, acute. Standard
broadly panduriform, as long as the keel; claw short,
slightly dilated at the base, ± 2 x ± 1,5 mm, often with
an inconspicuous central callosity; lamina 7—9 x 4—5
mm, without lobes or callosities, strongly striated, dorsal
surface densely pubescent; apex tapering to an acuminate
tip. Wing petals narrowly oblong, slightly shorter than the
keel, distinctly auriculate, pubescent along the lower
margin; apex rounded; sculpturing in ± 5 rows of inter-
costal lunae. Keel petals oblong, distinctly auriculate,
pubescent along the lower half, strongly striated; apex
obtuse. Androecium with the anthers markedly dimorphic;
basifixed anthers oblong, more than 2x longer than the
ovoid dorsifixed anthers; carinal anther slighdy larger than
the dorsifixed anthers. Gynoecium subsessile; pistil
oblong, densely pubescent; style long and slender, upper
part curved upwards. Pods subsessile, shortly oblong, 8—10
x 3—3,5 mm, only slightly inflated laterally, pubescent,
apex obtuse, upper suture distinctly verrucose,
12-15-seeded, the seeds on up to 1,5 mm long funicles.
Seeds suborbicular, ± 1,2 mm in diameter; testa brown,
densely tuberculate (Figure 5).
This species (named after Dr Annelis Schreiber of the
Botanische Staatssammlung Miinchen) was listed as L.
spec. (no. 18) in the Prodromus einer Flora von Siid-
westafrika (Schreiber 1970). L. schreiberi is a very distinct
new species and has no obvious affinities. The relatively
large flowers, pubescent and striated petals and single-
flowered inflorescences are useful diagnostic characters
(Figure 5). It is superficially similar to L. leptoloba, but
differs in the very small, inconspicuous stipules, the
panduriform (not broadly cordate) and conspicuously
striated, pubescent standard petal, with an acute (not
obtuse) apex and a dilated base, the more markedly
dimorphic anthers, the shortly elliptic (not oblong) pods
that are ± 12-15-seeded (not ± 30-seeded). It is also su-
perficially similar to L. falcata and related species, but
Bothalia 20,1 (1990)
15
differs from these and all other species of the section
Oxydium in the pubescent wing and keel petals (L.
arenicola Schltr. is the only species of Oxydium with
pubescent wing and keel petals but the morphology of this
species is totally different). Several collections of L.
schreiberi have been made in the north-western parts of
Namibia, where it appears to be quite common (Figure 2).
NAMIBIA. — 2013 (Unjab Mouth): Skedelkuspark (—AC), Venter
9025 (BLFU); 23 km SE of Torra Bay (-AD), Giess 8020 (M, PRE,
WIND); 14 miles [22,4 km] E of Torra Bay (-AD), Giess, Volk &
Bleissner 6277 (M, PRE, WIND); 15 miles [24 km] SE of Torra Bay
(—AD), Nordenstam 3789 (M). 2014 (Welwitschia): Welwitsch,
Damaraland (— BD), Galpin & Pearson sub Galpin 7547 (PRE, SAM);
koppies S of Ugab River Station (— DD), Muller & Loutit 1096 (M,
WIND). 2113 (Cape Cross): flats a few miles inland on road to Brandberg
West Mine (— BD), Oliver & Muller 6660 (PRE, partly); Cape Cross,
about 0,5 km from the coast (— DD), Giess 8707 (PRE, holo.; K, M,
WIND, iso.). 2114 (Uis): S of the Messumberge ± 12 miles [19,2 km]
E of the coast (—BA), Giess 3576 (PRE, WIND). 2214 (Swakopmund):
track from Goanicontes to Rossing Mountain (— DB), Kers 1311 (WIND).
2215 (Trekkopje): Farm Nordenberg, Swakop River at Tsavischab
(— CA), Kers 4 (WIND); Swakop River, along the track from Swakop
at Tsavischab to Farm Nordenberg and Karibib (— CA), Kers 1532
(WIND). 2315 (Rostock): Kuiseb River, SE of Hope Mine, on the Namib
plain between Hope Mine and Garob Mine (-CB), Kers 1580, 1586
(WIND).
ACKNOWLEDGEMENTS
I 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 registered 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 : 472 —481, 559 —613.
BURTT DAVY, J. 1912. Botanical investigations into gal-lamziekte.
Reports of the Veterinary Research Institute of South Africa 2:
181-196.
FIGURE 5. — Lotononis schreiberi.
A, inflorescence and flower
(note the absence of a
peduncle); B1 & B2, leaves in
adaxial view, showing the
variation in pubescence and
leaflet shape (note the small
stipules); C, calyx opened out
with the upper lobes to the left;
D, standard petal, showing the
panduriform shape of the
lamina, dilated claw and
central callosity; E, wing petal
(note vestiture); F, keel petal
(note vestiture); G, pistil; HI,
H2 & H3, pods (note verrucose
upper suture): HI & H2, in
lateral view, H3, in top view;
II, 12 & 13, anthers: II,
basifixed anther, 12, carinal
anther, 13, dorsifixed anther; J,
seed, showing the tuberculate
surface. All from Giess 8020
except Bl, C, D, E, F & G
from Kers 1586. Scales in mm.
16
Bothalia 20,1 (1990)
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.
SCHREIBER, A. 1957. Amphinomia DC. Mitteilungen aus der
Botanischen Staatssammlung, Miinchen 2: 286—289.
SCHREIBER, A. 1970. Fabaceae. In H. Merxmiiller, Prodromus einer
Flora von Sildwestafrika 60: 76—85. Cramer, Lehre.
VAN WYK, B-E., 1989. The taxonomic value of cyanogenesis in
Lotononis and related genera. Biochemical Systematics and
Ecology 17: 297-303.
VAN WYK, B-E. & SCHUTTE, A. L. 1988. Chromosome numbers
in Lotononis and Buchenroedera (Fabaceae— Crotalarieae). Annals
of the Missouri Botanical Garden 75: 1603—1607.
VAN WYK, B-E. & VERDOORN, G. H. 1989. A chemotaxonomic
survey of major alkaloids in Lotononis and Buchenroedera.
Biochemical Systematics and Ecology 17: 385—389.
WILMAN, A. 1946. Preliminary checklist of the flowering plants and
ferns of Griqualand West. Deighton Bell, Cambridge.
Bothalia 20,1: 17-22 (1990)
Studies in the genus Lotononis (Crotalarieae, Fabaceae). 13. Two new
species and notes on the occurrence of cleistogamy in the section Leptis
B-E. VAN WYK*
Keywords: cleistogamy, Fabaceae, flower dimorphism, Lotononis section Leptis, new species
ABSTRACT
The occurrence of flower dimorphism in the genus Lotononis (DC.) Eckl. & Zeyh. is reported for the first time. Cleistogamous
flowers have been observed in 12 species from four different groups of the sections Leptis (Eckl. & Zeyh.) Benth. and Oxydium
Benth. Morphological differences between chasmogamous and cleistogamous flowers are discussed and illustrated. The
phenomenon of flower dimorphism appears to be of limited taxonomic value but nevertheless supports the idea of an affinity
between the L. laxa, L. pungens and L. leptoloba groups. Two recently discovered new species of the L. leptoloba and
L. calycina groups, L. venosa B-E. van Wyk and L. acuticarpa B-E. van Wyk, are described.
UITTREKSEL
Die voorkoms van blomdimorfisme in die genus Lotononis (DC.) Eckl. & Zeyh. word vir die eerste keer gerapporteer.
Kleistogame blomme is by 12 spesies van vier verskillende groepe van die seksies Leptis (Eckl. & Zeyh.) Benth. en Oxydium
Benth. waargeneem. Morfologiese verskille tussen chasmogame en kleistogame blomme word bespreek en gefllustreer. Die
verskynsel van blomdimorfisme is klaarblyklik van beperkte taksonomiese waarde maar ondersteun nietemin die idee van
’n verwantskap tussen die L. laxa-, L. pungens- en L. leptoloba- groepe. Twee nuwe spesies van die L. leptoloba- en L.
calycina-gmtpt wat onlangs ontdek is, L. venosa B-E. van Wyk en L. acuticarpa B-E. van Wyk, word beskryf.
INTRODUCTION
The occurrence of flower dimorphism in the genus
Lotononis (DC.) Eckl. & Zeyh. is reported here for the
first time. Morphological differentiation between
chasmogamous and cleistogamous flowers appears to be
restricted to the sections Leptis (Eckl. & Zeyh.) Benth.
and Oxydium Benth. As presently circumscribed (Bentham
1843; Diimmer 1913), Leptis was recently shown to be an
artificial group and it was suggested that some species
would be much better placed in Oxydium (Van Wyk 1990).
The taxonomic value of cleistogamy is briefly discussed
and two recently discovered new species of Leptis sensu
lato are described below.
Cleistogamy has been reported from several genera of
the Fabaceae (Uphof 1938; Arroyo 1981). Precocious bud
pollination, with little or no effect on flower morphology,
is known to occur in Lotononis bainesii Bak.f. (Byth 1964)
and in the genus Dichilus DC. (Schutte 1988) and may
be more common in the Crotalarieae than was previously
recognized. Morphological differentiation between
cleistogamous and non-cleistogamous flowers of the same
species however, is less common (Arroyo 1981). Flower
dimorphism in the genus Argyrolobium Eckl. & Zeyh. was
discussed in detail by Harms (1909, 1917) but no reports
for other genera of the tribe Crotalarieae could be found.
FLOWER DIMORPHISM IN LOTONONIS
Dimorphic flowers were observed in 12 species of the
sections Leptis and Oxydium (Table 1). Unlike precocious
bud-pollination, which is difficult to observe (and which
probably occurs in many species of Lotononis ), flower
dimorphism is readily detected in herbarium material.
* Department of Botany, Rand Afrikaans University, P. O. Box 524,
lohannesburg 2000.
MS. received: 1989.06.06.
Bud-cleistogamy in the species listed in Table 1 results
in a marked reduction in the size of the corolla,
androecium and style but has only a slight effect on the
calyx. The corolla does not open, but is usually pushed
out of the calyx by the developing ovary. Figure 1 shows
a typical example of flower dimorphism in Lotononis — the
two flowers illustrated are from different branches of a
single plant. It is also possible to recognize (by the remains
of the style) those pods which were formed from
cleistogamous flowers. The size and shape of the pods and
TABLE 1. — Flower dimorphism in various groups of the sections Leptis
and Oxydium. The occurrence, observed frequency and degree of
differentiation are indicated
18
Bothalia 20,1 (1990)
FIGURE 1. — Flower and fruit dimor-
phism in Lotononis pungens.
Morphological differences
between flowers and pods
taken from the same individual
(Van Wyk 1626b, JRAU)) are
shown above, and details of a
cleistogamous flower from
Schutte 215 (JRAU) below.
Al— A6, chasmogamous flow-
er: Al, calyx opened out with
upper lobes to the left; A2,
standard petal; A3, wing petal;
A4, keel petal; A5, androe-
cium; A6, pistil. B1-B6,
cleistogamous flower: Bl,
calyx; B2, standard petal; B3,
wing petal; B4, keel petal; B5,
androecium; B6, pistil. Cl &
C2, pods: Cl, pod formed
from a chasmogamous flower;
C2, pod formed from a
cleistogamous flower (note the
small hooked stylar tissue and
the anthers adhering to the stig-
ma). Dl— D4, cleistogamous
flower: Dl, standard petal; D2,
wing petal (note sculpturing);
D3, keel petal; D4, pistil.
the number of seeds are apparently not affected, but the
old stylar tissue is short and hooked, often with one or
more anthers adhering to the stigma. Scanning electron
microscope studies have shown that pollen tubes grow right
through the anther wall and into the stigma, thereby firm-
ly attaching the anther to the stigma. Cleistogamy in
Lotononis seems identical to that described in species of
Lespedeza Mich. (Hanson & Cope 1955) and in
Omithopus L. (Wojciechowska 1972).
difference between cleistogamous and non-cleistogamous
flowers — intermediate stages are rare or absent. Many
factors are known to cause cleistogamy (Uphof 1938;
Erickson 1975) and it may be worthwhile to gain
experimental evidence for the mechanisms that influence
cleistogamy in Lotononis. Personal observations have
indicated that it is a seasonal phenomenon (at least in L.
laxa ) and that it may be induced by unfavourable climatic
conditions.
It is obvious that the 12 species listed in Table 1 are only
facultatively cleistogamic and that they show no particular
adaptation towards cleistogamy. The presence of petal
sculpturing for example, indicates that the normal develop-
ment of the flower bud is arrested at a relatively late stage
of development. Furthermore, the occurrence of both
flower forms at the same locality (and even on a single
specimen) shows that cleistogamy is not a permanent
condition. It is nevertheless significant that there is a clear
Cleistogamy is generally considered to have limited
value as a taxonomic character because of its variability
and the likelihood of convergence (Uphof 1938). In the
genus Lotononis, it has indeed led to taxonomic errors,
such as a confusion between L. calycina (E. Mey.) Benth.
and cleistogamous forms of L. tenella (Harvey 1862). The
former is not cleistogamous but the very short corolla
results in a superficial similarity with L. tenella. Early
bud-cleistogamy (i.e. that which leads to dimorphic
Bothalia 20,1 (1990)
19
flowers) does not appear to be randomly distributed in
Lotononis. A direct relationship between the section
Oxydium and some groups of the section Leptis is
indicated, and other characters were shown to support this
idea (Van Wyk 1990). Paradoxically, most of the species
with dimorphic flowers have acute or beaked keel petals,
a character that is more readily associated with outcrossing
because it promotes an increased efficiency in the
transfer of pollen. This remarkable versatility may partly
explain why L. laxa and L. tenella have a wider geo-
graphical distribution than most other perennial species
of Lotononis.
The two new species described below clearly illustrate
that cleistogamy has limited value as a taxonomic
character. L. venosa B-E. van Wyk is apparently not
cleistogamic but is here placed in the L. leptoloba group,
while L. acuticarpa B-E. van Wyk is the only species of
the L. calycina group known to display at least some
degree of flower dimorphism (see Table 1).
Lotononis venosa B-E. van Wyk, sp. nov., L.
leptolobae H. Bol. affinis sed habitu minore foliosiore,
foliis anguste oblongis vel linearibus (in L. leptoloba
obovatis) stipulis geminis disperse dispositis (in L.
leptoloba semper singularis), floribus leguminibusque
valde maioribus differt.
TYPE. — Cape Province, 3220 (Sutherland): De Hoop
in Klein Roggeveld, 06.09.1986, Oliver 8965 (PRE, holo.;
STE, iso.).
Small prostrate annual up to 0,2 m wide. Branches
densely leafy; twigs sparsely pilose with long spreading
hairs. Leaves digitately trifoliolate, very variable in size,
sparsely pilose with long spreading hairs; petioles slightly
winged, as long as the terminal leaflet or longer, (2—)
3 — 7( — 16) mm long; leaflets narrowly oblong to linear,
(2 — )5 — 12( — 14) x (0,5 — )1 — 2( — 2,5) mm, abaxially
sparsely pilose, adaxially glabrescent. Stipules large,
consistently present, single at each node or rarely paired
at some nodes, oblong to narrowly oblong and often
slightly falcate, similar to the leaflets or much larger,
(2— )6— 12(— 14) X (0,5 — )1,5 — 2,5( — 3,5) mm. Inflores-
cences subterminal and leaf-opposed towards the branch
ends, invariably single-flowered; peduncle usually ± as
long as the calyx, (3 — )6 — 8( — 12) mm long; bracts
relatively large, narrowly linear, 5-8 mm long; bracteoles
absent. Flowers large, 16-20 mm long, pale yellow with
grey venation; pedicel short, ± 2 mm long. Calyx almost
as long as the corolla, equally lobed, sparsely pilose; lobes
very long and slender. Standard very large, broadly ovate;
claw 4—5 mm long, very slightly dilated at the base, di-
lated part up to 1,8 mm wide; lamina acute, cordate at the
base, 13—15 x 13-16 mm, glabrous but with a line of hairs
dorsally along the middle. Wing petals oblong, much
longer than the keel, glabrous, apex obliquely truncate;
sculpturing in 4-5 rows of thin transcostal lunae and
lamellae. Keel petals small, semi-circular, acute but not
beaked, auriculate and pocketed near the base, glabrous.
Anthers dimorphic, basifixed anthers 2x longer than the
dorsifixed anthers, carinal anther similar to dorsifixed
anthers. Pistil 11—13 mm long; ovary oblong, ± 6 mm
long, densely pubescent; style sharply curved. Immature
pods oblong, slightly turgid, longer than the calyx, up to
18 X 5 mm, ± 30-seeded, upper suture distinctly
verrucose, sparsely pilose; funicles up to 2 mm long. Seed
unknown (Figure 2).
L. venosa is a distinct species known only from two
recent collections in the Roggeveld area (Figure 4). The
annual habit, general morphology, inflorescence structure
and flower morphology are very similar to that of L.
leptoloba. It differs, however, from this species in its
smaller, leafier habit, its narrowly oblong to linear leaves
(obovate in L. leptoloba), stipules which may be paired
(in L. leptoloba always single), and in the much larger
flowers and pods. Although there is no evidence of
cleistogamy in this species and mature pods are unknown,
it is placed in the affinity of L. leptoloba with some
confidence. The flowers turn from yellow to a bluish
colour when dried and the species may therefore be
mistaken for L. maximiliani Schltr. , but the shape of the
leaflets and the very large flowers (Figure 2) are quite
distinct.
FIGURE 2. — Lotononis venosa. A, flower in lateral view showing the
long peduncle and the vestiture of the calyx and standard petal;
B1 & B2, leaves in adaxial view, showing paired stipules (Bl)
and a single stipule (B2); C, calyx opened out, upper lobes to
the left (vestiture not shown); D, standard petal; E, wing petal;
F, keel petal; G, pistil; H, bract; I, androecium. All from Oliver
8965.
20
Bothalia 20,1 (1990)
FIGURE 3. — Lotononis acuticarpa. A, habit; B, leaf in abaxial view; C, calyx opened out, upper lobes to the left showing the fusion of the
lateral lobes (vestiture not shown); D, standard petal; E, wing petal; F, keel petal; G, pistil; HI & H2, mature fruit in lateral view (note
the size, shape and also the pointed, tapering apices). 11—13, anthers: II, basifixed anther; 12, carinal anther; 13, dorsifixed anther. J, flower
in lateral view; K, seeds in lateral view, showing tuberculate surfaces; L, peduncle with pedicel and bract. All from Van Wyk 1815. Scales in mm.
Bothalia 20,1(1990)
21
FIGURE 4. — The known geographical distribution of Lotononis venosa,
O ; and L. acuticarpa , ▲.
CAPE. — 3119 (Calvinia): ± 3,6 km from the Farm Blomfontein to
De Hoop (-DD), 13.09.1986, Schutte 259 (JRAU). 3220 (Sutherland):
De Hoop in Klein Roggeveld (—DC), 06.09.1986, Oliver 8965 (PRE,
holo.; STE, iso.).
Lotononis acuticarpa B-E. van Wyk, sp. nov., L.
calycinae similis sed habitu foliisque minore, floribus
valde minoribus angustioribus, praesertim forma magnitu-
dineque fructus differt. Fructus calycem valde superantia
(calycem aequantes in L. calycina), plani (non turgidi ut
in L. calycina ) et apices sunt acuti sursum curvati (non
obtusi deorsum curvati ut in L. calycina). Etiam similis
estL. adpressae N.E. Br. sed ab ilia specie habitu minori
annuo, superficie adaxiali glabro foliolorum et fructu valde
longiori acutiori subfalcato differt.
TYPE. — Transvaal, 2628 (Johannesburg): Springs Dis-
trict, 5 km from Devon radar station to Leandra (-BD),
20.02.1986, Van Wyk 1815 (PRE, holo.; JRAU, K, MO,
NBG, iso.). Figure 3.
Small procumbent annual up to 0,1 m high and 0,2 m
wide. Branches slender, spreading from a very short main
axis; twigs sparsely pilose. Leaves digitately trifoliolate,
sparsely pubescent, variable in size, those on the main axis
with long and slender petioles, those of lateral twigs with
the petiole 3 — 6( — 8) mm long; leaflets oblanceolate to
elliptic, (3— )5 — 10( — 14) x (1 — )2 — 2,5( — 3) mm, abaxially
sparsely pubescent, adaxially glabrous. Stipules consis-
tently present, single at each node, oblanceolate to elliptic,
2-4 mm long. Inflorescences leaf-opposed at each node,
fasciculate, 1-3-flowered; peduncle very short or absent;
bracts small, up to 2,5 mm long, often inserted slightly
above the pedicel base; bracteoles absent. Flowers very
small, narrow, 6-7 mm long, yellow; pedicel short, up
to 2 mm long. Calyx almost as long as the corolla, with
the lateral lobes on either side fused much higher up in
pairs, sparsely pubescent; lobes long and slender. Standard
oblong-ovate, 6—7 mm long; claw 1,5—2 mm long; lamina
acute, densely pubescent on the dorsal surface. Wing petals
narrowly oblong, slightly shorter than the keel, pubescent
towards the rounded apex; sculpturing in 3—4 rows of
transcostal lunae fading into a few transcostal lamellae
towards the auricle. Keel petals oblong, obtuse, auriculate
and pocketed near the base, pubescent along the lower edge
of the lamina. Anthers dimorphic, basifixed anthers more
than 2x longer than the dorsifixed anthers, carinal an-
ther intermediate in size. Pistil 5—6 mm long; ovary ob-
long, ± 3 mm long, densely pubescent; style short, almost
straight. Pods oblong, compressed, slightly falcate, twice
as long as the calyx, 8—12 X 2—3,5 mm, gradually taper-
ing to the acute apex, ± 6-seeded, upper suture ± smooth,
indehiscent or tardily dehiscent, inconspicuously pubes-
cent. Seed suborbicular in side view, up to 1,8 mm long;
funicles up to 2 mm long; testa brown, often yellowish
brown mottled with black, distinctly tuberculate (Figure
3). Chromosome number. 2n = 18!
L. acuticarpa is similar to L. calycina (E. Mey.) Benth.
but differs in the smaller habit and leaves, the much smaller
and narrower flowers and particularly in the shape and
size of the fruit. The fruit are much longer than the calyx
(as long as the calyx in L. calycina), flat (not turgid as
in L. calycina) and the apices are acute and curved upwards
(not obtuse and curved downwards as in L. calycina). It
is also similar to L. adpressa N. E. Br. but differs from
this species in the smaller and annual habit, the glabrous
adaxial surface of the leaflets and in the much longer, more
acute and slightly falcate fruit.
It is remarkable that this distinct species has escaped
the notice of plant collectors for so long. L. acuticarpa
was only recently collected for the first time and is now
known from three different localities in the south-eastern
Transvaal (Figure 4). It appears to be restricted to well-
drained stony soils and is locally abundant, at least at the
type locality.
TRANSVAAL. — 2628 (Johannesburg): Springs District, 5 km from
Devon radar station to Leandra (-BD), 20.02.1986, Van Wyk 1815 (PRE,
holo.; JRAU, K, MO, NBG, iso.), 22.02.1987, Van Wyk 2625 (M, NH,
S, STE). 2630 (Carolina): Mbabane, Kalkoenkranz, Goedemoed Farm
(-CC), 15.03.1987, Turner 1468 (PRE). 2729 (Volksrust): Frankfort,
Amersfoort, Bergvliet Farm (— BB), 01.04.1987, Turner 1624 (PRE).
ACKNOWLEDGEMENTS
I thank the directors and staff of the cited herbaria for
the loan of specimens. Dr H. F. Glen (Botanical Research
Institute, Pretoria) kindly translated the diagnoses. The
taxonomic study of Lotononis is a registered Ph. D. project
at the University of Cape Town.
REFERENCES
ARROYO, M.T.K. 1981. Breeding systems and pollination biology in
Leguminosae. In R.M. Polhill & P.H. Raven, Advances in legume
systematics 2 : 723 —769. Royal Botanic Gardens, Kew.
BENTHAM, G. 1843. Enumeration of Leguminosae, indigenous to
southern Asia, and central and southern Africa. Hooker’s London
Journal of Botany 2: 594—613.
BYTH, D.E. 1964. Breeding system and chromosome number in
Lotononis bainesii Baker. Nature 202: 830.
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.
ERICKSON, E.H. 1975. Variability of floral characteristics influences
honey bee visitation to soybean blossoms. Crop Science 15:
767-771.
22
Bothalia 20,1 (1990)
HANSON, C.H. & COPE, W.A. 1955. Reproduction in the cleistoga-
mous flowers of ten perennial species of Lespedeza. American
Journal of Botany 42: 624—627.
HARMS, H. 1909. Uber Kleistogamie bei der Gattung Argyrolobium.
Berichte der Deutschen Botanischen Gesellschaft 27: 85—96.
HARMS, H. 1917. Weitere Beobachtungen uber Kleistogamie bei
afrikanischen Arten der Gattung Argyrolobium. Berichte der
Deutschen Botanischen Gesellschaft 35: 175—186. HARVEY,
W.H. 1862. Leguminosae. In W.H. Harvey & O.W. Sonder, Flora
capensis 2: 47—66. Hodges & Smith, Dublin.
SCHUTTE, A.L. 1988. 'n Taksonomiese studie van die genus Dichilus
DC. (Fabaceae—Crotalarieae). M.Sc. thesis, Rand Afrikaans
University, Johannesburg.
UPHOF, J.C.Th. 1938. Cleistogamic flowers. The Botanical Review 4:
21-49.
VAN WYK, B-E. 1990. Studies in the genus Lotononis. 12. Four new
species of the L. falcata group, section Leptis. Bothalia 20:
9-16.
WOJCIECHOWSKA, B. 1972. Pollination and fertilization in Omithopus
sp. Genetica Polonica 13 : 37—52.
Bothalia 20,1: 23—29 (1990)
Studies in the genus Riccia (Marchantiales) from southern Africa. 15.
R. hirsuta and R. tomentosa , sp. nov., two distinct species previously
treated as one
O.H. VOLK* and S.M. PEROLD**
Keywords: Marchantiales, Riccia hirsuta, R. tomentosa, section Pilifer, section Micantes, section Pannosae, southern Africa, subgenus Thallocarpus ,
taxonomy, tetrad spores
ABSTRACT
The description of Riccia hirsuta (Volk & Perold 1986) is emended and new illustrations of the species are provided,
as the original description and illustrations were based on two distinct, but rather similar species. Examination of newly
collected material of R. hirsuta, showed it to belong to subgenus Riccia, section Pilifer (Volk 1983). Section Micantes (Volk
& Perold 1986) of which this was the type species, is therefore transferred to section Pilifer. R. tomentosa Volk & Perold,
sp. nov., described here, is at first glance somewhat similar to R. hirsuta in its habit, hence the earlier confusion in Volk
& Perold (1986). It also resembles R. hirsuta in its dorsal covering of long hairs and in the triangular scales apically split
into cellular strands. R. tomentosa, however, differs from R. hirsuta in the spongy (not compact) construction of the thallus
and in the spores permanently united in tetrads (not single). It belongs to subgenus Thallocarpus and is the type species
of the new section Pannosae.
UITTREKSEL
Riccia hirsuta (Volk & Perold 1986) word herbeskryf en herillustreer, aangesien die vroeere beskrywing en illustrasies
op twee verskillende, maar tog ietwat soortgelyke spesies gebaseer was. Die ondersoek van nuwe materiaal van R. hirsuta,
het getoon dat dit tot subgenus Riccia , seksie Pilifer (Volk 1983) behoort. Seksie Micantes (Volk & Perold 1986) waarvan
dit die tipe-spesie was, word derhalwe onder seksie Pilifer geplaas. R. tomentosa Volk & Perold, sp. nov., hier beskryf,
is by die eerste aanblik taamlik soortgelyk aan R. hirsuta wat groeivorm betref, vandaar die vroeere verwarring in Volk
& Perold (1986). Dit toon ook ooreenkoms met R. hirsuta in die dorsale bedekking van lang hare en in die driehoekige
skubbe wat apikaal in sellulere stringe verdeel is. R. tomentosa verskil egter van R. hirsuta in die sponserige (nie kompakte)
bou van die tallus en in die spore wat permanent in tetrades verenig is (nie enkel nie). Dit behoort tot die subgenus Thallocarpus
en is die tipe-spesie van die nuwe seksie Pannosae.
INTRODUCTION
When R. hirsuta Volk & Perold (1986), the figures of
which are referred to below as 1986, was described, it was
assumed that all five specimens then listed under this
species, definitely belonged here. Among these specimens,
only Oliver 8040 had mature spores. Even though there
were some indications that the material was not quite
homogeneous, the possibility of another, rather similar
species, existing in the same area seemed remote, and it
was assumed that all material belonged to one species.
During fieldwork by Perold in Namaqualand in
September/October 1987 and August/September 1988 and
by Oliver in July 1989, six new collections of densely hairy
specimens were made, each one with distinctly papillose
spores which on maturity remain in permanent, ±
globular tetrahedral tetrads. These collections belong to
a new species, R. tomentosa (see below). Fresh gatherings
of R. hirsuta, also with a dense dorsal hair covering but
with single, reticulately ornamented spores were also
made, thus providing more material for study.
Riccia hirsuta emend. Volk & Perold
Monoica (?), viridi-grisea, mediocris, perennis; frons
usque ad 10 mm longa, 2—4 mm lata, duplo ad triplo latior
* Botanische Anstalten d. Univ. Wurzburg D8700, Germany, BRD.
** National Botanical Institute, Private Bag X101, Pretoria 0001.
MS. received: 1989.02.20.
quam crassa, simplex vel furcata, obcuneata vel oblonga,
apice breviter emarginata sparsimque sulcata, antice
convexa, in sicco subplana ad concava, dense hirsuta ob
pilos longos (inde nomen speciei), marginibus plus minus
attenuatis; costa lata, crassa, subplana vel convexa, ad
margines sensim excurrens. Stratum aeriferum canalibus
aeriferis altis. Squamae grandes, marginem frondis
superantes, imbricatae, hyalinae, deltatae, apicibus in filis
liberis scissis, vel dentatis. Sporae triangulo-globulares,
polares, brunneae, 115—125 pm diametro, late alatae,
margine subtiliter crenato, imperfecte reticulatae,
granulatae; areolae in diametro sporarum 6-8, centrales
majorae. Chromosomatum numerus n = 8 (Bomefeld
1984).
Thallus monoicous (?), perennial, scattered, not in
rosettes, dorsal surface hirsute with thick pelt of shiny hairs
(Figure 2A), whitish along margins, greenish grey over
centre; medium-sized to large (Figures LA, C; 1986: LA);
branches simple or bifurcate, up to 10 mm long, 2, 0-4,0
mm wide, 1, 5-2,0 mm thick, i.e. about once to twice
wider than thick; oblong, broadening towards truncate apex
(Figure 2B); groove short, soon becoming flat; margins
subacute; flanks sloping outwards in a short wing (Figure
ID), greyish green, occasionally with some reddish pur-
ple flecks; ventral surface slightly rounded to plane, pale
green; when dry (Figure IB), sides partly inflexed, dorsal
surface grey, dusty from accumulation of sand grains
trapped between hairs.
24
Bothalia 20,1 (1990)
FIGURE 1. — Riccia hirsuta . Morphology and anatomy. A, old thallus
with sporangia; B, dry thallus, margins indexed; C, two young
thalli from culture; D, transverse section through mature thal-
lus; E, dorsal hair pillars in transverse section; F, horizontal sec-
tion at level of stomata (hatched) and basal cells of hairs (hairs
omitted) with air canals (stippled) and chlorenchyma also shown;
G, scale; H, chromosomes. A, B, H, Oliver 8040 (Type); C,
E-G, S.M. Perold 2182\ D, Oliver 8038a. A-G by Volk; H by
Bomefeld. Drawings by J. Kimpton. Scale bars on A— D = 1
mm; E— G = 100 Jim; H = 1 /am.
Anatomy : dorsal covering of free-standing, water-
repellent, hyaline, straight to bent, closely packed hair-like
cell pillars or filaments of nearly equal length, up to 1 200
gm long, occupying up to almost *;2 the thickness of the
thallus (Figure ID), consisting of four to six elongated,
thin-walled cells (Figures IE, 2C), up to 375 x 40-100
/tm, gradually tapering to an often abruptly bent apex; air
pores mostly 4-sided (Figure 2E), but varying from
3 -6-sided, in contact with about four epithelial cells and
leading to rather wide (40—80 p m), mostly 6-sided air
canals (Figure IF); assimilation tissue (chlorenchyma)
about 300—500 pm thick, almost ^5 — 1/4, the thickness
of thallus; storage tissue about 700 /im thick, consisting
of polygonal cells, up to 50 jam wide; in older resting
thalli, sometimes central core filled with fatty oil and
starch (Figure 1986: 1C3); rhizoids arising from ventral
epidermis and base of scales, hyaline, some smooth and
others tuberculate, up to 25 jam wide. Scales partly
extending above margin of thallus, overlapping apically,
triangular (Figures 1G; 2F; 1986: 1E1— 3), about 650 jam
wide at base and up to 1 500 jam high, hyaline, occasion-
ally with reddish purple cells at base; cells thin-walled,
in body of scale up to ± 180 x 50 jam, oblong-hexagonal,
apex dentate, or more usually cells elongated and
separating into a few short, variously bent or twisted
filaments (Figures 1G; 2F; 1986: LF1 — 4). Antheridia flask-
shaped, with much elevated necks (Figures 2D; 1986: 1G),
hidden by dorsal pillars. Archegonia with purple necks.
Sporangia arranged across width of thallus, up to 700 pm
wide, overlying tissue often tinged with purple, each
containing about 650 spores. Spores triangular-globular,
polar, deep dull brown to nearly black, semitransparent
to opaque, (95— )115— 125 (—130) pm in diameter, with
wing about 10 jum wide, granular, slightly undulating,
margin crenulate to somewhat eroded, at angles with a
pore or notched (Figures 2H, I; 1986: 3F); distal face
reticulate, with 3— 5(— 6) large central areolae, 25—38 pm
wide, completely or incompletely subdivided into smaller
areolae, about 12,5 pm wide, often with a papilla in the
middle (Figure 2H, I), occasionally areolae equally wide
and then 8—10 across diameter; central ridges thick and
high, outer ridges thinner and lower, sometimes extending
partly on to wing (Figure 1986: 3C, D); proximal face with
triradiate mark distinct, but poorly delineated, each facet
irregularly and rarely completely reticulate (Figures 2G;
1986: 3A, B). Chromosome number n = 8 (Bornefeld in
Volk & Perold 1986) (Figure 1H).
R. hirsuta is extremely rare and only known from the
plateau N of Leliefontein (Figure 3) (altitude ± 1 500 m
above sea level), in a winter rainfall area with less than
200 mm rain per annum. It grows in association with other
Riccia species, e.g. R. limbata Bisch., R. bullosa Link
ex Lindenb., R. cupulifera A.V. Duthie and R. schelpei
Volk & Perold, on clayey soil, at a seepage area or in
seasonally damp sandy places between small shrublets and
Restionaceae.
R. hirsuta is distinguished from other species in section
Pilifer by the very tall, shiny dorsal cell pillars and by the
triangular scales, apically split into filamentous, short,
loose, cellular strands.
SPECIMENS EXAMINED
CAPE. — 3018 (Kamiesberg): Kamiesberg, lower slopes east of
Rooibeig, damp ground between and under renoster bushes (—AC), Oliver
7240 p.p. (F; PRE); Kamiesberg plateau, north of Leliefontein, towards
Bothalia 20,1 (1990)
25
FIGURE 2. — Riccia hirsute. Morphology and anatomy, spore ornamentation. A, two young thalli; B, apex of thallus from above, densely hairy;
C, apices of hair pillars; D, part of transverse section, showing bases of hairs and young antheridial neck; E, two rows of 4-sided air pores,
from above, hairs partly removed; F, thallus flank with four scales, two central ones with appendages; G, proximal face of spore; H, distal
face; I, side view of distal face. A, B, D, Oliver 8040 (Type); C, E— I, S.M. Perold 2101. A,D, LM micrographs by Volk; B, C, E— I,
SEM micrographs by Perold. Scale bars on A = 1 mm; B— F = 100 jim; G— I = 50 /x m.
Draaiklip.on sandy, periodically moist soil, associated with other Riccia
species and Restionaceae, (—AC), Oliver 8038(a), 8040 (PRE, holo.);
on road between Kamieskroon and Leliefontein, 5 km north of Lelie-
fontein, at seepage area (—AC), S.M. Perold 2099—2101; 2182 (PRE).
REJECTION OF SECTION MICANTES VOLK & PEROLD
Section Pilifer Volk in Mitteilungen aus der
Botanischen Staatssammlung, Miinchen 19: 453 (1983).
Type species: Riccia albomarginata Bisch. ex Krauss*
Section Micantes Volk & Perold in Bothalia 16: 187 (1986). Type
species: Riccia hirsuta Volk & Perold.
R. hirsuta, as here emended, agrees in its anatomy and
spores with members of the section Pilifer and is there-
fore placed under that section. Section Micantes, of which
it previously was the type species (Volk & Perold 1986),
is now rejected.
* See Perold (1990) [one of the present authors (S.M.P.) is of the opinion
that the name R. albomarginata has been misapplied since Sim (1926)] .
Riccia tomentosa Volk & Perold, sp. nov.
Dioica (?), pallida, magna, perennis; frondes ad 18 mm
longae, ad 5 mm latae, ad 4 mm crassae, late liguliformae,
FIGURE 3. — Distribution map of Riccia hirsuta, ♦; and R. tomento-
sa, #, in southern Africa.
26
Bothalia 20,1 (1990)
FIGURE 4. — Riccia tomentosa. Morphology and anatomy. A, mature thallus; B, older thallus, flanks indexed towards base; Cl-4 transverse
sections through thallus branches (Cl, near apex; C2, C3 at different distances along length of B, C4 through oldest dying part); D, variable
length of filaments on dorsal surface; E, hair base with supporting cells; F, lacunae and epidermis; G, horizontal section near dorsal surface
viewed from below, hence hairs not visible: bottom right, epidermis with stomata (hatched); above, chlorenchyma with lacunae (stippled);
H, scale with filaments at apex; I, chromosomes. A, B, C2— 4, D— F, H, S.M. Perold & Crosby 2157\ Cl, Schelpe 7784', G, S.M. Perold
1495\ I, Le Roux & Fourie PRE-CH4494. A-H by Volk; I by Bomefeld. Drawings by J. Kimpton, after Volk. Scale bars on A-C =
1 mm; D, H = 500 /xm; E— G = 100 /im; I = 1 /zm.
Bothalia 20,1 (1990)
27
concavae, simplices vel furcatae, subgregariae vel
singulares; pagina superiora in sicco tomentosa (inde
nomen) pilis longitudine inaequali usque ad 2 700 /xm
longis usque ad 14 cellulis compositis vestita; costa semi-
lunata vel subplana in alas breves excurrens, alae in sicco
erectae; stomata distantes, aliquot cellulis circumdata;
lacunae aeriferae ad 280 /xm dilatatae; squamae
triangulares, ad 1 500 /xm longae, apicem versus sparsim
filamentosae. Sporae in tetradibus tetraedrica globularibus
permanenter conjunctae, 100-130-145 /xm diametro,
sulphureae vel fuscatae, subglobosae, dense papillosae,
semiopacae vel opacae. Chromosomatum numerus n =
8 (Bomefeld 1989).
TYPE. — Cape Province, 3018 (Kamiesberg): Pedros-
kloof, on road to Rooifontein, 2 km beyond Willem Stone
Bridge, on sandy soil overlying clay (-AA), 1987.09.24,
S.M. Perold 1495 (PRE, holo.), associated with other
Riccia spp. and Bryum spp.
Thallus dioicous (?), perennial, scattered or in crowded
patches, dorsal surface ‘shaggy -haired or tomentose
(Figure 4A, B; 1986: IB, 2A), silvery to dusty grey, often
with trapped sand particles between hairs; branches large,
up to 18 mm long, simple or bifurcate, segments ± 12
X 2-4-5 mm, narrower toward base (Figure 4C4),
oblong to ovate-oblong, apex slightly narrowed, shortly
emarginate, groove short and wide, middle part concave,
±3-4 mm thick, i.e. almost as wide as thick in section
(Figure 4C1), margins raised, obtuse, shortly winged
(Figures 4C2, C3; 5B), flanks sloping steeply upward and
outward, lower parts occasionally coloured red; ventral
face rounded to plane, pale green; when dry, whitish,
wide, dorsally deeply concave in centre, wings erect or
scarcely inflexed to somewhat reflexed.
Anatomy : dorsal surface covered by free, straight or bent
hair-like filaments or pillars (Figures 4D; 5D; 1986: ID),
up to 2 700 /xm long, tapering upwards from a broad base
and composed of up to 14 short or elongated thin-walled,
hyaline cells of variable length and thickness (Figure 5E),
often supported by slightly raised epidermal cells (Figure
4E, 1986: 2D); air pores circumscribed by several (5-7)
radially arranged, wedge-shaped cells (Figures 4G; 5F;
1986: 2G, H); assimilation tissue (chlorenchyma) ± 500
/xm thick, 1;8 - 1;6 the thickness of thallus, with sloping,
elongated, polygonal air chambers or lacunae up to 280
/xm wide (Figures 4F; 1986: 2E), 37— 62(— 112) /xm wide,
longest and widest in wings (Figure 5C3) surrounded by
isodiametric cells in plates, one cell thick and up to 20
cells high; storage tissue ± 500-600 /xm thick, 1;8- !/6
the thickness of thallus, cells angular, up to 50 /xm wide,
with small spaces between. Rhizoids arising ventrally and
from base of scales, forming a thick mat, some smooth,
mostly tuberculate, 12—25 /xm wide. Scales triangular,
large, up to 1 500 /xm long and base 1 250 /xm wide,
hyaline, often red at base, apically with filaments up to
1 000 /xm long (Figure 4H; 1986: 2F), cells in body of
scale 5-sided, walls straight to slightly sinuous, up to 112
x 25—42 /xm, smaller at base, 62 x 40 /xm; at one side
of scale, a row of rectangular cells, up to 180 x 40 /xm.
Antheridia and archegonia difficult to see, as obscured
by dense dorsal hairs. Sporangia bulging dorsally, ± 800
/xm wide, each containing ± 300 spore tetrads. Spores
thick-walled, permanently united in tetrads (Figure 6A,
B, E, F), 116-131-145 /xm in diameter, pale yellow to
ferruginous brown, thickly beset with papillae or
verruculae, 3—5 /xm wide and projecting as much, obtuse
or truncate, arising from nodes of scarcely visible (Figure
6C), to obvious reticulum (Figure 6D). Chromosome
number n = 8 (Bomefeld 1989).
R. tomentosa is endemic and rare, as it has only been
found at a few localities in Namaqualand (Figure 3), at
altitudes of 1 000-1 200 m above sea level, with winter
FIGURE 5. — Riccia tomentosa. Morphology and anatomy. A, two young thalli; B, transverse section through thallus branch; C, transverse section
through wing with wide lacunae; D, hairs from above; E, variable sizes of dorsal hairs; F, epidermis with air pores. A, S.M. Perold &
Crosby 2157\ B-D, S.M. Perold 1556\ E, S.M. Perold 1495 ; F, Le Roux & Fourie PRE-CH4494. A, by A. Romanowski; B-E, SEM micrographs
by S.M. Perold. Scale bars on A, B, D = 1 mm; C, E, F = 100 /xm.
28
Bothalia 20,1 (1990)
rainfall of less than 200 mm per annum. It grows on
reddish brown, sandy soil, overlying clay, at pH values
5,6; 6,6; 6,8 and 7,5, in open areas and in association with
Bryum argenteum Hedw. Its distribution is sympatric with
that of R. hirsuta.
R. tomentosa is dorsally tomentose, hence the specific
epithet, and differs from other southern African Riccia
species, e.g. R. hirsuta (Table 1), by the unique hair-like
outgrowths from many of the epidermal (not epithelial)
cells and by the papillose to verruculose tetrad spores. It
shares large triangular scales, apically splitting into
filamentous cellular strands with R. hirsuta.
SPECIMENS EXAMINED
CAPE. — 2917 (Springbok): Hester Malan Res., Carolusberg N, at
gate on western boundary (— DB), Le Roux & Fourie PRE-CH4494 (PRE);
Schelpe 7784 (BOL, PC, PRE). 3017 (Hondeklipbaai): Arkoep area, 15
km N of Kamieskroon, road towards Brak water, sandy clay flats, open
ground (— BB), Oliver 9196, 9197 (PRE). 3018 (Kamiesberg): 18 km NE
of Kamieskroon on road to Rooifontein, at rock outcrop, on soil (— AA),
Perold & Reid 1462 (PRE); 4 km on road to Rooifontein, after turnoff
from Kamieskroon/Leliefbntein road, sandy soil (— AA), Perold & Crosby
2157, 2158 (PRE); Pedroskloof, on road to Rooifontein, 2 km beyond
Willem Stone Bridge (— AA), S.M. Perold 1495 (PRE, holo.); 29 km
SE from Platbakkies on road to Kliprand, Farm Banke, rock outcrops,
sandy soil (— BC), S.M. Perold 1556 (PRE). Vogel C5446 (MJG),
Nuwefontein, Hondeklipbaai, may belong here, but its identity is uncertain
as it is sterile).
Among the species of Riccia presently known, R. tomen-
tosa occupies a unique position as it has both primitive
and derived characters. The spores of subgenus
Thallocarpus which remain in tetrads, are regarded as
primitive (Jovet-Ast 1987), as are also the ‘loose’ construc-
tion of the assimilation tissue with wide lacunae, and the
chlorophyllose epidermis which is pierced by air pores
(stomata) that are circumscribed and separated by a ring
of cells. Regarded as derived characters are the hairy
covering of the dorsal surface of the thallus (similar to that
in the section Pilifer, subgenus Riccia ) and the raised basal
cells of the larger hairs.
SECTION PANNOSAE
Pannosae Perold, sect, nov., subgen. Thallocarpus
(Lindb.) Jovet-Ast. Sporae permanenter conjunctae in
tetradibus; thallus dorsaliter pannosus ob pilos longos (inde
nomen).
Ripe spores permanently united in tetrads; thallus
dorsally felt-like (pannosus) because of long hairs.
TYPE. — R. tomentosa Volk & Perold.
Pannosae is a new section of subgenus Thallocarpus.
R. tomentosa is the type species and only known species
of this new section, which is characterized by spores per-
manently joined together in tetrads, as in other members
of subgenus Thallocarpus (Lindb.) Jovet-Ast, but differs
from all hitherto known species of this subgenus by a
thick felt-like (Lat. pannosus) covering of very long, multi-
cellular hairs.
CORRECTION OF PUBLISHED ILLUSTRATIONS
The existence of two similar-looking species, in the same
area, regrettably gave rise to mistakes not only in the
naming of the specimens, but also in the illustrations (Volk
& Perold 1986). Specimens from both species were used
for Figures 1 and 2 (pp. 188 & 189). For Figures IB, H
and 2A— H, Le Roux & Fourie PRE-CH4494 was used and
for Figure ID, Schelpe 7784 ; these two specimens were
collected from the same population, and are R. tomentosa-,
firstly, because the air pores are distant and secondly,
because the spores from Schelpe 7784, though immature
and diseased, nevertheless suggest that the ornamentation
FIGURE 6. — Riccia tomentosa. Spores. A, three tetrads; B, one tetrad; C, join mostly hidden by tubercles; D, reticulum between tubercles;
E, F, tetrads. A, E, S.M. Perold 1556\ B, C, F, S.M. Perold 1495\ D, S.M. Perold & Crosby 2157. A— D, SEM micrographs by S.M. Perold;
E, F, LM micrographs by Volk. Scale bars on A— D = 50 /rm; width of tetrads on E, F = ± 125 jim.
Bothalia 20,1 (1990)
29
TABLE 1. — Differences between R. hirsuta and R. tomentosa
R. hirsuta
R. tomentosa
1. spores single; ornamentation reticulate
2. hairs of dorsal covering crowded, ± equally long
3. base of hairs without supporting cells
4. epithelium with unspecialized air pores closely spaced, each epithelial
cell in contact with several air spaces
5. air canals short, fairly wide
6. thallus medium-sized, dorsally flat to ± convex
7. found in exposed locality at seepage, or in light shade at area under
shrublets
spores in tetrads; ornamentation papillate
dorsal hairs spaced, of variable length
base of some hairs with raised supporting cells
epidermis with well-spaced stomata, radially circumscribed by 5— 7
wedge-shaped cells; some cells without contact with air pores
air chambers long, wide, sloping
thallus large, concave dorsally
found in fully exposed localities in sandy, drier areas
could be papillate to tuberculate. Unfortunately, these clues
were subsequently ignored, when examining Oliver 8040
(the type specimen of R. hirsuta Volk & Perold emend.),
which has very crowded dorsal hairs and mature,
triangular-globular spores with ± incomplete reticulate
ornamentation on both faces, as illustrated in Figure 3 p.
190 (Volk & Perold 1986).
The captions should accordingly be corrected as follows:
for Figure 1, R. hirsuta A, Cl— 3, El— 3, FI— 4, G ( Oliver
8040, 8038b); R. tomentosa B, H (Le Roux & Fourie PRE-
CH4494); D, Schelpe 7784; and for Figure 2, R. tomen-
tosa (not R. hirsuta ) Le Roux & Fourie PRE-CH4494.
ACKNOWLEDGEMENTS
The authors wish to thank Dr habil. T. Bomefeld, Am
Reele 1, D-8706, Hochberg, W Germany for the chromo-
some counts and figures and the Curator of BOL,
University of Cape Town for the loan of the late Prof. E. A.
Schelpe’s specimen. Sincere thanks to Dr E.O. Campbell,
Massey University, New Zealand and Dr E.W. Jones,
Oxford, for reading the manuscript and for their valuable
criticism.
REFERENCES
BORNEFELD, T. 1989. The Riccia species of S and SW Africa.
Chromosome numbers and composition of the chromosome sets.
Nova Hedwigia 48 : 371—382.
JOVET-AST, S. 1975. Precisions sur les caracteres de deux Riccia du
sous-genre Thallocarpus. Revue Bryologique et Lichenologique
41: 449 -456.
JOVET-AST, S. 1987. Vers une classification phylog£n6tique des especes
du genre Riccia. The Bryologist 90: 321—330.
KRAUSS, F. 1846. Pflanzen des Cap- und Natal-Landes, gesammelt von
Dr. Ferdinand Krauss. Flora 29: 135, 136.
PEROLD, S.M. 1989. Spore-wall ornamentation as an aid in identifying
the southern African species of Riccia (Hepaticae). Journal of
the Hattori Botanical Laboratory 67: 109 —201.
PEROLD, S.M. 1990. Studies in the genus Riccia (Marchantiales) from
southern Africa. 16. R. albomarginata and R. simii sp. nov.
Bothalia 20: 31-39.
SIM, T.R. 1926. The bryophyta of South Africa. Transactions of the Royal
Society of South Africa 15: 1-475.
VAN ZINDEREN BARKER, E.M. 1956. South African pollen grains
and spores. Part II: 61—132. Balkema, Amsterdam.
VOLK, O.H. 1981. Beitrage zur Kenntnis der Lebermoose (Hepaticae)
aus Siidwestaffika (Namibia). II. Mitteilungen aus der Botanischen
Staatssammlung, Milnchen 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 1988. R. crenatodentata (Marchantiales) sp. nov. aus
Arabien. Studien liber Bryophyten aus Arabien 11. Nova Hedwigia
46: 27-35.
VOLK, O.H. & PEROLD, S.M. 1986. Studies in the genus Riccia
(Marchantiales) from southern Africa. 6. R. hirsuta, a new species
in a new section. Bothalia 16: 187—191.
Bothalia 20,1: 31-39 (1990)
Studies in the genus Riccia (Marchantiales) from southern Africa. 16.
R. albomarginata and R. simii, sp. nov.
S. M. PEROLD*
Keywords: Marchantiales, Riccia albomarginata, R. concava, R. simii, sp. nov., Riccia section Pilifer, southern Africa, taxonomy
ABSTRACT
A description of Riccia albomarginata Bisch. ex Krauss, augmented with the aid of new collections which most closely
match the relevant fragments of the type material, is presented. The type collection comprises two species. The fragments
that I consider to be in closest agreement with Krauss’s protologue are selected as lectotype. As far as could be established
the rest of the type material is probably referable to R. concava Bisch. As previously mentioned by Perold (1989b), Sim
(1926) and subsequent authors had applied the name 'R. albomarginata' to a different taxon. This taxon, R. albomarginata
auct. non Bisch., is now described and illustrated as R. simii, sp. nov. Its distribution and ecology are also noted.
UITTREKSEL
’n Beskrywing van Riccia albomarginata Bisch. ex Krauss, uitgebrei met behulp van nuwe versamelings wat die nouste
met die toepaslike fragmente van die tipeversameling ooreenkom, word gegee. Die tipeversameling bestaan uit twee spesies.
Die fragment wat na my mening die nouste met Krauss se protoloog ooreenstem, word as lektotipe aangewys. Sover vasgestel
kon word, verteenwoordig die res van die tipemateriaal waarskynlik R. concava Bisch. Soos reeds vermeld deur Perold
(1989b), het Sim (1926) en latere outeurs die naam 'R. albomarginata' op ’n ander takson toegepas. Hierdie takson, R.
albomarginata auct. non Bisch., word nou beskryf en geillustreer as R. simii, sp. nov. en die verspreiding en ekologie word
ook vermeld.
INTRODUCTION
The collections discussed below are in a poor to very
poor condition. The relevant descriptions by early authors
are very incomplete and lack reference to important
diagnostic features such as the free-standing dorsal cell
pillars. Furthermore, no illustrations were provided by
these authors. In addition, species of section Pilifer, to
which the species under discussion belong, are often very
difficult to distinguish, particularly in a dry state, as the
dorsal pillars remain collapsed and cannot be reconstituted
to examine their shape and size. Nevertheless, after a
thorough investigation of all available evidence and on the
basis of expertise gained through the study of numerous
specimens of species belonging to the section Pilifer, I have
arrived at the following conclusions: the collection of R.
albomarginata, annotated by Bischoff and held at BM
(presumably Krauss’s collection) is decidedly a mixed
collection; die parts alpha and beta referred to by Bischoff
on the specimen label (or varieties alpha and beta
according to Gottsche et al. 1846) belong to different
species, and var. beta is probably referable to R. concava
Bisch. ex Krauss; R. albomarginata has no cilia along the
thallus margins and had been incorrectly classified by
Gottsche et al. under their section Ciliatae.
Riccia albomarginata Bisch. ex Krauss in Flora 29:
135 (March 1846); Gottsche, Lindenberg et Nees ab
Esenbeek: 604 (Oct. 1846) p.p. as to var. alpha ; Stephani:
329 (1898) (he did not see the type collection).
TYPE. — Cape, In Promontorio Bonae Spei, Krauss
Julio 1838 (BM!, lecto. here designated: the specimen in
the middle of the herbarium sheet).
* National Botanical Institute, Private Bag X101, Pretoria 0001.
MS: received 1989.02.20.
Thallus monoicous (?), perennial, in crowded gregari-
ous patches, in partial rosettes or scattered, olivaceous
green to green when actively growing, somewhat velvety,
with hyaline scales extending above thallus margins
(Figures 1A; 2D); rather small (Figure 2A), branches once
to several times symmetrically or asymmetrically furcate,
medium divergent, 5—7 mm long, terminal segments 1—3
mm long (Figure 2C), 0,7— 1,8 mm wide and 0,6— 1,1 mm
thick, i.e. scarcely wider to nearly twice wider than thick
in cross section (Figure IE); Ungulate to oblong or linear,
apex rounded, emarginate, deeply grooved apically, soon
widely concave dorsally (Figure 2B); margins subacute,
flanks steeply rising, brown, occasionally with some
purple colouring, distally covered by fragile, hyaline scales,
basally often denuded of scales; ventral face gently
rounded, green to brown; when dry (Figure IB), dorsally
concave, greenish white to light brown, scurfy or streaked
with thin white threads of collapsed epithelial cell pillars,
margins proximaUy incurved, undulating, with crisp, white
or hyaline scales extending above parchment-like, brown
flanks; distally sometimes tightly inflexed with scales
meeting and clasped together along midline.
Anatomy of thallus: dorsal epithelium (Figures 1C; 2E)
consisting of 3— 4(— 5) elongated cells in free-standing,
fragile, hyaline pillars, ± 130-200(-230) /xm high, apical
cell conical, narrowing toward tip, or of uniform width,
sometimes bent, with apex rounded, 45—65 x 20—30 /xm,
second and third cells 42—60 x 32—37 /xm, basal cell(s)
25-37 x 30—40 /xm; from above, pillars quite densely
crowded together, fine and delicate, proximally collapsed,
air pores mostly obscured, small, 4-5-sided (Figure ID);
assimilation tissue ± 250—350 /xm thick in section, ±
!/3 the thickness of thallus and consisting of vertical
columns of 7 or 8 isodiametric to short-rectangular cells,
32-47 x 32 /xm, enclosing narrow air canals; storage
32
Bothalia 20,1 (1990)
FIGURE 1. — Riccia albomarginata . Morphology and anatomy. A,
thallus wet; B, thallus dry; C, transverse section through dorsal
cell pillars at margin and scale; D, air pores from above; E,
transverse section through thallus; F, scale. A, C, E, S.M. Perold
1979 ; B, S.M. Perold 2118; C, D, S.M. Perold 538; F, S.M. Perold
2031 p.p. Drawings by J. Kimpton. Scale bar on A, B, E = 1
mm; C, D = 50 jitm; F = 100 ^tm.
tissue ± 300 gm thick, occupying ventral part of thallus,
cells rounded, ±31 pm wide; rhizoids arising from ventral
epidermal cells and from base of scales, ± 25 ju,m wide,
some smooth, others tuberculate. Scales (Figures IF; 2F)
rounded, imbricate, hyaline, fragile, extending up to ±
150 (tm above thallus margins, 700— 800 x 400 fim, cells
in body of scales hexagonal, 60- 80(-100) x 40 /tm,
smaller at edge, cell walls straight, sometimes stained yel-
lowish, base occasionally with a little purple colouring;
basally scales absent. Antheridia numerous. Archegonia
with purple necks. Sporangia along length of branches,
single or occasionally in pairs, bulging dorsally and each
containing ± 300 spores. Spores (75— )80— 95(-105) pm
in diameter, triangular-globular, polar, brown to dark
brown, semi-transparent to opaque, with wing 5—1 pm
wide, wider at perforated angles, margin + smooth to
faintly crenulate; ornamentation reticulate and somewhat
similar, to rather dissimilar on the two faces, with ± 14
irregular areolae across diameter of distal face (Figure 3B),
complete or incomplete, up to 7 pm wide, walls thick,
slightly raised at nodes, otherwise smooth, convoluted
(Figure 3C) or anastomosing into wide ridges that radiate
outwards from centre (Figure 3E); proximal face with
triradiate mark poorly to well defined (Figure 3A); ±
30—35 small, completely or incompletely separated
areolae on each facet, walls thick, convoluted , smooth,
but raised at nodes (Figure 3D).
The above description and illustrations are based on
S.M. Perold 538 p.p., 1930, 1979 and 2118.
As can be seen from Figure 3B, C, E, the spore
ornamentation in R. albomarginata is quite variable,
ranging from smallish areolae and a few thick radiating
ridges, to mostly very thick, prominent ridges. On the
whole, the ornamentation is not markedly different from
that of R. concava (Perold 1989) or from the Zeyher
collection of R. albomarginata (see below) (the part
provisionally referred by me to var. beta), and the spores
of these taxa would only be distinguished with difficulty
by conventional light microscopical examination.
R. albomarginata has no outstanding vegetative
characters by which it can be readily recognized; however,
it is generally rather smaller than most other species in
section Pilifer. In the dry state, it can frequently be
distinguished by the light brown colouring of the dorsal
face and the incurved flanks, fringed with hyaline or crisp,
white scales that contrast strongly with the brown flanks,
to which, it is thought, the specific epithet refers. Stephani
(1898) described the scales as being prominent. Later
authors, such as Sim, apparently assumed that the word
‘prominent’ also meant ‘large’ and therefore erroneously
concluded that R. albomarginata is a species with large
scales. In some cultured specimens, the brown colouration
of the thalli and pale yellowish stain of the scale cell walls
are persistent, even after a year. Sometimes tarry smudges
are found at the flanks, as was also noticed in the var. alpha
part of the Krauss specimen. It is doubtful whether this
is the result of algal or fungal infestation.
R. albomarginata generally grows on rather coarse,
gravelly soil, overlying granitic or sandstone outcrops, at
altitudes between 200 and 1 300 m above sea level, and
receiving less than 200 to 400 mm of mostly winter rain
per annum. It is often found in association with other
Riccia species, e.g. R. nigrella DC. (which is rather
smaller and much darker brown dorsally) and with R.
concava Bisch., as well as with moss species, e.g. Barbula
crinita Schultz, Bryum spp. and Brachymenium spp. It is
Bothalia 20,1 (1990)
33
FIGURE 2. — Riccia albomarginata. Morphology and anatomy. A, young thallus, field-grown; B, young branch, cultured; C, older branch, cultured;
D, scales at margin; E, dorsal cells; F, part of scale. A, S.M. Perold 2031; B, C, E, S.M. Perold 1979 ; D, S.M. Perold 538; F, Zeyher
p.p. A— E, SEM micrographs; F, LM micrograph. Scale bar on A— D = 1 mm; E, F = 50 m.
not common and its distribution appears to be confined
to a few areas in the north-western and south-western Cape
(Figure 4), which Zeyher (the other collector referred to
by Gottsche et al.) is known to have also visited (Gunn
& Codd 1981). Although the label on his collection reads
‘Prom. b. spei’, Gottsche et al. reported it to be from ‘ter-
ra Capensi’, which could have been anywhere in the Cape.
Krauss’s type specimen is from Cape Town (= Caput bo-
nae spei).
SPECIMENS EXAMINED
CAPE. — 2918 (Gamoep): Carolusberg, Hester Malan Res., top of
mountain, nr old mine, at seepage area and flat granitic rock outcrop
(-CA), S.M. Perold 1424 p.p., 1425 p.p., 2040 p.p. (PRE). 3017
(Hondeklipbaai): Kamiesberg Pass, dry area above seepage (-BB), S.M.
Perold 1610 p.p. (PRE); Brakdam, 31 km S of Kamieskroon, dirt road,
at rock outcrops (-BD), S.M. Perold 2115 p.p. (PRE); 3018 (Kamiesberg):
22 km NE of Garies, Studer’s Pass, rock outcrop (-AC), S.M. Perold
2124 p.p. (PRE); 5 km E of Garies, on road to Studer’s Pass, rock outcrop
(-CA), S.M. Perold 2U8 p.p., 2722 (PRE). 3119 (Calvinia): 2 km from
Nieuwoudtville, on road from Vanrhynsdorp, above ditch, on soil between
sandstone rocks (-AC), S.M. Perold 1756 p.p. (PRE); Nieuwoudtville,
Farm Paardekraal, most northerly area of fynbos on escarpment (—AC),
C.M. van Wyk 1489 (PRE). 3218 (Clanwilliam): 17 km E of Clanwilliam,
along Pakhuis Pass, Leipoldt’s Grave (— BB), S.M. Perold 1930 (PRE);
Citrusdal, 20 and 21 km N of Hex River Estates, sandstone rock outcrops
above Olifants River (— BD), S.M. Perold 538, 2382, 2383 p.p. (PRE).
3219 (Wuppertal): Biedouw Youth Camp, 19 km along road to Wuppertal,
on soil over sandstone outcrops, nr waterfall (— AA), S.M. Perold 1891
(PRE); 2 km S of Algeria For. Sta. , on sandy soil on sandstone rock
outcrop (—AC), S.M. Perold 235 7 (PRE); 21 km from Algeria For. Sta.
on road to Cedarberg, opp. ruins of house, on sandy soil at base of rocks
(-AC), S.M. Perold 1979 (PRE). ? Promontorio Bonae Spei ? terra
Capensi (exact locality not known) Zeyher s.n. in Herb. G13U7 (G!).
DISCUSSION
The specimen taken to be the type specimen (BM ex
Herb. Hampe) has no annotations except ‘R. albo-
marginata (hyphenated by Bischoff but not by Krauss)
alpha et beta, Cap. b. sp.’ written in Bischoff s hand, as
well as his signature on the label. Three small thalli, or
groups of thalli, are mounted in three clumps on the
herbarium sheet and they represent two different species.
Bischoff had given no indication which specimen he
considered to belong to alpha or to beta and Krauss did
not refer to any varieties in the protologue. The thallus
on the extreme left of the sheet is slightly larger than the
others, white and ‘fluffy’ dorsally (not having been
pressed) and could belong to R. concava Bisch. ex Krauss.
The clump in the middle consists of three small thalli
which are slightly brownish, + scurfy dorsally and have
undulating white margins. I have identified it as R. albo-
marginata and have chosen it as the lectotype. The thallus
on the right of the sheet has been affixed with too much
glue and is unrecognizable.
The description by Krauss and that by Gottsche et al. ,
which was published only a few months later, are almost
identical and are clearly based on the same manuscript
by Bischoff. Gottsche et al. , however, distinguished two
varieties: var. alpha , which I consider to belong to R. albo-
marginata and var. beta maior, described as larger and
having a broader thallus and shorter lobes, which in my
opinion, could also represent R. concava Bisch. ex Krauss.
This var. beta is based on a Zeyher collection from the
Cape, held at G (ex Herb. Bisch.).
Gottsche et al. listed the collections as follows: ‘In
Promontorio Bonae Spei formam alpha leg. Dr Krauss
Julio 1838. Eiusdem formae et var. beta specimina in terra
Capensi lecta retulit cl. Zeyher (Herb. Bisch.)’, thus
acknowledging the presence of two taxa in the Zeyher
collection. The Zeyher collection, however, is also not
annotated to distinguish between var. alpha and var. beta.
The collection comprises about nine thalli or fragments
of thalli.
34
Bothalia 20,1 (1990)
FIGURE 3. — Riccia albomarginata. Spores. A, D, proximal face; B, E, distal face; C, distal face, side view. A, B, S.M. Perold 2382\ C, C.M.
van Wyk 1489\ D, E, S.M. Perold 538. Riccia concava. Spores. F, proximal face; G, proximal face, side view; H, I, distal face. F— I,
Zeyher p.p. A— I, SEM micrographs. Scale bar on A— I = 50 ftm.
The two fertile fragments and four somewhat larger thalli
could probably also be referable to R. concava. These thal-
li are, however, pressed flat, and are less robust than is
usual for this species; they have a somewhat glaucous
colour (as noted in the original description of R. conca-
va), rounded hyaline scales, which are apically more
prominently projecting (± 175 pm) beyond the margin,
and squashed dorsal cells. The remaining two (or three
?) smaller thalli in the same packet of this collection are
no longer green, but brownish, have flattened, sparse, fine
threads dorsally and ± undulating white, membranous
scales above purple-brown flanks. They are here identified
as R. albomarginata var. alpha. The single thallus in the
other packet of the Zeyher collection (and presumably the
one Stephani may have sectioned) is very thin, as it has
been pressed quite flat and is therefore unidentifiable.
The spores from the above Zeyher collection (Figure
3F— I) are closely similar in their ornamentation to those
of R. concava as noted above. They are 75—90 pm in
diameter, triangular-globular, polar, bright brown, semi-
transparent to opaque; wing ± 5 pm wide, margin almost
smooth; ornamentation reticulate: distal face with ± 13
areolae across diameter, ± 5 pm wide, walls thick,
smooth, raised at nodes; proximal face with triradiate mark
distinct, ornamentation similar to that on distal face
(compare with Figure 3, No. 14 in the series ‘Studies in
the genus Riccia from southern Africa’, Perold 1989a and
also Perold 1989b, Fig. 26). Volk (1983) also suspected
FIGURE 4. — Distribution map of R. albomarginata , # ; and R. simii ,
▲ , in southern Africa.
Bothalia 20,1 (1990)
35
that Stephani’s (1898) description and illustration of the
spores from Zeyher’s collection of R. albomarginata in
his previously unpublished leones hepaticarum (G), were
actually of R. concava spores.
In their description of R. concava, which follows
immediately on that of R. albomarginata (but which they
placed in a different section, ‘Subtus Squamatae’ and not
in the Ciliatae, Gottsche et al. noted that R. concava was
similar to R. albomarginata var. beta maior (and to R.
lamellosa ). This apparent similarity between a part of both
collections and R. concava, was also found in the present
investigation as noted above.
Because of the fragmentary nature and generally poor
condition of the type material and the adverse effects of
cleaning and pressing the Zeyher collection, only the habit,
scales and to some extent, the colour of the thalli, could
be checked for each species and were found to be correct.
Other characters, such as the vitally important shape and
size of the cells in the dorsal pillars and the shape of the
thalli in cross section, could not be checked at all,
unfortunately. Marginal cilia are definitely absent.
Later authors, such as Stephani (1898), who incidentally
had only seen Zeyher’s collection, again ignored the fact
that the collections were mixed, and treated them as a
single species with prominent scales, disregarding the
reference to small membranes, proximally not divided into
separate lamellae (fide Gottsche et al.). Sim (1926) applied
the name to a different taxon (see below), Amell (1957,
1963) to yet another, and Volk (1981, 1983) followed Sim;
the specimens from lie de l’Est (Crozet Archipelago),
assigned to R. albomarginata Bisch. ex G.L. & N. emend.
Sim by Jovet-Ast (1986) have not been seen by me, but
it is my opinion that their identity should be reconsidered.
CLASSIFICATION OF R. ALBOMARGINATA
R. albomarginata Bisch. belongs to section Pilifer Volk,
characterized by free-standing, multicellular dorsal
epithelial pillars and only lately recognized as a distinct
group by Volk (1983) and also by Schuster (1984, 1985)
as his subgenus Pteroriccia. Gottsche et al. (1846) had
incorrectly classified R. albomarginata under the Ciliatae,
which have unicellular marginal hairs, although R.
concava Bisch., which they regarded as similar to R.
albomarginata (var. beta, maior), was classified under a
different group, ‘Subtus Squamatae’. However, it appears
from BischofPs observations when referring to R. concava
(quoted by G.L. & N.), that the ‘small scales’ in the dry
plants could be taken for cilia!.
Stephani (1898) placed R. albomarginata in his Inermes
(without cilia) and also failed to report on the dorsal
epithelial cell pillars. Admittedly, in the pressed, dried
plants of the original material, they are very difficult to
recognize.
In his key to the Riccia species, Amell (1963) grouped
R. albomarginata and R. concava together with a ciliated
species, R. natalensis, partly following Sim (1926) who
had classified R. albomarginata, R. natalensis and his un-
traced species, R. coronata, together under the heading
‘epidermal cells, or some of them, elongated or mammil-
late and free’. It is therefore obvious that earlier authors did
not have a clear understanding of the difference between
unicellular marginal cilia and multicellular hairs covering
the entire dorsal face of the thallus.
MISAPPLICATION OF R. ALBOMARGINATA
From his own and Potts’s collections, Sim was familiar
with a Riccia species with large white scales to which he
applied the specific name ‘albomarginata’. This species is
further characterized by ‘upper pillars quite free from one
another. . .’ which he (Sim 1926) proceeded to describe and
illustrate, but without citing specimens. Sim could of
course examine fresh material (with the dorsal pillars intact
and not collapsed, as they are in long-dried specimens)
and this admittedly gave him an advantage denied the
earlier workers, who in any case, were totally unfamiliar
with such cell pillars.
Whether the specimens of R. albomarginata Bisch. cited
by Krauss and by G.L. & N. were less fragmentary and
in a better state of preservation in Sim’s time, is not known.
However, Sim evidently noticed signs of loose dorsal
pillars in the type specimens [although he overlooked them
in R. concava (Perold 1989)] and assumed that his and
Potts’s collections, presumably the only ones to his
knowledge with similar pillars, belonged to R. albo-
marginata.
Volk (1983) finds it ‘erstaunlich, dass Bischoff die Haare
des Epithels nicht erwahnt’, but I venture to state, that,
if Bischoff had indeed been dealing with R. albomarginata
auct. non Bisch., he, Krauss and Gottsche et al. could
not have failed to notice the thick velvety dorsal covering
of the thallus, which is still very evident in Sim 338, a
hundred years after collection and also after experimental
pressing by me.
Earlier, Stephani had remarked on the ‘ganz diinnes
Laub’ of R. albomarginata Bisch., which could have
referred to either var. alpha or var. beta, but certainly not
to the plants described by Sim as 2 mm thick in cross
ection. Even if subjected to prolonged pressing, such thick
lobes could not have been flattened to the almost paper-
like thinness of some of the type material.
Finally, the scales of R. albomarginata auct. non Bisch.
are very prominent, wavy, white, and apparently with some
striations on the cell surfaces (Figure 6F), unlike the
smaller (± 600 X 325-450 pm), clear, hyaline or white
membranous scales (Figure 2F) of the type. Sim may also
have been misled by Stephani’s comments that the scales
of Zeyher’s plants were prominent, extending much above
the thallus margins (but only up to ± 150 and 175 pm in
the two varieties alpha and beta respectively, as measured
by me), and that the species had been named for this reason
(see under R. albomarginata Bisch.). Sim did not describe
spores for his species.
The concluding remark in Sim’s (1926) description of
R. albomarginata (p. 10) viz.: 7?. villosa Steph.
(Brunnthaller 1913, p.p. 1—14), from Matjiesfontein, C.P.
(an arid locality) answers the description exactly’, raises
doubts whether he had thoroughly examined the type
specimens of either species, as R. albomarginata has
rounded scales and R. villosa has triangular scales.
36
Bothalia 20,1 (1990)
Amell (1963) gave only a very brief description of R.
albomarginata auct. non Bisch. The spore illustrations and
description (Amell 1963) were based on Volk 12433 p.p.
and 12462 p.p. (Amell 1957), collected in SWA/Namibia,
and reassigned to R. albovestita Volk (Volk 1981).
Regardless of what influenced Sim, it is now indis-
putable that he misapplied the specific name albo-
marginata to a species which is here named and described
as R. simii, sp. nov. The specific epithet, simii, has been
chosen in deference to Sim’s work, as he was the first to
describe and illustrate free-standing dorsal epithelial cell
pillars in a Riccia species.
Riccia simii Perold, sp. nov.
R. albomarginata auct. non Bisch.: Sim: 9 (1926); Volk: 453 (1983).
Thallus monoicus, perennis, magnitudine medius vel
magnus, in vivo smaragdinus, velutinus, squamis magnis,
hyalinis, marginis in sicco squamis supra paginam
dorsalem granularem inflexis. Frons usque ad 12 mm
longa, 1,8— 2,5 mm lata, 0,9— 1,3(— 1,5) mm crassa, in
sectione transversali duplo latiora quam crassa,
symmetrice vel asymmetrice furcata, oblonga vel obovata,
apice acuta, marginibus subacutis, apicem versus dorsaliter
sulcata, aliter plana, ad latera verticalis vel obliquiter
proximaliter devexa. Anatomia : epithelium dorsale 350 /zm
crassum, ex columnis liberis deminuentibus, 4-5-cellu-
laribus fragilibus compositum; aerenchyma 350 /zm
crassa, textura penaria 400 /zm. Squamae magnae, 1500
x 600—900 /zm, rotundatae, hyalinae, crebre imbricatae,
plus quam 250 /zm supra margines thalli eminentes. Sporae
(70-)82-105(-120) /zm diametro, deltoideo-globulares,
ala ± 5 /zm lata; omamento variabile, perfecte vel
imperfecte reticulato; pagina distali seriebus areolarum
omnibus vel solum exterioribus completis, 5-10 /zm latis,
parietibus irregulariter ramificantibus vel in cristis
radiantibus; pagina proximali nota triradiata distincta, in
quoque superficie 25—30 areolis parvis.
TYPE. — Cape, 3227 (Stutterheim): Perie Mission
Station, Kafffaria (-CC). T.R. Sim 338 ( PRE-CH 1035)
(PRE, holo.) alt. 2500 ft., 1888.
Thallus monoicous, perennial, in crowded gregarious
patches or scattered, bright green to emerald green,
velvety, with large hyaline scales (Figure 6D) extending
above and beyond thallus margins (Figures 5A; 6A);
medium-sized to large, branches simple or one to twice
symmetrically or asymmetrically furcate, medium to
widely divergent, up to 12,0 mm long segments 4,0— 5,0
mm long, 1,8— 2,5 mm wide, 0,9 — 1,3( — 1,5) mm thick,
i.e. ± twice wider than thick in section (Figure 5E); ob-
long to obovate (Figure 6B), apex acute (Figure 6C), dor-
sal face distally deeply grooved, soon becoming flat;
margins subacute, flanks steep to proximally sloping ob-
liquely upward and outward, green, sometimes flecked
with violet, covered with large scales; ventral face gently
rounded to flat, green; when dry (Figure 5B), margins
tightly inflexed, apically meeting along midline over white,
finely granular dorsal surface, scales white, imbricate,
wavy.
FIGURE 5. — Riccia simii. Morphology and anatomy. A, thallus wet;
B, thallus dry; C, transverse section through dorsal cell pillars;
D, horizontal section through basal cells of cell pillars with air
pores hatched, and through assimilation tissue with air canals
stippled; E, transverse section through thallus; F, scale. A, E,
S.M. Perold 1318, B, S.M. Perold 1346: C, S.M. Perold 505 ; D,
Smook 6631: F, C.M. van Wyk 1781. Drawings by J. Kimpton.
Scale bar on A, B, E = 1 mm; C, D = 50 /t m; F = 100 /zm.
Anatomy of thallus: dorsal epithelium (Figures 5C; 6E)
consisting of free-standing, 4-5-celled, gradually taper-
ing pillars, fragile, but basally somewhat thicker-walled,
Bothalia 20,1 (1990)
37
FIGURE 6. — Riccia simii. Morphology and anatomy. A, thalli in cultivation; B, branch from field-grown thallus; C, apex of same; D, scales
near apex; E, dorsal cell pillars; F, part of scale showing striations. A, J. Thompson 257; B— E, S.M. Perold 505; F, S.M. Perold 1348.
A, by A. Romanowski; B— E, SEM micrographs; F, LM micrograph. Scale bar on A = 1 mm; B— F = 50 jun.
hyaline, up to ± 350 pm long, apical cells 25—50 x
18—25 pm, intermediate cells 45— 75(— 80) x 25—35 pm,
basal cells 62—80 x 30—38 pm, mostly equally long, with
upper transverse walls in an interrupted horizontal line
across; from above, pillars forming a thick pelt, mostly
collapsed proximally, air pores small, 4— 6(— 8)-sided
(Figure 5D), obscured by cell pillars; assimilation tissue
± 350 pm thick in section, 1;3 the thickness of thallus
and consisting of vertical columns of up to 8 cells, ± 40
x 25 pm, enclosing narrow 4—5-sided air canals; storage
tissue ± 400 pm thick, 1;3 the thickness of thallus, cells
rounded to angular, closely packed, ± 50 ^im wide;
rhizoids arising from ventral epidermal cells, some
smooth, others tuberculate, 22 ^m wide. Scales (Figures
5F; 6F) nearly semi-circular, large, closely imbricate,
projecting more than 250 pm above thallus margins,
hyaline, up to 1500 x 600-900 /un, margins smooth, cells
often with somewhat striate surface, in body of scale
long-hexagonal or rectangular, 80 — 110( — 125) x
30— 35(— 40) pm, at margins ± 2 rows of smaller
brick-shaped cells. Antheridia with hyaline necks,
extending nearly 500 pm above dorsal surface. Archegonia
with purple necks, scattered along median part of thallus.
Sporangia mostly single, rarely up to 3 crowded together
in narrow basal part of thallus, dorsally bulging, each with
± 370-400 spores, but thalli only very occasionally
sporulating. Spores (70— )82-105(-120) ^m in diameter,
triangular-globular, polar, yellow or light brown, colour
deepening to mahogany brown or turning black on ageing,
semitransparent to opaque, with wing ± 5 pm wide,
margin finely crenulate, angles perforated; ornamentation
variable, completely or incompletely reticulate, similar or
dissimilar on two spore faces: distal face (Figure 7B, D,
F) with all, or only outer rows of areolae complete,
variable in size, 5-10 pm wide, irregularly shaped,
rounded or elongated, walls raised at nodes, sometimes
anastomosing to form ridges, irregularly branching and
twisting or radiating outwards from centre; proximal face
(Figure 7A, C, E) with triradiate mark clearly defined,
sometimes papillate, 25—30 complete or incomplete small
areolae, up to 5 /*m wide, on each of 3 facets, walls thin,
raised at nodes, sometimes sprinkled with papillae towards
wing.
R. simii can be distinguished from other species in
section Pilifer, subgenus Riccia, by the rounded, very
large, wavy, closely imbricate, hyaline scales, projecting
much beyond the thallus margins and by the velvety dorsal
surface of the thallus, with the tapering cell pillars apically
‘finer’ than those in other species, except for R. villosa,
which has triangular scales and is strictly confined to
the north-western, south-western and southern Cape. R.
simii is fairly common in the Orange Free State and
is also known from the south-western, southern, central
and eastern Cape and from Transkei (Figure 4). Sim (1926)
reported this species, R. simii (= R. albomarginata auct.
non Bisch.), from Transvaal and Natal, but this has
not been verified. Plants that he collected from Southern
Rhodesia (Zimbabwe), Sim 9068—9070, 9072 (PRE),
and identified as R. albomarginata Bisch. ex G.L. &
N. (Sim 1931), are in fact R. moenkemeyeri Steph.
Curiously, Sim had identified his collection, Sim 338
(PRE-CH1035), here selected as the holotype of R. simii,
as R. limbata, which has black scales. It was subsequently
identified as R. albomarginata, presumably by Duthie or
Amell.
R. simii is found on shallow soil overlying rock outcrops,
which can be sandstone, dolerite or even limestone. It
grows in association with other Riccia species, and
occasionally with Mannia capensis (Steph.) S. Amell, and
with small succulents, in both summer and winter rainfall
areas.
38
Bothalia 20,1 (1990)
Volk (1983) chose this species, under the name R. albo-
marginata, as the type species of his section Pilifer and
provided a detailed description.
SPECIMENS EXAMINED
O.F.S. — 2727 (Kroonstad): grassland, on Farm Caroline, 8 km SE of
Steynsrus (—DC), Smook 6631 (PRE). 2728 (Frankfort): Wonder-
fonteinspruit, 40 km from Bethlehem on road to Lindley (— CC), S.M.
Perold 1366 (PRE). 2729 (Volksrust): Farm Witkoppe, SE of Vrede
(— CB), Smook 6418 (PRE). 2826 (Brandfort): Brandfort (— CB), Duthie
5330 (BOL). 2827 (Senekal): Allemanskraal, 5 km from Ventersburg
(—AC), on dolerite, Volk 81/226, 1984/730 (M, PRE); Senekal, on koppie
behind town (— BC), S.M. Perold 1337, 1341 (PRE); Paul Roux, on flat
rocks on koppie behind town (— BD), S.M. Perold 1346 (PRE); 6 km
N of Clocolan, on road to Marquard, nr bridge, on flat weathered
sandstone rocks (-DC), S.M. Perold 1323 (PRE); on flat rocks on Farm
Holstein, on road to Ficksburg, 22 km E of Clocolan (—DC), S.M. Perold
1318 (PRE). 2828 (Bethlehem): on koppie at HF tower (-AB), S.M.
Perold 1360, 1363 (PRE); 14 km E of Paul Roux, on road to Bethlehem,
on weathered sandstone (—AC), S.M. Perold 1353 (PRE); 11 km E of
Fouriesburg, on slope next to road to Clarens (— CB), S.M. Perold 1304,
1306 (PRE); on rocky oucrop nr Golden Gate Highland Park, between
Clarens and Kestell (-DA), J.M. Perold 29 (PRE). 2926 (Bloemfon-
tein): Bloemfontein, Eagle’s Nest (-AA), Duthie 5456, 5461 (BOL);
Geo. Potts PRE-CH 1101 (PRE); Rayton Caravan Park, Volk 81/051,
811289a (M, PRE); Uitkykhoogte, along road from Reddersburg to van
Stadensrus via Helvetia (—CD), Van Rooy 2337, 2338 (PRE). 2927
(Maseru): 10 and 12 km S of Ladybrand, on shallow soil on road R26
(-AB), J.M. Perold 36, 39, 43, 44 p.p. (PRE). 3026 (Aliwal North):
NE of Zastron, edge of small pan in grassland on Farm Elandsberg
(— BB), Van Rooy 2403 (PRE); between Zastron and Wesselsdale, Farm
Olievenrand (-BB), Van Rooy 2416 (PRE). 3027 (Lady Grey): Zastron,
on soil under sandstone overhang on E slopes of Aasvoelberg (-AC),
Van Rooy 2515 (PRE).
TRANSKEI.— 3129 (Port St. Johns): Holy Cross Mission, 1,6 km
along road to Flagstaff, on soil over flat sandstone (-BA), Van Rooy
1817, 1818, 1823 (PRE); sandstone outcrops in grassland, 22 km from
Holy Cross Mission on road to Mkambati (— BB), Van Rooy 1708a (PRE).
CAPE. — 3025 (Colesberg): Colesberg Dist., on koppie between rocks
and shrubs (— CA), Hitchcock 5478 (BOL). 3027 (Lady Grey): on soil
over sandstone between Farms Rietfontein and De Kraal, 23 km S of
Lady Grey (— CC), Van Rooy 2602 (PRE); on soil over sandstone, Farm
Femgrove, on road to Jamestown, 14 km from Lady Grey (— CC), Van
Rooy 2685 (PRE); 17 km from Barkly East to Rhodes, on cave sand-
stone, at Rebelhoogte, near Farm La Colleen (-DC), Van Rooy 2766
(PRE). 3124 (Hanover): on soil at edge of dry stream on road to
Middelburg, 1 km S of Lootsberg Pass (-DB), S.M. Perold 949 (PRE).
3127 (Lady Frere): on shallow soil nr seepage nr Farm Clifford between
Barkly East and Rossouw (— AB), Thompson 257 (PRE). 3218
(Clanwilliam): on flat rock outcrop, above Platkloof River, Farm
Middelpos, on road to Goedverwag NW of Piketbeig (-DC), S. M. Perold
505 (PRE). 3224 (Graaff Reinet): Valley of Desolation, at lookout, on
flat rock outcrops (— BC), Smook 3908 (PRE). 3225 (Somerset East):
Cradock Mountain Zebra Park (—AD), Liebenberg 7632 (PRE). 3227
(Stutterheim): Perie Mission Sta. (-CC), Sim 338 ( PRE-CH 1035)
(holotype) (PRE). 3322 (Oudtshoom): 15 km from De Rust, on road
to Uniondale (— BC), Koekemoer 291 (PRE); 5 km from Dysselsdorp
on road to Kammanassie Dam (-CB), Koekemoer 292 (PRE). 3323
(Willowmore): Uniondale, Vetvlei (— CA), Duthie 22 (PRE-CH 1016)
(PRE); Langkloof Mts. (-DC), Duthie 5115 (BOL). 3420 (Bredasdorp):
De Hoop area, in pass on road from Wydgelee to De Hoop, between
fynbos on slope (—AD), Fellingham 746 (PRE); NW of Cape Infanta,
Potteberg Estates, Farm Witwater, on S facing limestone slopes with
fynbos (— BD), Oliver 8490 (PRE); De Hoop, Buffelsfontein, on moist
sandy patches between fynbos (-CB), C.M. van Wyk 1781 (PRE).
CORRECTION
Unbeknown to me and therefore previously omitted
(Perold 1989; Perold & Volk 1988), Krauss had also
published brief but validly published descriptions of Riccia
concava Bisch. and R. limbata Bisch. in Flora 29: 135,
136 in March 1846, thereby antedating the publication of
the description of these two species (and R. albomarginata)
by Gottsche et al. in Synopsis hepaticarum Oct. 1846, by
seven months (fide Grolle pers. comm.).
ACKNOWLEDGEMENTS
Sincere thanks are due to the curators of BM, G and
BOL herbaria for the loan of the type and other specimens.
I gratefully acknowledge information from Dr R. Grolle,
FIGURE 7. — Riccia simii. Spores. A, C, E, proximal face; B, D, F, distal face. A, B, J.M. Perold 39; C, D, Volk 81/289a; E, F, Sim 338. Scale
bar on A-F = 50 /jm. All micrographs by S.M. Perold.
Bothalia 20,1 (1990)
39
Friedrich Schiller University, Jena, concerning the Krauss
publication and the dates of the publication by Krauss and
by Gottsche et al., as well as the typification. I am also
grateful for discussions with, and advice from Messrs J.
van Rooy and F. Brusse as well as Dr H.F. Glen, who also
translated the Latin texts and suggested the specific epithet,
simii. Photographs were prepared by Mrs A. Romanowski
and the line drawings are by Ms J. Kimpton, for which
they are both sincerely thanked. Dr E.W. Jones, Oxford,
Dr E.O. Campbell, Massey University, New Zealand and
Prof. O.H. Volk, Wurzburg University, commented on the
manuscript and I owe them a large debt of gratitude. Prof.
Volk was also instrumental in obtaining the information
referred to above, from Dr Grolle.
REFERENCES
ARNELL, S. 1957. Hepaticae collected in South West Africa by Prof.
Dr O.H. Volk, Mitteilungen aus der Botanischen Staatssammlung,
Mtinchen 16: 262-272.
ARNELL, S. 1963. Hepaticae of South Africa , pp. 411. Swedish Natural
Science Council, Stockholm.
GOTTSCHE, C.M., LINDENBERG, J.B.G. & NEES ab ESENBECK,
C.G. 1844-1847. Synopsis hepaticarum, pp. 835. Hamburg.
Reprinted 1967.
GUNN, M. & CODD, L.E. 1981. Botanical exploration of southern
Africa , pp. 400. Balkema, Cape Town.
JOVET-AST, S. 1986. Riccia de 1’ Archipel des Crozet (lie de l’Est).
Cryptogamie, Bryologie et Lichenologie 7: 479—485.
KRAUSS, F. 1846. Pflanzen des Cap- und Natal-Landes, gesammelt und
zusammengestellt von Dr Ferdinand Krauss. Flora 29: 135, 136.
PEROLD, S.M. 1986. Pteroriccia Schust., should it be upheld?
(Ricciaceae). Bothalia 16: 63—64.
PEROLD, S.M. 1989a. Studies in the genus Riccia (Marchantiales) from
southern Africa. 14. R. concava section Pilifer. Bothalia 19:
161-165.
PEROLD, S.M. 1989b. Spore-wall ornamentation as an aid in identifying
the southern African species of Riccia (Hepaticae). Journal of
the Hattori Botanical Laboratory 67: 109—201.
PEROLD, S.M. In press. Studies in the genus Riccia (Marchantiales)
from southern Africa. 17. Three new species in section Pilifer:
R. elongata, R. ampullacea and R. trachyglossum. Bothalia.
PEROLD, S.M. & VOLK, O.H. 1988. Studies in the genus Riccia
(Marchantiales) from southern Africa. 9. R nigrella and the status
of R. capensis. Bothalia 18: 43—49.
SCHUSTER, R.M. 1984. Diagnoses of some new taxa of Hepaticae.
Phytologia 56: 65 —74.
SCHUSTER, R.M. 1985. Some new taxa of Hepaticae. Phytologia 57:
408-414.
SIM, T.R. 1926. The bryophyta of South Africa. Transactions of the Royal
Society of South Africa 15: 9, 10.
SIM, T.R. 1931. South African Bryophytes. Further notes. Transactions
of the Royal Society of South Africa 20: 15—17.
STEPHANI, F. 1898. Species hepaticarum. Bulletin de l ’ Herbier Boissier
6: 309-411.
STEPHANI, F. 1876—1907. leones hepaticarum. Ed. P. Geissler. IDS
Microform Publishers, Zug, Switzerland, publ. 1986.
VOLK. O.H. 1981. Beitrage zur Kenntnis der Lebermoose (Hepaticae)
aus Siidwestafrika (Namibia). II. Mitteilungen der Botanischen
Staatssammlung, Mtinchen 17: 245—252.
VOLK, O.H. 1983. Vorschlag fur eine Neugliederung der Gattung Riccia
L. Mitteilungen der Botanischen Staatssammlung, Mtinchen 19:
453-465.
Bothalia 20,1: 41-48 (1990)
New species of Erica (Ericaceae) from the Cape Province
E.G.H. OLIVER*
Keywords: Cape Province, Erica, new species, taxonomy
ABSTRACT
Three new species of Erica L. (Ericaceae) are described from the southern parts of the Cape Province. E. amicorum
E.G.H. Oliver, an endemic marsh species from the Langeberg near Riversdale, is one of the most distinctive species in
the genus on account of its almost free reflexed petals, reflexed stamens and exserted gynoecium. The mat-forming E.
tegetiformis E.G.H. Oliver is allied to E. senilis Klotzsch ex Benth. and E. eriophoros Guth. & Bol., all occurring at high
altitudes in the Cold Bokkeveld and Cedarberg. E. cunoniensis E.G.H. Oliver is allied to the E. squarrosa Salisb./E gysbertii
Guth. & Bol. complex and is very localized in the mountains of the western part of the Caledon Division near Rooi Els.
UITTREKSEL
Drie nuwe spesies van Erica L. (Ericaceae) van die suidelike dele van die Kaap Provinsie word beskryf. E. amicorum
E.G.H. Oliver, ’n endemiese moeras-spesies van die Langeberg naby Riversdal, is een van die merkwaardigste spesies in
die genus omdat dit byna vry teruggebuigde kroonblare, teruggebuigde meeldrade en ’n ontblote ginesium het. E. tegetifor-
mis E.G.H. Oliver, wat groot matte vorm, is naverwant aan E. senilis Klotzsch ex Benth. en E. eriophoros Guth. & Bol.,
en al drie kom voor op hoe plekke in die Koue Bokkeveld en Sederberge. E. cunoniensis E.G.H. Oliver is naverwant aan
die E. squarrosa Salisb./£. gysbertii Guth. & Bol. kompleks en het ’n beperkte verspreiding in die berge van die westelike
gedeelte van die Caledon distrik naby Rooi Els.
INTRODUCTION
The genus Erica in southern Africa has recently been
increased in size with the inclusion of the 15 species of
Philippia Klotzsch (Oliver 1987, 1988) and now contains
some 654 species. As such it is by far the largest genus
in the flora of southern Africa, indeed the continent of
Africa. The considerable increase in collections of plants
from the region of the Cape Flora during the last three
to four decades has produced a wealth of new and
interesting records. These include a number of distinct
undescribed species of which three are described here.
Erica amicorum E.G.H. Oliver, sp. nov., in genere
distinctissima et singularis propter lobos corollae
subliberos reflexos stamina erecta deinde reflexa et
gynoecium exsertum, sed aspectu typico specierum
sectionis Ceramiae.
Fruticulus prostratus repens. Rami ad 1,5 m longi glabri
intemodis longis cortice irregulari grisei. Folia 3-nata,
5-6 x 4-4,5 mm patentia ovata ad subcircularia apice
attenuato marginibus leviter invaginatis ad subnaviculata,
ciliata pilis parvis et pilis crassis glandulosis adaxiale
puberula abaxiale pubescentia pilis crispis, petiolo 0,8— 1,0
mm longo puberulo. Flores 1—2 in extremis brachy-
blastorum lateralium absolutorum vel 1—3 in extremis
ramulorum; pedicellum 3-5 mm longum ad basim
crassum glabrum vel juniores interdum puberulum;
bractea 0,5 -1,0 mm longa basalis vel submediana anguste
ovata vel elliptico-oblonga; bracteolae 2 oppositae subba-
sales ad submedianae 0,4 -0,7 mm longae. Calyx profunde
4-lobatus subliber, lobis 1,7 x 1,5 mm late ovatis initio
erectis demum reflexis ciliatis rosellis atrorubescentibus.
Corolla profunde 4-lobata, tubo 0,8 mm longa latissime
* Stellenbosch Herbarium, National Botanical Institute, RO. Box 471,
Stellenbosch 7600.
MS. received: 1989.04.03.
cyathiformis, lobis 2,9 x 2,2 mm, late naviculato-
ellipticis, patentibus demum reflexis, puberulis ciliatis pilis
parvis et 5—6 glandulis, rosellis atrorubescentibus.
Stamina 8 exserta initio erecta demum reflexa; filamenta
2 mm longa subteretia in annulo base conjuncta, glabra
apice dentata; antherae erectae dorsales, thecis 1,2 mm
longis obtusis parum prognathis scabridulosis muticis,
poro theca 3-plo breviore; pollen in tetradis. Ovarium
manifestum demum omnino exsertum, depresso-globosum
echinoideum pilis brevibus longis glandulosis tectis, disco
base; stylus exsertus filiformis 2,8-3,2 mm longus; stigma
simplex. Capsula globosa glanduloso-setosa, seminibus
ellipsoideis ad subsphaericis reticulatis. Figura 1.
TYPE. — Cape, Riversdale District, the Langeberg,
south slopes at head of Valsrivier, 609 m, 12 December
1987, Oliver, Schumann & Kirsten 9054 (STE, holo.; BM,
BOL, E, G, K, MO, NBG, NY, P, PRE, S).
Prostrate delicate creeping shrublet. Branches up to 1,5
m long spreading through restioid vegetation, glabrous,
brown, with long internodes, side branches often curved,
bark irregularly sloughing off in grey portions. Leaves
3-nate, 5-6 x 4-4,5 mm, patent, ovate to subcircular
with an attenuate apex and slightly inrolled margins to sub-
naviculate, ciliate with very short hairs and long stout
hairs, sparsely puberulous adaxially when young,
sometimes glandular-ciliate along the pseudomargin
adaxially, crisped pubescent abaxially and with a few stout
gland-tipped hairs along the midrib; petiole 0,8— 1,0 mm
long, puberulous when young, ciliate. Flowers 1— 2-nate
at ends of lateral absolute brachyblasts or 1- 3-nate at ends
of branchlets, sometimes mixed in a synflorescence;
pedicel 3-5 mm long tapering to the base, glabrous or
sparsely glandular villous, sometimes puberulous when
young, dark red; bract 0,5— 1,0 mm long, basal, occasion-
ally submedian, narrowly attenuate-ovate or elliptic-
oblong, with or without a sulcate tip; bracteoles 2,
opposite, subbasal to submedian, 0,4— 0,7 mm long elliptic-
42
Bothalia 20,1 (1990)
FIGURE 1. — Erica amicorum : A, flowering branch, x 0,5; B, leaf, abaxial view, x 6; C, two florescences, x 1; D, bud, x 6; E, flower
at anthesis, x 12,5; F, older flower with reflexed stamens, x 6; G, bract and bracteole, x 12,5; H, sepal, x 12,5; I, petal, x 12,5;
J, anther, front, side & back views, X 12,5; K, ovary, x 12,5; L, base of mature ovary and stamens, x 12,5; all drawn from the type
collection, Oliver, Kirsten & Schumann 9054 (STE).
oblong, acute, ciliate and with sessile glands. Calyx deeply
4-lobed, lobes almost free, 1,7 x 1,5 mm broadly ovate with
a sulcate apex, at first erect becoming reflexed, ciliate with
fine hairs and short stout glands, pink becoming dark red.
Corolla deeply 4-lobed, tube 0,8 mm long, very broadly
cyathiform, lobes 2,9 x 2,2 mm broadly naviculate-
elliptic, obtuse to truncate, spreading, becoming reflexed,
finely puberulous and finely ciliate and with 5—6 stout
short gland-tipped cilia, pink becoming dark red. Stamens
8, exserted, at first erect, becoming reflexed into the
reflexed corolla lobes, the alternate stamens reflexed over
the interstices; filaments 2 mm long, rather thick, subterete,
joined into a thick ring at the base, glabrous with a toothed
apex; anthers erect, dark brown dorsally attached near the
base, thecae 1,2 mm long, obtuse, slightly prognathous
or with downwardly pointing sharp base, scabridulous,
pore ± !/3 length of the theca, pollen in tetrads. Ovary
manifest becoming fully exserted, flattened globose, echi-
noid, covered with short to long glandular hairs with a
slight disk below; style exserted, filiform, 2,8— 3,2 mm
long; stigma simple. Capsule globose, glandular setose;
seeds ellipsoid to subspherical, reticulate. Figure 1.
This remarkable new species is unlike any other species
in the genus on account of having, in the mature flowers.
almost free corolla lobes which are spreading and then
reflexed, with the alternate stamens reflexed between these
lobes, and an exserted gynoecium. However, the species
appears to be allied to species in the section Ceramia
because of its habit, habitat preference and small pink flow-
ers and open-backed leaves. In particular it superficially
resembles species such as E. oligantha and E. planifolia
L., but is different in a number of unique characters.
The possession of almost free corolla lobes is very rare
in the subfamily Ericoideae. It occurs in a few
schizopetalous cultivars of European heathers. In southern
Africa only four species share such a divided corolla, i.e.
Erica lanuginosa Andr., E. occulta E.G.H. Oliver and E.
bodkinii Guth. & Bol., but not to the same extent as in
E. amicorum. Also they do not have the lobes reflexed
in the mature flowers. This condition renders the flowers
unique in that the complete androecium and gynoecium
are exserted, thus giving the impression of an erica flower
in which the corolla has been eaten away by some insect.
The mature flower is also unique in having those stamens
alternating with the corolla lobes totally reflexed between
the lobes. The stamens are also rather unusual in being
joined together at the base into a thickened collar. Fused
Bothalia 20,1 (1990)
43
or partially fused stamens are now no longer rare in the
genus Erica because of the recent inclusion of the genus
Philippia into Erica (Oliver 1987, 1988).
Another striking feature of E. amicorum is the arrange-
ment of the flowers which can be borne in a variety of
ways on the branches from simple 1— 3-nate terminal
florescences to 1-flowered florescences on lateral absolute
brachyblasts (Figure 2). These may then all be aggregated
together into a loose synflorescence on a lateral branch
(Figure 3). When the main branches are long, up to 1 m,
they bear numerous lateral branches which themselves
bear various combinations of florescences. In some cases
the simple 3 -flowered terminal florescence can look very
similar to a terminal synflorescence. Examination of a
florescence will show that flowers alternating with the
leaves belong to a simple florescence (Figure 2A) whereas
those opposite the leaves are terminal on absolute
brachyblasts (Figure 2B).
The species appears to be very restricted in its
distribution, occurring only on one mountain complex in
the Langeberg range north-east of Riversdale (Figure 4).
It is confined to low thick vegetation in moist places along-
side streams or in seepage zones. In the type locality it
produces matted plants intertwined among other moisture-
loving fynbos plants especially tussocks of the restiad,
Platycaulos acutus Esterhuysen, and therefore the
extricating of reasonable flowering branches for herbarium
purposes was time-consuming work. One branch
disentangled from several restiad clumps was 1,5 m long.
This species is named after my two friends, Dolf
Schumann and Gerhard Kirsten, who introduced me to
it and with whom the type collection was made.
CAPE. — 3321(Ladismith): Romanshoek, 600 m, 8. xii.86 (—CD),
Kirsten 1064 (NBG); 1067 (STE); ibid. Schumann 492 (PRE; STE); ibid.
Schumann 596 (STE); Langeberg, south slopes, headwaters of Valsrivier,
609 m, 12.xii.1987 (—CD), Oliver , Kirsten & Schumann 9054 (STE &
others as per type); Perdeberg Plateau, 975 m, 6.ix.83 (—CD), Viviers
1100 (STE). 3421 (Riversdale): Gavelsbos, 307 m, 2.ix.87 (-AB), De
Kock sub Bohnen 8811 (STE).
Erica cunoniensis E.G.H. Oliver, sp. nov. , in Sectione
Euryloma E. squarrosae Salisb. et E. gysbertii Guth. &
Bol. affinis sed ab eis floribus patentioribus, id est corolla
tubulocampanulata ad anguste cyathiformi non ovoideo-
urceolata ad tubuloconica, antheris adhaerentibus propter
pilos intertextos laterales non liberis glabris differt.
Fruticulus sparsus erectus ad 300 mm altus. Rami erecti
ad extremis foliosi, glabri sterigmatis infrafoliaceis, rami
veteres irregulariter porcati cicatribus foliorum notati.
Folia 4-nata arete disposita patentia ad recurva, 2,5— 3,5
mm longa, ciliata pilis longis validis ad apice caespitosis
seta unica longa atrorubenti terminali; petiolus 1 mm
longus appressus complanatus ciliatus. Flores 3-9
verticillati vel secundi ad extremis ramorum; pedicellus
3,5 -5,0 mm longus rubens pilis crispis glandulis subses-
silibus parvis; bractea 3 x 0,5 mm linearis ad lineari-
spathulata, bracteolae bracteis similes sed graciliores.
Calyx 4-lobatus, 5 mm longus, lobis longitudine 3/4 calycis
partes aequantibus, dimidio inferiore ovato ciliato glandulis
sessilibus dimidio superiore acuto elongato foliato sulcato
ciliato pilis longis glandulis sessilibus in parte interiore
basi. Corolla 6—9 x 3—4 mm variabilis tubulocampanu-
lata ad anguste cyathiformis viscida atrosanguinea; lobis
deltoideis longitudine ^6 corollae partes aequantibus,
erectis ad patentibus ad reflexis. Stamina inclusa; filamenta
5 mm longa apice dilatata et ciliata; anthera adhaerentes
propter pilos intertextos, dorsales, thecis anguste ovatis
l, 2 mm longis pilis crispis. Ovarium 1,6— 2,0 x 1,0— 1,2
mm obovoideum plus minusve emarginatum stipite
minimo; stylus 6—7 mm longus anguste cylindraceus
glaber; stigma plus minusve exserto subcapitato. Fructus
anguste obovoideus capsularis stipite parvo; semina sub-
sphaerica papillata. Figura 5.
TYPE. — Cape, Caledon Division, Rooi Els area,
Buffelstalberg, ridge running NW from the beacon, 670
m, 5 October 1986, Oliver 8974 (STE, holo.; BM, BOL,
E, G, K, MO, NBG, P, PRE, S, W).
Sparse erect woody shrublet up to 300 mm, rarely 500
mm, tall. Branches erect, leafy only towards the ends, gla-
brous with distinct infrafoliar ridges, bark in older
FIGURE 2. — Tw<o similar-looking inflorescences commonly found in E. amicorum'. A, a simple 3-flowered florescence condensed from the schematic
diagram Al; B, a compound synflorescence consisting of three 1-flowered florescences (in each florescence there are two aborted flowers)
condensed from the schematic diagram in Bl. Zigzag lines represent expanded absolute brachyblasts.
44
Bothalia 20,1 (1990)
FIGURE 3. — Synflorescence of five florescences on a lateral branch in E. amicorum : A, the branch as it appears on the plant; B, an enlarged
schematic explanation of the branch and five florescences, florescences 1—4 are equal to Figure 2 A and florescence 5 to Figure 2B, the
apparently single terminal florescence is a synflorescence made up of four flowers from two florescences. Zigzag lines represent expanded
absolute brachyblasts.
branches splitting into white infrafoliar flakes, oldest
branches red-brown, irregularly ridged with prominent
protruding leaf scars. Leaves 4-nate, closely arranged,
spreading-recurved, 2,5— 3,5 mm long, oblong-elliptic,
terminated by a long stout dark red seta, ciliate with long
stout white hairs tufted towards the apex, setae with
puberulous bases; petiole appressed, 1 mm long gradually
merging with the lamina, flat, ciliate, occasionally with
a few small subsessile glands. Flowers in a terminal group
of 3—9 at the ends of main branches, either verticellate
or secund; pedicel 3,5— 5,0 mm long, red with crisped hairs
and subsessile small glands; bract 3 x 0,5 mm, linear to
linear-spathulate subbasal; bracteoles like the bract but
more slender, median. Calyx 4-lobed mostly 5 mm long,
lobes 3/4 the length of the calyx, basal half flat, ovate, and
ciliate with sessile marginal glands, upper half acute
elongate foliar sulcate and ciliate with long hairs, sessile
glands on inner surface of the base otherwise glabrous,
green to completely red in dark-coloured flowers. Corolla
6—9 x 3—4 mm variable in shape from tubular-
campanulate to narrowly campanulate to narrowly cyathi-
form, shiny viscid oxblood-red sometimes very darkly so;
lobes deltoid, 1/6 the length of the corolla, erect or
spreading to reflexed. Stamens included; filaments 5 mm
long, considerably dilated at the apex, ciliate in the upper
half; anthers remaining in a ring around the style due to
interlocking lateral hairs, dorsally attached near the base,
thecae 1,2 mm long, narrowly ovate, golden brown with
back and filament apex dark red-brown, adaxial hairs
colourless, lateral and abaxial hairs reddish. Ovary 1,6—
2,0 x 1,0— 1,2 mm, obovoid slightly emarginate with a
short stipe, glabrous green; style 6—7 mm long, narrowly
cylindric, glabrous; stigma just exserted, subcapitate. Fruit
a narrowly obovoid capsule with a slight stipe and valves
splitting to the base; seeds subspherical , papillate. Figure 5.
This species is similar in overall respects to the species
of the Section Euryloma which is concentrated in the
Caledon District of the south-western Cape. However, it
is easily recognizable on account of the open, almost
cup-shaped flowers which have a dark red colour like
freshly dried oxblood.
E. cunoniensis is known from only a single mountain
peak in the far western portion of the Caledon District near
Rooi Els. This area is named after the indigenous rooi els
tree, Cunonia capensis L., hence the specific epithet. On
the same mountain there occur two other species in
Euryloma, E. gysbertii and E. cygnea Salter. Both
these species flower later in the summer and have
FIGURE 4. — Known distribution of E. amicorum, #; and E. cunonien-
sis, 4t.
FIGURE 5. — Erica cunoniensis : A, flowering branch, x 0,5; B, young leaf from below a florescence, x 12,5; C, old leaf, X 12,5; D, florescence,
x 1; E, flower, x 12,5; F, bract, x 12,5; G, sepal, x 12,5; H, anther, front, side & back views, x 12,5; I, gynoecium, X 12,5; all
drawn from the type collection, Oliver 8974 (STE).
pale pink, tubular conical flowers with a closed mouth to
the corolla. E. cygnea is also endemic to Buffelstalberg
but is unique in having nodding terminal inflorescences
and branches with bends like a swan’s neck.
E. cunoniensis also bears some resemblance to material
of E. squarrosa which does not occur in the same area
but further east on the mountains east and north of the
Grabouw Basin. Like E. gysbertii, E. squarrosa has
flowers with a closed narrow ovoid-urceolate corolla with
spreading lobes. In the latter two species, the basal portion
of each flower (pedicel, bract, bracteoles and calyx) is
much less hairy than in E. cunoniensis. The anthers of
E. cunoniensis are similar in shape to those of E.
squarrosa. In contrast to the free glabrous anthers of
E. squarrosa, however, those of E. cunoniensis are united,
due to lateral interlocking crisped hairs. E. gysbertii has
arcuate anthers typical of another group of species in the
section. The more open shape of the flowers is also
distinctive and resembles the flowers of E. lananthera L.
Bol. from the mountains around the mouth of the Palmiet
River but this species too has arcuate anthers and very
different leaves resembling those of E. retorta Montin.
E. cunoniensis was first collected by Thomas Stokoe in
1936. I first saw the species in 1966 as several scattered
shrubs on the dry northern slopes of Buffelstalberg. The
mountain was visited again in 1969 when a single plant,
just past flowering, was all that could be found. At the
time it was thought that the plant may have been a hybrid
between E. gysbertii and E. massonii L.f., both common
on these slopes but only flowering later in summer. Natural
hybrids between very different-looking putative parents
have previously been recorded in Erica (Oliver 1977,
1986). Lower down the slopes on an earlier occasion I had
noted a plant of clearcut hybrid origin between E. mas-
sonii and E. cygnea.
46
Bothalia 20,1 (1990)
A subsequent search failed to locate any plants of the
new species until a concerted effort was made together
with members of the A Team of the Botanical Society of
South Africa. The plants were eventually found on the
southern slopes, thus indicating that the original plants had
been chance strays. On the southern side of Buffelstalberg
the plants were growing in a single population on a cool
moist loamy slope with a south-western aspect. Subsequent
examination of the pollen showed it to be normal and well
developed.
Material of E. squarrosa in the British Museum and
Kew, which could form part of the type collection by
Masson, is similar to that of E. gysbertii, but unlike several
collections which have been identified as E. squarrosa in
Cape herbaria. I refer to this latter material as Species A,
which has been collected from the Hottentots Holland
Mountains between Landrostkop and the Triplets. It has
been recorded as a low compact shrublet with flowers ‘bril-
liant shiny crimson, dark at the throat’. In the flower colour
Species A is similar to E. cunoniensis but it has globose-
urceolate corollas with a very narrow mouth and the
anthers are glabrous, free and situated at the mouth of the
corolla. This, coupled with the open nature of the flowers
and semiconnate anthers, would indicate a totally different
pollinator for E. cunoniensis. However no pollinators were
observed by Rebelo and myself on Buffelstalberg. The
pollinator for E. gysbertii and E. squarrosa is a long-
proboscid fly giving the rhino-myiophilous pollination
syndrome (Rebelo, Siegfried & Oliver 1985).
E. cunoniensis
CAPE. — 3418(Simonstown): Hazel Peak (?) near Rooi Els, 600 m,
x.1950, (— BD), D.K. Davis SAM 65410 (SAM); Buffelstalberg near Rooi
Els, 670 m, x. 1966 (-BD), Oliver STE 31926 (PRE, STE); ibid, x.1969
(— BD), Oliver sub Baker 2974 (NBG); ibid. 5.x. 1986 (-BD), Oliver
8974 (BM, BOL, E, G, K, MO, NBG, P, PRE, S, STE, W); ibid.
18. ix . 1987, Schumann 560 (STE); mountains near Rooi Els, 1 066 m,
x.1936 (— BD), Stokoe 6380 (BOL); ibid. Stokoe SAM 68303 (SAM).
E. squarrosa Salisb. in Transactions of the Linnean Society
6: 380 (1802). Type: Cape, Lange Kloof [?], Masson s.n.
(possibly BM!).
CAPE. — 3419 (Caledon): Emerald Dome (-AA), Kruger & Haynes
729 (STE); French Hoek Kloof (-AA), Niven 151 (BOL, K, SAM);
Groenlandberg(— AA), Oliver 4172 (K, STE). Without precise locality:
Rivier Zondereinde (?AA), Masson 40 (BM, BOL, K). Without locality:
Masson s.n. (BM).
Species A
CAPE. — 3418 (Simonstown): Somerset Sneeukop (— BB), Brenan
14033A (K, NBG); Clarke sub NBG 2506/30 (NBG); Dyke sub Marloth
4359 (BOL); Esterhuysen 3613, 3614, 3/790 (BOL); Stokoe 6427 (BOL);
Williams 1155 (K, NBG); Triplets (— BB), Esterhuysen 8238 (BOL);
Landroskop (— BB), Stokoe sub SAM 54159 (SAM); Stokoe sub SAM 55177
(SAM); Valley Berg (-BB), White sub SAM 12743 (SAM). 3419
(Caledon): Victoria Peak (— AA), Esterhuysen s.n. (BOL); Kerfoot 5737
(STE).
E. gysbertii Guth. & Bol. in Flora capensis 4: 90 (1905).
Type: Cape, Stellenbosch Div., on the western foot of the
Hottentots Holland Mountains, 200 ft., Guthrie 3654
(BOL!).
CAPE.— 3418 (Simonstown): Somersfontein (-BB), Boucher 1003
(STE); Oliver 8996 (STE); Pringle Peak (-BD), Barker 8506 (NBG,
STE); Groot Hangklip (-BD), Boucher 747 (STE); path to Hangklip
(— BD), B. Guthrie sub F. Guthrie 3654 (BOL); Buffelstalberg (-BD),
Le Maitre 421 (STE); Rooi Els (— BD), Oliver 96 (STE).
Erica tegetiformis E.G.H. Oliver, stat. etnom. nov.,
E. senilis Klotzsch ex Benth. var. australis Dulfer: 32
(1963), non E. australis L.: 231 (1771). Type: Cape,
Worcester Div., Sonklip, N of Matroosberg, 6500 ft.,
17. i. 1959, Esterhuysen 28122 (W, holo.!; BOL!, K!, STE!).
Low compact mat-forming shrublet up to 1 m across
and 150 mm tall. Branches closely set, mostly prostrate,
often partially buried and rooting, with numerous erect
branchlets, sparsely and finely puberulous. Leaves 3- or
4-nate, erect, imbricate, 3, 4-3,8 mm long, oblong-elliptic
obtuse to subacute, sulcate, setose-ciliate with the cilia soon
becoming stublike, occasionally with some sessile glands
admixed, puberulous all over to almost glabrous; petiole
0,8 mm long, finely and shortly ciliate. Flowers 2-6-nate
at the ends of erect lateral mesoblasts; pedicel puberulous,
1, 2-2,0 mm long; bract median, 2, 5-2,8 mm long,
narrowly ovate or ovate-acute, sulcate in the upper 2/3, with
a puberulous base, ciliate with ± 10 long stiff hairs and a
longer apical one, white; bracteoles like the bract but
narrower, subapproximate. Calyx 4-partite, white; segments
2,7 x 1,0 mm, erect-spreading, narrowly ovate, subobtuse,
sulcate in upper ciliate with ± 12 long stiff hairs. Corolla
4-lobed, oblate-urceolate with 2 invaginations in the basal
half, finely puberulous, lobes broadly deltoid, spreading
to reflexed. Stamens manifest to partially exserted;
filaments2,3— 0,5 mm long, elongate-oblong, narrowed and
subsigmoid below the anther, glabrous or villous; anthers
dorsally attached near the base, thecae 1,3 mm long,
narrowly ovate-oblong, obtuse, hairy in front sometimes
with a few lateral hairs, basally crested, crests short, broad,
fimbriate, pore 2/3 — 3/4 the length of the theca. Ovary 1,1
x 1,3 mm, crown-shaped, 8-ridged, emarginate, puberulous
sometimes above only; style 4 mm long reducing in
thickness upwards, glabrous; stigma exserted, subcapitel-
late. Fruit shortly subcylindric; capsule sparsely puberulous;
seeds 0,8 mm long, irregularly ellipsoid , reticulate. Figure 6.
The unspecialized inflorescence, dense mat-forming
habit, globose urceolate flowers with lanceolate sepals and
distinctly visible corolla and the dark brown anthers serve
to distinguish E. tegetiformis.
This taxon was originally described by Dulfer (1964) as
a variety of the Cedarberg species, E. senilis Klotzsch ex
Benth., based on the single collection of Esterhuysen 28122
from Sonklip near Matroosberg. However, material recent-
ly collected and sent to me for identification, looked so
different from E. senilis that I did not link the material
with that species at the time and regarded it as a distinct
undescribed species. Only when checking on possible
alliances within the Section Chromostegia, particularly E.
senilis and E. eriophoros Guth. & Bol. did I realise that
Dulfer’s variety was in fact this new species and that it
bears only a superficial resemblance to E. senilis.
E. tegetiformis undoubtedly belongs in the Section Chro-
mostegia because of the prostrate habit and the compact
heads of flowers even though in the new species this latter
character is not very pronounced as is the case in E. senilis
in which the leaves below the inflorescence as well as the
bract and bracteoles, are enlarged and coloured giving an
involucrate appearance to the inflorescence. This feature is
Bothalia 20,1 (1990)
47
FIGURE 6. — Erica tegetiformis : A, flowering branch, x 0,5; B, leaf, x 12,5; C, two florescences, x 1; D, flower, X 12,5; E, bract, x 12,5;
F, sepal, x 12,5; G, anther, front, side & back views, X 12,5; H, gynoecium, x 12,5; all drawn from Oliver 9074 (STE).
far less developed in E. eriophoros and hardly so in
E. tegetiformis. Table 1 lists the characters showing
differences between the three species. Similarities with E.
senilis are not close, perhaps only in the relative size and
shape of the anther. There is more affinity with
E. eriophoros, but this in turn shows a closer affinity with
E. senilis in many characters.
Populations of this species were visited on Rocklands
Peak north of Ceres. Here the plants were growing among
very short restiads on open dry stony slopes at an altitude
of 1 440 m. At this altitude the plants must be subjected
to considerable cold in winter and even a heavy covering
of snow. In this regard they were effectively adapted
by being very low-growing and forming extensive compact
mats, hence the specific epithet. The long prostrate
branches frequently produce adventitious roots at intervals
along their length. From other collections the notes
give some indication of the habit and habitat: Stokoe
4518 — ‘straggling over sq. yds of bare ground’;
Esterhuysen 24328 — ‘rocky summit, sprawling amongst
rocks’; Esterhuysen 29853 — ‘trailing in habit sometimes
colonizing after a fire’. Figure 7.
E. senilis, which occurs to the north, does not produce
dense mats. It has prostrate as well as erect branches
forming a spreading shrub up to 300 mm tall and
grows in amongst denser ericoid/restioid vegetation.
E. eriophoros is rather rare. Until recently it was known
only from the type collection made at the end of last
century on Gydoberg in the southern Cold Bokkeveld.
Recent collections were made on Gideonskop in the
southern Cedarberg. From these few collections it is
difficult to assess the habit of the species. Plants growing
in the National Botanical Garden, Kirstenbosch, were
originally small, compact and erect, but have begun to
form lateral creeping branches. Mr Kotze, horticulturist
at Kirstenbosch, informs me that the plants on Gideonskop
were prostrate and mat-forming on a sandy open plateau.
E. tegetiformis appears to grow at slightly higher
altitudes and occurs south of the other two species, hence
Dulfer’s choice of epithet, which cannot now be adopted
at species level because of the European species,
E. australis L. The locality from Naudesberg in the
westernmost part of the Langeberg range is somewhat
unusual for species associated with the Cedarberg/
Bokkeveld region.
CAPE. — 3319 (Worcester): Rocklands Peak, vi.1955 (-AB),
Esterhuysen 24328 (BOL); ibid., 1 440 m, 19. xii.1987 (— AB), Oliver
9074 (BM, BOL, K, MO, NBG, PRE, STE); Baviaansberg, 2.i.l942
(—BA), Compton 12838 (BOL); ibid., 1 676 m, 4.xi.l962, Esterhuysen
48
Bothalia 20,1 (1990)
TABLE 1. — Characters distinguishing between Erica senilis, E. eriophoros and E. tegetiformis
29853 (BOL, K); ibid., i.1937, Stokoe 4518 (BOL); ibid., 1 828 m,
12. i. 1956, Stokoe in SAM 68303 (BOL, SAM, STE); Sonklip, north of
Matroosberg, 1 980 m, 17. i . 1959 (— BC), Esterhuysen 28122 (BOL, K,
STE, W); Conical Peak, xii.1940 (-BC), Stokoe 7832 (BOL); ibid, i.1941,
Stokoe SAM 55290 (SAM); Naudesberg, 13.xi.80 (-DB), Van Jaarsveld
& Bean 5819 (BOL, NBG, PRE, STE).
E. senilis Klotzsch ex Benth. in DC., Prodromus 7: 617
(1839); Guth. & Bol.: 235 (1905). Type: Cape, Ceder-
bergen, Drege 2966 (Bf, holo.!; K!, SAM!, W!).
FIGURE 7. — Known distribution of E. tegetiformis, #; E. senilis, ▲ ;
and E. eriophoros, O.
CAPE. — 3219 (Wuppertal): Sneeukop (— AA), Bolus 8627 (BOL,
STE); Heuningvlei (— AA), Emdon 146 (K, STE); Krakadouw (— AA),
Pocock 498, 593 (STE); Heuningvlei (— AA), Taylor 8548, 10475 (STE);
Hoogvertoon (—AC), Forsyth 39 (STE); Wolffberg to Gabriel’s Pass
(—AC), Kruger 926 (STE); Sneeuberg (—AC), Pocock 401 (STE); Taylor
6163 (STE); Middelberg Pass (-CA), Oliver 4016 (STE); Gideonskop
(-CB), Rourke 664 (NBG, STE); Bloukop (-CB), Williams 2123 (NBG,
STE); Tweefontein (-CC), Schlechter 10129 (BM, BOL, K); Heiveld
in Cold Bokkeveld (—CD), Hanekom 1212 (STE); Cederberg, without
precise locality, Drege 2966 (K, SAM).
E. eriophoros Guth. & Bol. in Flora capensis 4: 234
(1905). Type: Cape, Ceres Div., Gydouw Mountain in
Cold Bokkeveld, 6000 ft, Schlechter 10240 (BOL, holo.!;
K!, PRE!, SAM!).
CAPE. — 3219 (Wuppertal): Gideonskop (— CB), Williams 990 (NBG);
ibid. Kotze 787 (hort. NBG). 3319 (Worcester): Gydouw (— AB),
Schlechter 10240 (BOL, K, SAM).
REFERENCES
DULFER, H. 1964. Revision der siidafrikanischen Arten der Gattung
Erica L. 2. Annalen des Naturhistorischen Museums Wien 68:
25-177.
GUTHRIE, F. & BOLUS, H. 1905. Erica. In W.T. Thistleton-Dyer, Flora
capensis 4,1: 2—315.
LINNAEUS, C. 1771. Mantissa plantarum altera: 229-236. Stockholm.
OLIVER, E.G.H. 1977. The identity of Erica flavisepala. Bothalia 12:
195-197.
OLIVER, E.G.H. 1986. The identity of Erica vinacea and notes on
hybridization in Erica. Bothalia 16: 35—38.
OLIVER, E.G.H. 1987. Studies in the Ericoideae (Ericaceae). VII. The
placing of the genus Philippia into synonymy under Erica; the
southern African species. South African Journal of Botany 53:
455-458.
OLIVER, E.G.H. 1988. Studies in the Ericoideae (Ericaceae). VI. The
generic relationship between Erica and Philippia in southern
Africa. Bothalia 18: 1-10.
REBELO, A.G., SIEGFRIED, W.R. & OLIVER, E.G.H. 1985.
Pollination syndromes of Erica species in the south-western Cape.
South African Journal of Botany 51: 270—280.
Bothalia 20,1: 49-59 (1990)
Studies in the southern African species of Justicia and Siphonoglossa
(Acanthaceae): seeds
K.L. IMMELMAN*
Keywords: Acanthaceae, Justicia, seeds, .Siphonoglossa, South Africa, taxonomy
ABSTRACT
The seeds of all species of Justicia (except J. thymifolia (Nees) C.B. Cl.) and Siphonoglossa in the southern African region
(as defined by the Flora of southern Africa) were examined with the Scanning Electron Microscope. A wide range of different
seed surfaces were seen, many of which could be correlated with other characters to further reinforce grouping of species
into sections. However, some species that on other characters, would be placed together, had widely differing seed testas.
Gross surface morphology varied from smooth to colliculate, irregularly rugose, reticulate with the ridges variously arranged,
wheel-shaped (radiate or ammonite-like), with a central ridge or with multicellular barbed scales. The seeds were without
hairs, unlike some American species sometimes considered to belong in Justicia (Graham 1989). Micromorphology also
provided useful characters, with seeds being reticulate, papillate, minutely or deeply rugose or with one to many crystals
visible. Critical-point drying was sometimes helpful in viewing microstructure.
UITTREKSEL
Die sade van al die Justicia- en Siphonoglossa-spesies (behalwe J. thymifolia (Nees) C.B. Cl.) in die Suider-Afrikastreek
(soos deur die Flora of southern Africa omskryf) is met die aftaselektronmikroskoop bestudeer. ’n Wye reeks verskillende
saadoppervlakke is waargeneem, waarvan baie met ander kenmerke gekorreleer kon word om die groepering van spesies
in seksies verder te ondersteun. Sommige spesies wat op grand van ander kenmerke saam gegroepeer sou word, het egter
uiteenlopend verskillende saadtestas gehad. Makro-oppervlakmorfologie het varieer vanaf glad tot knopperig, onreelmatig
rimpelrig, netvormig met die riwwe op verskeie maniere gerangskik, wielvormig (straalvormig of ammonietvormig), met
’n sentrale rif of met meersellige stekelskubbe. Die sade was sonder hare, anders as by sommige Amerikaanse spesies wat
soms as lede van Justicia beskou word (Graham 1989). Mikromorfologie het ook nuttige kenmerke opgelewer, met sade
wat netvormig, gepapilleer, fyn of diep rimpelrig was of met een tot baie kristalle sigbaar. Kritieke-punt-droging was soms
nuttig by waameming van mikrostruktuur.
INTRODUCTION
Very few SEM investigations of the seed surfaces of
African members of the Acanthaceae have so far been
carried out. Those with relevance to the present study are
briefly reported on below.
Munday (1980), in revising the southern African species
of Monechma (Justiciae), found that, although all
seeds had a smooth testa, there was variation in their
size, shape and colour. A longitudinal ridge was charac-
teristic of one species [M. desertorum (Engl.) C.B.C1] and
mottled colouring of another [M. spartioides (T. Anders.)
C.B.C1.].
In his paper on Anisotes (also Justiciae) Baden (1981b)
found that the seeds often had a longitudinal ridge, and
he described the texture as rugose, tuberculate, verrucose
or smooth. In one species (A. guineensis Lindau) the seeds
were glandular. This species was unusual in other ways
also, such as having 4-porate pollen, and was placed by
Baden in a separate monospecific section.
Balkwill & Getliffe Norris (1984) used the fact that the
seed surface in Hypoestes was asperous, tuberculate or
smooth in their key to species.
Balkwill, Getliffe Norris & Schoonraad (1986) used the
microsculpturing of the seed surface in their investigation
of Peristrophe and concluded that its value at present was
* National Botanical Institute/Department of Agricultural Development,
Private Bag X101, Pretoria 0001.
MS. received: 1989.06.15.
mainly at species level. The testa of the species of the genus
was basically reticulate with minute papillae, and may or
may not have tubercules bearing one, two or three layers
of recoiled hooks. Details of the tubercule surface were
also used.
Hedren (1987) in his revision of Justicia capensis Thunb.
and its tropical African allies, described the seed surface
only as ‘tuberculate’ and did not use it to distinguish
species. In his paper on the Justicia mollugo group in
tropical Africa (1988), he described the seeds of most of
the species dealt with, but did not use them in his key.
Balkwill & Getliffe Norris (1988) re-appraised tribal and
subtribal limits within the Acanthaceae of southern Africa,
and considered the lack of hairs on the seed coat to be
definitive of Justiciae and some genera of Acanthoideae.
However, Graham (1989) included species of Justicia with
a variety of hair types, some even being tested by her and
found to be hygroscopic (in her sect. Anisostachya) .
Graham’s delimitation of Justicia was very wide, including
a number of previously segregate genera, and she
described some 20 seed types. In sect. Sarotheca some
species had seeds with hairs having recoiled barbs similar
to those seen by Balkwill et al. (1986) in Peristrophe. In
sect. Vasica some of the Old World species had smooth
testas, a character usually considered characteristic of
Monechma, which she accepted as a distinct genus.
METHODS
Seeds of each species and subspecies of Justicia in
southern Africa (except/, thymifolia), as well as Siphono-
50
Bothalia 20,1 (1990)
FIGURE 1. — Justicia sect. Harnieria p.p. : A-C, J. capensis : A, whole seed, x 30; B, detail of seed, x 360; C, whole seed, x 1200. Justicia
sect. Rostellularia subsect. Ansellia : D— G. D, E, J. crassiradix: D, whole seed, arrow indicates semicircular marginal ridges, X 70; E,
detail of seed, x 360. F, G, J. anselliana : F, whole seed (critical-point dried), x 70; G, detail of seed (critical-point dried), x 1600.
Bothalia 20,1 (1990)
51
glossa leptantha subsp. leptantha and Monechma mollis-
simum, were examined. Taxa covered in this study all occur
in southern Africa as defined by the Flora of southern
Africa, i.e. the area south of the Cunene and Limpopo
Rivers. Seeds of each were removed from herbarium sheets
and mounted directly onto 15 mm aluminium stubs for
viewing under the SEM. Two to six seeds per taxon were
viewed depending on the size of the seeds.
Where fresh fruiting material was available from the
nursery (National Botanical Institute, Pretoria), seeds were
also dehydrated, fixed and critical-point dried by the
following method:
1, specimens were dehydrated either in series of ascending
concentrations of alcohol or, more usually, in two changes
of 2,2-Dimethoxypropane (DMP) for up to two hours;
2, they were then transferred to 100% acetone for five
minutes;
3, critical point drying was done with liquid C02
evaporated off at 40°C and 80-90 atmospheres (critical
point for C02 is at 31,3°C and 72,8 atmos.);
4, the dried specimens were stored in a desiccator with
silica gel, or immediately mounted on aluminium stubs
for coating.
All seeds were glow-discharge coated with metallic gold
in an Eiko sputter coater. They were then viewed with an
MSM 4 Hitachi-Akashi (desk top model) SEM at kv 15
and photographed.
Delimitation and names of sections are according to
Graham (1989).
RESULTS AND DISCUSSION
Results of critical-point drying: it was noted in many
instances that material that had not been critical-point dried
showed structures of the testa such as radial cell walls,
ornamentation of these walls, and crystals in the testa,
better than fresh, critical -point dried material. This was
due to the collapse of the outermost cell walls, which made
it easier to see underlying structures. On the other hand,
structures such as fine papillae were sometimes only
visible when the material had been pre-treated before
viewing. This was the case with /. anselliana (Nees) T.
Anders. (Figure 1G), and it was felt that critical-point
drying would be a valuable procedure for those species
which, like / anselliana, had ‘amorphous’ testas, as found
in sects. Ansellia and Justicia p.p.
A wide variety of seed surface types was seen,
sometimes peculiar to the species, but in at least some
cases correlating with characters used to demarcate
sections.
Sect. Rostellularia subsect. Ansellia p.p. (/. crassira-
dix Burkill & Clarke, /. anselliana (Nees) T. Anders. , /.
anagalloides (Nees) T. Anders.): in /. crassiradix the seed
has a gross pattern of reticulate ridges and depressions on
the sides, whereas around the margin of the seeds the
ridges form a roughly semicircular pattern (Figure ID,
arrowed). The testa of two other southern African species,
/. anselliana (Figure IF) and /. anagalloides , which on
the basis of other characters should probably be placed
with /. crassiradix in subsect. Ansellia, do not show this
semicircular pattern. Instead, the ridges are randomly
placed on the seed surface. At higher magnifications, when
they were viewed without having been critical-point dried,
the seeds of all three species had an irregularly rugose
surface (Figure IE). When seed of /. anselliana was
critical-point dried, the cells in the depressions between
the ridges were found to each have a centrally situated
papilla (Figure 1G), while the cells on the ridge were
without papillae, having instead a minutely verrucose
surface. The papillae were longer than wide, with an acute
or truncate apex. It was not possible to establish whether
these papillae were present in the other two species, as
fresh material was not available.
Sect. Justicia p.p. (/. orchioides L.f., /. cuneata
Thunb., /. guerkeana Schinz, /. platysepala (S. Moore)
RG. Mey., /. thymifolia (Nees) C.B. Cl.): rugose testas
are also present in all examined species of Sect. Justicia
except /. bolusii C.B. Cl. (Figure 2A, B). Only the first
four species could be examined, as seeds of /. thymifolia
were not available. This irregularly rugose pattern is
apparent at both low and high magnifications. Seeds of
/. guerkeana were also critical-point dried for viewing.
In this case, unlike /. anselliana, the irregularly rugose
pattern remained, and no papillae were seen.
Sect. Justicia p.p. (/. bolusii C.B. Cl.): this species
(Figure 2C, D), which on other characters might be
considered to occupy an isolated position among the
southern African species of the genus, also has very
atypical seeds. These are almost smooth, being only very
slightly rugose even at high magnifications (Figure 2D)
and compare well with Monechma mollissimum (Nees)
P.G. Mey., which was also viewed for purposes of
comparison (Figure 2E, F). However, it was decided that,
despite similarities in the seeds and capsules, /. bolusii
could not be transferred to Monechma. It differs from
Monechma in its ‘spicate’ inflorescence and its tricolporate
pollen with entire margocolpi (Immelman 1989).
Sects. Raphidospora p.p. (/. campylostemon (Nees
T. Anders.), Tyloglossa p.p. (/. flava (Vahl) Vahl, /.
kirkiana T. Anders.), Rostellularia subsect. Ansellia p.p.
(/. minima A. Meeuse) and Harnieria p.p. (/. capensis
Thunb.): seeds of a number of species of Justicia from
various sections of the genus (listed above) are basically
colliculate or rugose with slightly protruding radial cell
walls forming a reticulate surface pattern. However, if
other characters of the species are taken into consideration,
they are not necessarily related, and there are also many
differences in detail in both the macro- and micro-patterns.
/. capensis (Figure 1A— C) and/, minima (Figure 3A— C)
have a randomly colliculate macro-pattern and, at high
magnifications, show a simple reticulate micro-pattern.
/. campylostemon (Figure 4A— C) has an irregularly
vermiculate macro-pattern, and the radial cell walls are
ornamented with a double row of minute projections,
which are more clearly visible without critical-point drying
(Figure 4C). /. flava, J. kirkiana, and /. petiolaris have
characteristic macro-patterns (see below) and crystals
visible in the testa.
Sect. Tyloglossa (/. flava (Vahl) Vahl, /. kirkiana T.
Anders, and /. petiolaris (Nees) T. Anders.): these three
52
Bothalia 20,1 (1990)
FIGURE 2. — Justicia sect. Justicia : A-D. A, B, J. orchioides : A, whole seed, x 55; B, detail of seed, x 1900. C, D, J. bolusii : C, whole
seed (composite figure), x 25; D, detail of seed, x 1800. E, F, Monechma mollissimum: E, whole seed, x 60; F, detail of seed, X 2300.
Bothalia 20,1 (1990)
53
FIGURE 3. — A-C, Justicia minima: A, whole seed, x 50; B, detail of seed, x 200; C, detail of seed, x 1200. Justicia sect. Hamieria p.p.:
D-G. D, E, J. protracta subsp. protracta: D, whole seed, X 50; E, detail of seed, X 1400. F, G, J. parvibracteata: F, whole seed, X
70; G, detail of seed, x 3500.
54
Bothalia 20,1 (1990)
species (Figure 5) form a well-defined section. They all
have a characteristic reticulate micro-pattern on the seed
surface and one large or many small crystals visible as
cubic or rectangular projections below the surface of the
outer wall (Figure 5B, D, F). Such crystals were not seen
in any other Justicia species examined. These crystals are
scarcely visible in critical -point dried material, but are
clearly seen when the material is viewed without pre-
treatment. The two yellow-flowered species (7. flava and
7. kirkiana) have a highly characteristic macro-pattern of
raised segments or bosses (colliculae) (Figure 5C, E),
termed ‘ammonite-like’ by Graham (1989), but which could
also be interpreted as three segmented rows or as segments
radiating from a central ‘hub’ like spokes of a wheel. This
pattern is not present in any of the subspecies of the blue-
flowered 7. petiolaris, where the gross pattern of the testa
is sometimes colliculate, and sometimes rather irregularly
rugose, often having a central longitudinal ridge.
Sect. Betonica (7. betonica L. and 7. montis-
salinarum A. Meeuse): both these species have a macro-
pattern of short sinuous ruminate ridges (Figure 6). The
ridges are often drawn out into dentate structures which
either have an areolate micro-pattern on their surface, as
in 7. betonica (Figure 6B) or they are irregularly striate
as in 7. montis-salinarum (Figure 6D). When broken, these
teeth can be seen to be hollow.
Sect. Raphidospora p.p. (7. glabra Koenig ex Roxb.):
the most complex pattern observed was that of 7. glabra,
which has seeds quite unlike those of any other Justicia
species (Figure 7A— E). It is probable, on the basis of
inflorescence and pollen characters, that it should be
grouped with 7. campylostemon, in sect. Raphidospora,
but the seed testas of the two species are quite different.
The testa of 7. glabra is drawn out into numerous long
flat multicellular scales (Figure 7B) with each cell of the
scale producing a sharp retrorse barb (Figure 7C— E). The
scales are probably a means of seed dispersal, analogous
to the barbed scales on the fruit of Bidens pilosa
(Asteraceae). 7. campylostemon, on the other hand, has
the seeds covered with irregular vermiculate ridges, each
of which has a reticulate micro-pattern on the surface.
Along each margin of the raised ridges of the reticulations
there is a row of small, blunt, tooth-like projections (Figure
4B, C).
Sect. Harnieria p.p. (7. protracta (Nees) T. Anders.,
7. odora (Forssk.) Vahl, 7. parvibracteata Immelman, 7.
dinteri S. Moore) as well as Siphonoglossa leptantha
(Nees) Immelman subsp. leptantha : these species have a
colliculate macro-pattern. They also share a micro-pattern
where each cell is defined with a single large rounded
papilla in the centre of each cell (Figures 3D— G; 8). These
papillae are visible in material that was not pre-treated be-
fore mounting, as well as in critical-point dried material
of S. leptantha subsp. leptantha (Figure 8A). Material of
Siphonoglossa viewed without pre-treatment had an
irregularly rugose testa.
FIGURE 4. — Justicia sect. Raphidospora'. A— C, J. campylostemon-.
A, whole seed (composite picture), x 32; B, detail of seed, x
725; C, detail of seed, x 3000.
In 7. dinteri two kinds of capsule are regularly produced,
a four-seeded normal capsule and a one-seeded smaller
capsule with four irregularly toothed wings. The seeds
from each type of capsule were compared, and the testa
found to be similar, though the seeds in the one-seeded
capsules were larger (Figure 8B, D, F, cf. Figure 8C, E).
Bothalia 20,1 (1990)
55
FIGURE 5. — Justicia sect. Tyloglossa : A— F. A,B,D, J. petiolaris: A, subsp. petiolaris, whole seed, X 30; B, subsp. incerta , detail of seed,
x 2700; D, subsp. petiolaris , detail of seed, X 50. C, F, J. kirkiana: C, whole seed, x 50; F, detail of seed, x 2000. E, J. flava, whole
seed, x 50.
56
Bothalia 20,1 (1990)
FIGURE 6. — Justicia sect. Betonica : A— D. A, B, J. betonica : A, whole seed, x 32; B, detail of seed, X 70. C, D, J. montis-salinarum : C,
whole seed, x 50; D, detail of seed, X 220.
Bothalia 20,1 (1990)
57
FIGURE 7. — Justicia sect. Raphidospora: A— E, J. glabra : A, whole seed (composite picture), x 55; B, scales on seed (composite picture),
x 190; C, apex of scale, x 1800; D, shaft of scale, x 1800; E, base of scale, x 725.
One other species was examined, which on flower and
pollen characters possibly belongs in sect. Hamieria. This
was J. capensis Thunb. (Figure LA— C), which was placed
in this section by Graham (1989). The seeds, however, are
different from the papillate seeds of the other species in
the section, being colliculate with a strongly reticulate
micro-pattern.
CONCLUSION
Both macro- and micro-pattern of the seed testa may
be considered of major importance in placing many species
in their sections, and delimiting some sections in Justicia,
but need to be used with caution as they do not always
correlate with other features. They are also suggestive of
a relationship between some species of Justicia sect. Har-
nieria and Siphonoglossa.
ACKNOWLEDGEMENTS
This work was done as part of a Ph.D. thesis in the
Department of Botany, University of Natal, Pietermaritz-
burg. I would like to thank Mrs S. Perold of the National
Botanical Institute, Pretoria, for her assistance with the
Scanning Electron Microscope and my supervisor. Dr F.
Getliffe Norris, for her help and advice in the writing up
of my thesis.
58
Bothalia 20,1 (1990)
FIGURE 8 — Siphonoglossa leptantha subsp. leptantha : A, detail of seed (critical-point dried), X 2000. Justicia stcLHamieria p.p.. B E
j dinteri\ whole seed from one-seeded capsule, x 60; C-F, detail of seed, C, from four-seeded capsule, X 900; D, from one-seeded
capsule, X 370; E, from four-seeded capsule, x 3000; F, from one-seeded capsule, X 1000.
Bothalia 20,1 (1990)
59
REFERENCES
BADEN, C. 1981a. The genus Macrorungia (Acanthaceae), a taxonomic
revision. Nordic Journal of Botany 1: 143—153.
BADEN, C. 1981b. The genus Anisotes (Acanthaceae). Nordic Journal
of Botany 1: 623 —664.
BALKWILL, K. & GETLIFFE NORRIS, F. 1984. 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.
BALKWILL, K., GETLIFFE NORRIS, F. & SCHOONRAAD, E. 1986.
Thxonomic studies in the Acanthaceae: testa microsculpturing in
southern African species of Peristrophe. South African Journal
of Botany 52: 513-520.
GIBSON, D.N. 1972. Studies in American plants HI. Fieldiana (Botany)
34,6: 57-87.
GRAHAM, V.A.W. 1989. Delimitation and infra-generic classification
of Justicia (Acanthaceae). Kew Bulletin 43: 551—620.
HANSEN, B. 1985. Taxonomic revision of the SE Asian species of
Isoglossa (Acanthaceae). Nordic Journal of Botany 5: 1—13.
HEDREN, M. 1987. The Justicia capensis species group (sect. Hamieria ,
Acanthaceae) in tropical Africa. Kew Bulletin 43: 349—359.
HEDREN, M. 1988. The taxonomy of the Justicia mollugo group ( Justicia
sect. Hamieria, Acanthaceae). Bulletin du Jardin botanique
national de Belgique 58: 129—158.
IMMELMAN, K.L. 1987. A study of the genera Justicia L. and
Siphonoglossa Oerst. (Acanthaceae) in southern Africa. Ph.D.
thesis. University of Natal, Pietermaritzburg.
IMMELMAN, K.L. 1989. Studies in the southern African species of
Justicia and Siphonoglossa (Acanthaceae): palynology. Bothalia
19: 151-156.
MANNING, J.C. & GETLIFFE NORRIS, F. 1985. Taxonomic studies
in the Acanthaceae: a reappraisal of the genera Duvemoia and
Adhatoda in southern Africa. South African Journal of Botany
51: 475-484.
MUNDAY, J. 1980. The genus Monechma Hochst. (Acanthaceae tribe
Justiciae) in southern Africa. M.Sc. thesis, University of the
Witwatersrand, Johannesburg.
Bothalia 20,1: 61-66 (1990)
Studies in the southern African species of Justicia and Siphonoglossa
(Acanthaceae): indumentum
K.L. IMMELMAN*
Keywords: Acanthaceae, Justicia, Siphonoglossa , trichomes, indumentum, southern Africa, taxonomy
ABSTRACT
The trichome types present on all species of Justicia and Siphonoglossa in the southern African region (as defined by
the Flora of southern Africa) were examined with the dissecting and with the Scanning Electron Microscope. Both glandular
and eglandular trichomes were observed, the former comprising both sessile, peltate glands and stalked glands of various
lengths. Eglandular trichomes are either straight or sharply bent (anvil-shaped), and comprise two to many cells. They are
either smooth or have raised ornamentation on the cell walls. The presence and morphology of the trichomes on various
organs is recorded and measured, and the range of trichomes seen was illustrated. The possible significance of trichome
type and ornamentation in the taxonomy and ecology of the genera is discussed.
UITTREKSEL
Die trigoomtipes aanwesig op al die Justicia- en Siphonoglossa-spesies in die Suider-Afrikastreek (soos deur die Flora
of southern Africa omskryf) is met die disseksiemikroskoop en die aftaselektronmikroskoop ondersoek. Geklierde sowel
as klierlose trigome is waargeneem: eersgenoemdes behels beide sittende, skildvormige kliere en gesteelde kliere van verskeie
lengtes. Klierlose trigome is 6f reguit 6f skerp gebuig (aambeeldvormig) en bestaan uit twee tot baie selle. Hulle is 6f glad
6f het verhewe omamentasie op die selwande. Die aanwesigheid en morfologie van die trigome by verskeie oigane is aangeteken
en gemeet, en die verskeidenheid trigome wat waargeneem is, is gefllustreer. Die moontlike belang van trigoomtipe en
omamentasie in die taksonomie en ekologie van die genusse word bespreek.
INTRODUCTION
The indumentum of all species of Justicia and
Siphonoglossa found in southern Africa (as defined by the
Flora of southern Africa ) was examined. Numerous
authors, including Ahmad (1974a, b; 1978), Munday
(1980), Baden (1981), Balkwill & Getliffe Norris (1985),
Hansen (1985) and Manning & Getliffe Norris (1985), have
found pubescence and trichome types important as
indicators of relationships in various genera of the
Justicieae.
METHODS
Several specimens of each taxon were examined with
the dissecting microscope and the eglandular and stalked
glandular trichomes were measured using a micrometer
eyepiece.
Indumentum type present was recorded from the stem,
the leaf lamina and petiole/leaf base area, from the bracts,
the inner and outer surface of the calyx and from the
capsule. Material of one specimen of each species or
subspecies recognized by Immelman (1987), except J.
crassiradix, was examined under the SEM. The leaf, outer
and inner surface of calyx as well as the bract and the leaf
base/petiole area were scanned. Approximately 2x3 mm
of leaf tissue (and also of the bracts where these were large)
was removed from herbarium sheets. This was mounted
directly onto 15 mm diameter aluminium stubs and glow-
discharge-coated with ± 400 A° of metallic gold in an Eiko
sputter coater. They were then viewed directly in an MSM
4 Hitachi-Akashi (desk top model) SEM at kv 15,
* National Botanical Institute/Department of Agricultural Development,
Private Bag X101, Pretoria 0001.
MS. received: 1989.06.23.
and photographed using a Mamiya 6x7 camera and Ilford
FP 4 125 ASA film. The film was developed in Microdol
X.
Where fresh material was available this was critical-point
dried before being examined under the SEM. This was
found to be preferable to using dried material as the
trichomes, especially glandular trichomes, are then not
collapsed and their structure can be more accurately
observed. The material was dehydrated in 2,2-dimethoxy-
propane (DMP), fixed in FAA, and critical-point dried
in a Balzer’s Union critical-point drier using liquid carbon
dioxide. It was then mounted, viewed and photographed
as described above. Trichome types were defined and
named according to the terminology used by Munday
(1980). Presence or absence of the different types was
recorded for the different organs mentioned above (Table
1). Figures were prepared illustrating the range of trichome
types seen (Figures 1 & 2).
MORPHOLOGY
Two basic types of trichomes were found: 1, eglandular
and 2, glandular.
1. Eglandular trichomes (Figure 1 and type A’ on Table
1) are either straight (varying from long to short) or
gradually to sharply bent. They consist of (1?)2 to many
uniseriate cells with or without ornamentation.
In some species, e.g. J. parvibracteata, the trichomes
are two-celled (Figure 1G). J. montis-salinarum A.
Meeuse appeared to have one-celled trichomes, but the
trichomes were so heavily ornamented that it was not
possible to see whether they were 1- or 2-celled (Figure
1H). In most other species one to many additional cells
62
Bothalia 20,1 (1990)
FIGURE 1. — Eglandular trichomes, all x 420, except where otherwise stated. A, C, J. protracta subsp. rhodesiana, Germishuizen 974: A, from
leaf showing swollen basal cells; C, from calyx. B, J. cuneata subsp. hoerleiniana , from leaf, with ornamented and non-ornamented heads,
Dinter 6401; D, J. capensis, from leaf showing small branch on trichome, Immelman 375; E, J. odora, from leaf showing ornamented
trichome surface, x 3000, Meeuse 9530; F, J. kirkiana, from petiole, x 300, Brummitt & Banda 8539; G, J. parvibracteata, from leaf,
Esterhuizen 1101 ; H, J. montis-salinarum , from leaf, Van Wyk 5536. I, K, J. protracta subsp. protracta, Balkwill s.n. : I, from bract showing
striate joint at cell wall; K, from leaf near petiole, showing eglandular and glandular trichomes in two separate layers, x 120. J, J. orchioides
subsp. glabrata, from bract, Muir 1101.
Bothalia 20,1 (1990)
63
TABLE 1. — Trichome types present in taxa of Justicia and Siphonoglossa : A, eglandular trichomes; B, peltate glandular trichomes; C, stalked
glandular trichomes
are present, these being progressively narrower than the
basal cell. The upper cells are frequently set at an angle
to the one below, so that the trichome is gradually or
sharply bent or even appressed to the surface of the organ
on which they occur. The sharply bent condition (Figure
1A) has been described as anvil-shaped by Munday (1980).
The basal cell of the eglandular trichomes is thicker and
set into the epidermis. It is sometimes surrounded by a
ring of swollen epidermal cells (e.g. J. protracta (Nees)
T. Anders. — Figure LA). In anvil-shaped trichomes, it
is usually the second cell which is set at an angle to the
basal cell. The terminal cell is usually pointed but in J.
cuneata subsp. hoerleiniana (P.G. Mey.) Immelman it is
swollen, with one end rounded and the other more pointed
(Figure 1B). The terminal cell of J. cuneata subsp. hoer-
leiniana is either smooth or ornamented. Rarely, a
branched trichome was encountered, as in J. capensis
Thunb. (Figure ID). The cells of the upper tiers are usually
ornamented, except in those trichomes found on the inner
surface of the calyx lobes. The ornamentation is striate
to papillose (Figure IE, cf. Figure IF) but, as these types
intergrade, no attempt was made to draw a distinction
between them. The area of the joint between two cells is
smooth and not ornamented except for a ring of vertical
striations (Figure II).
In a number of taxa the eglandular and glandular trichomes
form discrete layers with the eglandular being the longer,
e.g. in J. protracta subsp. protracta (Figure IK).
2. Glandular trichomes : two main types were recorded,
peltate and stalked glands.
2.1 Peltate glands (Figure 2A— C; type ‘B’ in Table 1)
These glands consist of a solitary basal cell set as a
wedge into the epidermis (Figure 2B). The head, set on
the basal cell, is broad and flattened, and is probably four-
celled. No sections were cut to confirm this, but the head
is marked with two lines at right angles (probably cell
walls, Figure 2A). The top of the head is sometimes
marked with a feint ring where the lines cross. In J.
anagalloides (Nees) T. Anders. (Figure 2C), each cell of
the head has a single papilla on the outer edge. This feature
is unique among the species of Justicia seen in this study.
2.2 Stalked glands (Figure 2D— G; type ‘C’ in Table 1)
The basal cell of stalked glands is similar to that of the
peltate glands.
64
Bothalia 20,1 (1990)
FIGURE 2. — Glandular trichomes, all x 600, except where otherwise
stated. A, B, D, J. flava, Immelman 249: A, leaf, peltate gland;
B, leaf base, peltate gland showing wedge-shaped basal cell; D,
bract, stalked gland with long stalk, X 240. C, J. anagalloides ,
leaf base, peltate gland showing small protuberances at outer
margin of each cell; E, J. petiolaris , calyx, stalked gland with
long stalk; F, J. protracta subsp. protracta , inner surface of calyx,
stalked gland with short stalk, Balkwill s.n. ; G, J. betonica, calyx,
stalked gland with short stalk, Immelman 178.
The stalk varies in length (0,02 mm in 7. protracta subsp.
rhodesiana (S. Moore) Immelman to 1,56 mm in J.
petiolaris (Nees) T. Anders, subsp. petiolaris ) and in the
number of cells, and is not ornamented. Long-stalked
glands on certain organs are characteristic of some species,
e.g. the bracts of 7. kirkiana, J. petiolaris subsp. petiolaris
(Figure 2E), and sometimes of 7. flava (Vahl) Vahl, have
very long glandular trichomes.
Below the head is a collar cell, which is shortly
cylindrical and usually narrower than the rest of the stalk
(Figure 2D).
The glandular heads are of various types, and are
occasionally species-specific. In 7. betonica L., for
instance, the glands on the petiole (when present) have
elongated clavate heads, while those on the calyx have the
heads broad at the base and narrowing towards the apex
(Figure 2G). In some species the head is longitudinally
faintly striate (?cell walls). It is either the same width as
the stalk and longer than wide (Figure 2E) or much wider
than the stalk and wider than long (pin-shaped glands)
(Figure 2D, F, G).
The number of cells in the head is uncertain, as no
sections were cut, but accounts in the literature (Ahmad
1978) record from 1 to 17 cells in trichomes of the family.
TAXONOMIC VALUE
On the basis of indumentum morphology, Siphonoglossa
could not be distinguished from Justicia. It was also not
possible to delimit sections within these genera, or in the
related genus Monechma, solely on indumentum features.
Some of the trichome types Munday (1980) found in
Monechma are very rare or absent altogether from the
species of Justicia seen in this study. These types are
glandular papillae with the tip not wider than the stalk,
T-shaped eglandular trichomes, flattened uniseriate
eglandular trichomes, and dendroid trichomes. Anvil-
shaped trichomes having broad flattened shoe-like heads
were rare in Justicia but relatively common in Monechma.
However, as none of these trichomes were exclusive to
either of the genera, they cannot be used to distinguish
them at generic level.
At species and subspecies level, however, the trichomes
in Justicia provide many useful characters, and are
sometimes characteristic of one or a few species. The
following species, for example, have very long-stalked
glands on the bracts: 7. kirkiana, 7. flava (Figure 2D),
7. dinteri and 7. peuolaris subsp. petiolaris (Figure 2E).
In the case of 7. kirkiana and 7. dinteri, these glands give
the bracts a cottony appearance. Some species, such as
7. parvibracteata and 7. montis-salinarum, are puberulous,
with a dense indumentum of short, heavily ornamented
trichomes. One species pair among the southern African
species of Justicia is most easily distinguished on pubes-
cence characters. Because differences also exist in other
characters it seems reasonable to maintain these taxa at
specific rank. They are 7. platysepala, which has leaves
with sparse to dense pubescence, whereas in the closely
related 7. guerkeana the leaves are usually completely
glabrous. In 7. guerkeana the leaves are also three-veined
from the base and there is a difference in distribution.
Bothalia 20,1 (1990)
65
The pubescence is the most reliable morphological
feature for distinguishing certain subspecies. Because there
was also a degree of geographical separation, they were
recognized at subspecific rank. 7. cuneata, for example,
has the following three subspecies: subsp. latifolia is
glabrous except on the stems and on the inside of the calyx
lobes, subsp. cuneata is densely puberulous on all
vegetative parts as well as the exterior of the calyx, and
subsp. hoerleiniana has a dense covering of broad-headed
anvil-shaped trichomes. Another example, / protracta
subsp. protracta, has multicellular eglandular trichomes
averaging 0,47 mm in length (on the leaf lamina), whereas
in subsp. rhodesicma the whole plant is densely puberulous
with short, two-celled eglandular trichomes, of which the
average length (on the leaf lamina) is 0,04 mm. J.
orchioides subsp. orchioides has stiff opaque white
trichomes on stem and leaves while subsp. glabrata is
glabrous.
In Justicia, four taxa from arid areas have dense to
moderately dense long pubescence: J. dinteri, J. ansel-
liana, J. guerkeana and J. cuneata subsp. hoerleiniana.
The last taxon, a woody shrub, has an indumentum type
unique in the genus, though present in the related
Monechma. The very dense eglandular trichomes have
large swollen shoe-shaped heads which sit at 45—90° to
the stalks.
In some taxa, the indumentum is reduced or even lacking
(except for the sessile glands, see Table 1). Examples are
J. orchioides subsp. glabrata, J. crassiradix and / odora.
ECOLOGICAL SIGNIFICANCE
A number of probably closely related pairs of Justicia
species or subspecies are known of which one member
occurs in a more arid environment, and possesses an
indumentum markedly different from that of the other.
These are / montis-salinarum and/, betonica, J. protracta
and /. dinteri, J. protracta and 7. parvibracteata, J.
protracta subspp. protracta and rhodesiana and /. cuneata
subspp. cuneata and hoerleiniana. The last taxon occurs
in the most arid environment of any of the southern African
taxa of Justicia, in the Sperrgebiet of southern Namibia.
Johnson (1975), after reviewing the evidence for
the presence and density of pubescence in geographical
areas of different moisture stress, concludes that ‘It
thus seems clear that ecogeographic relationships of
pubescence must involve factors in addition of moisture’.
It is even possible, he suggests, that the presence of
trichomes increases the rate of transpiration, and that
their function may rather be to act as a light shield in
areas with intense radiation. Uphof & Hummel (1962)
make a similar suggestion, though they stress that
experimental proof is lacking. The bent shape and heavier
ornamentation of trichomes of some species would
seem to add to the effectiveness of such trichomes as
light shields. They state: ‘... the faculty to function as a
screen against the light would depend on the presence of
a large number of light-dispersing centres. The latter may
be present in the form of knobs on the cell-wall or on the
cuticle only or in that of more or less sharp bends in the
trichomes.’
This provides a possible explanation for the
predominance of short dense pubescence, consisting of
bent trichomes having heavy ornamentation, in many of
those species of Justicia occurring in arid or semi-arid
areas.
Species with such a dense puberulous indumentum in
which the trichomes are shorter and more heavily
ornamented than is generally found in species from moister
areas include / cuneata subsp. cuneata (W Karoo), /
protracta subsp. rhodesiana (Namibia, Botswana, N
Transvaal, Zimbabwe), 7. parvibracteata (N Cape) and /
montis-salinarum (Soutpansberg of the Transvaal). When
examined under a dissecting microscope these species all
appear minutely puberulous. When the trichomes are
examined under the SEM, they are seen to be reduced to
(1)2 cells. If the trichome is 2-celled, then the upper cell
is at an angle to the basal cell. The whole surface is
heavily ornamented with papillae, which are noticeably
denser and more projecting than in other species of the
genus.
Monechma, a genus predominantly of dry areas, was
found by Munday (1980) to have many species with
puberulous leaves, especially those occurring in the
southern part of Namibia. There are, however, exceptions
to this tendency, as in Justicia, where / orchioides subsp.
glabrata from the Karoo is glabrous.
Taxa with reduced or even lacking indumentum (except
for the sessile glands) (see Table 1) are found in both
arid regions (e.g. 7. orchioides subsp. glabrata from the
Karoo), and in moist, even marshy, habitats, e.g. 7
crassiradix, J. odora and sometimes 7 capensis.
It is considered probable that the function of the
trichomes in reflecting light is more significant than in
water conservation, and their shape and ornamentation
may be important in this regard. However, no clear-cut
picture has emerged, where a single type of indumentum
(or even its absence) could be consistently correlated with
aridity or geographical distribution.
CONCLUSIONS
Two basic types of trichomes were found: eglandular
and glandular. In most species both types are present. It
was not possible to distinguish genera or sections on the
basis of indumentum morphology. At species and
subspecies level, however, trichomes provide many
taxonomically useful characters. No clear-cut correlation
was found between indumentum and habitat, but a
predominance of short, dense pubescence was noted in
many Justicia species from arid and semi-arid areas.
ACKNOWLEDGEMENTS
This work was based on a Ph.D. thesis done in the
Department of Botany, University of Natal, Pietermaritz-
burg. I would like to thank Mrs S. Perold of the National
Botanical Institute, Pretoria, for her assistance with the
Scanning Electron Microscope, Ms Gill Condy for her
illustrations, and my supervisor, Dr F. Getliffe Norris, for
her help and advice in the writing up of my thesis.
66
Bothalia 20,1 (1990)
REFERENCES
AHMAD, A.D. Q. 1974a. Cuticular studies in some Nelsonioideae
(Acanthaceae). Botanical Journal of the Linnean Society 68:
73-80.
AHMAD, A.D. Q. 1974b. Cuticular studies in some species of Mendoncia
and Thunbergia (Acanthaceae). Botanical Journal of the Linnean
Society 69: 53—63.
AHMAD, A.D. Q. 1978. Epidermal trichomes of Acanthaceae. Blumea
24: 101-117.
BADEN, C. 1981. The genus Macrorungia (Acanthaceae), a taxonomic
revision. Nordic Journal of Boumy 1: 143—153.
BALKWILL, K. & GETLIFFE NORRIS, F. 1985. Taxonomic studies
in the Acanthaceae: the genus Hypoestes in southern Africa. South
African Journal of Botany 51: D3-144.
HANSEN, B. 1985. Taxonomic revision of the SE Asian species of
Isoglossa (Acanthaceae). Nordic Journal of Botany 5,1: l—O.
JOHNSON, H.B. 1975. Plant pubescence: an ecological perspective.
Botanical Review 41: 233-258.
MANNING, J.C. & GETLIFFE NORRIS, F. 1985. Taxonomic studies
in the Acanthaceae: a reappraisal of the genera Duvemoia and
Adhatoda in southern Africa. South African Journal of Botany
51: 475-484.
MUNDAY, J. 1980. The genus Monechma Hochst. ( Acanthaceae tribe
Justiciae) in southern Africa. M.Sc. thesis, University of the
Witwatersrand, Johannesburg.
UPHOF, J.C.Th. & HUMMEL, K.K. 1962. Plant trichomes. In
Encyclopaedia of plant anatomy 10,1: 1—206. Nicolasse, Berlin.
Bothalia 20,1: 67-90 (1990)
Notes on African plants
VARIOUS AUTHORS
ASTERACEAE
A NEW SPECIES OF PTEROTHRIX (GNAPHALIEAE) FROM THE NORTHERN CAPE
Pterothrix tecta Brusse, sp. nov.
Frutex clonalis usque ad 400 mm altus et 3,5 m latus,
aromaticus. Caules principals tortiles, atrocinerei, usque
ad 40 mm diametro, infra in arena infossi. Rami secundarii
usque ad 5 mm crassi, cortice tenui, aureo-badio vel badio
vel passim rubescenti, longistrorsum fissurato, glabri vel
aureo-pelliculato. Rami tertiarii robusti, primum lutei et
albo-tomentosi dein aurei, glabri et pelliculati. Folia
altema, sessilia, linearia, 1—11 x 0,7— 2,2 mm, ad basin
latissima, primum ascendentia dein patentia, marginibus
involutis. Pagina superior canaliculata, albo-lanuginosa.
Pagina inferior glabrescens, praesertim ad basin folliculata;
costa praecipue ad basin prominente. Apex obtusus,
mucrone minimo, inconspicuo. Capitula solitaria,
terminalia, sessilia, discoidea, cylindrica, 7-8 x 2, 5-3, 5
mm, 11—13-flora. Bracteae involucri 8— 10-seriatae,
interiores exterioribus longiores, apicibus pellucidis,
aeneis, cucullatis. Bracteae intimae glabrae, lanceolatae,
6 x 0,8— 1,0 mm. Bracteae aliae ovatae, 1,5— 6,0 x
1.5— 2,0 mm, stereomis integris, ad apices versus cum
areolis conspicue albo-tomentosis (ut in Amphiglossa
tomentosa (Thunb.) Harv.). Receptaculum favosum,
nudum. Flores omnes hermaphroditi, omnes fertiles, circa
7 mm longi. Corolla desuper alba vel lilacina, circa 5,5
mm longa, apicem versus quinquelobata; tubus
cylindricus, 5 mm longus, vertice purpurascens, infime
pallide virescens; lobi deltoidei, 0,5 —0,7 mm longi, acuti,
patentes. Stamina quinque; filamentum 1,30-2,55 x
0,08-0,17 mm; collum filamenti 0,26-0,47 mm longum,
ad basin 0,08—0,11 mm latum, cellulis 9— 11-seriatis; thecae
1,37—2,00 mm longae; appendix apicalis lanceolata,
0,27—0,43 X 0,11—0,17 mm, apice obtuso; caudae
0,40—0,68 mm longae, infra pubescentes (Figura 4); pollen
spinulosum, 21-29 jum diametro, 3-colporatum. Stylus
3.5 - 4,5 x 0,12-0,20 mm, ad apicem versus biramosus,
ramis 0,87-1,22 mm longis, truncatis. Stylopodium
0,19—0,37 mm latum. Nectarium 0,20—0,29 mm altum,
0,25—0,31 mm latum. Ovarium 1,0-1, 8 mm longum,
glabrum, cremeum, quinquecostatum; apex annulo
incrassato. Setae pappi liberae, plumosae, sed ad basin
hamatae, 3,5— 4,5 mm longae.
TYPUS. — Cape Province, 2822 (Glen Lyon): Hay
District, Witsand, some 70 km SW of Postmasburg. Farm
Witsands 250. Approximately 2 km W of Doornaar
homestead. White Kalahari Sand outlier, just W of the
Langeberg. Clump-forming shrub, up to 400 mm tall, and
up to 3,5 m wide. Florets white or pale mauve from above.
Involucral bracts a golden colour. Main stems gnarled,
dark grey, up to 40 mm diam. , buried in sand below. Bark
poorly developed. Leaves with camphor odour when
crushed. Occurs in sparsely grassy dune shrubland, with
Lopholaena cneorifolia and Crotalaria virgultalis, in
locally level place on E slope, near summit of rocky hill,
overlain with white dune sand. Locally common. Alt.
1 230 m (-CB). F. Brusse 5629, 26.11.1989 (PRE, holo.;
AD, B, BAF, BH, BM, BOL, BR, BRI, C, CAN, CANB,
COI, E, EA, G, GH, GRA, HBG, J, K, L, LD, LG, LISU,
LMA, M, MEL, MO, NBG, NH, NSW, NU, O, P, R,
S, SRGH, U, UC, UPS, US, W, WAG, WIND, Z, iso.).
Figure 1.
Clonal shrub up to 400 mm tall and 3,5 m across,
aromatic. Main stems gnarled, dark grey, up to 40 mm
diam., buried in sand below. Secondary branches up to
5 mm thick, bark thin, golden chestnut brown to reddish-
brown to reddish in places, longitudinally fissured,
glabrous or with golden flakes. Tertiary branches robust,
yellow and white-tomentose when young, golden-yellow,
glabrous and peeling when older. Leaves alternate, sessile,
linear, 1—11 x 0,7— 2,2 mm, broadest at base, ascending
but spreading in upper half, margins involute. Upper
surface canaliculate, white-woolly. Lower surface
FIGURE 1. — Pterothrix tecta Brusse, habit. F. Brusse 5629 , holotype.
68
Bothalia 20,1 (1990)
FIGURE 2. — Pterothrix tecta Brusse, flowering twigs at the type locality,
with the tomentose patches on the involucral bracts clearly visible,
F. Brusse 5629.
glabrous, epidermis sloughing-off and folliculate especially
at base; midrib raised particularly at base. Apex obtuse,
mucro very small, inconspicuous. Flower heads solitary,
terminal, sessile, discoid, cylindrical, 7-8 x 2,5 -3,5
mm, 11— 13-flowered. Involucral bracts 8— 10-seriate,
becoming progressively longer inwards, tips galeate
(hooded), pellucid, bronze. Innermost bracts glabrous,
lanceolate, 6 x 0,8 -1,0 mm. Remaining bracts ovate,
1,5— 6,0 x 1,5— 2,0 mm, stereomes entire, with
conspicuous white-tomentose patches at apices (as in
Amphiglossa tomentosa (Thunb.) Harv.). Receptacle
honey-combed, nude. Florets all hermaphrodite and fertile,
about 7 mm long. Corolla white or pale mauve from above,
about 5,5 mm long, five-lobed towards apex; tube cylin-
drical, 5 mm long, purplish above becoming pale greenish
below; lobes deltoid, 0,5— 0,7 mm long, acute, spreading.
Stamens five; filament 1,30-2,55 x 0,08—0,17 mm;
filament collar 0,26—0,47 mm long, 0,08—0,11 mm wide
at base, 7—11 cells across; thecae 1,37—2,00 mm long;
apical appendage lanceolate, 0,27—0,43 x 0,11—0,17 mm,
apex obtuse; tails 0,40—0,68 mm long, pubescent below,
well exceeding filament collar base (Figure 4); pollen
spinulose, 21—29 gm diam., 3-colporate. Style 3,5— 4,5
x 0,12—0,20 mm, two-branched towards apex.
FIGURE 3. — Pterothrix tecta Brusse, close-up of some flower heads
at the type locality, with the conspicuous white tomentose patches
on the involucral bracts clearly visible, F. Brusse 5629.
FIGURE 4. — Pterothrix tecta Brusse, photomicrograph of the pubescent
anther tails, extending well beyond the filament collar base, LAW.
MacDonald 76/43, paratype. Bar = 0,3 mm.
branches 0,87—1,22 mm long, truncate. Stylopodium
0,19—0,37 mm wide. Nectary 0,20—0,29 x 0,25—0,31
mm, 5-buttressed like a Diosma fruit. Ovary 1,0— 1,8 mm
long, glabrous, cream, 5-ribbed, apex with thickened ring.
Pappus bristles free, plumose, but base barbed; 3,5— 4,5
mm long.
The present new species belongs to the genus Pterothrix,
but this genus is closely related to the genus Amphiglossa,
and a problem of generic assignment thus arose. However,
the two genera can be distinguished by the following key:
Heads homogamous, composed entirely of bisexual disc
florets. Anther tails pubescent below, or tipped with well
developed tail antlers Pterothrix DC.
Heads heterogamous, composed of marginal, female ray florets
and bisexual disc florets. Anther tails glabrous or simple,
without well developed tail antlers Amphiglossa DC.
Pterothrix tecta resembles Amphiglossa tomentosa
(Thunb.) Harv. in the almost identical complement of
involucral bracts; all except the innermost row, possess
Bothalia 20,1 (1990)
69
a very conspicuous median patch of white tomentum
(Figures 2 & 3). However, A. tomentosa is a true
Amphiglossa, with a few ray florets present in each head,
and with glabrous anther tails, that extend for only a short
distance beyond the base of the filament collar. P. tecta ,
on the other hand, lacks any female or ray florets, and
the anther tails are pubescent below,, with well developed
tail antlers, that extend well beyond the filament collar base
(Figure 4). A. tomentosa is a plant with wiry twigs,
whereas P. tecta is one with comparatively robust twigs.
The authentic material of Pterothrix flaccida Schltr.
nom. nud. (Hutchinson 1917) has the same conspicuous
tomentose patches on the bracts, has heads that clearly
contain a few ray florets each, and has anther tails that
are glabrous. The material is therefore clearly assignable
to Amphiglossa tomentosa (Thunb.) Harv., and the name
P flaccida Schltr. nom. nud., should not be validated.
Pterothrix tecta is distinct from all the other species in
Pterothrix, by the broader involucral bracts, each with a
FIGURE 5.— Pterothrix spinescens DC., photomicrograph of the
pubescent anther tails, only slightly exceeding the filament collar
base, H.G. Flanagan 1454 (PRE). Bar = 0,3 mm.
conspicuous white-tomentose patch. The other species all
have lanceolate involucral bracts, which are glabrous or
irregularly tomentose. P tecta is also distinct from all other
species of Pterothrix in possessing anther tails that extend
well beyond the filament collar base (Figure 4), whereas
those of all other species only just exceed the base of the
filament collar, exemplified by the widespread and
common P. spinescens DC. (Figure 5).
The following key serves to distinguish all the presently
known narrow-leaved species of Pterothrix — P cymbi-
folia Harv. is the only broad-leaved species known (Harvey
1865):
la Anther tails well exceeding filament collar base. Except for
the innermost row, involucral bracts about 2 mm broad,
each with a conspicuous median white-tomentose patch
(as in Amphiglossa tomentosa ). Florets 11—13 per head
P. tecta Brusse
lb Anther tails only just exceeding filament collar base. Involucral
bracts lanceolate, all about 1 mm broad, glabrous or
irregularly tomentose. Florets less than 10 per head:
2a Stylopodium bulbous, clearly off-set from style. Leaves
fascicled P. perotrichoides (DC.) Harv.
2b Stylopodium not clearly off-set from style, running into style
imperceptibly. Leaves alternate, not fascicled:
3a Plant spiny. Capitula with 4—7 florets each . . P. spinescens DC.
3b 'Plant not spiny. Capitula with 3 florets each . . P. thuja Merxm.
Like P spinescens, P. tecta is aromatic, and gives off
a camphor odour when the leaves are crushed.
Pterothrix tecta grows in clones up to 3,5 m across, but
only grows up to 400 mm above the sand level (Figure
6). The clones may constitute single plants with most of
the large stems buried in sand. The species was not found
in pure sand areas but only in sand fills between sand-
stone outcrops. This may indicate that this plant may be
rooted in rocky soil, with the main stems buried in the
loose white sand, and the aerial shoots visible above the
soil level as in Figure 6. However, no excavation was
carried out to confirm this. The plant was not found in
rocky places without sand, or where the sand fill was not
substantial. The species occurred on the east side of
Witsand, in level sand fills on east slopes near the summits
of these rocky hills covered in white sand. These places
were actually gently sloped in a westerly direction, not
level (Figure 6).
Attempts to find this plant at other sand dune-covered
rocky hills nearby, namely at Prynnsberg (north of Kheis
FIGURE 6. — Pterothrix tecta Brusse, a clone of plants, looking eastward
with the Langeberg in the distance, F. Brusse 5629.
70
Bothalia 20,1 (1990)
on the Orange River) and Donkieberg, on the Farm
Waterford (on the main Griquatown— Groblershoop road)
proved fruitless. These latter dunes were composed of red
sand, unlike that of Witsand. It therefore seems likely that
this new species occurs only at Witsand, some 70 km
south-west of Postmasburg.
CAPE PROVINCE.— 2822 (Glen Lyon): Witsand, Hay. 28°32’S :
22°28’E. On white sand dunes. A common bush with dull yellow/brown
flowers (— CB). I.A.W. MacDonald 76/43 , 26.11.1976 (KMG, PRE).
REFERENCES
HARVEY, W.H. 1865. Compositae. In W.H. Harvey & O.W. Sonder,
Flora capensis 3 : 275 -276. Hodges & Smith, Dublin.
HUTCHINSON, J. 1917. Plants collected in the Percy Sladen Memorial
Expeditions, 1908—1911. Compositae. Annals of the South African
Museum 9: 379.
F. BRUSSE
MS. received: 1989.04.20.
FABACEAE
STUDIES IN THE GENUS LOTONONIS (CROTALARIEAE). 10. L. ESTERHUYSENIANA ,
A NEW SPECIES FROM THE SOUTH-WESTERN CAPE
INTRODUCTION
The taxonomic position of an undescribed species with
a superficial similarity to species of the section Leobordea
(Del.) Benth. is considered in this paper. At first, the
morphology did not seem unusual, but on closer
examination it became clear that the species did not fit
readily into any of the existing sections of Lotononis (DC.)
Eckl. & Zeyh. The significance of the new species
described below is that it indicates a direct relationship
between the L. pentaphylla group [presently part of the
section Lipozygis (E. Mey.) Benth., see Van Wyk 1989]
and the section Leobordea. In terms of appearance
however, it is rather insignificant.
A summary of similarities and differences between the
new species and various groups of Lotononis is given in
Table 1. The comparison shows that it has more in common
with the sections Leobordea and Lipozygis than with Leptis
(Eckl. & Zeyh.) Benth. It differs from Leobordea mainly
in the alternate leaf arrangement and the smaller number
of ovules and seeds. The character states also agree well
with those of Lipozygis (L. pentaphylla group) except
for the structure of the calyx and inflorescence. The
only way to retain the present diagnostic value of the
opposite leaf arrangement (for Leobordea) and the capitate
inflorescence (for the L. pentaphylla group) would be to
exclude the new species from these two groups. The very
small number of ovules and the unusual combination of
other characters are of sufficient diagnostic value to form
a new monotypic section. This seems the most practical
solution.
Lotononis esterhuyseniana B-E. van Wyk, sp. nov. ,
distincta sine affinitatibus manifestis. Similis est L.
platycarpae (Viv.) Pichi-Serm. aliisque speciebus sectionis
Leobordeae, sed ab illis foliis alternis in ramis floriferis
atque ovulis valde paucioribus (1 vel 2, non 5 vel ultra
ut in Leobordea) differt. Etiam similis est L. laticipi
B-E. van Wyk aliisque speciebus gregis L. pentaphyllae
(sectionis Lipozygis), ab illis inflorescentia racemosa (non
capitata), lobo carinale calycis parvo, leguminibus
maioribus compressis atque calyce non inflato non
legumen maturum includente, differt. Etiam similis est L.
microphyllae Harv. (sectionis Leptidis), sed ab ilia habitu
annuo, fabrica calycis, forma longitudineque vexillae et
numero parvo ovulorum differt.
TYPE. — Cape Province, 3219 (Wuppertal): Ceres
District, Stompiesvlei, Swartruggens (in sand near pan,
4000 ft.), 19.11.1961, Esterhuysen 29341 (BOL, holo.; C,
K, MO, iso.). Figure 7.
The species is named after Miss Elsie Esterhuysen of
the Bolus Herbarium, who collected and distributed the
only material known so far. Through her numerous
collections of rare and unusual species. Miss Esterhuysen
has made a very significant contribution to the phyto-
geography and taxonomy of Lotononis.
Prostrate annual up to 0,4 m wide. Branches sparsely
leafy, minutely hirsute. Leaves invariably 3-foliolate,
(5 — )8 — 12( — 15) mm long; petiole as long or longer than
the terminal leaflet; leaflets relatively small, obovate,
(2— )5— 7(— 9) x (1 — )2 — 4( — 5) mm, base cuneate, apex
rounded to truncate, minutely but densely pubescent on
both surfaces. Stipules single at each node, small, lanceo-
late to ovate, up to 3 x + 1,5 mm, minutely pubescent
on both surfaces. Inflorescences in terminal and leaf-
opposed racemes, (3— )6— 12(— 15)-flowered; peduncle
short, (2— )3— 6(— 12) mm long; bracts small and incon-
spicuous, lanceolate, + 1 mm long; bracteoles absent.
Flowers small, 7—8 mm long, yellow; pedicel up to 2 mm
long. Calyx subequally lobed but with the lower lobe much
narrower and slightly shorter than the upper four lobes;
lateral sinuses a little shallower than the upper and lower
ones; lobes narrowly triangular, acute. Standard sub-
orbicular, much shorter than the keel; claw short, ± 1,5
mm long; lamina ±4x3-4 mm, without lobes or cal-
losities, abaxially pubescent along the apex. Wing petals
oblong, much shorter than the keel, distinctly auriculate;
pubescent along the apex; sculpturing in ± 4 rows of inter-
costal lunae, fading into transcostal lamellae towards the
auricle. Keel petals broadly oblong, wider towards the
obtuse apex, only slightly auriculate; claw ± 3 mm long;
lamina 5-6 x ±3 mm, pubescent on most of the upper
half. Androecium long and narrow; anthers dimorphic;
basifixed anthers oblong, almost twice as long as the
small ovoid dorsifixed anthers; carinal anther similar to
dorsifixed anthers. Gynoecium shortly stipitate; pistil very
small, ovoid-oblong, pubescent, with 1 or 2 ovules; style
with the basal part straight, broad and pubescent, the upper
part short, slender, glabrous. Pods very small, ovoid,
± as long as the calyx, 3—3,5 X 2—2,5 mm, shortly stipi-
tate, compressed (not inflated), densely pubescent, inde-
Bothalia 20,1 (1990)
71
FIGURE 7. — Lotononis esterhuyseniana. A, flowering twig, showing the racemose inflorescence and a leaf in adaxial view; B, leaf and stipule
in abaxial view; C, calyx opened out with the upper lobes to the left; D, standard petal; E, wing petal; F, keel petal; G, pistil. HI, H2
& H3, pods: HI, in longitudinal section, showing the seeds; H2, in lateral view; H3, in top view. II, 12 & 13, anthers: II, long basifixed
anther; 12, carinal anther; 13, dorsifixed anther. J1 & J2, bracts; K, seed, showing the almost smooth surface; L, flower in lateral view.
All from Esterhuysen 29341. Scales in mm.
72
Bothalia 20,1 (1990)
TABLE 1. — Similarities and differences between L. esterhuyseniana and various other groups of the genus Lotononis
hiscent (?); upper suture minutely verrucose, 1 or 2-
seeded. Seeds suborbicular, ±1,2 mm in diameter, testa
pale orange-brown, sparsely and minutely tuberculate
(Figure 7).
L. esterhuyseniana is a distinct species with no obvious
affinities. It is similar to L. platycarpa (Viv.) Pichi-Serm.
and other species of the section Leobordea but differs from
these in the alternate arrangement of the leaves on
flowering twigs and also in the much smaller number of
ovules (more than five in section Leobordea). It is also
similar to L. laticeps B-E. van Wyk and other species of
the L. pentaphylla group (section Lipozygis), but differs
from these in the racemose inflorescence (not capitate),
the small carinal lobe of the calyx, the larger and
compressed pods and in the calyx, which is not inflated
and which does not enclose the pod at maturity. It also
resembles L. microphylla Harv. (section Leptis) but differs
from this species in the annual habit, the calyx structure,
the shape and length of the standard petal and the small
number of ovules.
The geographical distribution of L. esterhuyseniana
supports the suggested affinity with the L. pentaphylla
group (presently section Lipozygis) and the section
Leobordea rather than with the section Leptis. The single
known locality is shown in Figure 8.
CAPE. — 3219 (Wuppertal): Ceres District, Stompiesvlei, Swartruggens
(—DC), Esterhuysen 29341 (BOL, holo.; C, K, MO, iso.).
FIGURE 8. — The known geographical distribution of Lotononis
esterhuyseniana.
Bothalia 20,1 (1990)
73
ACKNOWLEDGEMENTS
I thank Dr H. F. Glen (National Botanical Institute,
Pretoria) for the Latin translation and the directors and
staff of the cited herbaria for the loan of specimens. The
taxonomic study of Lotononis is a registered Ph. D. project
at the University of Cape Town.
REFERENCE
VAN WYK, B-E. 1989. Studies in the genus Lotononis (Crotalarieae,
Fabaceae). VIII. A new species of the L. corymbosa group and
notes on the taxonomy of the section Lipozygis. South African
Journal of Botany 55: 528—532.
B-E. VAN WYK*
* Department of Botany, Rand Afrikaans University, P. O. Box 524,
Johannesburg 2000.
MS. received: 1989.04.12.
STUDIES IN THE GENUS LOTONONIS (CROTALARIEAE). 11. A NEW SPECIES OF THE SECTION LEOBORDEA
FROM NORTH-WESTERN NAMIBIA
INTRODUCTION
The section Leobordea (Del.) Benth. is geographically
the most widespread section of Lotononis (DC.) Eckl. &
Zeyh. L. platycarpa (Viv.) Pichi-Serm. occurs throughout
Africa and eastwards to Pakistan but the other species of
the section are all restricted to southern Africa.
Bentham (1843) listed the small subsessile flowers, the
subsessile leaf-opposed inflorescences and the small
carinal lobe of the calyx as diagnostic characters for the
section, but also mentions the dichotomous branches.
Detailed studies of the genus as a whole have shown that
the length of the pedicels, the inflorescence structure and
the calyx structure are rather variable, and that most of
the diagnostic characters of Leobordea also occur
sporadically in other sections. It is suggested here that the
opposite arrangement of leaves on flowering branches is
the only reliable character to distinguish Leobordea from
other sections of Lotononis. In the section Leptis (Eckl.
& Zeyh.) Benth., some leaves may occasionally be
subopposite, but the leaves are not invariably opposite as
in Leobordea. It is important to note that the distinction
only applies to flowering nodes. Basal leaves that are
formed during the vegetative phase are alternate (also in
Leobordea). This is true for all species of Lotononis except
L. lenticula (E. Mey.) Benth. (section Oxydium Benth.)
where the basal nodes (but not the flowering nodes) have
opposite leaves.
The new species described below provides an interesting
example of convergence in Leobordea. Schreiber (1970)
confused the species with the superficially similar L.
stipulosa Bak. f. The many-flowered fascicles of the two
species are remarkably similar in appearance due to the
presence of large leaflike structures around the base of
the inflorescences. In the new species, these are enlarged
bracts and not enlarged stipules as in L. stipulosa.
Lotononis bracteosa B-E. van Wyk, sp. nov. L.
stipulosae Bak. f. valde affinis sed habitu minori, foliis
basalibus oppositis (L. stipulosa foliis basalibus altemis),
foliolis minoribus, stipularum magnitudine formaque, quae
oblongae vel ovatae ad 5 x 3 mm sunt, inflorescentiam
non tegentes (in L. stipulosa late cordatae, plerumque valde
maiores quam 5x3 mm, inflorescentiam tegentes),
differt. Stipulae inflorescentiam subtendentes costam
unicam, dum illae L. stipulosae venas plures e basi habent.
Etiam bracteis 2—4 mm latis, late ellipticis vel obovatis
(in L. stipulosa infra 2 mm latis, lanceolatis differt).
L. stipulosa Bak. f. sensu Schreib. : 85 (1970).
TYPE. — Namibia, Outjo District, mountains 14 miles
[22,4 km] east of Torra Bay, Giess, Volk & Bleissner 6198
(PRE, holo. ; M, PRE, WIND, iso.).
Prostrate or procumbent herbaceous annual, often small
and short-lived. All mature parts densely pubescent.
Leaves trifoliolate, densely pubescent on both surfaces.
Leaflets elliptic to oblanceolate, (4 — )6 — 11( — 16) x
(l,5-)2-4(-5) mm; base cuneate; apex acute. Stipules
broadly oblong to ovate, up to 5 x 3 mm. Inflorescences
sessile, umbellate, (2— )5— 8(— 12)-flowered; bracts
conspicuous, broadly elliptic to broadly obovate, 3— 4 x
2—4 mm; apex acute to obtuse, mucronate; base cuneate
to cordate. Flowers subsessile, 6—8 mm long. Calyx not
inflated, densely pubescent, with the upper and lateral lobe
on either side fused higher up in pairs, the lower lobe
slightly narrower and shorter; lobes usually broadly
acuminate. Standard ovate to oblong, usually shorter than
the keel, densely pubescent. Wing petals oblong, not much
shorter than the keel, pubescent along the lower edge of
the lamina; apex obtuse to acute; sculpturing upper basal
and upper left central, in 4 rows of intercostal lunae, fading
into thin transcostal lamellae towards the auricle. Keel
petals half oblong-elliptic to oblong, densely pubescent;
apex obtuse. Pods sessile, scarcely longer than the
persistent calyx, broadly obovoid to broadly oblong, only
slightly inflated, upper suture ± smooth, ± 8-seeded.
Seeds suborbicular, testa minutely and densely tuberculate
(Figure 9).
Closely related to L. stipulosa Bak. f. but differs in the
smaller habit, the smaller leaflets and in the size and shape
of the stipules, which are oblong to ovate, up to 5 x 3
mm and not covering the inflorescence (broadly cordate,
usually much more than 5x3 mm and covering the
inflorescences in L. stipulosa). The stipules subtending
the inflorescences have a single midrib, whereas those of
L. stipulosa have several veins from the base (Figure 9).
It also differs in the 2-4 mm wide, broadly elliptic to
obovate bracts (less than 2 mm wide and lanceolate in L.
stipulosa). The bracts are visible and conspicuous, not
hidden by the large and foliaceous stipules as in L.
stipulosa (Figure 9). There is no vegetative phase as in
L. stipulosa, where the basal parts of the branches are
without inflorescences and the basal leaves alternate. In
L. bracteosa, inflorescences are formed at the first nodes
and the basal leaves are opposite.
74
Bothalia 20,1 (1990)
FIGURE 9. — Lotononis bracteosa. A, habit. B1 & B2, leaves: Bl, abaxial view; B2, adaxial view. C, calyx opened out with the upper lobes
to the left. D1 & D2, standard petals: Dl, abaxial view showing vestiture; D2, adaxial view. E, wing petal; F, keel petal. G1 & G2, pistils:
Gl, from young flower; G2, from older flower. HI & H2, pods: HI, top view; H2, pod after dehiscence as viewed from inside, showing
funicles. II, 12 & 13, anthers: 11, small dorsifixed anther; 12, carinal anther; 13, long basifixed anther. J, androecium; K1 to K4, bracts,
showing the broadly ovate shape; L, seed in lateral view showing the densely tuberculate surface. Lotononis stipulosa. M, leaf from flowering
node showing the foliaceous stipules; N1 to N4, bracts, showing the lanceolate shape; O, calyx opened out with the upper lobes to the
left. All from Giess, Volk & Bleissner 5772 except A from Giess, Volk & Bleissner 6198, Bl, D2, Gl, G2, T1 & T2 from Kers 1513; M,
N4 & O from Miller 7748; N1 from Davies 715; N2 from Eyles 2020; N3 from Biegel 2923. Scales in mm.
Bothalia 20,1 (1990)
75
FIGURE 10. — The known geographical distribution of Lotononis
bracteosa , ■; and L. stipulosa, #.
Ihlenfeldt, De Winter & Hardy 3229 (M, PRE); 27 miles [43,2 km] south-
east of Torra Bay, at Koichab River (—AD), Nordenstam 3795 (S); Farm
Wereld-End, 5 miles [8 km] west of house (-BB), Giess, Volk & Bleissner
6208 (M, WIND); Farm Wereld-End, on Petrified Forest to Torra Bay
Road, at the coast (-BB), Kers 1513 (WIND); Middle-Huab (— DB),
Milller & Loutit 1153 (WIND). 2014 (Welwitschia): Damaraland, just
south of Twyfelfontein (— CA), Craven 989 (WIND); Welwitschia, Gai-as
(— CC), Milller & Loutit 1248 (WIND). 2214 (Swakopmund): Swakop-
mund District, 22 miles [35,2 km] east of Hentiesbay (—BA), Giess,
Volk & Bleissner 5772 (M, WIND). 2215 (Trekkopje): Swakop River,
Welwitschia flat, along the track from Farm Nordenberg to Swakop River
at Tsavischab (— CA), Kers 8 (WIND). 2315 (Rostock): Walvis Bay
District, Walvis Bay to Kuiseb River to Gamsberg Road, 20 miles [32
km] before the Gorob Mine track (—BA), Kers 1287 (WIND); Namib
Desert Park, Hotsas (-BA), Milller 224 (M, WIND).
ACKNOWLEDGEMENTS
Thanks are due to Dr H. F. Glen (National Botanical
Institute, Pretoria) for the Latin translation and the
directors and staff of the cited herbaria for the loan of
specimens. The taxonomic study of Lotononis is a
registered Ph. D. project at the University of Cape Town.
L. bracteosa is known only from the north-western parts
of Namibia, where it appears to be quite common. Figure
10 shows that L. bracteosa and L. stipulosa are geo-
graphically isolated and that the latter does not occur in
Namibia.
SWA/NAMIBIA. — 1812 (Sanitatas): Kaokoveld, river course 6 miles
[9,6 km] south of Orupembe (—BA), Giess & Leippert 7524 (M, NBG,
PRE, WIND). 2013 (Unjab Mouth): Outjo District, mountain 14 miles
[22,4 km] east of Torra Bay (—AD), Giess, Volk & Bleissner 6198 (PRE,
holo.; M, PRE, WIND, iso.); 23 km south-east of Torra Bay (—AD),
Giess 8020A (WIND); ± 13 miles [20,8 km] east of Torra Bay (—AD),
REFERENCES
BENTHAM, G. 1843. Enumeration of Leguminosae, indigenous to
southern Asia, and central and southern Africa. Hooker's London
Journal of Botany 2: 594—613.
SCHREIBER, A. (1970). Fabaceae. In H. Merxmiiller, Prodromus einer
Flora von Siidwestafrika 60. Cramer, Lehre.
B-E. VAN WYK*
* Department of Botany, Rand Afrikaans University, P. O. Box 524,
Johannesburg 2000.
MS. received: 1989.04.12.
STUDIES IN THE GENUS LOTONONIS (CROTALARIEAE). 14. THREE NEW SPECIES OF THE SECTIONS
TEUNA AND POLYLOBIUM
INTRODUCTION
As part of preparatory studies for a synoptic revision
of Lotononis (DC.) Ecld. & Zeyh., several new species
have been described in previous parts of this series. The
section Telina (E. Mey.) Benth. and the L. angolensis
group of the section Polylobium (Eckl. & Zeyh.) Benth.
were treated in parts 2 and 3 respectively (Van Wyk 1988,
1989). Recent studies however, have shown the presence
of another two new species of Telina and one new species
of the L. angolensis group. These are described below.
Section Telina (E. Mey.) Benth.
Lotononis filiformis B-E. van Wyk, sp. nov., L.
azureae Eckl. & Zeyh. valde affinis sed habitu longissimo
gracili sparsim ramoso, foliolis gracilibus linearibus, lobis
calycis oblongis (non triangularibus), fructu minore
angustiori et praesertim pube minute strigillosa ramulorum
foliorum calycis fructusque (sparsim pilosa in L. azurea)
differt.
TYPE. — Cape, 3321 (Ladismith): northern slopes of
Outeniqua Mountains, between Farms Bonniedale and
Woeska (-DD), 14.10.1988, Vlok 2030 (PRE, holo.;
JRAU, K, MO, NBG, SAAS, STE, iso.).
Very sparse perennial herb, up to 1 m wide. Branches
long, slender, sparsely leafy, glabrescent, reddish-brown;
twigs minutely strigillose. Leaves digitately trifoliolate;
petiole (4— )7— 14(-22) mm long; leaflets linear, thick in
texture, (4— )8— 22(— 26) x (0,5— )1— 1,5(— 2) mm, incon-
spicuously strigillose on both surfaces. Stipules
consistently present, single or paired at each node,
narrowly lanceolate, (2 — )4 — 8(— 10) mm long. Inflores-
cences leaf-opposed, slender, long-pedunculate, 25-120
mm long, invariably single-flowered; bracts small, up to
1,5 mm long; bracteoles absent. Flowers relatively large,
8-12 mm long, blue; pedicel 3-8 mm long. Calyx 8-10
mm long, subequally lobed, minutely strigillose; lobes
narrowly oblong. Standard broadly ovate to suborbicular,
7—12 mm long, minutely pubescent on the middle part of
the dorsal surface, deep blue with yellow at the base. Wing
petals oblong, longer than the ke€l; apex rounded;
sculpturing in 2-3 rows of mostly transcostal lunae and
lamellae. Keel petals semicircular, obtuse, auriculate and
pocketed near the base. Anthers dimorphic. Pistil 9—11
mm long; ovary linear, 6—9 mm long, pubescent; style
erect. Pods narrowly oblong, 14—20 x 3-4 mm, laterally
much inflated, ± 5-8-seeded, minutely strigillose, low-
er suture sunken, upper suture verrucose, dehiscent. Seed
suborbicular, ± 2 mm in diameter; testa reddish brown,
dark brown or almost black, densely tuberculate; funicles
up to 1,2 mm long (Figure 11).
76
Bothalia 20,1 (1990)
FIGURE 11. — Lotononis filiformis. A, flower in lateral view. B1 & B2, leaves, showing the stipules and strigillose vestiture: Bl, abaxial view;
B2, adaxial 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. HI & H2, fruit, showing the densely strigillose vestiture and verrucose upper suture: HI, lateral view; H2, top view. I,
androecium. J1 & J2, seeds: Jl, hilar view; J2, lateral view. Lotononis azurea var. lanceolata. K, flower in lateral view; L, leaf in adaxial
view; Ml & M2, fruit in lateral and top view. All from Van Wyk 2857 except HI, H2, Jl & J2 from Vlok 2030 and K, L, and M from
Acocks 13713. Scales in mm.
Bothalia 20,1 (1990)
77
L. filiformis is very closely related to L. azurea Eckl.
& Zeyh. but differs in the very long, slender and sparsely
branched habit, the slender linear leaflets, the oblong (not
triangular) calyx lobes, the smaller narrower fruit and
particularly in the minutely strigillose vestiture of the
twigs, leaves, calyx and fruit (not sparsely pilose as in L.
azurea). It is also geographically isolated from the latter
and has a more western distribution, known only from the
northern slopes of the Outeniqua Mountains (Figure 12).
The narrow calyx lobes are very similar to those of L.
azurea var. lanceolata Harv. (based on Ononis villosa
Thunb.), with which L. filiformis has previously been
confused (Figure 11). I have examined the Thunberg type
specimen of Ononis villosa, but the long, sparse and
spreading hair covering of the latter is very different from
the dense and minutely strigillose vestiture of L. filiformis.
I am much indebted to Mr J. H. J. Vlok of Saasveld,
George for the opportunity to study the species in situ.
The almost climbing habit is quite unlike that of any other
species of Lotononis — the long, slender branches are
supported by the surrounding vegetation and when not in
flower, the plants are almost invisible.
CAPE.— 3321 (Ladismith): Mossel Bay Division, Vryers Berg (—DC),
Muir 2042 (BOL); northern slopes of Outeniqua Mountains, between
Farms Bonniedale and Woeska (-DD), Vlok 2030 (PRE, holo.; JRAU,
K, MO, NBG, SAAS, STE, iso.). 3322 (Oudtshoom): Oudtshoorn
District, Moeras River between Oudtshoom and Robinson’s Pass (— CC),
Bolus 11767 (BOL, GRA); lower northern slopes of the Outeniqua
Mountains, near Zebrafontein in Doomrivier Wilderness area (— CC),
Van Wyk 285 7 (JRAU, K, MO, NBG, PRE, STE); 4,2 miles [6,72 km]
WNW of Camfer Station (-CD), Acocks 23250 (STE).
Lotononis complanata B-E. van Wyk, sp. nov. , L.
variae (E. Mey.) Benth. similis sed habitu foliisque
minore, apicibus foliolorum recurvatis, stipulis singu-
laribus (plerumque geminis in quoque nodo in L. varia),
pube foliorum plus dense strigillosa (foliis ± glabris
in L. varia), inflorescentiis semper unifloris (saepe 2- vel
pluri-floris in L. varia), floribus minoris et praesertim
fructu breve oblongo compresso indehiscenti (longiore
valde inflato dehiscenti in L. varia et speciebus omniis
aliis sectionis Telinae). Fructus etiam multiseminales
ad 30 seminibus (± 6— 12-seminales in speciebus aliis).
FIGURE 12. — The known geographical distribution of Lotononis
filiformis, O; L. complanata, and L. subulata, A.
funiculis sunt usque ad 4 mm longis (usque ad 2 mm in
speciebus aliis).
TYPE. — Cape, 3319 (Worcester): Riebeeck-Wes, in
Elandsberg Nature Reserve (-AC), 15.06.1988, Vlok 1941
(PRE, holo.; K, NBG, iso.).
Dwarf suffrutescent perennial up to 0,1 m high and 0,2
m wide. Branches slender, procumbent from a thick
central rootstock; young twigs densely strigillose. Leaves
digitately trifoliolate, relatively small, sparsely pubescent,
variable in size, those on the main axis with long and
slender petioles, those of lateral twigs with the petiole
(4 — )6 — 15( — 22) mm long; leaflets narrowly to broadly
oblanceolate, variable in length, (2 — )4 — 10( — 15) x
(1 — )2 — 3( — 3,5) mm, adaxial surface glabrous, abaxial
surface minutely pubescent; apex acute, recurved. Stipules
invariably single at each node, small, narrowly lanceolate,
up to 3 mm long. Inflorescences subterminal or leaf-
opposed on short lateral branches, invariably single-
flowered; peduncle slender, variable in length, (16-45)
mm long; bract small, oblong, up to 1 mm long; bracteoles
absent. Flowers relatively small, 8—12 mm long, blue;
pedicel short, 1—2 mm long. Calyx 5—7 mm long,
subequally lobed, minutely pubescent; lobes narrowly
triangular. Standard large, suborbicular, 7-11 mm long,
with a line of minute hairs dorsally along the middle. Wing
petals longer than the keel, oblanceolate; apex obliquely
obtuse; sculpturing in 3—4 rows of mostly intercostal lunae
and lamellae. Keel petals semicircular, somewhat acute,
auriculate and pocketed near base. Anthers dimorphic.
Pistil short; ovary oblong, ± 7 mm long, minutely
pubescent; style short, erect. Pods very broadly oblong,
10—21 x 5— 7(— 8) mm, compressed, indehiscent, minute-
ly pubescent, apex broad and rounded, 6-30-seeded,
upper suture very conspicuously verrucose. Seed kidney-
shaped, 1,5 mm in diameter, on very long funicles; funicles
up to 4 mm long; testa brown, densely tuberculate (Figure
13).
This species is similar to L. varia (E. Mey.) Benth. but
differs in the smaller habit and leaves, the recurved leaflet
apices, the single stipules (usually paired at each node in
L. varia), the more densely strigillose vestiture of the
leaves (± glabrous in L. varia), the invariably single-
flowered inflorescences (often 2- or more-flowered in L.
varia), the smaller flowers and particularly in the shortly
oblong, compressed and indehiscent fruit (longer, laterally
much inflated and dehiscent in L. varia and in all other
species of the section Telina). Differences between the two
species are shown in Figure 13. The shape of the pods
[complanatus = flattened out] is very unusual for the
section Telina and the tooth-like projections along the
upper suture are more strongly developed than in most
if not all other species of Lotononis. The large number
of seeds per pod and the very long funicles are also unique
features not found in other species of the section Telina.
Only a single collection (without precise locality details)
was known prior to the discovery of the species in the
Elandsberg Nature Reserve (Figure 12). This discovery
is another example of the valuable contributions Mr J. H.
J. Vlok has made to the phytogeography of rare Cape
78
Bothalia 20,1 (1990)
FIGURE 13. — Lotononis complanata. A, flower in lateral view. B1 & B2, leaves, showing the slightly recurved leaflet apices and strigillose
vestiture: Bl, leaf from central branch in adaxial view; B2, leaf from lateral branch in 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. H1-H4, fruit, showing the distinctive shape,
densely strigillose vestiture and verrucose upper suture: HI & H2, lateral view; H3, top view; H4, transverse section. I, androecium.
J1 & J2, seeds: Jl, hilar view; J2, lateral view. Lotononis varia. K, flowers in lateral view; L, leaf in abaxial view; M, stipule. N1-N3,
fruit: Nl, lateral view; N2, top view; N3, transverse section. All from Vlok 1941 except H1-H4, Jl & J2 from Van Wyk 2879 , K from
Esterhuysen 35558, L, M, N1-N3 from Barker 7164. Scales in mm.
Bothalia 20,1 (1990)
79
legumes. L. complanata is known only from one
population and it may be worthwhile to assess the
population structure from time to time.
CAPE. — 3319 (Worcester): Riebeeck-Wes, in Elandsberg Nature
Reserve (-AC), 15.06.1988, Vlok 1941 (PRE, holo.; K, NBG, SAAS,
iso.), 07.10.1988, Van Wyk 2879 (BOL, GRA, JRAU, K, MO, NBG, PRE,
S, SAAS, STE). Without precise locality, Bowie s.n. (BM, K).
plicato in L. bainesii ) differt. A L. listii lobis calycis longis
angustis (late triangularibus in L. listii), inflorescentiis
usque ad 4-floris (rare minus quam 6-floris in L. listii)
et ramulis foliis calyce fructuque distincte pubescentibus
(ramulis foliis calyce fructuque ± glabris in L. listii)
differt. A L. marlothii habitu floresque valde maiore, et
fructu valde maiore plicato pubescente (fructu parvo rec-
to glabrescenti in L. marlothii) differt.
Section Polylobium (Eckl. & Zeyh.) Benth. (L. angolensis
group)
Lotononis subulata B-E. van Wyk, sp. nov., L.
bainesii Bak. f., L. listii Polhill et L. marlothii Engl, valde
affinis. A L. bainesii lobis calycis longis angustis (non late
triangularibus), racemis sparsim paucifloris (non subum-
bellate multifloris) et fructu semper plicato (rare tantum
TYPE. — Transvaal, 2627 (Potchefstroom): Parys, near
bridge over Vaal River on Potchefstroom Road (-CD),
01.05.1989, B-E. van Wyk 2884 (PRE, holo.; JRAU, K,
MO, S, iso.).
Prostrate herbaceous perennial, up to 1 m in diameter.
Branches slender, prostrate, spreading from a woody
caudex, often rooting at the nodes to form dense mats,
FIGURE 14. — Lotononis subulata.
A, flowering branch, showing
the inflorescence structure,
leaves and paired stipules; B,
pedicel with bracteoles; 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. HI & H2, fruit, showing
the distinctive shape and
vestiture: HI, lateral view; H2,
top view. 11—13, anthers: II,
basifixed anther; 12, carinal
anther; 13, dorsifixed anther. J,
seed in lateral view (slightly
immature). K, L & M, calyces
and fruit of related species
(note differences in the shape
of the calyx lobes and fruit):
K, Lotononis listii ; L, Lotono-
nis marlothii', M, Lotononis
bainesii. A-G, 11—13 from
Krynauw 61', HI, H2 & J from
Louw 1691. Scales in mm.
80
Bothalia 20,1 (1990)
thinly and minutely pubescent. Leaves digitately
trifoliolate, very variable in size and shape, adaxially
glabrous, abaxially thinly and minutely pubescent, at
length glabrescent, somewhat fleshy; petiole slender,
(5 — )7 — 24( — 60) mm long; leaflets variable in size and
shape, linear, narrowly elliptic to oblanceolate, the
terminal one (4 — )10 — 28( — 36) x (1 — )2 — 5( — 7) mm, the
lateral ones similar but smaller. Stipules up to 5 x 3 mm,
dimorphic (the one large and foliaceous, the other similar
or more often much smaller), ovate to narrowly lanceolate;
base cordate; apex acute; minutely pubescent. Inflores-
cences leaf-opposed or subterminal on lateral branches,
slender, 20—120 mm long, sparsely 1-4-flowered; bract
linear to narrowly oblanceolate, 2—3 mm long; bracteoles
linear, ± 1 mm long. Flowers 8—10 mm long, yellow.
Calyx 4—6 mm long, with the upper and lateral lobe on
either side fused higher up in pairs, minutely but distinctly
pubescent; lobes very long and narrow, the free part up
to 3 mm long. Standard orbicular, ± as long as the keel.
Wing petals oblong, only slightly shorter than the keel.
Keel petals elliptic; apex obtuse. Anthers dimorphic. Pistil
12—14 mm long; ovary oblong-linear, 8—10 mm long,
pubescent; style short. Pods 10—15 X 2— 4 mm, invariably
folded like a concertina, ± 12-seeded, upper suture +
smooth, indehiscent, distinctly pubescent at maturity.
Seeds small, ± 1 mm in diameter, testa nearly smooth
(Figure 14).
L. subulata is closely related to L. bainesii Bak. f., L.
listii Polhill and L. marlothii Engl, and may be confused
with these species, particularly when mature fruit are not
available. As shown in Figure 14, however, it can easily
be distinguished from these species by the shape and size
of the calyx alone. The long and narrow calyx lobes have
suggested the specific epithet. The inflorescence structure
and the shape and vestiture of the fruit are also useful
diagnostic characters. L. subulata differs from L. bainesii
in the shape of the calyx lobes (not broadly triangular),
the sparsely few-flowered racemes (not subumbellately
many-flowered) and in the consistently plicate fruit (only
rarely plicate in L. bainesii). It can be distinguished from
L. listii also by the shape of the calyx lobes, the fewer
(up to 4) flowers per inflorescence (rarely less than
6-flowered in L. listii) and in the distincdy pubescent twigs,
leaves, calyx and fruit (twigs, leaves, calyx and fruit +
glabrous in L. listii). From L. marlothii it differs in the
much larger habit and flowers and in the much larger,
plicate and pubescent fruit (fruit small, straight and
glabrescent in L. marlothii).
Compared to its close relatives, L. subulata has a much
more restricted distribution and has been recorded only
from the banks of the Vaal River (south-western Transvaal,
eastern Orange Free State and the northern Cape Province,
Figure 12). The robust habit and obvious similarities with
L. bainesii suggest that it may be worthwhile to investigate
the agronomic potential of the new species. Despite the
similarity between L. subulata and L. listii and the
occurrence of both species at some localities, Wilman
(1946: 52) recognized L. subulata as a distinct species.
TRANSVAAL. — 2627 (Potchefstroom): Potchefstroom District,
Scandinavia Drift, bank of Vaal River (-CC), 06.03.1948, Louw 1691
(PRE), 25.04.1979, Krynauw 61 (PRE); Schoemansdrift road, near turn-off
to Venterskroon (—CD), 20.02.1979, Ubbink 865 (PRE); Parys (—CD),
04.1907, Potts 553 (BLFU); Parys, near bridge over Vaal River on
Potchefstroom Road (—CD), 01.05.1989, B-E. van Wyk 2884 (PRE, holo.;
JRAU, K, MO, S, iso.). 2724 (Taung): Intemierungslager Andalusia
(— DD), 28.12.1941, Giess 103 (M), 05.1942, Volk 228 (WIND).
CAPE.— 2824 (Kimberley): bank of Vaal River at Warrenton (-BB),
13.10.1936, Acocks & Hafstrom 1279 (KMG, PRE); Schmidt’s Drift, bank
of Vaal River (-CA), 03.1935, Wilman s.n. sub KMG 3294 (BOL, KMG);
Riverton (—DA), 10.1917, Wilman s.n. sub BOL 15660 (BOL).
ACKNOWLEDGEMENTS
I wish to thank Mr J. H. J. Vlok (Saasveld Forestry
Research Centre, George) for his keen interest in the
Lotononis project and for the valuable material of rare
species. The directors and staff of the cited herbaria are
thanked for the loan of specimens and Dr H. F. Glen for
kindly translating the diagnoses. The taxonomic study of
Lotononis is a registered Ph. D. project at the University
of Cape Town.
REFERENCES
VAN WYK, B-E. 1988. Studies in the genus Lotononis (Crotalarieae,
Fabaceae). III. A new species of the L. angolensis group (section
Polylobium) from the northern Cape Province. South African
Journal of Botany 54: 628—630.
VAN WYK, B-E. 1989. Studies in the genus Lotononis (Crotalarieae,
Fabaceae). 2. Three new species of the section Telina from the
Cape Province. Bothalia 19: 1—5.
WILMAN, A. 1946. Preliminary checklist of the flowering plants and
ferns of Griqualand West : 52. Deighton Bell, Cambridge.
B-E. VAN WYK*
* Department of Botany, Rand Afrikaans University, P. O. Box 524,
Johannesburg 2000.
MS. received: 1989.06.06.
LILIACEAE/ASPHODELACEAE
THE CORRECT AUTHOR CITATIONS OF ALOE BOW1EA AND A. MYR1ACANTHA (ALOOIDEAE)
INTRODUCTION
Available botanical indexes and catalogues indicate a
discrepancy regarding the authorship of the names Aloe
bowiea and A. myriacantha (Table 2). These two species
were originally described by Haworth (1824, 1827), but
in Roemer & Schultes, Systema vegetabilium, both were
transferred to Aloe (Schultes & Schultes 1829).
Monotypic generic status was later suggested for A.
bowiea as Chamaealoe africana (Haw.) Berger (1905)
whereas Stapf (1933) transferred A. myriacantha to
Leptaloe. Recently Obermeyer (1973) and Smith (1983)
suggested that C. africana should best be regarded as a
synonym of Aloe bowiea. Reynolds (1947) sank Leptaloe
Stapf under Aloe and included A. myriacantha in the
section Graminialoe. The aim of this paper is to establish
Bothalia 20,1 (1990)
81
TABLE 2. — Comparison of author citations given by a selection of botanical publications for Aloe bowiea and A. myriacantha , respectively
* Aloe bowiea incorrectly listed as an orthographic variant, viz. Aloe bourea.
** A. bowiea incorrectly listed as an orthographic variant, viz. Aloe bowieae.
the correct author citations of Aloe bowiea and A.
myriacantha, respectively.
DISCUSSION
After Linnaeus’s death in 1778, the botanical part of the
editions of the Systema naturae was frequently published
in amended form as Systema vegetabilium (Stafleu &
Cowan 1983). The sixteenth edition of Systema vegeta-
bilium was published in seven volumes which appeared
in print between 1817 and 1830. Volumes 1-4 were
authored by Johann Jakob Roemer (1763—1819) and Josef
August Schultes (1773-1831); volumes five and six by J.A.
Schultes, except for Umbelliferae (6: 315-628) which is
by K.RJ. Sprengel, and special entries which are by other
authors; and volume seven by J.A. Schultes and Julius
Herman Schultes (1804-1840).
Although Roemer died in 1819, Stafleu & Cowan (1983)
justifiably suggest that citation of edition 16 (volumes 1-7)
of Systema vegetabilium should be to ‘Roemer and
Schultes, Syst. veg'. It is, however, clear that Roemer could
not have contributed to nomenclatural or taxonomic
changes effected in volume seven of Syst. veg. since it was
published 10 years after his death. Furthermore, on the
title page of volume 7,1, the authors are clearly given as
Jos. Augusto Schultes and Jul. Herm. Schultes, Roemer
not being mentioned. The preface of this part is also
attributed to J.A. and J.H. Schultes only.
However, it is insufficient to refer to J.A. Schultes and
J.H. Schultes only when citing authority for taxonomic
and nomenclatural changes made in volume seven of Syst.
veg. J.A. Schultes had two sons who had identical Christian
names, namely Julius Herman (Stafleu & Cowan 1985)
and the use of J.H. Schultes or Schultes fil. only as co-
author along with J.A. Schultes is therefore ambiguous
and incorrect. These two brothers can be distinguished
by birth dates, J.H. Schultes I having lived from 1804 to
1840 and J.H. Schultes II from 1820 to 1887. J.A. Schultes,
J.H. Schultes (1804-1840) and J.H. Schultes (1820-1887)
are abbreviated as Schult., J.H. Schult. and J.H. Schult.
II, respectively (Stafleu & Cowan 1985). Since J.H.
Schult. II was only nine years old when Syst. veg. 7,1 was
published, it is clear that Schult. and J.H. Schult. should
be held responsible for new combinations and nomina nova
which were published in volume seven of Syst. veg.
Part one of the latter volume included the treatment of
Linnaeus’s Class VI Hexandria Monogynia with Aloe
sensu Linnaeus (1753) being monographed on pp. 631-715.
The name Aloe bowiea was published here. This species
was originally described under the name Bowiea africana
by Haworth (1824, 1827). The specific epithet africana
had, however, previously been validly published for a tall-
stemmed species of Aloe (Miller 1768). For this reason,
B. africana had to be renamed. B. myriacantha Haw., the
only other species described in Bowiea Haw., was also
transferred to Aloe in Syst. veg. 7,1. Since the specific
epithet myriacantha had not previously been used in Aloe,
it was correctly retained. Aloe bowiea should therefore be
ascribed to Schult. & J.H. Schult. only. In the case of Aloe
myriacantha the author of the basionym, namely Haworth,
should be cited in parentheses followed by Schult. & J.H.
Schult., the authors who effected the transfer (Greuter et
al. 1988, Article 49).
From the above discussion, it is clear that in all the
publications listed in Table 2 the author citations of Aloe
bowiea and A. myriacantha are listed incorrectly. These
two species should be cited as follows: Aloe bowiea Schult.
& J.H. Schult.; Aloe myriacantha (Haw.) Schult. & J.H.
Schult.
ACKNOWLEDGEMENT
I am indebted to Prof. A.E. van Wyk for critically
reading the manuscript.
REFERENCES
BAKER, J.G. 1880. A synopsis of Aloineae and Yuccoideae. Journal
of the Linnean Society, Botany 18: 148-241.
BAKER, J.G. 1896. Aloe. In W.T. Thiselton-Dyer, Flora capensis 6,2:
302—329. Reeve, London.
BENTHAM, G. & HOOKER, J.D. 1883. Genera plantarum 3,2 : 776.
Reeve, London.
82
Bothalia 20,1 (1990)
BERGER, A. 1905. Uber die systematische Gliederung der Gattung Aloe.
Botanische Jahrbiicher filr Systematik, Pflanzengeschichte und
Pflanzengeographie 36: 42—68. Engelmann, Leipzig.
BERGER, A. 1908. Liliaceae-Asphodeloideae-Aloineae. In A. Engler,
Das Pflanzenreich 4.38.3.2 (Heft 33): 120, 166. Engelmann,
Leipzig.
BOND, P. & GOLDBLATT, P. 1984. Plants of the Cape flora. A
descriptive catalogue. Journal of South African Botany
Supplementary Volume 13: 1—455.
BORNMAN, H. & HARDY, D.S. 1971. Aloes of the South African veld.
Voortrekkerpers, Johannesburg.
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.
GREUTER, W. et al. 1988. International Code of Botanical Nomen-
clature. Regnum Vegetabile 118.
HAWORTH, A.H. 1824. Decas secunda novarum plantarum
succulentarum. Philosophical Magazine 64: 298—302.
HAWORTH, A.H. 1827. Description of new succulent plants.
Philosophical Magazine 67: 120—126.
JACKSON, B.D. 1895. Index Kewensis 1. Clarendon Press, Oxford.
JACOBSEN, H. 1954. Handbuch der sukkulenten Pflanzen, Band I,
Abromeitiella bis Euphorbia. Gustav Fischer Verlag, Jena.
JACOBSEN, H. 1977. Lexicon of succulent plants, 2nd edn. Blandford
Press, Poole.
JEPPE, B. 1977. South African aloes, 2nd edn, 2nd impr. Purnell, Cape
Town.
KUNTH, C.S. 1843. Enumeratio plantarum. Part 4. Enumeratio
Asphodelearum. Sumptibus J.G. Cottae, Stuttgart and Tubingen.
LINNAEUS, C. 1753. Species plantarum, Vol. 1, 1st edn. Impensis
Laurentii Salvii, Stockholm.
MILLER, P. 1768. The gardener’s dictionary, 8th edn. Rivington,
London.
OBERMEYER, A. A. 1973. Aloe, Chamaealoe, Haworthia, Astroloba,
Poellnitzia and Chortolirion (Liliaceae). Bothalia 11: 119.
REYNOLDS, G.W. 1947. Genus Leptaloe Stapf. Restoration to Aloe Linn.
Journal of South African Botany 13: 99-105.
REYNOLDS, G.W. 1950. The aloes of South Africa. The Trustees of
the Aloes of South Africa Book Fund, Johannesburg.
REYNOLDS, G.W. 1954. The aloes ofNyasaland. The Nyasaland Society
and African Book Centre of Nyasaland, Blantyre.
REYNOLDS, G.W. 1966. The aloes of tropical Africa and Madagascar.
The Trustees, The Aloe Book Fund, Mbabane.
REYNOLDS, G.W. 1982. The aloes of South Africa, 4th edn. Balkema,
Cape Town.
SALM-DYCK, J.M.F.A.H.I. 1834. Hortus dyckensis, ou. Catalogue des
plantes cultivees dans les jardins de Dyck. Chez Amz, Diisseldorf.
SALM-DYCK, J.M.F.A.H.I. 1836. Monographia generum Aloes et
Mesembryanthemi, Aloe bowiea. Fasc. 1, fig. 24 (Sect. 14. fig.
1). Diisseldorf.
SCHULTES, J.A. & SCHULTES, J.H. 1829. Classis VI, Hexandria
Monogynia Genera, 1417. Aloe. In J.J. Roemer & J.A. Schultes,
Systema vegetabilium 7,1: 631—715. Sumptibus J.G. Cottae,
Stuttgardtiae.
SMITH, G.F. 1983. Die taksonomiese status van Aloe bowiea Roem.
& Schult. f. Unpublished B.Sc. project. University of Port
Elizabeth.
STAFLEU, F. A. & COWAN, R.S. 1983. Taxonomic literature 4: P— Sak,
2nd edn. Bohn, Scheltema and Holkema, Utrecht. ( Regnum
Vegetabile 110).
STAFLEU, F.A. & COWAN, R.S. 1985. Taxonomic literature 5: Sal— Ste,
2nd edn. Bohn, Scheltema and Holkema, Utrecht. ( Regnum
Vegetabile 112).
STAPF, O. 1933. Leptaloe albida. Botanical Magazine 156: t.9300.
G.F. SMITH*
♦Department of Plant Sciences, Potchefstroom University for C.H.E.,
Potchefstroom 2520.
MS. received: 1989.06.29.
POACEAE
TWO NEW SPECIES OF STIPAGROSTIS (ARISTIDEAE) FROM THE DUNE-NAMIB DESERT, NAMIBIA
Stipagrostis seelyae De Winter, sp. nov.
Stipagrostis seelyae De Winter, sp. nov. , S. sabulicolae
(Sect. Stipagrosti ) affinis, sed inter alia inflorescentia brevi
ovato-oblonga ramis maturis reflexis, foliis brevioribus
curvatis spinescentibus, aristarum basi supra articulum
glabra et lemmate non ultra basim aristarum producto
differt.
Suffrutex perennis ligneus, plus minusve 1 m altus;
culmi basi simplices, sursum multo ramosi et fasciculati;
nodi glabri, in culmis principalibus paribus distante et in
ramis fasciculatis arete dispositi. Foliorum laminae
plerumque 40—70 mm longae, spinescentes, teretes,
canaliculatae, curvatae. Inflorescentia laxa, foliis superata,
20-50 x 20-30 mm; rami mature reflexi, in fasciculis
altemantibus 6-10 disposita. Spiculae 11—12 mm longae.
Glumae inaequales, superiores inferiores superantes.
Lemma 3-nervatum, tubulosum, arist?r"rp ham nrhcuia
turn; aristae 3, omnes plumosae, aequales; callus
acuminatus, pilosus.
TYPE. — Namibia, 2315 (Rostock): Namib Naukluft
Park, Mniszechi’s Vley, 23°43'S, 15°29'E, 23.10.1985,
Seely 2156 (PRE, holo.; G, K, M, MO, NBG, P, S, WIND,
Z). Figure 15.
Stipagrostis seelyae has previously been referred to in
literature as Stipagrostis sp. cf. namaquensis (Nees) Trin.
& Rupr. (Yeaton 1988; Boyer 1989). S. seelyae is named
after Dr M.K. Seely, Officer in Charge of the CSIR,
Namib Research Station at Gobabeb, who has made
extensive observations on this species and collected the
type material.
Perennial, robust, glabrous suffrutex (Figure 16), ± 1
m high, forming stiffly erect, sparse to dense tufts from
short woody rhizomes. Culms woody, straw-coloured or
purplish, persistent, up to 5 mm in diameter, rigid, erect,
unbranched below, fasciculately and much branched above
with the branches and spiny leaves forming dense erect
clusters. Nodes glabrous, approximate in geminate pairs
spaced widely apart on the unbranched lower portion of
the culms, the lower node of each pair visible, upper
covered by the leaf sheath of the lower node; nodes
numerous, close together, spaced 2-10 mm apart towards
the uuca oi tii*. i . 1 ‘’-vddpn by the leaf sheaths.
Leaves in pairs, !>U-14U nun apai t on '• .tidied
culms; spaced 2—10 mm apart towards the ends of the
branches. Leaf sheaths glabrous, smooth, straw-coloured,
basal ones reduced to cataphylls. Ligule a dense fringe of
very short hairs. Auricles inconspicuous, sparsely hairy
or glabrous. Leaf blades rigid, terete, canaliculate,
spinescent, slightly curved, stiffly erect or spreading,
(15)40—70(140) x ± 2 mm; glabrous and smooth
abaxially; adaxially strongly ribbed, shortly and densely
bristly on the ribs.
Bothalia 20,1 (1990)
83
FIGURE 15. — Stipagrostis seelyae. Holotype specimen, Seely 2156
(PRE), showing fasciculate branching pattern, x 0,3. Photo: A.
Romanowski .
Inflorescence a somewhat loose terminal panicle,
overtopped by the leaves (Figure 17A), 20—50 x 20—30
mm, glabrous except for the densely woolly points of
attachment of one or two of the basal branches; branches
6—10, single or in clusters alternating on the main axis,
each cluster consisting of 2— 3 branchlets each bearing one
to three spikelets; clusters subtended by a swollen pulvinus
and reflexed when mature. Spikelets one-flowered, 11—12
mm long including the awns, straw-coloured, the lemma
below the articulation occasionally purplish (Figure 17B,
C). Glumes somewhat curved, straw-coloured, firmly
chartaceous with thinner margins; lower glume 4,0-4, 5
mm long, ovate-lanceate, 3-nerved, the lateral nerves much
shorter than the midnerve which does not extend to the
apex, apex obtuse, lacerate; upper glume 8-9 mm long,
3-nerved, all the nerves extending nearly to the apex, apex
acute. Lemma chartaceous, tubular, ± 6 mm long up to
the base of the awns and including the callus; callus
densely hairy, acuminate, about 1,5 mm long, the hairs
long at the base of the lemma, sharply diminishing in
length down to the fine sharp point; articulation ± 1 mm
below the branching point of the awns; awns 3,
disarticulating together with the conical hollow apex of
the lemma, spreading and finally reflexed when mature,
equally developed, about 5 mm long, densely plumose
with silvery spreading hairs, plumes lanceate in outline
with the apex subobtuse. Palea 1,4 mm long, chartaceous,
glabrous, two-nerved, the nerves strongly developed but
not produced into a distinct keel, apex obtuse to
emarginate, margins inrolled. Lodicules 2, 1,3 -1,5 mm
long, narrowly obovate, one-nerved, membranous and
hyaline but subfleshy and denser towards the base. Stamens
3; filaments hairlike, elongating at anthesis; anthers ± 4,5
mm long, yellow, linear. Pistil ± 3 mm long; ovary ovoid,
glabrous; styles 2; stigmas plumose, very narrowly oblong
in outline. Caryopsis narrowly subturbinate, (3)2— 3(4)
mm long, shallowly grooved adaxially, and rounded
abaxially; hilum and embryo about ^3 of the length of the
grain.
NAMIBIA. — 2315 (Rostock): Namib Naukluft Park, Mniszechi’s Vley,
eastern edge of Namib dunes immediately south of the Kuiseb River
(— CB), 07.12.1983, Seely 2154: 23.10.1985, Seely 2155, 2156 (PRE); 8
km W of Tsondap Vlei (-CD), 30.5.1969, Jensen 238 (PRE); south of
Natab (— CA) 18.4.1969, Jensen 170 (PRE). 2415 (Sossusvlei): Sossus
Vley (-CB), 29.6.1974, Giess 13433 (PRE, WIND).
Affinities : even though the vegetative and inflorescence
characters of Stipagrostis seelyae differ fairly strongly
from those of S. sabulicola (Pilg.) De Winter as set out
in the diagnosis, the spikelets show great similarity, both
in size and general configuration and S. sabulicola is
considered to be the only species closely allied to. S.
seelyae. S. seelyae is placed in the Section Stipagrostis
because of the positioning of the articulation near the apex
of the lemma. The similarity in habit with S. namaquensis
can probably be ascribed to convergent adaptation to the
arid and sandy habitat occupied by these two species.
Distribution and ecology: Dr Seely summarizes her
extensive knowledge of the distribution and ecology of S.
seelyae , as follows: ‘It occurs extensively in the eastern
Namib dunes on the middle and upper dune slope. It does
not occur in the interdune valleys when these are not sandy
FIGURE 16. — Stipagrostis seelyae growing on a sandy dune slope,
showing suffrutescent habit. Copy of a colour slide taken by M.K.
Seely.
84
Bothalia 20,1 (1990)
FIGURE 17. — Stipagrostis seelyae,
Seely 2156 (PRE). A, single
inflorescence overtopped by
leaves, x 0,8; B, spikelet
showing the strongly unequal
glumes, the three-awned
lemma and spreading, plumose
awns, x 4,5; C, lemma
showing acute, hairy callus,
articulation below branching
point of awns and reflexed
awns, x 4,5. Photos: A, by A.
Romanowski; B, C, by M.
Koekemoer.
nor does it occur on the less stable, shifting dune crest.
When the full suite of dune grass species is present, S.
gonatostachys (Pilg.) De Winter and S. ciliata (Desf.) De
Winter occur on the gravelly interdune and Centropodia
glauca (Nees) T. A. Cope and Cladoraphis spinosa (L.f.)
S.M. Phillips on the low to middle dune slope. Usually
Stipagrostis lutescens (Nees) De Winter occurs in the
interdunes when they are sandy and on the base of the
dunes themselves. At the very tops of the dunes,
particularly in the eastern part of its range, S. sabulicola
occupies the shifting dune crest. Where S. seelyae and
S. lutescens are absent in the western part of the dune
fields, S. sabulicola occurs throughout.’
This generalized account of the zonation of the grassy
elements of the vegetation, places the ecological niche
occupied by S. seelyae in broad perspective. The
composition of the suite of species with which S. seelyae
is associated, however, does vary from site to site. Yeaton
(1988) reports S. seelyae (5. cf. namaquensis (Nees) Trin.
& Rupr.) as associated with S. sabulicola, S. lutescens
and Centropodia glauca at Bushman’s Circles. On this site
S. seelyae occupies an intermediate position between S.
sabulicola on the dune crest and C. glauca on the dune
base. At the Far East Dune site both S. seelyae and C.
glauca are absent from the suite whereas at Noctivaga
Dune the three species present are clearly zoned over the
dune slope. Centropodia glauca occupies the base of the
dune and is replaced upslope by S. seelyae on the middune
slope, which in turn is replaced at the top of the dune by
S. sabulicola. Yeaton (1988) points out that species growing
where sand movement is rapid, will only be able to sur-
vive if they are able to produce sufficient seed before be-
ing buried. Plants with strong rhizomes are more
successful at establishing in moving sands and their
vegetative growth may be stimulated by sand deposition.
Moreover, because seeds are rapidly buried in
swift-moving sand, plants surviving in such habitats do
so predominantly vegetatively. In most of these respects
S. seelyae seems to be extremely well adapted to its sandy,
windswept environment. For its known distribution see
Figure 18.
This distinct species with its very limited distribution
range (Figure 18) occupies an even more specialized niche
in the dune habitat than most of the other members of the
genus Stipagrostis which are restricted to the Namib dune
environment. Further intensive study of the relationships,
evolution and ecological adaptation of plants of the shifting
dunes of the Namib is needed to gain an understanding
of the functioning of individual species in the ecosystem.
14° 16° 18° 20°
j»
FIGURE 18. — Distribution of Stipagrostis seelyae. Dots represent
herbarium records; hatching indicates distribution as observed
by Seely.
Bothalia 20,1 (1990
85
Stipagrostis pellytronis De Winter, sp. nov.
Stipagrostis pellytronis De Winter, sp. nov., S.
hochstetterianae affinis, sed foliis expansis marginibus
cartilagineis; intemodiis vaginis basalibus lanatis, callo
spiraliter torto bifido differt.
Perennis caespitosa in anno primo florens, turn annuum
similis. Vaginae basales foliorum indumento lanato dense
tectae vaginae collo accedentes, persistentes, coronam
lanatam perennem formantes. Nodi annulum elevatum
lanatum tecti. Foliorum laminae expansae, nervis
marginatis costas cartilagineas formantes. Inflorescentia
spicata, singula vel plures ex vaginis aphyllis supremis
oriens, ex vagines ad maturitatem exserta, ex spiculis ±
9 sessilibus unifloris constans. Gluma inferior superiorem
longitudine superans. Lemma admodum infra medium
articulatum; callus ad maturitatem spiraliter tortus, bifidus;
aristae 3, tantum centrale longa plumosa, lateralibus multo
brevioribus filiformibus glabris.
TYPE. — Namibia, 2315 (Rostock): (-CD) Namib
Naukluft Park, Tsondab Vlei, on south-feeing scree slope,
28.4.1971, Jensen (Seely) s.n. in Herb. Giess 11519 (PRE,
holo. ; G, K, M, MO, NBG, WIND). Figure 20.
The specific epithet refers to the woolly indumentum
on the basal leaf sheaths which ends abruptly at the collar
of the sheath thus forming a sock-like structure. The Greek
Lexicon by Plassow gives the following: ‘Pellytron, a sort
FIGURE 19. — Stipagrostis pellytronis. Perennial tufts growing on gravel
derived from Tsondap sandstone. Photo: copy of a colour slide
by M.K. Seely.
of bandage or buskin worn by runners next to the foot and
ankle.’ Elsewhere also referred to as ‘a sock’.
Caespitose perennial 300-400 mm high (Figure 19),
frequently flowering in the first year and then often very
small and resembling an annual. Culms single (first year)
or several from a perennial base, simple with a single
apical inflorescence, or branched from the upper nodes
bearing one to several inflorescences on each branch.
Nodes 3—5 per culm, covered by a dense, raised ring of
woolly hairs. Intemodes (10— )50(-70) mm long, pale
yellow, lower ones densely, upper ones gradually more
sparsely, covered with woolly crisped hairs, glabrescent.
Leaves mainly basal, upwards on the culms gradually
reduced to the sheaths which envelop the immature
inflorescences. Leaf sheaths with ± 12 ribs (nerves)
abaxially, smooth adaxially, shorter than the intemodes;
the basal sheaths covered with a dense woolly indumentum
which in the perennial plants forms a wool-covered
persistent basal crown, upper sheaths glabrous (Figure
21A). Collar on the basal leaves densely woolly, sharply
demarcated from the glabrous leaf blade; in the upper
leaves inconspicuous and glabrous. Ligule a straight line
of stiff bristles of irregular length. Leaf blade very
narrowly lanceate, 3—5 mm wide, expanded, (10— )40
(—110) mm long, firm in texture, glabrous, basal blades
well developed, gradually diminishing in size upwards on
the culms, absent from the inflorescence sheaths; marginal
nerves conspicuous cartilaginous ribs; midrib incon-
spicuous; other nerves fine, numerous, equally strongly
developed.
Inflorescence (Figure 21B) a short spike, borne singly
or several together enclosed in apical sheaths on the culms
or culm branches, and where more than one spike is
present, their subtending sheaths also enclosed by the
apical sheath when immature, exserted on a slender
rhachis, when mature; ± 40 mm long excluding the awns,
each spike composed of ± 9 sessile spikelets, arranged
alternately on a slender, strongly ribbed rhachis. Spikelets
with a single bisexual flower. Glumes (Figure 22B) pallid,
chartaceous; lower exceeding the upper in length, 12—16
mm long, boat-shaped, lanceate, 3-5-nerved, central
nerve reaching the apex, lateral nerves evanescent, tapering
to a bifid or lacerate membranous apex; upper glume
10—12 mm long, 3-nerved, central nerve reaching the apex,
lateral nerves slightly shorter, boat-shaped; the elongate-
truncated, cartilaginous prolongation of the rhachilla or
‘callus-placenta’ adnate to inside base of the upper glume
Lemma chartaceous, tubular, glabrous, including the callus
+ 12 mm long, articulated slightly below the middle;
callus ±2,5 mm long, strongly spirally twisted when
mature, tapering to a glabrous bifid point, densely covered
with short, white antrorse bristles which are 0,5— 1,0 mm
long at the base of the lemma, diminishing in length towards
the apex (Figure 22 A), awns 3, disarticulating and felling
with the upper half of the lemma; central awn densely
plumose with spreading silvery hairs, 25—70 mm long,
plume narrowly obovate in outline, tapering to a glabrous
or sparsely hairy lower third, apex of plume obtuse; lateral
awns glabrous, filiform, 13-25 mm long, spreading. Palea
a broadly-oblong, semi-chartaceous, truncate scale, ± 1,5
mm long, the nerves represented by two approximate
thickened ribs flanking a shallow depression. Lodicules
2, ± 1,5 mm long, asymmetrically lanceate, fleshy at the
86
Bothalia 20,1 (1990)
FIGURE 20. — Distribution of Stipagrostis pellytronis. Dots represent
herbarium records; hatching indicates distribution as observed
by Seely.
base, with several indistinct nerves. Stamens 3; anthers
yellow, linear, ± 7 mm long. Pistil ± 6 mm long; ovary
glabrous, ovate-oblong; styles 2; stigmas plumose.
Caryopsis narrowly turbinate, oblanceate in outline,
3—3,5 mm long, smooth, pale brown; hilum a longitudinal
line as long as the grain; embryo slightly more than ^3
the length of the grain.
NAMIBIA. — 2315 (Rostock): Namib Nauklufit Park, Tsondap Vlei
(-CD) 23°54'S, 15°24'E, in sand at base of dune near river, 11.4.1971,
Hofineyer 45 (WIND); grass at edge of gravel mountain, 30.5.1969, Jensen
(Seely) 236 (PRE); lower reaches of Tsondap River (—DC) 23°54'S,
15°21E, 22.9.1979, Ward 256 (PRE).
S. pellytronis is unique in the genus because of the very
well developed spirally twisted, bifid callus of the lemma.
In spite of the unique combination of these features, an
affinity with S. hochstetteriana (Boeck. ex Hack.) De
Winter and especially the variety secalina, is proposed
for the following reasons: the general configuration of the
spikelet and particularly the inversion of the length of the
glumes (the lower being longer than the upper), the
spiciform inflorescence, the plumose central and the
glabrous filiform lateral awns, as well as the position of
the inflorescence subtended by sheaths at the apex of the
culms. The placing of S. pellytronis in the Section
Schistachne of Stipagrostis, the section in which S.
hochstetteriana is placed, presents no particular difficulty
since the allocation of species to the section is based solely
on the positioning of the articulation. Both species are
articulated near the middle of the lemma. Previous to the
discovery of S. pellytronis, a bifid callus was known only
in S. obtusa (Del.) Nees, where both pungent and bifid
calli have been recorded in the same species. The unique
configuration of the callus is regarded as representing a
specific rather than a sectional difference.
The distribution of the species is apparently limited to
exposures of Tsondap Sandstone within the central Namib
FIGURE 21. — Stipagrostis pellytro-
nis. A, basal part of a first year
seedling showing woolly
sheaths and annular woolly
covering on the nodes, X 1; B,
inflorescence, X 1. From the
holotype, Jensen (Seely) s. n. in
Herb. Giess 11519 (PRE).
Photo: A. Romanowski.
Bothalia 20,1 (1990)
87
FIGURE 22. — Stipagrostis pellytronis. A, lemma showing articulation
in middle of the body of the lemma and the spirally twisted, hairy,
bifid callus; B, glumes, the lower exceeding the upper; both x
7. From the holotype, Jensen (Seely) s.n. in Herb. Giess 11519
(PRE). Photo: M. Koekemoer.
dune sea. The distribution map (Figure 20) indicates the
area in which it has been collected or observed. Its
presently known most southerly occurrence is at Sossus
Vlei where Tsondap Sandstone outcrops are found. Its
distribution further south is uncertain but it may occur in
other areas in the eastern part of the dunes, where Tsondap
Sandstone outcrops on the surface. The absence of both
S. pellytronis and S. seelyae from the western part of the
dune complex cannot be explained. The two species extend
in an easterly direction, apparently as far as suitable
habitats occur.
ACKNOWLEDGEMENTS
Dr D.B.J. Killick is thanked for the translation of the
diagnoses and diagnostic descriptions into Latin. For the
information on distribution and ecology of the new species,
I am indebted to Dr M.K. Seely who also collected the
type material.
REFERENCES
YEATON, R.I. 1988. Structure and function of the Namib dune grass-
lands: characteristics of the environmental gradients and species
distributions. Journal of Ecology 76 : 744 —758.
BOYER, 1989. Some characteristics of the plant communities of three
dunes situated across a climatic gradient in the Namib Desert.
Madoqua 16: 141—148.
B. DE WINTER*
* National Botanical Institute, Private Bag X101, Pretoria 0001.
MS. received: 1989.12.04.
ASCLEPIADACEAE
CORONA LOBE VARIATION AND THE GENERIC POSITION OF ASCLEPIAS MACRA
INTRODUCTION
Generic concepts in parts of the tribe Asclepiadeae have
in the past varied greatly. This was partly due to a lack
of insight into the evolutionary trends underlying the
variation seen in coronal morphology in the tribe.
Baillon (1890), unable to establish well circumscribed
genera in the tribe Asclepiadeae, sank most of the African
members into one large super-genus to which he applied
the name Asclepias. N.E. Brown (1902, 1907-1909) had
a better understanding of the variation he encountered and
produced a workable classification that has served us well
for over 80 years. He relied heavily on corona lobe
morphology and genera were often separated from each
other on single characteristics. In Brown’s system Asclepias
was distinguished from all other genera in the tribe sole-
ly on the possession of a corona lobe sinus. Recent work
in Asclepias sensu N.E. Brown (Nicholas 1981) has high-
lighted the feet that distantly related groups have occasion-
ally followed the same evolutionary pathways ending up
with coronal characteristics that are analogous rather than
homologous. Species with such analogues have in the
past been lumped together into genera that form more or
less workable units but do not reflect underlying evolu-
tionary realities or affinities. Three distinct evolutionary
trends can be seen in corona lobe morphology: 1, reduc-
tion to a blob-like structure. Continuation of this trend
leads to the eventual disappearance of the lobe; 2, or-
namentation, namely increasing ornamentation by the
production of wings and/or proximal, distal and sinal
appendages (Figure 23). These can be produced singly,
together or in various combinations; 3, saccation or the
production of a corona lobe sinus, which may result from
the development of wings and/or appendages (Figure 23).
In Flora capensis the genus Pachycarpus was dis-
tinguished from others in the tribe Asclepiadeae by its
slipper-like corona lobes that may possess two parallel
wings proximally on the upper surface of the keel. In
addition, the distal end of the keel may become either
extended or ornate.
The species long known as Asclepias macra Schltr. was
originally described by R. Schlechter in 1895(a) under the
name Gomphocarpus suaveolens Schltr. Later that same
88
Bothalia 20,1 (1990)
FIGURE 23. — Corona lobe variation in Pachycarpus section
Campanulati showing trends towards both ornamentation and
saccation. Al, A2, P. campanulatus var. sutherlandii, Gerrard
1298, (K). Al, gynostegial column; A2, corona lobe; A3, corona
lobe of the type of P gerrardii, now considered synonymous with
P. campanulatus var. sutherlandii, Gerrard 1299, (K). B, P.
campanulatus var. campanulatus, Sanderson s.n., (K). C— F,
corona lobe: C, P. linearis, Baur 381, (K); D, P rostratus,
Haygarth ex Wood 7543, (K); El, E2, P. stelliceps. Bolus 12117,
(K); F, P. suaveolens, Schlechter 4109, (K). All drawings x 3.
year he sank Gomphocarpus into synonymy under
Asclepias (Schlechter 1895b) and transferred all the known
species of Gomphocarpus, including G. suaveolens, to this
genus (Schlechter 1896a). Having done this, Schlechter
then noticed that the name Asclepias suaveolens had
already been used by Leconte and cited by Decaisne in
De Candolle’s Prodromus (1844). He therefore proposed
the new name Asclepias macro for the taxon (Schlechter
1896b). This latter name was adopted by N.E. Brown in
Flora capensis and has been used in all subsequent
literature. However, Leconte’s name was never validly
published and according to Decaisne (1844) is no more
than an annotation on a herbarium sheet in the Museum
National d’Histoire Naturelle, Phandrogamie, Paris (P).
As a result Gomphocarpus suaveolens is the correct
basionym and Asclepias macro should be considered a
synonym. Ongoing research into the tribe Asclepiadeae
has highlighted the fact that the taxon originally described
as G. suaveolens should be placed in the genus Pachy-
carpus E. Mey. section Campanulati (Schltr.) A. Nicholas
& D.J. Goyder.
Brown (1907) excluded Asclepias suaveolens from the
genus Pachycarpus because it possessed a distinct corona
lobe sinus, although he did notice its similarity to this
genus — ‘this remarkable species is so exceedingly like
Pachycarpus gerrardi, N.E. Br., as to be easily mistaken
for that plant until the corona is examined’. However, if
one examines the corona lobe variation in Pachycarpus
section Campanulati, the progression towards a corona
lobe sinus can be clearly seen (Figure 23). P. suaveolens
represents the end point of the evolutionary trend towards
saccation by species in this section. If the plant is viewed
in its entirety, there is little doubt that its affinities lie with
this section and not with Asclepias sensu Flora capensis
(Table 3). The trend towards sinus production has also oc-
curred in section Pachycarpus, a fact hinted at by Bullock
(1953), who included species like P. lineolatus (Decne.)
Bullock and P. schweinfurthii (N.E.Br.) Bullock in the ge-
nus. These species were previously placed in Asclepias
because they possess a corona lobe sinus.
Pachycarpus section Campanulati is distinguished from
all other taxa within the tribe Asclepiadeae by the character
combination of linear to narrowly lanceolate leaves with
revolute margins, single erect stems that bear nodding
inflorescences and campanulate flowers with a hairy ovary.
Obscure characters shared with section Campanulati,
such as the beak-like shape of the anther wings and the
unusual multicellular brown hairs which may sometimes
be found on the ovary, also clearly establish the affinity
of P. suaveolens with this section.
Pachycarpus E. Mey. Section Campanulati (Schltr. )
A. Nicholas & D.J. Goyder, comb. nov.
Asclepias L. Section Campanulatae Schltr. in Botanische Jahrbucher
21: 9 (1896a). Pachycarpus E. Mey. Section Trichocodon D.M.N. Smith:
300 (1983); Smith: 399—439 (1988). Type.— Pachycarpus campanulatus
(Harv.) N.E. Br., chosen here.
Pachycarpus suaveolens (Schltr. ) A. Nicholas & D.J.
Goyder, comb. nov.
Gomphocarpus suaveolens Schltr. in Botanische Jahrbucher 20: 38
(1895a). Asclepias suaveolens (Schltr.) Schltr.: 9 (1896a). Asclepias macra
Schltr.: 456 (1896b); N.E. Br.: 670-671 (1907). Types. —Transvaal 2529
(Witbank): Olifants River (-CD), Schlechter 4109 (K!, lecto., chosen
here; BM, GRA, NH, PRE! isolecto.). 2530 (Lydenburg): Elandspruit-
berg (-AA), Schlechter 4006 (syn., not seen by the authors).
DESCRIPTION
Perennial herb. Rootstock not seen. Stem single, erect
to suberect, 205-275(-400) mm high, bifariously
strigose, younger parts pilose. Leaves erect or slightly
spreading, linear to narrowly lanceolate, (45— )65— 105
(-152) x (1,8 — )2, 5-5,0 mm, apex acute, base minutely
truncate (almost minutely auriculate) to attenuate, sparsely
hairy, margins revolute; petiole 2,0— 4,5 (—7,0) mm long.
Inflorescence umbelliform, semipendulous to pendulous,
terminal, rarely axillary, 1(— 3) per plant, 5— 10-flowered;
peduncles 20— 27(— 67) mm long. Flowers 17—24 x 9—15
mm; pedicel 15—20 mm long. Sepals lanceolate to ovate,
(0,4-) 5, 0-8,0 x l,8-2,8 mm, hirsute. Corolla subglo-
bose campanulate, petals fused for 2/3 their length,
11,5—16,0 x 7,0— 9,5 mm, occasionally with long multi-
cellular brown hairs outside; lobes (3,5— )5, 0—7,0 x
7,0— 9,5 mm, apices rounded and reflexed. Gynostegial
Bothalia 20,1 (1990)
89
TABLE 3. — Morphological comparison of Pachycarpus suaveolens with Asclepias sensu Flora capensis and Pachycarpus section Campanulati
stalk 0,5 -0,8 mm. Corona lobes fused basally, lobes
compressed cucullate, 3,2— 4,0 x 4,8— 5, 2(— 6,0) mm;
appendages more or less level with style apex, proximals
usually just over-topping it; proximal appendages broadly
falcate, almost unguiform, 1,0— 2,0 x 1,5— 3,0 mm, distal
appendage finger-like, bifid apically with ± 1,3 mm long
cleft, 0,7— 1,6(— 2,4) x (0,8— )1, 2— 2,0 mm; keel round-
ed; sinus a central slit, puberulous inside, ± 1,8 mm deep.
Anthers: anther wings beak-like, 0, 6-1,0 x 1,5— 2,1 mm;
anther appendages tongue-like, 1,3— 2,0 x 1,2— 1,5 mm,
decumbent on the style apex, apex rounded. Style apex
truncate with a central depression and 5 crenulate, almost
flanged outer lobes, whitish, 3,0— 4,0 mm in diameter.
Pollinarium : pollinia golf-club-shaped, 0,5— 1,0 x 1,1 —
1,25 mm; translator arms minutely winged at
junction with corpusculum, (0,4 — )0, 6-0,8 mm long;
corpusculum fusiform, 0,14 — 0,2( — 0,3) x 0,2— 0,4(— 0,6)
mm. Ovaries covered in long, brown, multicellular hairs.
Fruit & seed not seen. Figure 24.
TRANSVAAL. — 2529 (Witbank): Olifants River (-CD), Schlechter
4109 (BM, GRA, K, NH, PRE). 2628 (Johannesburg): Dersley Golf
course near Geduld (-AB), Flugge-de-Smit ex Moss 18178 (PRE). 2629
(Bethal): Spitskop, Ermelo (— BD), Scheepers 15043 (PRE). 2630
(Carolina): near Bosses (— ?), Burtt Davy 2956 (K).
The morphological affinities of Pachycarpus suaveolens
lie most closely with P campanulatus var. sutherlandii
N.E. Br. However, it can be distinguished from this and
all other taxa in section Campanulati by its distinct,
centrally placed corona lobe sinus. The key published by
Smith (1988) has been revised to include P suaveolens and
is presented below.
la Style apex with margins extended laterally beyond or
vertically above the anther appendages:
2a Style apex forming 5 lobes which extend horizontally past
the anther appendages; appearing stellate from above
P. stelliceps
2b Style apex extending vertically above the anther appendages,
terminating in 5 small erect lobes; never appearing
stellate from above P rostratus
lb Style apex with margins never extended laterally beyond
or vertically above the anther appendages:
3a Anther appendages 5, 0-7,0 mm long P linearis
3b Anther appendages 0,5-4, 0 mm long:
4a Corona lobe saccate-cucullate without large proximal
wings on the adaxial surface of the keel ... P. suaveolens
4b Corona lobe slipper-shaped with large proximal wings
on the adaxial surface of the keel:
5a Corona lobes 3, 0-8,5 mm long. Inflorescences
3— 10-flowered P. campanulatus var. sutherlandii
5b Corona lobes 9,5-17,0 mm long, inflorescences never
more than 5-flowered
P. campanulatus var. campanulatus
Pachycarpus suaveolens is a rarely collected southern
Transvaal endemic (Figure 25). Like the other species in
section Campanulati, this attractive plant is found in
annually burnt or cut grasslands. It flowers between
December and January and as such appears to have a rather
short flowering period. Only the two specimens collected
by Rudolf Schlechter have altitudes recorded — these being
2 040 m and 1 460 m. The rootstock, fruit and seeds have
neither been recorded nor collected. Flower colour has
not been recorded either and it is difficult to tell from the
dried specimens what colour they would have been.
FIGURE 24. — Pachycarpus suaveolens, Schlechter 4109, (K): A, whole
plant x 0,5; B, transverse section through leaf showing revolute
margins x 3; C, flower with corolla splayed x 1,5; D, gynostegial
column showing shape of corona lobes x 2; E, pollinarium x 7,5.
90
Bothalia 20,1 (1990)
FIGURE 25. — Presently known distribution of Pachycarpus suaveolens.
Map scale 4 mm - 100 km.
evidence of an attempt at self pollination. Had the
specimen not fallen prey to a plant press one wonders if
this attempt would have been successful.
In the field P. suaveolens must look a fine sight with
its single, almost erect stem bearing a collection of eight
or more densely crowded campanulate flowers in a nod-
ding inflorescence. Figure 26.
ACKNOWLEDGEMENTS
The authors would like to thank the directors of both
the Royal Botanic Gardens, Kew and the National
Botanical Institute, Pretoria for the opportunity to
undertake this work. Thanks also go to all those institutions
that allowed us to consult their specimens and use their
facilities. Mrs Grey-Wilson (RBG) and Mrs Romanowski
(BRI) are thanked for the art work and photography respec-
tively. We are also grateful to Dr Brummitt for his help,
and to the referees for their positive contribution.
REFERENCES
However, it is likely that they are brownish with a hint
of green and purple. One dissected flower ( Flugge-de-Smit
ex Moss 18178 ) showed a pollinium (in situ) from which
a mass of pollen tubes had started to germinate, probably
FIGURE 26. — Photograph of Pachycarpus suaveolens showing the habit
of the plant. Note the single erect stem with its many-flowered,
pendulous inflorescences. Photograph of Flugge-de-Smit ex Moss
18178, PRE.
BAILLON, H.E. 1890. Histoire des plantes 10: 221—304. Hachette, Paris.
BROWN, N.E. 1902. Asclepiadeae. In W.T. Thiselton-Dyer, Flora of
tropical Africa 4,1: 231-503. Lovell Reeves, London.
BROWN, N.E. 1907—1909. Asclepiadeae. In W.T. Thiselton-Dyer, Flora
capensis 4,1: 518—1036 & 1129—1133. Lovell Reeves, London.
BULLOCK, A. A. 1953. Notes on African Asclepiadaceae III. Kew
Bulletin 1953: 329-362.
DECAISNE, J. 1844. Asclepiadeae. In A.P. Candolle, A. Candolle &
C. Candolle, Prodromus systematis naturalis regni vegetabilis 8:
490-665.
NICHOLAS, A. 1981. Taxonomic studies in Asclepias L. (Asclepiadeae)
with particular reference to the narrow-leaved species in southern
Africa. M.Sc. thesis. University of Natal, Pietermaritzburg.
SCHLECHTER, R. 1895a. Beitrage zur Kenntnis sudafrikanischer
Asclepiadeen. Botanische Jahrbiicher fdr Systematik, Pflanzen-
geschichte und Pflanzengeographie 20: 1—56.
SCHLECHTER, R. 1895b. Asclepiadaceae Elliotianae. Journal of
Botany, British and Foreign 33: 300—307 & 333—339.
SCHLECHTER, R. 1896a. Die Drdge’schen Asclepiadaceen im Ernst
Meyer’schen Herbarium. Botanische Jahrbiicher fiir Systematik,
Pflanzengeschichte und Pflanzengeographie 21, Beiblatt 54:: 1—14.
SCHLECHTER, R. 1896b. Revision of extra-tropical South African
Asclepiadaceae. Journal of Botany, British and Foreign 34:
311-315, 417-421 & 449-458.
SMITH, D.M.N. 1983. Section Trichocodon. InO.M. Hilliard & B.L.
Burtt, Notes on some plants of southern Africa, chiefly from
Natal: X. Notes from the Royal Botanic Garden Edinburgh 41,2:
300.
SMITH, D.M.N. 1988. A revision of the genus Pachycarpus in southern
Africa. South African Journal of Botany 54: 399—439.
A. NICHOLAS* and D.J. GOYDER**
* National Botanical Institute, Private Bag X101, Pretoria 0001. Presently:
South African Liaison Botanist, Royal Botanic Gardens, Kew, Richmond,
Surrey TW9 3AB, England, UK.
** The Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey TW9
3AB, England, UK.
MS. received: 1989.05.23.
Bothalia 20,1: 91-103 (1990)
Vegetative morphology and interfire survival strategies in the Cape
Fynbos grasses
H.P. LINDER* and R.P. ELLIS**
Keywords: Cape flora, ecology, fire, fynbos, growth form, habit, Poaceae, survival strategies
ABSTRACT
It is shown that there is a wide range of structural variation in the habit of the Arundineae and Ehrharteae of the fynbos
of the Cape Floristic Region (Cape Province, South Africa). Structural differences in the bases of the fynbos grasses have
been classified into four groups: swollen, knotty tillering, weak and annual. Variation in the position of the innovation buds
occurs with one group having basal perennating buds, implying that all the culm material is annual, while the second group
has cauline innovation buds, leading to the development of a divaricate perennial herb. The recognition of caducous, mesic
(orthophyllous) and sclerophyllous leaf blades is also possible, based on leaf morphology and anatomy. These variations
in growth forms allow the classification of the Cape grasses into five guilds adapted for survival in the dense fynbos vegetation
that develops between the well-spaced fires in these heathlands. The following guilds have been recognized: competition
avoiders that grow on rock ledges and outcrops where competition from shrubby vegetation is reduced; reseeders, that survive
the protracted interfire period as seed; geophytes, that survive this period as underground organs; coppicers, that survive
as small plants; and competitors, that grow tall by means of cauline innovation buds, and so are able to compete with the
shrubby heath vegetation.
UITTREKSEL
Daar word aangetoon dat daar ’n wye reeks strukturele variasie in die groeiwyse van die Arundineae en Ehrharteae van
die fynbos van die Kaapse Floristiese Gebied (Kaapprovinsie, Suid-Afrika) bestaan. Strukturele verskille in die basisse van
die fynbosgrasse is in vier tipes ingedeel: geswolle, knoesterig stingelsuiervormend, swak en eenjarig. Daar bestaan variasie
in die posisie van die verjongingsknoppe, met een groep wat basale oorwinteringsknoppe het, wat impliseer dat al die
halmmateriaal eenjarig is. Daarteenoor kom die verjongingsknoppe by die tweede groep op die stingel voor, wat aanleiding
gee tot die ontwikkeling van ’n uitgespreide meerjarige kruid. Die erkenning van vroegafvallende, mesiese (ortofiele) en
sklerofiele blaarskywe is ook moontlik, op grand van die blaarmorfologie en -anatomie. Die Kaapse grasse kan op grand
van hierdie variasie in groeivorms in vyf gildes geklassifiseer word wat aangepas is om te oorleef in die digte fynbosplantegroei
wat tussen die goedgespasieerde brande in hierdie heideveld ontwikkel. Die volgende gildes word erken: kompetisievermyders
wat op rotslyste en klipriwwe groei waar kompetisie van struikagtige plantegroei verminder is; hersaaiers wat die langdurige
periodes tussen brande as saad oorleef; geofiete wat hierdie periode in die vorm van ondergrondse organe oorleef; stomplootplante
wat as klein plante oorleef; en kompeteerders wat deur middel van verjongingslote aan die stingels, hoog uitgroei en sodoende
met die struikagtige fynbosplantegroei kan kompeteer.
INTRODUCTION
The southern and south-western parts of the Cape
Province of South Africa possess a distinct flora, usually
called the Cape Flora (Good 1974; Taylor 1978; Goldblatt
1978; Takhtajan 1986). Goldblatt (1978) delimited the
geographical area of this flora, and called it the Cape
Floristic Region, with the major vegetation type being
‘fynbos’. This region is virtually synonymous with the
Fynbos Biome (Rutherford & Westfall 1986). This Cape
Flora contrasts with the floras of adjacent biomes both
in physiognomy and composition. Floristically, apart from
the normally common Asteraceae and Fabaceae, it is
characterized by the families Proteaceae, Ericaceae,
Iridaceae and Restionaceae, with the Mesembryanthe-
maceae and Crassulaceae dominating in the more arid
regions (Goldblatt 1978; Bond & Goldblatt 1984). The
levels of endemism are remarkably high, both at species
(68%) and at generic (20%) level. The Poaceae, although
generally ranking highly in most floras, is only the 13th
largest family in the Cape Floristic Region, even with the
* Bolus Herbarium, University of Cape Town, Rondebosch, 7700.
** National Botanical Institute/Department of Agricultural Development,
Private Bag X101, Pretoria 0001.
MS. received: 1989.05.18.
inclusion of the exotic pooids introduced from Europe
(Bond & Goldblatt 1984).
The Poaceae of the Cape Floristic Region are poorly
understood taxonomically and ecologically. The last
complete critical taxonomic revision was by Stapf (1899)
for the Flora capensis, while Chippindall (1955) produced
a guide to their identification. Conert (1970, 1971) revised
the generic limits of Danthonia, which resulted in
several new genera being recognized for the region.
Recently Gibbs Russell and Ellis (Gibbs Russell 1987a,
b; Ellis 1987a, b; Gibbs Russell & Ellis 1987, 1988)
have started a programme on Ehrharta, and a co-ordinated
programme on the arundinoid grasses of southern
Africa is under way. Davidse and Ellis have worked
on Tribolium and Prionanthium (Davidse 1988; Ellis
1989), Barker and Ellis on Pentameris and Pseudo-
pentameris (Barker 1986; Ellis 1985a, b,c,d, 1986),
while Linder & Ellis (1990) have commenced a programme
on Pentaschistis , Poagrostis and allied genera. Linder (in
prep.) has reviewed the phytogeographical patterns
inherent in the grasses of the Cape Floristic Region,
showing that several taxa may help elucidate the origins
and evolution of the Cape Flora. The distribution, breeding
systems and eco-physiology of the Cape grasses have not
been studied, and the only ecological information appears
92
Bothalia 20,1 (1990)
to be incidental comments in descriptions of vegetation
types (i.e. Taylor 1978; Kruger 1979).
The Cape Floristic Region is ecologically very diverse.
The climate is essentially Mediterranean, with most of the
rain falling in winter, whereas the summers are dry. The
amount and seasonal distribution of the precipitation is
influenced greatly by the mountain ranges running more
or less parallel to the coast. Along the coastal slopes the
rainfall is about 1 000 mm p.a. , with the drier summers
being ameliorated by mist and regular showers. The inland
leeward slopes are much drier, with rainfall often less than
250 mm p.a. , and with a very hot and dry summer (Fuggle
& Ashton 1979). The soils are generally deficient in
nutrients, and the combination of dry summers and low
soil nutrient status results in slow growth rates for the
vegetation as a whole. The vegetation is a shrubland or
heathland, from 0,5 to 3 m tall, with a very small
herbaceous component, and virtually no annual
component (Taylor 1978; Kruger 1979; Campbell 1985).
Along the wet coastal ranges the vegetation is dense, with
a high basal cover and no bare ground, while along the
arid inland slopes the shrubs are well scattered, with ample
bare ground which is colonized by annual plants after good
winter rains. Although the fynbos vegetation is pyrophytic,
fires are spaced well apart, with at least four years between
fires, and at most about 40 years, due to the slow biomass
build up. The modern average would probably be some
15 years between fires, although it appears likely that in
the past the fire cycle could have been much longer (Van
Wilgen 1987).
Habit and growth form in the Poaceae have received
remarkably little attention in the literature, and
consequently there is no generally useful terminology
available to describe the vegetative structures. The terms
‘tussock’ and ‘clump’ refer to rather similar type structures,
and some of the more curious, aerially branching
plants have no ready terms. The Cape Poaceae show
a remarkable diversity of habits, from the classical
caespitose tussock grass to a complex growth form
somewhat similar to a divaricate herbaceous plant.
These habits appear to be adapted to a range of habitats,
both spatial and temporal, in the Cape Floristic Region.
They may be highly informative on the ecology and
selective restraints operative in the area, and show that
these grasses ate closely integrated into this unique
system, but they have received very little research
attention.
Bond & Goldblatt (1984) list almost 200 species
of Poaceae for the Cape Floristic Region. Of these,
virtually all the endemic species belong to the
Arundinoideae in the tribes Arundineae and Ehrharteae.
The Pooideae is represented largely by exotic taxa and
the Bambusoideae is absent from this biome. Panicoideae
and Chloridoideae, although present in fairly large
numbers of individuals, have only one endemic species
between them, and are scarce in ‘typical’ fynbos
vegetation, but they tend to dominate the grassy fynbos
types of the eastern Cape (Campbell 1985; Cowling
1984). The present study is based on the Arundineae,
Ehrharteae and Pooideae, indigenous and largely
endemic to the Cape Floristic Region. Nomenclature
follows that of Gibbs Russell et al. (1985), except for
Pentaschistis, where the nomenclature of Linder & Ellis
(1990) is followed.
METHODS
The distributions of the Poaceae species in the Cape
Floristic Region were determined from herbarium records
using the PRECIS system of the National Botanical Insti-
tute, Pretoria (Gibbs Russell 1985a), and the collections
of the Bolus Herbarium, University of Cape Town.
Whether these grasses were exotic or indigenous
was determined following Gibbs Russell et al. (1985).
Vegetative morphology was also examined on these
herbarium specimens. The habit and growth forms of these
grasses was classified by recognizing four different base
types, two positions for the innovation buds and three
different types of leaf morphology. This classification was
tested during extensive field work in the spring and
summer of 1987, and in the spring of 1988 (August-
December). While engaged in these field observations it
became apparent that functional correlations exist between
the vegetative morphology and the response of these
fynbos grasses to fire, and more particularly, to the
protracted periods without fire. These long interfire
periods in the pyrophytic fynbos vegetation are unusual
and the vegetative morphology, together with the fire
history data for specific localities led to the recognition
of several interfire period survival strategies exhibited by
different fynbos grass species. Five different strategies
were recognized and this classification was also tested in
the field during this period.
Study sites are indicated on Figure 1. The age of the
vegetation since the last fire was determined from records
of the Chief Directorate of Nature Conservation, from
records held in the local offices of Nature Conservation
officials, from observations by mountaineers and farmers,
and from personal experience of the post-fire successions.
Detailed leaf, culm and base anatomical studies were
made from material freshly preserved in FA A in the field,
and the vouchers are housed either at PRE (for leaf
anatomy) or at BOL (for base and culm anatomy). The
results on the anatomy of the bases and culms will be
reported separately (Linder, Thompson & Ellis in prep.).
Many of the leaf anatomy results have already been
published (Ellis 1980a, b, 1982a,b, 1985a— d, 1986,
1987a, b, 1988a, b), whereas studies on Pentaschistis leaf
anatomy are in preparation (Ellis & Linder in prep.).
Data for a comparison of the Poaceae growth forms of
the Cape Floristic Region with those of other biomes was
based on species lists available from Gibbs Russell,
developed for the analyses of southern African flora and
biomes (Gibbs Russell 1985b, 1987c). Growth forms of the
savanna and grassland grass species were assigned on the
basis of herbarium material. These results are summarized
in Figure 2 and on p. 102.
OBSERVATIONS AND DISCUSSION
Base morphology
With the exception of the bamboos, most perennial
grasses have herbaceous culms that die back to the base
annually and are replaced by shoots arising from axillary
basal buds (Gould 1968). The perennial grass ‘plant’
resulting from several seasons of growth is therefore, made
up of several to many lateral shoots initiated at the base.
Bothalia 20,1 (1990)
93
FIGURE 1. — The south-western Cape
Province, South Africa. The
Cape Floristic Region, as
defined by Goldblatt (1978) is
outlined by the line, while
sample sites are indicated by
the dots.
This results in the formation of clones which can reach
a spread of 200 m as in Festuca rubra (Harberd 1961).
By varying the length and thickness of the internodes of
this basal portion of the culm, various structures can be
developed. In the Cape grasses four different types of base
can be distinguished (Figure 3).
1 Bulbous or geophytic bases
The base is modified into a distinct storage organ. This
consists of the lowermost culm internode(s) which is
generally swollen and tuberous, bulb- or corm-like, and
which is usually covered by the fibrous bases of the old
leaf sheaths (Bums 1946). The development of these bulbs
and corms is rare in the Poaceae (Clayton & Renvoize
1986; Watson & Dallwitz 1988), and occurs in relatively
few, unrelated species (Clark & Fisher 1987). In the
Cape arundinoids several different structures are found.
Bulb-like structures, formed from swollen leaf bases,
Coppicers
Competition avoiders
I: . I Competitors
Geophytes
Ephemeral reseeders
Exotics
FIGURE 2. — Relative occurrence of growth forms of Poaceae in the
Fynbos, Savanna and Grassland Biomes of southern Africa.
occur in Pentaschistis viscidula, P argentea, Merxmuel-
lera rufa, and others. In Pentaschistis aristidoides a stout
horizontal rhizome is formed, while in Ehrharta villosa
portions of the stolons become swollen to form
pseudobulb-like structures. These structures are deeply
sunken below the level of the soil surface, and are often
woolly from the densely hairy leaf sheaths, thus protecting
the shoot bases from excessive evaporation or temperature
changes (Tsvelev 1976). This could also be a protection
against fire damage, or a defence against predators. Plants
with these bases rarely form dense tufts, usually bearing
a single terminal cluster of leaves and few fertile culms.
Bulbous bases are unusually common in the Cape
arundinoid grasses, examples being Merxmuellera rufa,
M. decora, Ehrharta capensis, E. longifolia, E. dura,
Pentaschistis aristidoides, P. viscidula, P. argentea and
P. velutina in the Arundinoideae and Festuca scabra in
the Pooideae.
2 Knotty tillering bases
The base consists of tightly aggregated clusters of very
short internodes, positioned at or just below the soil
surface. The whole structure is usually tangled with old
leaf bases, which probably play a protective role for
the basal buds. There is no clearly differentiated storage
organ. The plants are capable of coppicing from this base
after fire, and form tussocks (e.g. Pentaschistis pallescens,
P pyrophila and Merxmuellera arundinacea) . If the
tillering base is underground, the resulting tussock is often
tightly caespitose, as in P eriostoma, P. pyrophila and
Merxmuellera stricta.
3 Weak bases
In this category only a few perennial culms arise from
a basal node. The basal internode(s) is not swollen into
storage organs, but sometimes associated swollen rhizomes
occur (e.g. Ehrharta setacea and E. villosa). The plants
developed from weak bases are usually loosely tufted,
short-lived perennials, with branching culms which are
decumbent at the base. This stooling (Gould 1968) gives
a ‘cushion’ type of growth. Selected examples of this type
of base are Pentaschistis pallida form B, P. densifolia,
94
Bothalia 20,1 (1990)
P. rosea subsp. rosea, P. alticola, P. aspera, P. acinosa,
Ehrharta rupestris and E. setacea. These weak bases are
very similar to annual bases in that the rooting system is
weak, and no thickened woody structures are found.
4 Annual bases
In grasses with annual bases the primary shoot arises
directly from the base, and few lateral innovation shoots
develop further up the culms. The base never becomes
woody. Annual grasses are rare in the mountains on the
most nutrient-poor soils (Kruger 1979), but are better
diversified on the arid margins of the Cape Floristic Region
( Pentaschistis airoides subsp. airoides, P. aristifolia, P.
capillaris, Urochlaena pusilla, Ehrharta brevifolia and E.
pusilla ). Biennials and triennials are found in the
mountains. These also have the annual type of base
FIGURE 3. — Variation in the base
structures of the fynbos
arundinoid grasses. A, knotty
tillering base, Pentaschistis
pyrophila\ B, geophytic base,
Merxmuellera rufa\ C, disc-
like tillering base, Pentaschistis
ampla\ D, weak base, P.
densifolia-, E, annual base, P
airoides subsp. airoides. All x
0,4.
(Pentaschistis rosea, P. pseudopallescens), showing the
link between these and weak bases.
Innovation bud position
The culm is the axis on which the inflorescence as well
as the leaves are borne. Each culm is terminated by an
inflorescence, and the next season’s growth is produced
by lateral shoots arising from the nodes. These lateral
shoots develop from innovation buds, which are enclosed
and protected by the prophyll, a two-keeled modified leaf,
which is located adaxially at the base of each lateral shoot
(Clark & Fisher 1987). Different growth habits are
determined by the position of these perennating buds and
their resultant lateral shoots. Two basic variants can be
recognized (Figure 4).
Bothalia 20,1 (1990)
1 Basal innovation shoots
Basal innovation shoots develop from buds at the base
of the plant. These produce culms which are annual, dying
back to the upper tillering node after completion of
flowering. All intemodes above this node perish, and those
below are persistent. This is the common situation in grass-
es (Gould 1968; Tsvelev 1976), and this tillering produces
erect or decumbent lateral culms and the caespitose habit,
usually as a result of intravaginal branching. After flower-
ing the culms and leaves senesce, and are eventually
replaced by a new basal innovation shoot. These basal in-
novation shoots are usually associated with knotty tiller-
ing bases. Plants with bulbous bases also innovate from
the base, but these are deeply subterranean.
Grasses with this type of lateral shoot innovation
essentially have an annual above-ground component.
95
Although generally grass leaves senesce after only a few
months, many of the Cape grasses have leaves that persist
for at least a year. The underground parts are strongly
perennial, and may continue to grow for many years,
reaching ages of up to 1000 years in Festuca ovina (Harberd
1962). This type of growth form is well adapted to the
regular annual or biennial fires characteristic of the
subtropical savannas and grasslands. In the Cape Floristic
Region examples of this growth form are Pentaschistis
glandulosa, P. pyrophila, Ehrharta calycina, and the
species listed as having bulbous bases.
2 Cauline innovation shoots
Cauline innovation shoots are produced from nodes
higher upon the culms. These culms are generally long-
lived, and the above-ground component is perennial,
resulting in an ‘evergreen’ plant. This is an unusual
FIGURE 4. — Variation in the position
of the innovation buds in the
fynbos arundinoid grasses. A,
caulescent innovation buds in
a ‘competitor’, Pentameris
squarrosa\ B, basal innovation
buds along creeping rhizomes,
Pentaschistis galpinii ; C, basal
innovation buds, forming a
caespitose plant, Pentaschistis
curvifolia\ D, innovation buds
at the ends of thin erect culms,
in an old plant of Pentaschistis
colorata, forming a cushion.
All x 0,4.
96
Bothalia 20,1 (1990)
FIGURE 5. —Transverse sections of the leaves of the fynbos arundinoid grasses, illustrating variation in leaf type. A— E, mesic or orthophyllous
leaf type; F— I, sclerophyllous leaf type. A, Ehrharta brevifolia , with large chlorenchyma cells and intercellular air spaces; B, Pentaschistis
papillosa , with discrete sclerenchyma girders and strands; C, Tribolium uniolae, chlorenchyma cells relatively large and angular but air
spaces reduced, intermediate type; D, Festuca scabra , typical pooid mesic anatomy with very diffuse mesophyll and widely spaced vascular
bundles; E, Chaetobromus involucratus, mesic leaf anatomy; F, Ehrharta setacea , compact mesophyll of isodiametric cells; G, Pentaschistis
eriosloma , compact mesophyll, bundles close together and continuous hypodermal sclerenchyma; H, Pentameris macrocalycina , compact
mesophyll of small cells and few intercellular air spaces, sclerenchyma girders well developed; I, Merxmuellera rufa, compact sclerenchyma
typical of sclerophyllous leaf type. All x 250.
Bothalia 20,1 (1990
97
situation in herbaceous grasses (Gould 1968). Branching
culms are very rare in the Pooideae, but there are
exceptions in the Andropogoneae (Clayton & Renvoize
1986), Paniceae and particularly the woody Bambuseae
(Roshevits 1937). The branching may be profuse with
secondary and even tertiary branches. The bases of the
species with cauline innovation shoots are either of the
weak or of the knotty tillering type. These species also
appear to be able to innovate from the base, and so survive
fires by coppicing.
Three different patterns appear to occur in the cauline
innovating species. In one group, the culms are more or
less erect, and the branches well spaced. This results in
a ‘divaricate herb’, which might function in allowing the
plant to grow taller than species with only basal innovation
shoots (i.e. Pentaschistis aspera, P. acinosa, P. scandens,
Pseudopentameris macrantha, P brachyphylla, Pentameris
thuarii and P squarrosa). In the second group the culms
are more or less decumbent (at least at the base), and the
branches mostly occur near the base of the culms. These
plants develop into cushions, often low on the ground.
Good Cape examples of this are Pentaschistis densifolia,
P. rosea subsp. purpurascens and P. alticola. In the third
group the plant is initially caespitose, with basal innovation
shoots. As the plant becomes older, the innovation shoots
are found further from the base of the plant, and a distinct,
and sometimes quite long, stem section is found between
the woody rootstock and the branching system. This
transforms the plant from caespitose to tangled. This
pattern is shown most clearly by P. colorata, but is also
seen in Cape plants of the widespread Themeda triandra.
Leaf morphology
The leaf anatomy and morphology of grasses is very
important taxonomically, but ecologically three leaf forms
may be distinguished in the Cape Floristic Region (Figure
5).
1 Caducous leaf blades
In some species of Ehrharta the leaf blade is either early
deciduous, or is represented by a small mucro at the apex
of the sheath. The leaf sheaths are generally well
developed, and together with the culms constitute the
photosynthetic organs of the plants. These suffrutescent
species are remarkable structural mimics of the
Restionaceae.
2 Sclerophyllous leaves
Sclerophyllous leaves (Ellis 1988a) persist for more than
one growing season, and possibly even for several years.
These are tough, leathery leaves, typically permanently
inrolled and filiform or acicular with no abaxial stomata
and very compact mesophyll of small isodiametric
chlorenchyma cells with minute intercellular air spaces.
Sclerenchyma tissue is abundant and may be either
lignified or with cellulose walls (Figure 5F— I). This type
of leaf appears to be analogous to the sclerophyllous leaves
which are characteristic of the vegetation of the Cape
Floristic Region, particularly with vegetation growing on
the nutrient-deficient sands of the mountains. This type
of blade structure probably helps to ‘safeguard’ the scarce
nutrient resources. All species of Merxmuellera and
Pentameris, as well as many species of Pentaschistis have
this type of blade.
3 Mesic leaves
Mesic leaves do not persist for more than one growing
season. They are typically soft and expanded with abaxial
stomata and diffuse mesophyll, and with an extensive air
space system. Sclerenchyma tissue is only associated with
the vascular bundles as strands or girders (Figure 5A-E).
This leaf type is common on soils with better nutrient
status. The pooid exotics all have mesic type leaves, and
indigenous taxa with mesic leaves (e.g. Chaetobromus and
Tribolium) are lowland taxa associated with intermediate
nutrient status soils. However, many species of Pentaschis-
tis with mesic leaves are widespread in the Cape
mountains. These mesic leaves of the Cape fynbos grasses
appear to be analogous to the ‘orthophyllous leaves’ of
Campbell (1985) although the first term refers to habitat
and the second to the texture of the leaves.
Interfire survival strategies
The vegetation of the Cape is a shrubby heathland,
dominated by Restionaceae, Ericaceae and Proteaceae
(Taylor 1978). These fynbos plants are evergreen, generally
highly divaricated, and they form a relatively dense
shrubbery of 1—3 m tall. The vegetation burns on a
variable cycle of between four and 40 years, as vegetation
less than four years old rarely has adequate fuel to burn
(Van Wilgen 1987). Grasses in general are well adapted
to withstand fire, and tropical grasses in particular, become
moribund if not burnt regularly. These tropical grasses
generally are subject to annual, biennial or triennial fires
and their growth forms and habits undoubtedly are
determined largely by this regular incidence of fire.
Ecological studies on the vegetation of the Cape Floristic
Region have generally emphasized the various methods
by which the flora survives the sporadic fynbos fires (e.g.
Manders & Cunliffe 1987). However, in the case of the
grasses, in addition to surviving the fires themselves,
survival in the dense, shrubby fynbos vegetation that
develops during the long interfire periods appears to be
a major constraint. Immediately after fire, perennial
grasses generally are an important component of the
regenerating fynbos vegetation, even though this family
is virtually absent in mature fynbos prior to burning. After
a bum the vegetation gradually becomes denser, generally
passing through dominance by the Restionaceae, then the
Ericaceae, and finally the Proteaceae (Kruger 1977). This
mature, dense vegetation excludes herbaceous species, and
so also the grasses. There appear to be five ways in which
the grass species of the Cape have responded to this
situation.
1 Competition avoiders (Table 1)
These species exploit habitats where taller woody plants
cannot grow. Such habitats are either in streams, or in rock
crevices and ledges. In these harsh conditions the grasses
compete with mosses, a few herbaceous evergreen species
and geophytic orchids. The protection from fire offered
by these habitats appears to be incidental. Some species
are totally restricted to these habitats (e.g. Pentaschistis
98 Bothalia 20,1 (1990)
TABLE 1. — Distribution of growth form and survival strategy in the competition avoider Cape grasses. The vegetation types are approximate and follow
Moll et al. (1984)
rigidissima, P. acinosa). Other species colonize soil
exposed after fire, but soon succumb to the rapidly
growing shrubby vegetation. Pentaschistis densifolia, P.
malouinensis, and often P eriostoma fall into the latter
category.
Typically, the avoiders are short, cushion forming plants
with weak bases, cauline innovation shoots, and
sclerophyllous leaves. They are generally restricted to
higher altitudes in the mountains (possibly because of the
availability of crevices). Pentaschistis eriostoma and P
malouinensis are both rather widespread, and ecologically
flexible. P. eriostoma , in particular, can vary from a typical
vegetation avoider at higher altitudes, to a classical tussock
grass on the arid fringes of the Cape Flora. Some
vegetation avoiders deviate from the above description by
the possession of mesic leaves, and often also basal
innovation shoots. Pentaschistis densifolia and P pusilla
are arundinoid examples, while most of the ‘avoider’
pooids also fall into this category ( Anthoxanthum
dregeanum, A. tongo and Brachypodium distachyon).
The only true graminoid hydrophyte in the Cape is
Pentaschistis capensis which is found over a wide altitude
range, but always growing on rocks and stones in perennial
streams, often draped over waterfalls. For this species it
is then not clear whether it is avoiding drought or
competition from woody vegetation.
2 Reseeders (Table 2)
This group can be defined as those taxa that survive the
interfire period as seed. The seed germinates after fire and
grows into annual or short-lived perennial plants that
survive for 1-3 years before being forced out by the
recovering fynbos vegetation. The grass plants die, leaving
their seed till the next fire, which may not be for 25 years,
and possibly longer. Whether the species survive as seed
banks, or whether they depend on regular dispersal is not
clear. If they survive as seed banks, the mechanisms by
which the seed is protected for such an extensive period
requires attention.
Within the Cape Floristic Region several different
reseeder strategies exist. True annuals (ephemerals) are
found only on the arid fringes of the Region. This includes
the West Coast Strandveld (Moll et al. 1984) and the
Succulent Karoo (Acocks 1988). In these vegetation types
soil nutrient levels are higher, but low rainfall prevents
the development of a dense basal cover, so that there is
ample open ground between the scattered bushes, where
annual grasses can grow following adequate rainfall. Pen-
taschistis airoides subsp. airoides, P aristifolia, P capil-
laris, Urochlaena pusilla, Stipa capensis, Ehrharta
brevifolia, E. longiflora, E. pusilla, Tribolium utriculosa
and T. echinata are annual arundinoid species of these arid
regions. This is possibly the closest analogue to the true
Mediterranean conditions of southern Europe and northern
Africa to be found in the winter rainfall region of the Cape.
These are not strictly postfire annuals and usually function
as interfire annuals as well.
Fynbos postfire reseeders are not true annuals, in that
they do not germinate annually, although the plants
themselves only grow for a single growing season.
Pentaschistis pseudopallescens, P. rosea and possibly
Bothalia 20,1 (1990)
99
P. alticola are biennial or triennial, which flower in the
second year after fire. They occur above 1 000 m on
nutrient-poor and leached soils in the western Cape moun-
tains, and it is possible that these slow development times
may be due to the poor soil nutrient status, and the harsh
growing conditions. P. pallida form B and P triseta are
reseeders which flower in the first year and are found at
lower altitudes, in warmer conditions, and on less leached
soils. Both groups appear to be facultative perennials,
depending on the severity of the summer droughts. If the
plant survives the summer, and is not shaded out by the
growing vegetation, it may flower for a second year. These
species may totally dominate the vegetation after the fires.
They all have weak or annual bases, basal innovation
shoots and mesic leaves.
3 Geophytes (Table 3)
This group of species behaves like typical geophytes,
appearing early in the first winter or spring after fire and
flowering proftisely soon after commencing growth. By
the second post-fire season they are not as p-ominent and
flower less profusely. When they become overshaded by
the regenerating woody vegetation, they appear to survive
the protracted interfire period by means of their swollen,
subterranean storage organs. It appears as if flowering
depletes the base, but that subsequent photosynthesis may
replenish the storage products (Hodgkinson & Williams
1983). However, research is still required to follow the cy-
cle of flowering and photosynthate accumulation and trans-
location, both in these grasses and in analogous groups,
such as Watsonia in the Iridaceae. The factors controlling
flowering and dormancy are not understood. The stored
products allow the plants to flower almost
immediately after fire, while the reseeders apparently have
to spend the first year accumulating sufficient reserves to
support a flowering episode.
The Cape flora is remarkably rich in geophytic grasses.
These species tend to dominate areas that are frequently
burnt or bush-cut, like fire-breaks. This survival strategy
is undoubtedly effective and the methods of overcoming
damage by herbivores are poorly understood. These geo-
phytic grasses are probably an important food source for
mole-rats, a possibility which needs to be investigated
(Lovegrove & Jarvis 1986). Examples are Pentaschistis
viscidula, P argentea, P. velutina, Merxmuellera rufa, M.
decora and Ehrharta longifolia. Morphologically, they all
have bulbous, subterranean bases, basal innovation shoots
and sclerophyllous leaf blades, although these leaves are
unusual in having abaxial stomata. Pentaschistis aristi-
doides probably also belongs to this group, but is unusual
in that the base is developed as a rhizome. Ehrharta dura,
E. microlaena, E. capensis and Festuca scabra differ from
the typical type described above by their mesic leaf
anatomy.
4 Coppicers (Table 4)
These species survive the interfire period as persistent,
almost dormant, moribund plants. After a fire, they
coppice from the base and then flower late in the first
spring or summer. The plants then persist above ground
without, or with very little, subsequent flowering. They
are often evident in the mature vegetation, persisting in
the vegetative state as weak moribund tussocks. Presum-
ably, if the interfire period is too long, the plants will
eventually die. This form is then usually found at higher
TABLE 2. — Distribution of growth form and survival strategy in the ephemeral reseeder Cape grasses. The vegetation types are approximate and follow
Moll et al. (1984)
100 Bothalia 20,1 (1990)
TABLE 3. — Distribution of growth form and survival strategy in the geophytic Cape grasses. The vegetation types are approximate and follow
Moll et al. (1984)
TABLE 4. — Distribution of growth form and survival strategy in the coppicing Cape grasses. The vegetation types are approximate and follow
Moll et al. (1984)
Bothalia 20,1 (1990)
101
altitudes where the mature vegetation is not very dense,
and where recovery rates are slower.
Pentaschistis pyrophila and Pentameris obtusifolia are
typical examples utilizing this survival strategy. They have
knotty tillering bases, basal innovation shoots and
sclerophyllous leaf anatomy. Other species which utilize
this strategy, but not exclusively so, are Pentaschistis
colorata, P. tortuosa, P. eriostoma and P. ampla. Penta-
schistis curvifolia, P malouinensis and Ehrharta calycina
may also best be placed into this group, despite their weak
bases.
This group overlaps to some extent with other strategies,
and is often difficult to define. Many species that form
large, persistent tussocks in the more open vegetation at
lower altitudes are probably best placed into this group,
despite the absence of direct evidence of coppicing after
fire. This would include Pentaschistis rupestris, P. pallida
form F, P glandulosa, P. tomentella, P. cirrhulosa, P
calcicola, Merxmuellera arundinacea and M. cincta.
These lower altitude species are generally not shaded out
during the interfire period, but this is because of the sparse
vegetation in which they grow, rather than any special
morphological adaptations that they may have.
5 Competitors (Table 5)
This small, specialized group of grasses appears to be
able to compete with low mature fynbos. The plants have
branching culms capable of almost indefinite growth. Af-
ter flowering has terminated the growth of a culm, a lateral
branch continues growth from an upper leaf innovation
bud. New leaves are continually formed near the top of
the culms with the basal portions being covered by
senesced leaf sheaths. These plants may either be tall (up
to 2 m) and erect, as in Pseudopentameris macrantha,
Pentameris macrocalycina and P. thuarii, or trailing
through the vegetation as in Pentameris squarrosa,
Pentaschistis scandens or some of the Ehrharta species.
These species do not appear to gain in biomass with
successive seasons (probably due to limiting nutrients) but
gain in height with a reduction in culm diameter and leaf
size. In old plants the leaves may be less than the length
of those produced in the first post-fire season, but the plant
may have doubled in height. Flowering is most frequent
in the first year after fire and gradually diminishes in later
years.
The group of suffrutescent species belonging to the
Villosa and Ramosa groups of Ehrharta are also
competitors, mostly at higher altitudes. These are
remarkably similar to the Restionaceae, with reduced,
caducous leaves and photosynthetic culms.
Members of this specialized group all have the weak type
of base and cauline innovation shoots. The leaves
vary from sclerophyllous for most of the higher altitude
taxa, or caducous (associated with suffrutescent culms)
in some species of Ehrharta, to almost mesic in some
TABLE 5. — Distribution of growth form and survival strategy in the competitor Cape grasses. The vegetation types are approximate and follow
Moll et al. (1984)
102
Bothalia 20,1 (1990)
low-altitude competitors. It is not clear how long these spe-
cies are capable of persisting in unburnt vegetation, as
some show signs of senescence. However, this may depend
on the density of the local variant of fynbos. It is this group
which shows the greatest deal of convergence with the Res-
tionaceae.
Comparison with other biomes
The spectra of interfire survival strategies found in the
Cape are compared in Figure 2 with those of the Grassland
and the Savanna Biomes as defined by Gibbs Russell
(1985b, 1987c). Direct comparison is difficult, as the
enormously different physiography of the different biomes
may bias for different growth forms. In addition, the
relative frequency of fires in the other biomes suggests
that tall and dense interfire vegetation is not a factor, as
fire intervals are never as long as 10 years. However, a
comparison is still illustrative of how diverse the Cape
grasses are in vegetative morphology and ecology in
comparison with tropical grasses. From Figure 2 it is
abundantly clear that correlated with the extended fire
intervals in the fynbos is the occurrence of the ‘competitor
strategy’, which is totally absent from the other biomes.
Competition avoiders in the other biomes are all
hydrophytes, but in the Cape Floristic Region only one
species is a hydrophyte, the rest being lithophytes restricted
to ledges and crevices. These are, therefore, true vegetation
avoiders, compared with the tropical hydrophytes.
Geophytes are remarkably common in the Grassland
Biomes as well as the fynbos, and the underground storage
of photosynthate in grasses may be worthwhile
investigating in more detail. Reseeders are understandably
more significant in the Savanna Biome than in both fynbos
and grassland where rainfall is usually not limiting.
However, the relatively large number of exotic pooids in
the fynbos tends to obscure this fact.
CONCLUSIONS
The Poaceae endemic to the fynbos show a range of
structural and morphological adaptations which allow them
to survive in a variety of niches in the Cape vegetation.
Some growth forms allow direct competition with the
shrubby Cape Fynbos, while others allow plants to sur-
vive the dense interfire vegetation either as seed, as geo-
phytes or as small dormant plants. The models postulated
here should contribute to the development of a better un-
derstanding of the unique selective forces operative in the
fynbos.
The Poaceae in the Cape show a vegetative plasticity
not expected from grasses. Unusual growth forms have
previously been documented in some arundinoid grasses
such as Steyermarkochloa (Davidse & Ellis 1985) and
Arundoclaytonia (Davidse & Ellis 1987) but, apart from
the woody bamboos, extensive cauline branching is
basically unknown in the family. The arundinoids do have
a wide range in habit, from annuals to reed-grasses like
Phragmites (Renvoize 1981). It is therefore not unexpected
that the arundinoids have developed the specialized habits
to cope with the Cape Fynbos, whereas the indigenous
pooids and other subfamilies have been relatively
unsuccessful in the Cape. Exotic pooids, on the other hand,
are particularly successful in the Cape but, noticeably, not
in the natural flora but in areas of physical disturbance.
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Bothalia 20,1: 105-111 (1990)
The ecology of the False Bay estuarine environments, Cape, South
Africa. 1. The coastal vegetation
M. O’CALLAGHAN*
Keywords: estuaries, False Bay, vegetation
ABSTRACT
The vegetation in and around eleven estuaries flowing into False Bay was surveyed during 1980 and 1981. Use was made
of colour aerial photographs and a combination of dominance and phytosociological techniques. Of the communities established,
three are aquatic and four are described as emergent or wetlands. Of the terrestrial communities, five are described as fynbos
and four occur on coastal sands. One community consists solely of alien plants. The communities thus classified generally
compare well with those discussed by other workers in the area. However, differences due to the destruction and disturbance
of the vegetation are commented upon.
UITTREKSEL
’n Opname van die plantegroei in en random elf riviermondings wat in Valsbaai invloei, is gedurende 1980 en 1981 gemaak.
Kleurfotos en ’n kombinasie van dominansie- en fitososiologiese tegnieke is gebruik. Van die gemeenskappe waarvan die
teenwoordigheid vasgestel is, is drie akwaties en vier word as vleilande beskryf. Van die landgemeenskappe, word vyf as
fynbos beskryf en vier kom op kussand voor. Een gemeenskap bestaan slegs uit uitheemse plante. Die gemeenskappe aldus
geklassifiseer, vergelyk goed met die wat deur ander werkers in die gebied bespreek word. Verskille wat egter aan die vemietiging
en versteuring van die plantegroei toegeskryf kan word, word bespreek.
INTRODUCTION
An urban landscape (village, town, city) can be seen
as a functional system requiring major inputs and outputs
(energy, commodities, waste) to remain viable (Bartowski
1982). Rivers which traverse this system often provide
convenient intra- and inter-system transport routes for these
inputs and outputs. Rivers and other wetland areas are
therefore in great demand and often over-utilized in urban
environments. The loss of natural functioning of wetlands
is an indication of the conservation mindedness of, or
quality of planning by, the developers. Degradation of
urban wetlands is usually due to: 1, alteration of
hydrological regime; 2, reduction of the quality of water;
3, physical destruction of the system or parts thereof (Day
1987).
A river typically flows through a number of different
environmental zones. Within each zone, local factors (for
example flow rate, water depth, substrate) can vary
considerably, resulting in a variety of wetland types along
the river (seeps, bogs, emergent wetlands, submerged
wetlands, swamps, vleis)(0’Keefe 1986).
It is generally accepted (see for example Goodman 1987)
that wedands function to improve the quality of the riverine
environment (sediment trapping, flood attenuation,
nutrient sink, production of food). The variety of wetland
ecosystems found along a river should ensure that the
riverine and related environments at the end of its course
have a high quality. This can be seen by the high
productivity estimates for estuaries in numerous texts (e.g.
Whittaker 1970). However, even though wetlands are
resilient in response to natural disturbance, they are
susceptible to human influence. Furthermore, the
man-induced effects on the functioning of a riverine
ecosystem are cumulative as one proceeds down the river
course. Any deterioration of the riverine ecosystem will
be reflected in the quality of the estuarine environment.
This is shown in the present study by analysing the
vegetation in and around the estuaries of False Bay.
STUDY AREA
False Bay (Cape, South Africa) is a large square-shaped
bay with open access to the sea (Figure 1). The eastern
and western sides of the Bay are bound by the Hottentots
Holland and Cape Peninsula mountain ranges respectively
W'Ve
* National Botanical Institute, RO. Box 471, Stellenbosch 7600.
MS. received: 1989.02.01. FIGURE 1. — The location of the rivers entering False Bay.
106
Bothalia 20,1 (1990)
and the northern shore is formed by the low sandy Cape
Flats. The coastline, from Cape Point to Cape Hangklip,
is approximately 121,5 km long.
The natural vegetation consists of mountain fynbos
communities on sandstone-derived soils, particularly on
the western and eastern sides of the Bay. The Cape Flats
consist predominantly of calcareous sand supporting
strandveld vegetation (Taylor 1980).
Many factors affect the Bay, some of which were
discussed at a seminar entitled ‘The future management
of False Bay’ (Gasson 1980). Pressures are being placed
on the False Bay environment in the form of utilization
by an ever-increasing local population for recreation,
residence and industry; and by encroaching alien plants,
especially Australian acacias. As such, there is need for
well-planned management policies. However, available
data are not sufficient for a complete understanding of the
diversity and functioning of the False Bay environment.
The formulation of well-planned management policies for
this Bay is limited by the absence of adequate environ-
mental data (Gasson 1980).
According to the 1:50 000 Topographical Sheets (3418AB
& AD; 3418BA; 3418BB; 3418BD), eleven named
rivers flow into False Bay. They are, from west to east
(Figure 1): Buffels (West) (34°19'S; 18°28'E); Elsies
(34°10'S; 18°26'E); Silvermine (34°07'S; 18°27'E); Sand
(34°05'S; 18°28'E); Zeekoe (34°06'S; 18°30'E); Eerste
(34°05'S; 18°46'E); Lourens (34°06'S; 18°49'E); Sir
Lowry’s Pass (34°09'S; 18°52'E); Steenbras (34°12'S;
18°49'E); Rooiels (34°18'S; 18°49'E); Buffels (East)
(34°20'S; 18°50'E).
Much information concerning the biotic and abiotic
features of these rivers has been published by Bickerton
(1982), Cliff & Grindley (1982), Grindley (1982),
Heinecken (1982a, b), Heinecken etal. (1982), Morant &
Grindley (1982). The catchment area of these rivers
exceeds 1100 km2 and the mean annual run off is estimated
as 200 x 106 m3/y. Residential and industrial effluent
from at least six sewage works and an unknown number
of storm water systems discharge into False Bay via these
rivers. The rivers contain a number of dams, especially
in the Hottentots Holland Mountain area, and water is
extracted directly from some of the rivers for irrigation.
They also receive runoff from forms, containing fertilizers,
pesticides and other form wastes. The Kuils River summer
flow consists almost entirely of treated sewage effluent.
Although the vegetation of False Bay has been discussed
(Taylor 1980), no mention has been made of the vegetation
in and around the estuaries.
METHODS
Physiographic/physiognomic units were demarcated on
colour aerial photographs of 1979 (University of Natal Job
326/79) for each of the 11 river mouths. In total, approxi-
mately 83,6 ha was studied. The area studied at each river
mouth varied from 3,1 ha (Elsies) to 25,8 ha (Sand). Each
study site was selected to include aquatic, littoral and
terrestrial vegetation.
The vegetation was sampled during 1980 and 1981.
Releves were not used. Each physiographic/physiog-
nomic unit was regarded as a sampling unit. The sampling
units were therefore not artificially bound, but each
consisted of an entire physiographic/physiognomic unit.
Where necessary, physiographic/physiognomic units were
combined or further divided to obtain some degree of
uniformity in the vegetation of each sampling unit.
The dominant and prominent species were recorded for
each unit, together with a Braun-Blanquet cover-abundance
value (see Braun-Blanquet 1965) for each recorded spe-
cies. These floristic data were sorted by tabulation into
communities using the TABSORT suite of computer pro-
grams (Boucher 1977).
RESULTS
Sampling units and vegetation types
A total of 301 species was recorded from 81 sampling
units. Of these, 60% had a single occurrence in the data
set. Undersampled vegetation types were not resampled.
Of those species that occurred more than once, one could
not be identified to the species level. In total, 44 specimens
could not be identified to the species level.
Fourteen of the sampling units contained only one
species. These monospecific communities were confined
to submerged aquatics, emergent reeds and alien shrubs.
Table 1 is a summary of the vegetation types found. The
name of each vegetation type was selected for the sake
of convenience only, usually according to the two dominant
species. A complete list of species and sorted phyto-
sociological tables are available on request.
The average numbers of species recorded for each
vegetation type were: Submerged Aquatic Communities,
1,0; Emergent and Wetland Communities, 5,2; Cassine
barbara Communities, 20,2; Non-littoral Dune Com-
munities, 14,8; Littoral Dune Communities, 6,5.
Vegetation maps of each river mouth were prepared, but
have been published elsewhere (Bickerton 1982; Cliff &
Grindley 1982; Grindley 1982; Heinecken 1982a, b;
Heinecken et al. 1982; Morant & Grindley 1982).
DESRIPTION OF COMMUNITIES
(Note that alien plants are not mentioned in the description
of the communities. They are discussed separately under
3.3)
1 Submerged Aquatic Communities
Three aquatic species were recorded in the study area:
Potamogeton pectinatus, Ruppia cirrhosa and R. maritima.
Each of these species forms monospecific stands and these
should be regarded as distinct communities. Potamogeton
pectinatus forms tall dense subsurface stands in fresh to
slightly brackish waters. Ruppia spp. are found in
shallower water: R. maritima forms sparse communities
Bothalia 20,1 (1990)
107
TABLE 1.— Names of communities and the rivers at which they occur
Community
Rivers
1 Submerged Aquatic Communities 4 10 11
1.1 Potamogeton pectinatus Submerged Community 4
1.2 Ruppia cirrhosa Submerged Community 10
1.3 Ruppia maritima Submerged Community 4 11
2 Emergent and Wetland Communities 12345678 10 11
2.1 Phragmites australis Emergent Community 3 4 6 7 10 11
2.2 Paspalum vaginatum Wetland Community 2 3 6 7
2.3 Scirpus maritimus Mixed Community 4 6 8
2.4 Typha capensis Emergent Community 12 5 7
3 Terrestrial Communities 123456789 10 11
3.1 Cassine barbara Fynbos Communities 9 10
3.1.1 C. barbara -Leucodendron salignum Dry Mountain Community 9 10
3.1. 1.1 L. salignum— Thamnochortus gracilis Mountain Community 9
3.1.2 C. barbara— Poly gala myrtiflora Coastal Community 10
3.1.3 C. barbara-Rhus lucida Riparian Community 9 10
3.1. 3.1 R. lucida— Pelargonium angulosum Riparian Community 9 10
3.2 Tetragonia decumbens Coastal Sand Communities
3.2.1 T. decumbens— Sideroxylon inerme Mature Hind-dune Community
3.2.2 T. decumbens— Senecio halimifolius Moist Dune Community
3.2.3 T. decumbens -Metalasia muricata Mid-dune Community
3.2.4 T. decumbens— Agropyron distichum Fore-dune Community
3.3 Acacia cyclops Monospecific Community
12345678 10 11
3 6 11
4 5 6 11
1 2 3 4 5 6 7 10 11
3 5 6 7 8 10 11
1 2
Rivers: 1 = Buffels (west); 2 = Elsies; 3 = Silvermine; 4 = Sand; 5 = Zeekoe; 6 = Eerste; 7 = Lourens; 8 = Sir Lowry’s Pass; 9 = Steenbras;
10 = Rooiels; 11 = Buffels (east).
in areas of weak salinity; R. cirrhosa requires higher
salinities and can form relatively dense stands. Within this
study area, aquatic plants were found only at Sand, Rooiels
and Buffels (East). These systems are either very large
(Sand) or relatively undisturbed. If the study areas were
increased to extend along the rivers, it is likely that sub-
merged aquatics would also be found in some of the other
systems.
It should be noted that the aquatic environments are
dynamic and imposed management procedures which
affect these communities are ongoing in these estuaries
(Bickerton 1982; Cliff & Grindley 1982; Grindley 1982;
Heinecken 1982a, b; Heinecken et al. 1982; Morant &
Grindley 1982).
1.1 Potamogeton pectinatus Submerged Community: this
community was found only at Sand where it is confined
mainly to the middle reaches of the vlei. It thrives in this
brackish to fresh, nutrient- rich water to form dense
meadows that are ‘managed’ by weed cutting (Morant &
Grindley 1982).
1.2 Ruppia cirrhosa Submerged Community: this species
is restricted to the shallow areas above the road bridge at
the Rooiels River.
1.3 Ruppia maritima Submerged Community: these
communities are found at the Sand and the Buffels (East)
Rivers, to a depth of 800 mm in clear water.
2 Emergent and Wetland Communities
The only truly tidal estuary along the False Bay coast
is found at the Steenbras River. The other estuaries have
strong fluvial input during winter and the mouths usually
close during summer. However, the banks of the Steenbras
River are covered by boulders, and wetland vegetation does
not occur. All the other estuaries have wetland communi-
ties which, to some extent, reflect the prevailing water
regimes. All the wetlands are poorly developed and most
show signs of disturbance. They generally contain only
one or a few species and do not display zonation patterns.
Two of the communities have a tall emergent growth form
and two may be flooded seasonally or tidally.
2.1 Phragmites australis Emergent Community: Phrag-
mites australis usually forms monospecific stands, up to
2,5 m tall, on substrates where silt deposition occurs. This
species grows optimally in water with a salinity of less
than 15 °/oo, although it can withstand higher salinities
(Benfield 1984).
In the estuaries studied, Phragmites forms a fairly dense
stand at the Rooiels River with other wetland species such
as Chenolea diffusa , Juncus acutis, Sarcocomia spp. ,
Scirpus nodosus, Sporobolus virginicus and Stenotaphrum
secundatum. At the Silvermine, Sand, Eerste and Buffels
(East) estuaries, Phragmites forms very dense mono-
specific stands. These emergent communities generally
grow in areas away from the river mouth where there is
little or no salt water penetration. They are emergent in
varying water depths, especially during the rainy season,
or may be exposed to some tidal fluctuation if the mouth
is open.
2.2 Paspalum vaginatum Wetland Community: this
community is found on coarse saline sands and withstands
lengthy periods of inundation. It forms monospecific
stands at the Elsies Estuary. At the Eerste River estuary,
Cotula coronopifolia and Scirpus nodosus are found in this
vegetation type, whereas at the Lourens Estuary, two
communities are present, one being monospecific and the
other containing various Clijfortia and Cyperus species.
Paspalum vaginatum is regarded by some as an alien grass
108
Bothalia 20,1 (1990)
(Bond & Goldblatt 1984). In some areas, this community
also contains numerous alien herbs and shrubs.
2.3 Scirpus maritimus Mixed Community: these plants are
generally found on non-saline sandy soils that contain
some organic matter. Many of them survive inundation
in an emergent form. At the Sand River, this community
includes other Cyperaceae, Sarcocomia natalensis,
Paspalum vaginatum, Typha capensis and Triglochin
bulbosa. At the Sir Lowry’s Pass River, Cliffortia
lanceolata, Triglochin bulbosa and Juncus rigidus are
included. At the Lourens River estuary, Scirpus maritimus
forms a monospecific stand.
2.4 Typha capensis Emergent Community: this community
is emergent in slow-running, fresh water. The species
forms monospecific stands at the Elsies and Lourens
estuaries, the latter being a wetland not in direct contact
with riverine flow. At the Buffels (West) and Zeekoe
Rivers, numerous other species such as Samolus
valerandii, Berula thunbergii, Triglochin striata, Senecio
halimifolius, Scirpus nodosus, Zantedeschia aethiopica
and Polygonum salicifolium also occur.
3 Terrestrial Communities
The terrestrial vegetation comprises two basic types. The
first is distinguished by the presence of Cassine barbara
and is found on soils derived from Table Mountain
Sandstone. It occurs in the mountainous areas along the
eastern and western shores of the Bay. The second type
of terrestrial vegetation contains Tetragonia decumbens.
This vegetation grows on recent calcareous sands along
the northern shore of the Bay and in the river valleys where
this soil type has accumulated.
3.1 Cassine barbara Fynbos Communities: within this
study area, fynbos occurs only at the Steenbras River
mouth and Rooiels River mouth, predominantly on
shallow sandstone soils. Boucher (1978) records C. barbara
on littoral dunes, on limestone and in riverine scrub in
the Cape Hangklip area. It would seem that this species
is tolerant of a wide range of habitat factors and its
value as a diagnostic species is, therefore, suspect.
Nevertheless, Taylor (1969) found this species in a
subassociation of upland fynbos. Other species common
to this community ( Leucospermum conocarpodendron,
Widdringtonia nodiflora) help to distinguish this fynbos
community.
3.1.1 C. barbara— Leucadendron salignum Dry Mountain
Community: this community is indicative of Dry Mountain
Fynbos in this area (see Boucher’s (1978) community
3. 2. 2.1). The L. salignum— Thamnochortus gracilis
variation (3.1. 1.1) was found at the Steenbras River and
seems to be a form characteristic of slightly moister
conditions.
3.1.2 C. barbara- Polygala myrtiflora Coastal Community:
this vegetation was found at the Rooiels River and does
not compare well with any of Boucher’s (1978)
communities. It contains Protea compacta, Phylica
ericoides, Chondropetalum microcarpum and Myrica
quercifolia. A better sampling technique might indicate
a combination of Boucher’s (1978) Acid Sand Fynbos and
South Coast Strand veld. However, C. Boucher (pers.
comm.) suggests that, in this region, P compacta does
not occur naturally west of the Palmiet River Mouth and
this community might be a dune/limestone community (see
3.1).
3.1.3 C. barbara- Rhus lucida Riparian Community: this
is found in the narrow valleys of the Steenbras and Rooiels
Rivers with Psoralea pinnata, Podalyria calyptrata and
others. The R. lucida— Pelargonium angulosum variation
(3.1. 3.1) is usually found at the coastal or riverine edge
of the community.
3.2 Tetragonia decumbens Coastal Sand Communities:
these communities were found at all the rivers in this study
area.
3.2.1 T. decumbens— Sideroxylon inerme Mature Hind-
dune Community: this community was found at the
Silvermine, Eerste and Buffels (East) Rivers (although it
was also noted outside the boundary of the study area at
the Buffels (West) River) and compares well with Boucher
(1978), Taylor (1969) and Taylor (1980). The structure of
this community is highly variable. In protected areas, it
forms a low forest (up to 5m) with a closed canopy of S.
inerme, generally with an open understorey of grasses,
herbs and shrubs such as Cussonia thyrsiflora and
Chasmanthe aethiopica. In more exposed areas, this
community is usually wind-cropped to form a dense
shrubland from 0,5 m to 2 m.
T. decumbens is seldom found in this community and
Ehrharta villosa might have been a better diagnostic
species for these (3.2) communities.
3.2.2 T. decumbens— Senecio halimifolius Moist Dune
Community: this community compares well with Taylor
(1980) and was found at the Sand, Zeekoe, Eerste and
Buffels (East) Rivers, mainly in dune slacks and poorly
drained dune areas. It varies in height from 0,8 m to 1,5
m and can contain other species such as Scirpus nodosus,
Zantedeschia aethiopica, Typha capensis, Nidorella
foetida, Juncus kraussii, as well as shrubs and herbs more
typical of dune vegetation in this area.
3.2.3 T. decumbens— Metalasia muricata Dune Com-
munity: this community is typical of deep coastal sands
where the above conditions do not prevail. Its structure
varies from a relatively sparse low-growing shrubland up
to 2 m. It was found on the non-littoral dunes near all the
rivers except at the Steenbras where dunes do not occur.
It can be subdivided into various classes and associations
(see below).
3.2.4 T. decumbens— Agropyron distichum Fore-dune
Community: this community consists of psammophilous
pioneer vegetation which is commonly found on the
fore-dunes. It is dominated by a sparse or open cover of
grasses and herbs. It compares well with Boucher’s (1978)
‘ Ehrharta- Ficinia Strand Pioneers’ and Taylor’s (1969)
‘Pioneer Mixed Dune Fynbos’ to which Taylor gives the
status of subassociation.
3.3 Acacia cyclops Monospecific Community: alien shrubs
and grasses were found throughout most of the emergent,
Bothalia 20,1 (1990)
109
wetland and terrestrial vegetation units. Acacia cyclops is
particularly common in the vegetation on coastal sands
whereas Pennisetum clandestinum is found in these as well
as wetland communities. Alien plants recorded (according
to Bond & Goldblatt 1984) include Acacia cyclops, A.
longifolia, A. meamsii, A. saligna, Paraserianthes
lophantha subsp. lophantha, Ammophila arenaria. Aster
subulatus, Atriplex vestita, Avena sativa, Briza maxima,
Bromus diandrus, Chenopodium ambrosioides, C. murale,
Datura stramonium, Eucalyptus globulus, E. lehmannii,
Lagurus ovatus, Malva parviflora, Myoporum serratum,
Paspalum vaginatum, P. urvillei, Pennisetum clandesti-
num, Populus canescens, Pinus pinaster, P. pinea, Pinus
sp., Solanum nigrum, Sonchus asper, S. oleraceus,
Trapaeolum majus, and Urtica urens.
Where present, these species dominate the vegetation
to varying degrees; usually causing a loss in species
richness and cover of the natural vegetation. At the Buffels
(West) and Elsies Rivers, A. cyclops has, in places, ousted
the natural vegetation to form monospecific stands.
COMPARISON OF COMMUNITIES WITH THOSE OF
BOUCHER (1987)
Boucher (1987) carried out a survey of the western Cape
coastal forelands. Included in his study area was a part
of the False Bay coast, approximately from the Zeekoe
to Eerste Rivers. The communities he found within 1 km
of this coast are listed in Table 2.
From the above table, classes 5.1 and 5.2 are equivalent
to coastal sand communities. Class 5.4 indicates halophytic
marshes; class 5.6 indicates non-riverine (inter-dune)
wetlands; class 5.7 is found along water courses.
When comparing the present study with that of
Boucher’s (1987) study, the following points are evident:
according to Boucher (1987), the association 5. 1.1. 2 C is
found on the dunes along this coast. This should therefore
be equivalent to the above T. decumbens-Agropyron
distichum Community. However, sampling procedures
used for the present study do not allow for classification
to the level of association. Also, other species in this
community seem to indicate a combination of Boucher’s
Cladoraphio-Hebenstretion cordatae (5. 1.1. 2) alliance and
an association designated Senecioni-Ammophiletum
arenariae (5. 1.1.1. A), belonging to the alliance
Cladoraphio-Senecion elegantis (5. 1.1.1). Boucher regards
this last association as artificial owing to the dominance
of the alien grass Ammophila arenaria.
The strong presence of Metalasia muricata in the T.
decumbens—Metalasia muricata Community would,
according to Boucher (1987), place this community in the
association Senecioni-Metalasietum muricatae (5.1.1.1.B).
This association again falls into the alliance Cladoraphio-
Senecion elegantis (5.1. 1.1) which he did not record within
100 m of the False Bay coast.
The remainder of the terrestrial dune communities along
this coast were classified by Boucher (1987) as belonging
to the class Ehrhartetea calycinae (5.2). In the present
study, Ehrharta villosa was recorded as being co-dominant
with Tetragonia decumbens in the dune communities. It
TABLE 2. — List of communities within 1 km of the False Bay Coast
(after Boucher 1987)
is possible that E. villosa was confused with E. calycina
in some areas. Neither Sideroxylon inerme nor Cussonia
thyrsiflora were recorded by Boucher (1987). The above
community described as T. decumbens— Sideroxylon
inerme Community therefore has no counterpart in his
system. Boucher et al. (1986) suggest that the class
Arctothecetea populifoliae be divided into two orders:
Arctotheco-Cladoraphietalia cyperoidis and Arctotheco-
Passerinetalia rigidae. According to these latter authors,
the geographical separation occurs at Cape Point, with the
former order to the west. However, Boucher (1987)
includes vegetation sampled along the False Bay coast in
the former order. The differential species for these orders
were not common in the present study. Considering the
preponderance of Sideroxylon inerme and Cussonia
thyrsiflora on the dunes of the southern and eastern Cape
(pers. obs.), if this community belongs to the class 5.1,
it is likely to be found in the order Arctotheco-Passerine-
talia rigidae (Boucher et al. 1986), even though it was
110
Bothalia 20,1 (1990)
found on the western shores of the Bay. However, if
Ehrharta calycina was misidentified (as E. villosa), this
community might fall into the class Ehrhartetea calycinae
(5.2).
The community found on moist dune slacks ( T.
decumbens—Senecio halimifolius Community) would
belong to Boucher’s (1987) class 5.6 Scirpetea noddsi. This
class consists of two associations: Scirpo-Linetum
africanum and Scirpo-Nidorelletum foetidi. The major part
of the community described in this study falls within the
latter association; according to Boucher, it is found near
Swartklip (in the vicinity of the Eerste River). The former
association seems to be poorly represented in the present
study and, contrary to Boucher’s (1987) findings, was found
at the Buffels (East) River and not at the rivers near Mitch-
ell’s Plain.
The wetland vegetation Boucher (1987) recorded along
this coast belongs to the classes Sarcocomietea pillansiae
(5.4) and Polygonetea salicifoliae (5.7). The former
describes halophytic salt marshes and the latter describes
the vegetation fringing water courses.
The estuaries along the False Bay coast are relatively
small and do not have a regular tidal fluctuation nor saline
input. Halophytic wetlands are therefore poorly developed.
Nevertheless, these vegetation types did occur historically
(O’Callaghan 1990) and some of the species of these
communities are present. Boucher’s (1987) differential
species Sarcocomia pillansii was found only at Sandvlei
whereas one of his dominants, Sporobolus virginicus, has
a wider distribution.
The emergent community described above as the
Phragmites australis Community is recognized by Boucher
(1987) as the association Junco-Phragmitetum australis
(5.4. 2.C) in the order Sarcocomio-Juncetalia kraussii. The
Paspalum vaginatum Wetland Community indicates
Boucher’s association 5.4.2.B: Junco-Chondropetaletum
tectorum.
The Scirpus maritimus Mixed Community does not have
a counterpart in Boucher’s system. He recorded this
species only once and another species in this community
( Triglochin bulbosa) is noted as being differential and
widespread for his class 5.4.
The Typha capensis Emergent Community indicates the
association Junco-Nidorelletum foetidi (5.4. 2. A) in the
order Sarcocomio-Juncetalia kraussii. However, the
presence of Polygonum salicifolium indicates the class
Polygonetea salicifoliae, in particular, the association
Polygono-Juncetum capensis (5.7. A). Typha capensis was
recorded in the present study from the inter-dune wetlands,
adjacent to permanent water and alongside streams. It is
therefore likely that this community represents a mixture
of associations found in the classes Sarcocomietea
pillansiae (5.4), Scirpetea nodosi (5.6) as well as
Polygonetum salicifoliae (5.7), but mainly 5.6 and 5.7.
CONCLUSIONS
The studies by Boucher (1978, 1987) and Taylor (1969,
1980) were carried out in relatively pristine vegetation and
over far larger areas than the present study. The
communities they established are, therefore, a better
indication of the natural vegetation of the area. Although
the vegetation units established by the current study are
generally similar to their communities, there are problems
in matching, particularly with the dune and wetland vege-
tation types. These difficulties may be due to differences
in sampling techniques, but they could also be due to
disturbance and destruction of the natural vegetation in
the present study area.
O’Callaghan (1990) has shown that much vegetation
around these estuaries has been destroyed by residential,
recreational and industrial developments. In addition, alien
plants and trampling have brought about a loss of species
richness and natural plant cover.
The wetland vegetation types are particularly difficult
to match with those discussed by Boucher (1987). Although
the above influences affect these vegetation types,
additional influences such as the quality of water in
the rivers and the disturbance of natural flow regimes also
play a role. It seems that, where they exist, the wetlands
around the rivers entering False Bay are in a poor
condition.
Gasson & MacKinnon (1983) have estimated the volume
of outfall entering the False Bay catchment. The domestic
outfall is expected to increase from 146,525 Ml/d to 261,5
Ml/d, i.e. by 78,5% between 1982 and 1990. The volume
of outfall for industrial waste was 7,34 Ml/d in 1982 and
it is not unreasonable to assume that these volumes will
also increase dramatically. By 1990, more than 500
Ml/d of polluted water might be flowing into False Bay,
approximately 60% via the rivers.
Gasson & MacKinnon (1983) state that the standard of
effluent disposal is generally above the acceptable norms
set by the authorities. However, some treatment works may,
at times, release effluent which is below standard (e.g.
Kuils River). Furthermore, the record and control of ef-
fluent entering the rivers via routes other than through
municipal treatment works is scarce.
Whether this effluent enters directly or indirectly by
surface run off, it affects the vegetation around the rivers:
species richness is reduced and dominance by a single
species increased, which results in an increase of structural
homogeneity of the wetlands around these estuaries. This,
together with the physical destruction of the wetland
environments for development will cumulatively affect the
ecology of the estuaries and the quality of water entering
the Bay. It seems that a well-planned interdepartmental
management scheme (see Malan 1982) for False Bay,
including catchment areas, is indicated.
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4). In A. E.F. Heydom & J.R. Grindley, Estuaries of the Cape,
part II: synopses of available information on individual systems.
CSIR Research Report No. 413.
O’CALLAGHAN, M. 1990. The ecology of the False Bay estuarine
environments, Cape, South Africa. 2. Changes during the last
fifty years. Bothalia 20: 113—122.
O’KEEFE, J.H. 1986. Ecological research on South African Rivers — a
preliminary synthesis. South African Scientific Programmes
Report No. 121.
TAYLOR, H.C. 1969. A vegetation survey of the Cape of Good Hope
Nature Reserve. M.Sc. thesis, University of Cape Town.
TAYLOR, H.C. 1980. Plant communities along the False Bay coast. In
B. Gasson, Proceedings of symposium on the future management
of False Bay: 59-67.
WHITTAKER, R.H. 1970. Communities and ecosystems. MacMillan,
New York.
Bothalia 20,1: 113-121 (1990)
The ecology of the False Bay estuarine environments, Cape, South
Africa. 2. Changes during the last fifty years
M. O’CALLAGHAN*
Keywords: estuaries, False Bay, management
ABSTRACT
Aerial photographs taken between 1936 and 1987 of the eleven rivers flowing into False Bay were studied. Various techniques
were used to obtain quantitative values and value judgements for the changes that had taken place. It was found that large
increases in industrial, residential, recreational areas and alien plant cover have taken place at the expense of open sand
and natural vegetation types. The rivers along the eastern shore of the Bay are relatively undisturbed. Those along the western
shore are more disturbed but still contain some noteworthy environments. The most detrimental changes have occurred around
the rivers of the Cape Flats, along the northern shore.
UITTREKSEL
Lugfotos vanaf 1936 tot 1987 van die elf riviere wat in Valsbaai uitmond, is bestudeer. Verskeie tegnieke is gebruik om
kwantitatiewe waardes en waardebeslissings vir die veranderinge wat plaasgevind het, te verkry. Daar is bevind dat groot
toenames in nywerheids-, woon-, ontspanningsgebiede en uitheemse plantegroei plaasgevind het ten koste van oop sand en
natuurlike plantegroeitipes. Die riviere aan die ooskus van die Baai is betreklik onversteur. Die van die westekant is meer
versteur, maar daar het tog enkele natuurlike omgewings langs die riviermondings behoue gebly. Die skadelikste veranderinge
het langs die riviere van die Kaapse Vlakte aan die noordelike kuslyn plaasgevind.
CONTENTS
Introduction 113
Methods 114
Delineation of mapping units 114
Evaluation of changes 114
1 Relative environmental importance (I) 114
2 Relative environmental state (E) 116
3 Co-efficient of change (A) 116
4 Conservability (C) 116
Changes observed in different environments 116
Water and wetlands 117
Dune vegetation 117
Riverine scrub and fynbos 117
Factors causing changes 118
Alien vegetation 118
Residential development 118
Recreational development 118
Industrial development 118
Discussion of individual estuaries 118
Buffels (West) 118
Elsies 119
Silvermine 119
Sand 119
Zeekoe 119
Eerste 119
Lourens 119
Sir Lowry’s Pass 120
Steenbras, Rooiels and Buffels (East) 120
Comparison of estuaries 120
Conclusions 120
References 121
* National Botanical Institute, RO. Box 471, Stellenbosch 7600.
MS. received: 1989.02.01.
INTRODUCTION
Patterns of development vary in different environments.
A typical pattern in a terrestrial environment might range
from uniform natural landscapes to a mosaic of natural
and man-made landscapes. As the demand for land
increases, the landscape is inexorably converted from a
natural to a man-made landscape (Schroevers 1982).
Gasson (1980), at a symposium discussing the manage-
ment needs of False Bay, divided the area into five zones:
the deep water area of the Bay itself; the near-shore zone;
the intertidal zone; the back-shore zone; and the
hinterland. Rivers are an important continuous connection
from the hinterland, through the back-shore and intertidal
areas, to the near-shore environment and, through
dispersal, to the Bay itself. However, very little, if any,
mention was made of the rivers flowing into False Bay
at this symposium.
The quality of much of the water flowing into this
bay has been reduced by developments in the hinter-
land, as shown by the deterioration of the estuarine
environments (O’Callaghan 1990). However, the coastal
environment is also under developmental threat, and has
been for many decades. Calls have been made for an
integrated management policy for the False Bay area
(Malan 1982; Gasson 1980). Very little has, as yet, been
achieved.
Along any coastline, the greatest diversity of habitats
is usually to be found in and around estuaries. Not only
are the coastal terrestrial environments present, but
wetland habitats, saline habitats and combinations of these
are also to be found. Unfortunately, estuarine areas are
sensitive to man-induced disturbances and are often in
great demand for various types of development. Being in
114 Bothalia 20,1 (1990)
TABLE 1. — Aerial photographs of estuaries used for this study. Scale ± 1:10 000. Housed at Trigonometrical Survey, Mowbray, unless otherwise stated
close proximity to a major city, the False Bay estuaries
are no exception. The aim of this study is to ascertain and
quantify, as an aid to management, the changes that have
occurred around the estuaries of False Bay, as described
by O’Callaghan (1990).
METHODS
Delineation of mapping units
Aerial photographs, from 1938 to 1979 (see Table 1),
of each of the estuaries were studied to delineate various
mapping units. It was ensured that, if present, the mapping
units would be distinguishable on all the aerial photo-
graphs. After they had been selected, the homogeneity
and practicability of each unit was checked by field
observation.
The following mapping units were thus defined: 1, water;
2, rock; 3, sand and pioneer vegetation; 4, dune scrub
(Boucher 1987; Moll etal. 1984; Taylor 1980); 5, wetlands;
6, riverine scrub; 7, fynbos (Boucher 1978, 1987; Moll et
al. 1984; Taylor 1969, 1980); 8, alien vegetation (Stirton
1978); 9, agricultural lands; 10, residential areas; 11, recrea-
tional areas; 12, industrial areas; 13, roads and rail.
Either two or three photographs, depending on the
amount of change that had taken place over the years, were
selected to form a temporal series for each estuary. A
Bausch and Lomb Zoom Transfer Scope (Model
53-05-04-23) was used to draw the outlines of the mapping
units for each estuary from colour aerial photographs of
1979, thus producing a land use map for 1979. A copy was
made of this map and the preceding photograph of the
temporal series was superimposed thereon. The land use
map was adapted by redrawing the mapping units so as
to correspond with the preceding photograph. This process
was repeated until land use maps had been drawn for all
the estuaries and all the aerial photographs of a temporal
series.
An area extending approximately 250 m on either side
of the river mouth and 1 km inland from the shore was
selected. The area of each mapping unit within this
selected area was measured and corrected according to
methods set out in O’Callaghan (1985).
The results of this procedure are shown in Table 2. From
this table one can observe the changes in land use that have
taken place at each estuary. These changes can be
measured and compared. They can be related to develop-
mental patterns in different parts of the environment.
However, integrated measurements and environmental
values are often required to compare different areas and
to facilitate decision making.
Evaluation of changes
Numerous techniques exist whereby some measure of
relative environmental importance can be determined
(Linston 1975). These usually require much expertise, time
and money; commodities which are not always readily
available to those who make decisions about the environ-
ment. For this study, these methods were simplified,
making use of some of the principles of more complex
techniques.
The procedures used can be divided into four stages:
1, the determination of the relative environmental
importance (I) of each mapping unit; 2, the determina-
tion of relative environmental states (E) for each estuary;
3, the determination of a co-efficient of change (A) which
is a measure of how much the environmental state of each
estuary has changed over the years; 4, the determination
of a conservability co-efficient (C) which integrates the
above.
1 Relative environmental importance (I)
The assigning of values to parts of the environment is
a subjective process. Values can vary according to
characteristics of the evaluators (Sandbach 1980).
Numerous techniques have been developed to overcome
this subjectivity (Fuggle 1983) and the use of specialist
panels (Helmer 1963) is widely accepted. As these
procedures are costly, a compromise is made by using a
small panel and critically analysing the results.
TABLE 2. — Changes in area (ha) of each mapping unit from each aerial photograph of a temporal series
Bothalia 20,1 (1990)
115
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116
Bothalia 20,1 (1990
TABLE 3. — The relative environmental importance factor (I) of each
mapping unit
Three biologists involved in coastal management and two
botanists involved in coastal ecology were asked to evaluate
the mapping units without the study area being named.
Each assessor was asked to give each unit a relative value
from —10 to 10 according to how an increase in the unit
would affect the functioning of the estuary and immediate
environs (a negative value would mean a detrimental effect
on the environment). The values thus obtained were
averaged for each unit to obtain a relative environmental
importance factor (I) for each mapping unit (Table 3).
Table 3 shows that the panel used for this study
determined that the most important beneficial change that
can occur at an estuary is an increase in wetland
vegetation. Wetland vegetation is generally accepted as
being beneficial to the functioning of an estuary,
performing functions such as flood attenuation, silt
trapping, reduction of excess nutrients, provision of
breeding sites, production of food, and others (see
Walmsley & Botten 1987).
An increase in wetland areas is related to water flow.
An increase in water surface area was regarded as the
second most important beneficial change in an estuary.
Provisos concerning the quality of the water and reasons
for its increase were attached to this selection.
Increases in dune scrub and other terrestrial vegetation
types were given similar ranks with respect to their
beneficial I values. Increases in other natural environments
(e.g. rocks and sand) were perceived as being less
beneficial.
All developments which result in a loss of natural
environments were taken to be detrimental to estuarine
functioning. An increase in industrial areas was regarded
as the the most detrimental. Not only does this type of
development physically destroy the natural environment,
but it also produces pollutants and utilizes water in various
ways.
An increase in farming activities was also regarded as
highly detrimental. The effects of agriculture on the
estuarine environment are similar to those mentioned
above.
Residential developments (including road and rail
developments) and infestation by alien plants were regarded
as having detrimental effects of a similar magnitude.
An increase in recreational areas was regarded as the least
destructive, although it still had a relatively high negative
value. The magnitude of these negative values can be
related to the rate and extent to which these developments
bring about a detrimental effect on the functioning of
estuaries.
2 Relative environmental state (E)
As each mapping unit has a relative importance value,
the relative environmental state of each study site can be
determined for each year, according to the area of each
mapping unit present. This was accomplished by first
adding eleven to each relative importance factor to obtain
positive values. Secondly, each of these positive values was
multiplied by the area of the mapping unit for each estuary
for each year. Thirdly, all the values thus obtained were
summed for each estuary for each year. This sum was
divided by the total corrected area studied at each estuary
for each year to result in a measure of the environmental
state (E) of each estuary for each year of observation (Table
4).
3 Coefficient of change (A)
When planning management policies, it is important to
have an indication of the environmental state and the
amount of change which has taken place at the study site
(as shown above). However, it is particularly useful,
especially for conservation policies (Roome 1984), to
determine the present state of the environment in terms
of its historical state and thus the amount of change that
has taken place relative to each site. This was accom-
plished by establishing a coefficient of change (A) for each
study site. For each mapping unit, the area present in
1938/44 was subtracted from the area present in 1979. This
difference was divided by the sum of the total areas of the
study site for these two years and multiplied by the relative
environmental importance factor (I) for each unit. The
values thus obtained were summed for each study site. This
resulted in a coefficient of change presented in Table 4.
4 Conservability (C)
Even though very little detrimental change might have
occurred at an estuary during this period of photography,
it is possible that the estuary is not well suited to con-
servation because very little natural environment was
present at the time the first aerial photographs were taken.
Calculations thus far do not take the original state of the
estuary into account.
This was rectified by establishing a factor arbitrarily
termed ‘conservability’ (C). This was achieved by obtain-
ing positive values for each coefficient of change by
arbitrarily adding five to each value of A (the values for
A varied from.O to -3,35). This positive value was
multiplied by the environmental state (E) of each study
site for 1938/44. The values of C are presented in Table 4.
CHANGES OBSERVED IN DIFFERENT ENVIRONMENTS
As can be seen from Table 2, profound changes have
occurred in land use around the estuaries entering False
Bay.
Bothalia 20,1 (1990)
Water and wetlands
Water surface area has shown a 17,6% increase during
this period of photography. This could be seen as an
indication of an increase of water entering the rivers from
hard surface run off and sewage works. However, the
factors affecting the surface area of water in an estuary
are numerous and varied. This area can be affected by the
season and time of photography with respect to tides and
annual rainfall variability in the catchment. This variability
can be illustrated by observing the photographs of 1987.
These photographs show that at the Rooiels River, an
increase in water surface area has resulted in a loss of open
sand. The opposite has occurred at the Buffels (East) River
where there has been an increase in open sand areas.
Direct comparisons of water surface area from different
photographs is not recommended unless these variables
can be correlated. On the other hand, wetland areas are
affected by water flow characteristics. Changes in the
extent and distribution of wetlands reflect long-term
changes in hydrological characteristics. Even though there
has been extensive physical destruction of wetlands
(especially at the Sand River), wetlands in this study area
have only decreased by 7,4%. With the exception of
Silvermine, Sand (physical destruction) and Buffels (West),
(at Steenbras wetlands absent), wetlands have increased
at all the rivers. This indicates a long-term increase in the
amount of water flowing into these rivers.
By 1979, a wetland of 7,94 ha had become established
at the Elsies River estuary. Prior to 1910, this river flowed
directly to the sea, or might have formed a small lagoon,
usually in winter, which persisted for varying periods
(E. McKie pers. comm.; D. de Villiers pers. comm.). The
building of rail and road embankments across the river
mouth (± 1910) restricted water flow, although water could
still enter the sea via a culvert at the southern end of the
beach and by seepage. By 1958, the road had been rebuilt
and the base and embankments were presumably
improved. It is likely that this development would have
further reduced seepage, and, together with increased run
off from hard surfaces, caused the wetland to become
established.
Around the Silvermine River estuary, there has been a
reduction of 1,89 ha in wetland areas between 1944 and
TABLE 4.— The environmental state (E) of each study site for each
photograph of a temporal series, the coefficient of change for each
study site (A) and the conservability (C) of each study site (see
text for explanations)
117
1979 (Table 2). During earlier years, the lower part of this
river formed a marshy wetland (Dickson 1974; Burman
1962). However, the river in these parts has been
channelled in recent years (Heinecken 1982a).
Wetland vegetation has disappeared from the study site
at the Sand River. The manipulations which changed the
configuration of this vlei are summarized by Morant &
Grindley (1982). By 1961, 32 ha of vegetated wetlands had
been destroyed by dredging in this vlei and, by 1981, more
than 100 ha of wetlands had been destroyed by further
dredging, and residential and recreational developments.
There has been an increase of 0,50 ha in wetland
vegetation at the Zeekoe River estuary (Table 2). This river
was constructed in 1942 as an overflow channel for
Zeekoevlei (Bickerton 1982). By 1944, an estuarine lagoon
had formed on the beach. The coastal road (Baden-Powell
Drive) had restricted this lagoon to the hind-dune area by
1958 and the development of the sewage works in the ear-
ly 1960s (Summers et al. 1976) increased the water flow
in this river, thus allowing the wetlands to increase.
The increase in wetlands measured for the Eerste River
estuary between 1938 and 1958 could be related to the
decrease in water surface area for this period (Table 2).
However, there was a large increase in water surface area
between 1958 and 1979, caused by increased run off due
to developments in the catchment and the discharge of
treated sewage effluent.
Prior to a bridge being built across the Rooiels River
mouth in the early 1950s, this river followed a meandering
course across the floodplain (Heinecken 1982b). This
bridge relocated the main flow channel towards the
northern side of the floodplain. A meander was re-
established on the seaward side of the road and the
damming effect of this bridge brought about a slight
increase in wetland area (Table 2).
In 1938, the Buffels (East) River formed a large
backshore lagoon. The considerable reduction of this
lagoon was probably facilitated by the enlargement of the
Buffels Dam in 1972 (Heinecken et al. 1982).
Dune vegetation
During this period, units 3 (sand and pioneers) and 4
(dune scrub) have decreased by approximately 50% each.
The most widespread reason for this decrease is the growth
of alien plants, which were found at all the rivers by 1979.
At the Cape Flats estuaries (Zeekoe, Eerste, Lourens),
industrial development is the most important factor
contributing to the demise of dune vegetation. At
Silvermine, Sand, Rooiels and Buffels (East), residential
developments were most important with recreational de-
velopments at Sir Lowry’s Pass. These developments are
discussed in greater detail below.
Riverine scrub and fynbos
These vegetation types were found mainly around rivers
on the eastern shores of the Bay. Little development has
taken place around these rivers and fynbos and riverine
scrub have hardly been affected.
118
Bothalia 20,1 (1990)
Less fynbos was present near the estuaries of the western
shore in 1938. Around these rivers (Buffels (West), Elsies,
Silvermine), fynbos has decreased by 60,4%, mainly as
a result of alien plant invasion and residential development.
Fynbos was not found around the river mouths of the Cape
Flats.
FACTORS CAUSING CHANGES
Alien vegetation
By 1979, alien plants, mainly Australian acacias, were
found at every study site (Table 2). Acacias were in-
troduced into the Cape between 1845 (Shaughnessy 1980)
and 1870 (Roux & Middlemiss 1963).
Official bodies, local authorities as well as central
government organizations, planted alien acacias at
numerous places along this coast, among others: 1885,
Eerste River Mouth (Shaughnessy 1980); 1893, Retreat
Station, north of the Sand River (Shaughnessy 1980); 1936,
Zeekoe River, vicinity of mouth (Bickerton 1982); 1942,
Sand River coast (Shaughnessy 1980).
In addition to the above plantings, Opie (1967) maintains
that aliens were planted by a farmer at the Buffels (West)
River during the mid-19th century to stabilize deep
sands in the area (Coke 1963). Acacias might also have
been planted at the lower Elsies River by the De Villiers
family who have farmed the area since the 1870s. Although
this farmer planted an unknown Hakea species (probably
H. sericea) at the Elsies River and acacias on the western
coast of the Peninsula, no record of the planting of
acacias at the Elsies River exists (D. de Villiers pers.
comm.).
The presence of alien plants at other sites is likely to
have occurred by natural encroachment (Glyphis et al.
1981). At the lower Silvermine River, major encroachment
seems to have occurred from the south where these plants
are still used for hedges, windbreaks and shade in Fish
Hoek town, established in 1918 (Burman 1977). At the Lou-
rens River estuary, alien plants appear to have entered the
study site along the river course from neighbouring farms.
At the lower Steenbras River, alien plants were first found
on disturbed areas near the road, while at the Rooiels and
Buffels (East) estuaries, the presence of these plants can
be related to the advent of residential development.
Residential development
In the area studied, there has been a 238% increase in
residential areas, most of this development has taken place
on the hillsides along the western and eastern shores of
the Bay.
The Buffels (West) River lies within the Cape of Good
Hope Nature Reserve, proclaimed in 1938. Consequently,
development has been restricted to the creation of picnic
sites around the tidal pool to the south of the river.
Much residential development has taken place around
the lower Elsies River (93% increase), Silvermine River
(186% increase) and Sand River (61% increase). These
developments took place mainly at the expense of fynbos
and the open sand/pioneer vegetation, although the latter
had often been dominated by alien vegetation prior to
residential development. At the Sand River, most of the
development has taken place on filled-in wetland
vegetation.
No residential development has taken place around the
estuaries of the central Cape Flats, but an increase in
housing of 272% and 312% has occurred respectively at
the Lourens and Sir Lowry’s Pass estuaries, mainly at the
cost of dune scrub and agricultural areas. Some minor
developments have taken place at the lower Rooiels and
Buffels (East) Rivers. These are mainly in the form of
seaside cottages.
Recreational development
Recreational developments are closely allied to
residential development. Some minor development has
taken place around the Buffels (West) and Elsies Rivers.
There has been a major increase in the percentage
recreational area at the Silvermine River in the form of
the golf club. The golf course covers approximately 3 ha
of what was previously farmland, open sand and dune
scrub. The 504% increase in recreational area at the Sand
River has been in the form of tended parklands on what
was previously wetlands. A caravan site of 3,8 ha has been
developed at the lower Sir Lowry’s Pass River at the
expense of dune scrub and open sand. Although the steep
topography around the Steenbras River is not suited to
residential development, recreational development has
proceeded in this area and, in 1979, consisted of 1,3 ha
of the study site. Minor developments in the form of picnic
sites have taken place at the lower Rooiels River.
Industrial development
Major industrial development seems to be restricted to
the rivers of the Cape Flats, away from the major
residential and recreational areas. At the Sand River, a
small nursery has been established near the head of the
vlei, hardly compatible with residential and recreational
development. The major industrial developments have
been in the form of sewage treatment works: 22,10 ha at
the lower Zeekoe River and 26,09 ha at the lower Eerste
River. The 8,65 ha of industrial development at the Lourens
River is in the form of a buffer and security zone around
a chemical and explosives factory. Although very few
buildings have been erected in this area, it is prone to
environmental destruction and invasion by alien plants.
DISCUSSION OF INDIVIDUAL ESTUARIES
Buffels (West)
The low conservability of this estuary can be explained
by a single factor: the encroachment of alien plants. Early
aerial photographs and historical records show that this
river valley consisted of a sand plume, and sediment
interchange took place with the near-shore environment
(Heinecken et al. 1982). Sparse dune pioneers and dune
scrub would have been found on this sand. Between 1944
and 1978, open sand and pioneers decreased by 71,5%;
dune scrub decreased by 88,2% ; and fynbos decreased by
58,3%. During this period, alien plant cover increased by
185,8%.
Bothalia 20,1 (1990)
119
The removal of alien plants from this reserve is an
ongoing policy of the local authorities, but with varying
success rates. MacDonald et al. (1989) quote reports which
state that this area was completely cleared of aliens in 1945
and again in 1959. However, the 1979 photographs show
that the area was again heavily infested. The 1987 colour
aerial photographs show that approximately 2 ha of alien
plants had been removed from this site and dune pioneers
and scrub had re-established. These developments will
serve to increase the conservability of this river.
Elsies
At this estuary, major impacts have been due to the
encroachment of alien plants (114,9% increase) and
residential developments (including roads: 108,2%
increase). Open sand and pioneers, and dune scrub have
decreased by 67,8% and 78,4% respectively. These changes
should have had the effect of somewhat decreasing the
conservability. However, a wetland has formed in areas
which were previously sand and dune vegetation. As
wetlands were given a higher relative importance factor
(I) (Table 3), this development has ameliorated the
decrease in conservability.
Photographs taken during 1987 show that little change
has taken place at this site. However, residential develop-
ment is taking place in the vicinity which could bring about
an increased run off from hard surfaces, further affecting
the wetland.
Silvermine
Around the lower part of this river, open sand and
pioneer vegetation, and fynbos has decreased by 79,9%
and 40,9% respectively. Residential and recreational
developments have increased by 186,5% and 157,4%
respectively. However, much of this development has taken
place in areas which were previously dominated by alien
plants. This has led to a net decrease in alien plant cover
of 0,4% over the period covered by the aerial photography
(in 1958, alien plant cover was 38,5% greater than in 1979).
Furthermore, dune scrub has established in areas which
were previously open sand. These environmental changes
ensure that the conservability of this area remains relatively
high.
Aerial photographs of 1987 show that a further 5 ha of
dune vegetation has been destroyed by residential
developments.
Sand
The major development that has taken place at the Sand
River is a 527 % increase in recreational areas. This has
led to the almost total elimination of wetland vegetation
and a 79,7 % decrease in dune scrub. Some of the residen-
tial (60,7% increase) and recreational development has
taken place in areas that were previously dominated by
alien vegetation, resulting in a decrease in alien vegeta-
tion of 96,1%.
The conservability of this estuary is comparable to those
of the other four estuaries of the Cape Flats (Zeekoe,
Eerste, Lourens, Sir Lowry’s Pass). The reasons for the
low value of C for the Sand River are, however, unique,
due primarily to recreational developments.
Zeekoe
This estuary has the lowest conservability and the
highest coefficient of change. The major reasons for this
are the establishment of sewage settling ponds (22,1 ha)
and a 126,6% increase in alien vegetaion. Dune scrub and
open sand and pioneers have decreased by 66,7% and
42,9% respectively.
The construction of the settling ponds destroyed large
areas of dune scrub, but they are no longer used, except
possibly in emergencies. They have become an important
coastal habitat for water birds and recommendations have
been made for these ponds to be proclaimed as a nature
reserve (Curtin et al. 1975; Cooper et al. 1976; Brummer
1981; Bickerton 1982). If this proclamation were to take
place, these ponds might be regarded as open water. This
estuary would then be the only one along False Bay with
a positive coefficient of change, resulting in a con-
servability of 80,8 % . This would be an artificially induced
beneficial change and might serve to mask detrimental
changes in the area.
The 1987 photographs show that approximately 4 ha of
land have been cleared, presumably for low-cost housing.
This has brought about a loss of dune scrub and some alien
vegetation.
Eerste
The reasons for the low conservability of this estuary
are similar to those discussed for the Zeekoe River estuary:
the establishment of a sewage works of 26,09 ha. This
development resulted in a decrease in dune scrub and open
sand and pioneers of 81,3% and 74,1% respectively. Alien
shrubs have long been established in this area
(Shaughnessy 1980) and the construction of this sewage
works served to decrease alien vegetation by 43,9%.
Another factor serving to increase the conservability is
a 668,7% increase in estuarine water surface area.
Although tidal and climatic factors might have led to this
observed increase of water surface area on the photo-
graphs, large areas of the catchments of the Eerste River
and Kuils River (a tributary) have become developed over
the last decades (Grindley 1982). Increased run off from
hard surfaces as well as effluent discharge from at least
five sewage works in this catchment have undoubtedly
increased the amount of water flowing into the sea via this
estuary.
At this river, photographs taken in 1987 show that
approximately 2 ha of alien plants have been cleared in
what was previously classified as industrial areas.
Lourens
There has been a 4 468,4% increase in industrial area
and a 272,6% increase in residential area around this
estuary. In addition, there has been a 74,2% demise of
dune scrub. These developments would serve to decrease
the conservability. However, they have resulted in a net
120
Bothalia 20,1 (1990)
decrease of alien vegetation (59,4%) and agriculture which
has disappeared from this area. There has also been a
slight increase in open sand and wetland vegetation, factors
which would increase the conservability.
Nevertheless, the conservability of this estuary was
relatively low when aerial photography began in 1938. The
conservability of this estuary remains low, even though
the coefficient of change (A) is lower than for most of the
estuaries of the Cape Flats.
The 1987 photographs show that some eradication of
alien plants has taken place within industrial areas. But
outside of these areas, approximately 1 ha of open sand
has been consolidated by alien shrubs.
Sir Lowry’s Pass
Even though the coefficient of change for this estuary
is close to zero, this estuary does not have a high
conservability. This is because much of the area had been
developed by the time of onset of photography for this
study. In 1938, most of this area was used for agricultural
purposes. By 1979, these agricultural areas had been
converted to residential areas. Between 1938 and 1961,
dune scrub had become established on much of the open
sand and pioneer vegetation. By 1979, this dune scrub had
largely been destroyed for recreational purposes.
Steenbras, Rooiels and Buffels (East)
Most of the changes that took place around these
estuaries were discussed above. These changes are rela-
tively minor, as can be seen from the high con-
servability values. However, they all show slight increases
in alien plant cover, residential and/or recreational areas,
with decreases in the natural dune vegetation.
Photographs taken in 1987 show that these developments
are ongoing. Some recreational and residential
development has taken place at the Steenbras and Buffels
(East) Rivers. There have also been slight increases in alien
scrub at the Rooiels and Buffels (East) Rivers. Soon after
these latest photographs were taken, major upgrading of
the main thoroughfare across the Rooiels River was com-
menced. The stage could be reached when these
developments may destroy the attractiveness of these areas,
i.e. the natural environment. This, unfortunately, is largely
the pattern observed along the western shores of the Bay.
COMPARISON OF ESTUARIES
The effects of development can be determined by using
the descriptive methods discussed under Methods:
evaluation of changes. However, quantitative measure-
ments, or environmental values, are often needed to
compare different areas and to facilitate decision-making.
Table 3 shows that the environmental state of all the
estuaries decreased during the period spanned by aerial
photography. When the first aerial photographs were taken,
the most easterly rivers of the Cape Flats (Lourens and
Sir Lowry’s Pass) had the lowest E value ( 9,59 and 6,22
respectively) whereas the highest E value was found at the
Buffels (East) River (16,17). In 1979, this range for E values
was similar (6,19 at Sir Lowry’s Pass; 15,45 at Rooiels).
However, the environmental state of the other rivers of the
Cape Flats (Sand, Zeekoe, Eerste) had deteriorated so
as to be in accordance with Lourens and Sir Lowry’s
Pass. The state of the Buffels (West) River had also
deteriorated drastically.
This deterioration of the environmental states of these
rivers is further demonstrated by the coefficient of change
(A). The value of A for all the rivers is negative for the
period covered by the aerial photography. The most change
has taken place at the Zeekoe River (A = —3,35). The
changes that have taken place at the lower Rooiels River
are minor and A is given as zero.
Although conservability (C) is a relative factor, it is
possibly the most important as it integrates all the factors
calculated above. The lowest value of C is seen at the
Zeekoe River estuary (23,13) where much industrial
development and invasion by alien plants has taken place.
The Rooiels River estuary has the highest value for C
(77,50). Only a small amount of recreational and
residential development has taken place at this estuary and
the environment retains a healthy state.
The estuaries of the Cape Flats (Sand to Sir Lowry’s
Pass) have the lowest C values. The C values for the
estuaries on the western side of the Bay (Buffels (West)
to Silvermine) are lower than those on the eastern side
of the Bay (Steenbrass to Buffels (East)). This is under-
standable as, being closer to the city of Cape Town, the
western side of the Bay would be in greater demand for
housing and other developments.
CONCLUSIONS
Aerial photographs are readily available for large parts
of the coast. For False Bay, these photographs span a
period between 1938/44 and 1987. The environmental
coefficients presented here are easily and rapidly
calculated. They give quantitative indications of the state
of the environment and can be used to facilitate
decision-making and policy formulation. For example,
using the data presented above, management of the most
easterly rivers of False Bay should be orientated
towards conservation while smaller sanctuaries could be
established at the Elsies and Silvermine Rivers,
particularly the wetland areas. The rivers of the Cape Flats
should be managed as developed areas and attempts should
be made to remove alien plants from the Buffels (West)
River.
A similar exercise can be carried out for the entire
coastline of False Bay. Communication between local
governmental bodies could establish an integrated manage-
ment policy for this coast with little expenditure in terms
of time, effort and equipment.
The results of such an exercise must, however, be
analysed critically. Without some understanding of the
reasons for changes in these environmental factors,
erroneous conclusions can be reached. For example, a
large part of the observed increase in water surface area
at the Eerste River is due to an increased run off from
developments in the catchment. The observed increase in
water surface area at the Rooiels River is more likely due
to climatic phenomena. Furthermore, if it was decided that
Bothalia 20,1 (1990)
121
residential developments were more detrimental to the
environment than agricultural developments, the environ-
mental state (E) of the Sir Lowry’s Pass River would have
decreased from 1938 to 1961. Any statement concerning
the importance of any part of the environment is subjective
and could thus be a source of error. Although data could
be collected by non-specialists, it is recommended that
environmental evaluation be carried out by specialists or
evaluation panels (Fuggle 1983).
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BOUCHER, C. 1978. Cape Hangklip area. II. The vegetation. Bothalia
12 : 455 -497.
BOUCHER, C. 1987. A phytosociological study of transects through the
western Cape coastal foreland. South Africa. Ph.D. thesis, Univer-
sity of Stellenbosch.
BRUMMER, T.B. 1981. A development plan for the Zeekoevlei complex.
M.Sc. thesis. University of Stellenbosch.
BURMAN, J. 1962. Safe to the sea. Human & Rousseau, Cape Town.
BURMAN, J. 1977. The False Bay story. Human & Rousseau, Cape Town.
COKE, M. 1963. An ecological study of the extent of alien infestation
of the Good Hope Nature Reserve and its spread over the years
1952-1961. Botany Department, University of Cape Town.
Unpublished.
COOPER, J., SUMMERS, R.W. & PRINGLE, J.S. 1976. Conservation
of coastal habitats of waders in the south-western Cape, South
Africa. Biological Conservation 10: 239-247.
CURTIN, R., AQUADRO, D., HILL, R & UPSHER, S. 1975. Manage-
ment proposals for Zeekoevlei. School of Environmental Studies,
University of Cape Town. Unpublished.
DICKSON, D. 1974. A short history of Clovelly Country Club. Clovelly
Country Club. Unpublished.
FUGGLE, R.F. 1983. Environmental evaluation. In R.F. Fuggle & M.A.
Rabie, Environmental concerns in South Africa : 483—515. Juta,
Cape Town.
GASSON, B. ed. 1980. Proceedings of seminar on The future manage-
ment of False Bay. Cape Town.
GLYPHIS, J.P., MILTON, S.J. & SIEGFRIED, W.R. 1981. Dispersal
of Acacia cyclops by birds. Oecologia 48: 138—141.
GRINDLEY, J.R. 1982. Report No. 16: Eerste (CSW 6). In A.E.F.
Heydom & J.R. Grindley, Estuaries of the Cape, Pan II: synopses
of available information on individual systems. CSIR Research
Report No. 415.
HEINECKEN, T.J.E. 1982a. Report No. 13: Silvermine (CSW 3). In
A.E.F. Heydorn & J.R. Grindley, Estuaries of the Cape, Part
II: synopses of available information on individual systems. CSIR
Research Report No. 412.
HEINECKEN, T.J.E. 1982b. Report No. 8: Rooiels (CSW 10). In A.E.F.
Heydom & J. R. Grindley, Estuaries of the Cape, Pan II: synopses
of available information on individual systems. CSIR Research
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HEINECKEN, T.J.E. BICKERTON, I.B. & MORANT, P.D. 1982.
Report No. 12: Buffels (wes)(CSW 1), Elsies (CSW 2), Sir
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55 : 56 -75.
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'
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Bothalia 20,1: 123-125 (1990)
Miscellaneous note
VARIOUS AUTHORS
NOTES ON THE DISTRIBUTION AND HABITAT OF ALOE BOWIEA (LILIACEAE/ASPHODELACEAE: ALOOIDEAE),
AN ENDANGERED AND LITTLE KNOWN SPECIES FROM THE EASTERN CAPE
A. bowiea is an extremely rare species only known from
a limited area in the eastern Cape. Under conditions of
normal rainfall it is a dwarf rosulate leaf succulent (sensu
Newton 1974), reaching a height of 300 mm at the
flowering stage.
Morphologically and taxonomically A. bowiea is
insufficiently known. A study on these aspects is in
progress and will be reported elsewhere.
HABITAT
Geographical situation
Aloe bowiea is known only from the three study sites
discussed below (Figure 1). Locality 1 is located south
of Uitenhage, between Uitenhage and Despatch, on the
form ‘Jachtvlakte’. Locality 2 is situated at Coega, approxi-
mately 25 km east of Uitenhage, on ‘Maasward’, a private
nature reserve bordering the Uitenhage— Sundays River
mouth road. Locality 3 is on the farm ‘Brakkefontein’, 20
km from the Kirkwood turn-off on the Uitenhage-Jansen-
ville road. These localities will subsequently be referred
to as Uitenhage, Coega and Kariega, respectively.
Although the population of A. bowiea at Uitenhage, still
thriving in 1983, is now considered extinct, this locality
will be included in this paper. Hitherto this species was
only known from the latter locality.
Climate
The climate at the three study sites is warm temperate
and conforms to the semi-arid valley climate recognized
by Cowling (1983a) for the river valleys of the south-eastern
Cape.
At Uitenhage the average annual rainfall is 435 mm with
monthly maxima in April, August and November and
minima in the summer and winter months. Temperatures
are moderate with all months being between 13° and 23 °C.
Coega and Kariega (Figure 2: Hermitage and Hillside
Farm respectively) experience a warmer and more arid
FIGURE 1. — Map showing the
regional limits of the eastern
Cape (after Lubke et al. 1986)
and the three known localities
of Aloe bowiea. In recent years
the species has become extinct
at locality 1.
124
Bothalia 20,1 (1990)
UITENHAGE 132 ml I8,I°C 435 mm HERMITAGE 1122 m) I8,3°C 373 mm
HILLSIDE FARM 1101 ml I8,6“C 339 mm
FIGURE 2.— Walter-Lieth climate diagrams of selected stations. Data
from Weather Bureau (1984).
climate. Rainfall is generally erratic with an average annual
figure of 373 mm at Hermitage and 339 mm at Hillside
Farm. Both sites have a bimodal distribution of rain, with
peaks in autumn and late spring. At these sites the mean
monthly temperatures show a slightly greater range
(13-24°C) with maximum temperatures exceeding 40°C
and minima dropping below 0°C.
Vegetation
Field observations at all the localities have shown that
Aloe bowiea is restricted to the Sundays River Scrub
variation of Valley Bushveld [Veld Type 23 sensu Acocks
1988)] of the Uitenhage, Coega and Kariega regions in the
south-eastern Cape. Acocks (1988) did not specifically
mention the vegetation surrounding Uitenhage, Coega and
Kariega, but at all localities it would, on the ground of
shared Euphorbia species, fit into his Sundays River Scrub
variation. On the map which accompanied Acocks (1988)
the vegetation of this area is given as Valley Bushveld.
Other authors have referred as follows to the vegetation
with which A. bowiea is associated: Thicket (Jessop &
Jacot Guillarmod 1969), Addo Bush (Penzhorn & Olivier
1974), Valley Bushveld (sensu Olivier 1977, 1981; Moll et
al. 1984), Subtropical Transitional Thicket (Cowling 1984;
Everard 1987), Subtropical Transitional Thicket, including
Valley Bushveld (sensu Lubke et al. 1986) and Subtropical
Thicket (Van Wyk et al. 1988a, 1988b). This vegetation
type is confined mainly to the valleys of rivers which flow
east and drain into the Indian Ocean, and currently covers
approximately 15% of the eastern Cape (Everard 1987).
These valleys are relatively hot and dry in comparison to
the intervening ridges.
The Valley Bushveld vegetation at both the Uitenhage
and Coega localities included in the present study forms
a dense and, where undisturbed, almost impenetrable
thicket. It agrees with the differentiating characters given
by Everard (1987) for Mesic Succulent Thicket in having
a mean total cover of about 90% , an average canopy height
of about 2,5 m and a large proportion of succulents (more
than 20%). At the Kariega locality the vegetation is less
dense (average total cover of approximately 60%) with a
slightly lower shrub canopy (2,0— 2,5 m) and succulents
comprise about 30% of the species. This locality falls
within the Xeric Succulent Thicket suborder. Succulents
associated with A. bowiea include, at Uitenhage: Aloe
africana Mill., A. striata Haw., Bulbine caulescens L.,
B. latifolia Roem. & Schult., Euphorbia stellata Willd.,
E. meloformis Ait., E. globosa (Haw.) Sims and
Pachypodium succulentum (L. f.) A. DC.; at Coega: Aloe
ferox Mill., Gasteria maculata (Thunb.) Haw., Haworthia
xiphiophylla Bak., Euphorbia meloformis, E. clava Jacq.,
E. ledienii Berger and Pachypodium succulentum and at
Kariega: Bulbine latifolia, Haworthia sordida Haw. (M.B.
Bayer pers. comm.), and Haworthia sp. cf. cooperi Bak.
At all three localities species of Cotyledon L. , Crassula
L. and the Mesembryanthemaceae occur in large numbers.
Although the vegetation at the three localities where Aloe
bowiea is known to occur can be broadly classified as
thicket, plants of this species have never been encountered
in the shade of surrounding vegetation. It seems to be
restricted to ecological niches where the valley bush opens
up naturally to form a less dense karroid/grass community.
These breaks in the canopy layer do not appear to be
man-induced (at least in recent times) since they also occur
in nature reserves which lie within valley bush, such as
Springs Reserve at Uitenhage and Maasward Private
Nature Reserve at Coega. As a result of litter decomposi-
tion the soils on which thicket occurs usually contain high
levels of organic matter. Aloe bowiea was not recorded
from these humic soils.
ENDEMISM
Everard (1985) lists only three species of Aloe as being
endemic to the eastern Cape, namely, A. africana, A. tid-
marshii and A. bowiea. On the other hand Cowling (1982,
1983b) states that five of the 25 species of Aloe indigenous
to the eastern Cape are endemic to this region. For A.
bowiea Cowling (1982, 1983b) prefers to uphold the name
Chamaealoe africana. The discrepancies in the literature
with regard to the number of Aloe taxa indigenous or
endemic to the eastern Cape (Cowling 1982, 1983b;
Everard 1985; Gibbs Russell & Robinson 1981; Holland
1978) can be explained by the different delimitation of the
eastern Cape by the different authors.
CONSERVATION
According to Everard (1985) the eastern Cape currently
has 662 threatened taxa. Of these, 485 are listed in the
category uncertain whether safe or not. This conserva-
tion status category is defined as one for plants that are
so little known that there is an even chance that they could
prove to be safe. Everard (1985) listed Aloe bowiea in this
category. Field observations during recent years have,
however, revealed only three localities for this species.
Moreover, the population of A. bowiea at Uitenhage is now
extinct and at both Coega and Kariega the populations are
vulnerable and declining. With the exception of the Coega
locality, where A. bowiea occurs in a privately owned
nature reserve, none of the known populations are
conserved. This species takes readily to cultivation, but
it is horticulturally unattractive and poorly represented in
Bothalia 20,1 (1990)
125
succulent plant collections. The distribution of known
populations of A. bowiea is shrinking fast. For example,
whereas in 1983 the population at Uitenhage comprised
141 individuals, no plants could be found at this locality
in 1988.
CONCLUSION
Recent field observations have shown that Aloe bowiea
has a very limited geographical distribution and that it is
now extinct at Uitenhage, the only previously known
locality. At the other two localities, Coega and Kariega,
the populations of A. bowiea are endangered and should
be added as such to the list published by Everard (1988).
It is recommended that populations be securely fenced
to prevent game animals and domestic stock from grazing
individuals of this species. This would allow plants to
flower and set seed and could ultimately lead to the
expansion of existing populations and the establishment
of new ones. Unless immediate action in the form of land
acquisition and the education of land owners are taken,
this unique and localized species of the karroid vegetation
of Valley Bushveld may soon be extinct in nature.
ACKNOWLEDGEMENTS
Discussions with Prof. G.J. Bredenkamp contributed
substantially to the formulation of a conceptual basis for
this paper. Many of the observations reported here were
made while the first author was an undergraduate student
at the University of Port Elizabeth and he therefore makes
acknowledgement to this institution.
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WEATHER BUREAU 1984. Climate of South Africa. Climate Statistics
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Government Printer, Pretoria.
G.F. SMITH* and A.E. VAN WYK**
* Department of Plant Sciences, Potchefstroom University for C.H.E.,
Potchefstroom 2520.
** Department of Botany, University of Pretoria, Pretoria 0002.
MS. received: 1989.06.29.
'
.
La
Bothalia 20,1: 127-130 (1990)
OBITUARY
MARY DAVIDSON GUNN (1899-1989)
The death of Mary Gunn on 31 August 1989 marked
the passing of the last of a trio known with affection in
the South African botanical world as the Three Graces.
Together with her colleagues, taxonomist Inez Verdoorn
(1896—1989) and botanical artist Cythna Letty (1895 —
1985), Mary Gunn formed a remarkable team, each
member of which left an indelible mark on their respective
field of botany over a working period totalling more than
150 years.
Mary Gunn was bom in Kerriemuir, Scotland on 15 May
1899 (Figure 1). Her father served in a Scottish regiment
in South Africa during the Anglo-Boer War and on
returning home decided to immigrate to this country where
he settled in Pretoria.
After attending the Gymnasium School, Mary Gunn
joined the Division of Botany as a clerk with a ‘knowledge
of typing’ as her letter of appointment indicates. It is clear
from this letter that appointments were expeditiously
executed at the time (Figure 2).
However, the new appointee was not constantly busy and
to keep herself occupied, she would dismantle and exa-
mine the intricate workings of her wrist watch. Caught
in the act by her sometimes irascible chief Dr Illtyd Buller
Pole Evans, her immediate superior was instructed to give
the young Scottish lass the books to take care of to fully
utilize her working hours. This was to prove an inspired
directive.
Mary was immediately interested in her new task
although the collection of books was so small that there
were not enough to fill even one shelf in the small room
of the Division headquarters in ‘Vredehuis’ (Figure 3). On
her return home each day she was enthusiastically
encouraged by her father to describe each of the books
she had catalogued. This daily ritual probably played a
large part in the development of the remarkable memory
which was to be one of her greatest attributes in the years
to come.
The young librarian (Figure 4) with no formal training,
was so excited by the world of botanical books to which
she had so fortuitously been introduced that she began
studying catalogues to establish which old works were
available and how she could acquire them. She rapidly be-
came an authority on old botanical literature and ascribed
her future years of dedication largely to the pleasure der-
ived from some of her first acquisitions. Heading the list
was P.J. Bergius’ Descriptiones plantarum ex capite bo-
nae spei. This book, which was purchased in the 1920s
from a German bookseller for a small sum, is inscribed
W.H. Harvey 1838. But the real thrill about the acquisi-
tion was the fact that it had belonged to a young German,
Georg Forster, who with his father had sailed on the ‘Reso-
lution’ with Captain Cook on his second circumnavigation
of the globe in 1772.
The delight of obtaining this type of Africana was to
remain with Mary Gunn throughout her life. Many other
noteworthy publications were to be purchased in the years
to come. The majority of these early works were published
privately and the limited copies were usually presented
to patrons and friends. Colour printing was unknown and
illustrations had to be hand-coloured in water colours.
Fortunately Mary Gunn realized the value of works of this
nature many years before it was generally appreciated in
other parts of the world.
Because of very limited funds, various ways and means
had to be devised to acquire these treasures and Mary
frequently had to resort to her considerable persuasive
powers to fulfil her wishes. In her own words ‘I frequent-
ly had to go hat-in-hand asking for money from influen-
tial people so that I could buy what I wanted!’.
Possibly one of the most notable and valuable works
acquired was Redoute’s Les Liliacees. The four volumes
were purchased in England for £200 and transported to
South Africa in the baggage of General J.C. Smuts after
he had attended the signing of the treaty of Versailles in
1919. General Smuts was a great friend of Pole Evans and
a frequent visitor to the Division of Botany.
FIGURE 1.— Mary Davidson Gunn (1899—1989).
128
Bothalia 20,1 (1990)
OF SOUTH
"i -Arx&rlaa Street
hi'-stcria.
for <i U'i..;c;tr y swaia*., -:it >.ith <; «cr.o«3'.dg8
:o:r typing txint* ... 2 C-.ice :;f the C:i*f, Division
ox trio : r *.*■* . -at . Tbs isi'.ri *1 remtme ratio:
‘ l'-‘t &• i«as txissTi b/o r— ^ -or king day. ’i.-.ur name
ixaa css', fvrnis;. : j, t ■ a.s by U.e Public Tice Qoaa.in-
si oners a. u,lr you are. ao.jiroua of off a Hag your
.services, will you iciauSy call ar.u see ics . of
if18 *&3 ;'i v'* ;ion Tba of ^-33 ij situated *«&t ; .
b«lo:v i he ;/;ii axi '-'X : Idln^a „
M< f- —
’ y ~''
$-»*&: 'AGSICl’LTtrKB «
FIGURE 2. — Letter of appointment of Miss Gunn to the Division of
Botany in September 1916.
FIGURE 4. — The young librarian.
FIGURE 3. — Women Staff- Botany
& Plant Pathology, 1919. Back
row L to R: Misses V.
Bottomley, E. Tenant, K.
Lansdell, A. Bottomley, Z.
Findley, Dr E.M. Doidge,
Miss S.M. Stent. Sitting L to
R: Miss I. Verdoorn, Miss
King, Mrs Stocks, Misses I.
Tambline, E. Linde, M.D.
Gunn, K. Vos.
It is of interest to enumerate a few of the other, today
almost priceless treasures which, through meticulous
research and personal determination and endeavour, Mary
Gunn was able to acquire for the library:
In G. Rondelletii . . . methodicam pharmaceuticum offici-
nam animadversiones (1605) M. de l’Obel
Florilegium (1612) E. Sweert
Exoticarium plantarum (1670) J. Breyne
Horti medici amstelodamensis (1697—1701) J. Commelin
Plantae selectae (1750—1773) C.J. Trew
Geraniologia (1787—1788) C.L. L’Heritier
leones plantarum rariorum (1781—86) and Oxalis (1794)
N.J. Jacquin
The Botanists repository (1797—1815) H.C. Andrews
(Figure 5)
Medicinal plants of the Cape (1857) C.W.L.Pappe
Bothalia 20,1 (1990
129
purchased personally by Mary Gunn in England before
1939 and held by the booksellers until the end of World
War 2 when they could be safely delivered to Pretoria.
Dr Illtyd Pole Evans, chief of the Division of Botany
from 1913 to 1939 had a profound influence on the
botanical fields in which he became involved and he
similarly influenced and inspired many of his staff. His
continual queries about botanical explorers led Mary Gunn
into her second field of specialization, namely biographical
research of early botanists. Information about travellers
such as Thunberg, Burchell, Masson, Ecklon and many
more was diligently collected from all comers of the globe.
A cherished ambition was finally realized in 1981 when
the major part of all this information was used for the
publication Botanical exploration in southern Africa. This
work, produced in collaboration with her one-time chief
Dr Leslie Codd, was the culmination of more than 65 years
of research.
In spite of her redoubtable knowledge this was to be
her only really significant personal publication. With Enid
du Plessis she introduced and edited The Flora capensis
of Jakob and Johann Philipp Breyne. This was a
publication of original 17th and 18th century water-colour
drawings contained in an album which is one of the
Africana treasures of H.F. Oppenheimers’s Brenthurst
Collection.
FIGURE 5. — Title page of the Botanists Repository , one of the priceless
treasures in the Mary Gunn Library.
Genera of South African plants (1838) Harvey, generally
considered to be the first scientific work published in
South Africa. This book, purchased in Johannesburg for
± R9,00 is of special interest as it contains handwritten
notes by the author.
Notable series acquired included a full set of Curtis ’s
Botanical Magazine published from 1790 onwards and the
exceptionally valuable Loddiges’ Botanical Cabinet. These
beautiful leather-bound, gold-leaf-edged volumes were
Apart from these two works her publications were
limited to tributes or notes on botanical colleagues or
personalities. She had always hoped to publish an account
of the adventurous Colonel Robert Gordon who was
commandant of the Dutch Garrison at the Cape from 1780
until 1795 when he committed suicide after surrendering
to the British. The history of this colourful soldier and
naturalist was particularly dear to her heart. When she
realized, in her eighties, that she would never be able to
utilize her information it was passed on to other historians.
This assistance was acknowledged by Raper and Boucher
in their two volumes on Gordon which were published in
1988. Her generosity in passing on the results of a
FIGURE 6. — Official opening of the
Mary Gunn Library, 15th
January 1970. From left to
right: Dr B. de Winter, Deputy
Director, Botanical Research
Institute; Dr J.W. Geyer, Chief
Director of Research, Depart-
ment of Agriculture; Miss
M.D. Gunn and Dr L.E.
Codd, Director, B.R.I.
130
Bothalia 20,1 (1990)
lifetime of research was one of the most noteworthy
attributes of this remarkable woman. Numerous
publications such as Aloes of South Africa (Reynolds),
Trees of southern Africa (Palmer) and There was a man
(Gutsche) bear witness to this quality.
In dedicating Volume 32 of The Flowering Plants of
Africa series to Mary Gunn in 1957/58, editor Dr Leslie
Codd writes: Librarian, authority on Africana, who during
her 40 years of service in the Department of Agriculture,
has with unremitting devotion built up a comprehensive
botanical library in the Division of Botany, who has
assisted so many research workers in tracing literature
relevant to their problems and who has since its inception
fostered the best interests of "The Flowering Plants of
Africa'.
After retiring officially in 1954 she continued working
in a temporary capacity until 1973 but until a few years
before her death she still enjoyed spending a few hours
a week in the privacy of her office. Here she would peruse
her personal collection of books and be available to readily
offer advice or search for information for anyone who
needed her expertise.
In 1976 she was honoured with the award of The Bolus
Medal which recognizes outstanding achievement in the
field of botany by those without formal training in this field
of science. But her highest honour was undoubtedly the
dedication of the Mary Gunn Library in 1969 (Figure 6).
It was a fitting tribute and now stands as a memorial to
someone who virtually single-handedly built up the library
(Figure 7) and served it with devotion and distinction for
almost 60 years.
Mary Gunn possessed charm, coupled with a sharp,
sometimes caustic wit, a sense of humour, steely deter-
mination and a high regard for those she felt warranted
respect. These qualities were admired by many who knew
her personally or who had corresponded with her.
Although quite happy to live alone she enjoyed good
company and was a loyal and loving friend to those dear
to her.
FIGURE 7. — View of the Mary Gunn
Library.
Tributes to Mary’s colleagues Inez and Cythna closed
with quotations from the latter’s book of poems Children
of the hours. No suitable quotation from that work could
be found to close this tribute to the last of the Three
Graces, but what could be more apt than fellow Scot
Thomas Carlyle’s words:
All that mankind has done, thought, gained or been:
it is lying as in magic preservation in the pages of books.
They are the chosen possession of men.’
PUBLICATIONS OF M.D. GUNN
GUNN, M.D. 1940. Joseph Burtt Davy. Journal of the South African
Forestry Association 5: 10.
— 1951. A.E. Roupell. Africana Notes and News 8: 55.
— 1967. E.M. Doidge. Bothalia 9: 251-253.
— 1969. E.P. Phillips. Bothalia 10: 1-3.
— 1970a. J.W. Bews. Standard Encyclopaedia of South Africa 2: 301.
— 1970b. Joseph Burtt Davy. Standard Encyclopaedia of South Africa
2: 605.
— 1971a. A. Dieterlen. Standard Encyclopaedia of South Africa 4: 38.
— 1971b. M.K. Dinter. Standard Encyclopaedia of South Africa 4: 41—42.
— 1971c. I.B. Pole Evans. Bothalia 10: 131—135.
— 1972. M.G.A. Henrici. Bothalia 10: 503-508.
— 1973. J. Hutchinson. Bothalia 11: 1—3.
CODD, L.E. & GUNN, M.D. 1979. Plant collecting pioneers in the
Barberton area. Veld & Flora 65: 98-101.
— 1982a. More early references to Cape plants. Veld & Flora 68: 93-94.
— 1982b. The collecting activities of Anton Rehmann (1840—1917) in
South Africa. Bothalia 14: 1-14.
— 1983. More notes on plant collectors. Veld & Flora 69: 145-147.
— 1984a. J.D.C. Lamb and the ‘Herbarium Lambii’. Veld & Flora 70:
62-63.
— 1984b. Some more notes on plant collectors. Veld & Flora 70: 67—68.
— 1985a. Additional biographical notes on plant collectors in southern
Africa. Bothalia 15: 631-654.
— 1985b. More early illustrations of Cape plants. Veld & Flora 71: 86-88.
GUNN, M.D. & CODD, L.E. 1984. Botanical exploration of southern
Africa. Balkema, Claremont, Cape.
GUNN, M.D. & DU PLESSIS, E. 1978. The Flora capensis of Jakob
and Johann Phillipp Breyne. Brenthurst Press, Johannesburg.
LETTY, C.L. & GUNN, M.D. 1975. E.G. Rice. The Transvaal Gardener
45: 65.
D.M.C. FOURIE
Bothalia 20,1: 131-132 (1990)
Book Reviews
FIELD GUIDE TO THE WILD FLOWERS OF THE WITWATERS-
RAND AND PRETORIA REGION, by BRAAM VAN WYK and SASA
MALAN. 1988. Struik Publishers , Oswald Pirow St, Foreshore, Cape
Town 8001. ISBN 0 86977 814 5. Pp. 352 with full-colour photographs
arranged in more than 130 plates and line drawings by Anne Pienaar.
Size 210 x 147 x 27 mm. Price: soft cover ± R35,00 + GST.
This invaluable book is undoubtedly one of the most ambitious and
useful plant identification guides ever published in southern Africa. It
covers 763 species, almost half the total number present in the Highveld
region of the southern Transvaal.
The problem of identifying wild flowers in a floristically rich area
is solved in a refreshingly new and practical way. Species are arranged
in six colour groups, so that the flower colour is used as the first clue
to identity. Some species with variable flower colours are included in
more than one colour group. Concise descriptions and line drawings of
leaves are provided. Grasses (so often ignored in general field guides)
are treated in a separate chapter. Useful hints on the identification of
this difficult family are supplemented by line drawings which give the
more salient features of the 74 species presented. The arrangement of
photographs and clever layout (descriptions are consistently given directly
opposite each illustration) make the book very easy to use. Since the
shape and venation of leaves are not always visible in the full-colour
plates, line drawings are provided. The photographs are generally of an
excellent quality, even those featuring very unphotogenic subjects! By
including several obscure and rather inconspicuous species (I am referring
particularly to the inclusion of green and even brown ‘flowers’) the authors
have greatly enhanced the value of the book.
The detailed text is written in an easy-to-read style and technical terms
are carefully explained to the layman. Useful general information is given
in the introductory chapter, describing the principles of classification
and identification, as well as the flora and vegetation of the field guide
area. This is followed by guidelines on how to use the book, after which
the most important families are briefly described. I agree with the authors
that the recognition of families is the first step towards competence in
plant identification, despite the initial difficulties for a person without
any botanical training. The family descriptions and the useful glossary
in the back of the book should go a long way towards solving this problem.
The authors have, in my opinion, provided all the necessary informa-
tion for a teach-yourself course in general botany.
Technically, the book is well designed and presented. Hirt & Carter
have done a commendable job on the reproduction of plates and the
colours (even the blues and purples) are as true as one can reasonably
expect. The text is remarkably free of errors. I did however, wonder about
the spelling of Kohoutia (Rubiaceae), which is given as Kohautia in all
standard references. The binding of one of my copies has started to de-
teriorate, but this may well be due to rough handling.
Apart from being an excellent identification guide, the book also con-
tains a wealth of general botanical information for the layman. I am sure
that it will contribute to a greater appreciation for a flora which is suffering
from the impact of a human population exceeding five million. I strong-
ly recommend this book to anyone who has become aware that, to quote
from the preview: ‘there is more to the Highveld vegetation than grass,
maize, cosmos, wattles and gum trees’. Indeed, no-one in this part of
the world should be without a copy.
B-E. VAN WYK*
* Department of Botany, Rand Afrikaans University, P.O. Box 524,
lohannesburg 2000.
THE HERBARIUM HANDBOOK, by L. FORMAN and D. BRIDSON
1989. Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB,
England. Pp. 214, 93 references, 52 figures. Size 245 x 155 x 15 mm.
Price: £12.
This publication was compiled from the lecture notes of various authors
who have contributed to the International diploma course in herbarium
techniques held annually at Kew since 1987. The handbook deals mainly
with the technical aspects of herbarium work including the preparation,
housing, preservation and organization of herbarium collections and
related subjects.
It comprises thirty-nine chapters divided into five sections. These
include an introduction which addresses the question ‘What is taxonomy?’
and then covers the development, purpose and types of herbaria. Other
sections cover the herbarium building, collections and materials;
herbarium techniques and management; additional techniques, for
example photography, the use of maps, gazetteers and computers; plant
collecting and finally three chapters on the role of the herbarium in
supporting economic botany, ecology and conservation.
The publication, appropriately depicting on its cover the wooden
latticework and straps of a plant press, is undoubtedly the best of
its kind to date. It combines the collective, internationally gained
expertise of no less than twenty-four herbarium botanists from Kew
and draws on a wealth of knowledge from the 103 references cited.
Previous publications have not succeeded to quite the same extent
in addressing the full spectrum of herbarium functions. Aspects
included in the handbook which have received little or no coverage
in comparable works include the dissection of floral organs and the
preservation of the results; the use of photography in fieldwork;
the photographic copying of herbarium sheets; the preparation of
checklists of geographical regions; collecting for seedbanks; collecting
living material; guidelines for dealing with visitors to the herbarium
and, to a lesser extent, the re-arrangement of herbarium collections
according to new publications. The book also includes a list of
seventy-three families together with the plant parts essential for their iden-
tification; and a comprehensive list of how to handle herbarium
specimens in order to ensure minimum damage, and specialist
collecting techniques for pteridophytes, bryophytes, fungi and lichens.
This publication achieves what it sets out to be (a good all-purpose
handbook for herbarium curators) by fulfilling the following criteria: it
is simultaneously concise and explicit, discussing in practical terms the
pros and cons of a range of herbarium techniques in a manner that makes
for easy application by even the most inexperienced of herbarium wor-
kers; recommendations and cautions are offered where appropriate; refer-
ences for additional reading are provided at the end of each chapter;
finally, the superior quality of the paper and binding are in keeping with
a book that will have to stand up to constant use. One does, however,
suspect that the index is not as comprehensive as could be expected.
When searching for information on paper acidity, for example, no
reference to this could be found in the index, even though the subject
is dealt with in the text.
Despite the many strengths of this work, there is one obvious
shortcoming, namely the chapter on computers. Although offered as an
introduction to the use of computers in herbaria, this area is receiving
much attention worldwide and should have been dealt with in more detail,
particularly the aspect of computerized herbarium databases.
In conclusion, one must congratulate the editors on an outstanding
reference work through which the staff at the Royal Botanic Gardens,
Kew, endeavour to share their knowledge and expertise gathered
during years of involvement with herbarium curation and management.
Although the book concentrates largely on the procedures followed
at the Kew Herbarium in a European context, these are essentially capable
of universal application with little or no modification. Viewed from
a southern African context, this book will certainly play an important
role in bringing to curators of the region those essential techniques
which are barely touched upon in university and technikon curricula
and which, except in the few larger herbaria, are not passed on to
apprentice curators. One can therefore look forward to a general improve-
ment in the standard of curation in these herbaria. It is predicted
that this handbook will become the standard source of reference for
curators for many years to come.
T.H. ARNOLD
132
THE GENUS WATSONIA, Annals of Kirstenbosch Botanic Gardens
Volume 19, by PETER GOLDBLATT and various artists. 1989. National
Botanic Gardens , Private Bag X7, Claremont, 7735. Pp. 148, 27 water-
colour plates and 44 black-and-white figures, 52 distribution maps. ISBN
0 620 12517 9, ISSN 0 258-3305. Price: R50,00 + GST.
This monographic account of the southern African endemic genus
Watsonia Mill., is the second major revision by Goldblatt to be published
in the Annals of Kirstenbosch Botanic Gardens. The first. Volume 14,
The Moraeas of southern Africa , 1986, was reviewed in Bothalia 17,2:
298 (1987).
As in the Moraea revision, a double-columned text layout is used, and
there are numerous watercolour and black-and-white illustrations. The
watercolour painting of one of the included species, IV dubia Eckl. ex
Klatt, is repeated on the dust jacket. The chapter headings and contents
are essentially similar to those of the previous work.
Some pleasing improvements were noted: space-saving (and cost-saving)
has been achieved by placing the colour plates in groups of eight, at
intervals throughout the volume. In the systematics section, the text for
each taxon therefore usually follows directly upon that for the previous
taxon. The plates and figures for each are of necessity now numbered,
and referenced in the text. The figures are also now captioned. The
volume’s smaller A4 size format allows for easier handling. The taxo-
nomic index has been rendered easier to use by listing the genus name
in full for each entry.
Sadly, as in the Moraea revision, some minor errors have crept in,
such as incorrect or transposed collectors’ numbers, and incorrectly cited
grid references, as on pp. 23, 104, 114, 116. However, these are only likely
to be noticed by herbarium workers and do not detract at all from the
value of this publication.
A more serious error is the omission of the Latin diagnosis from the
text for the new species, W. confusa Goldbl., rendering it invalidly
published.
The contents can be divided into two parts, the first of which includes
an introductory chapter, with Table 1 listing the two subgenera, three
sections, three subsections and 52 species recognised, plus a brief
distribution and genera! habitat description for each. Then there is a
history of the genus from Linnaean to present times, followed by a chapter
discussing important morphological characters and cytology, accompanied
by Table 2 listing chromosome counts for the 31 taxa studied to date.
A discussion of generic relationships and phylogeny follows. Watsonia ,
together with four other small genera, is placed within its own tribe,
Watsonieae, and is separated from its two sister tribes within the subfamily
Ixioideae, Ixieae and Pillansieae, by, inter alia, the corm being of axillary
rather than apical origin, and the deeply divided style branches.
Relationships within the Watsonieae are discussed and presented in the
form of a cladogram, and the major species groups recognised within
Watsonia are treated in a similar fashion. The subgeneric classification
presented here is new, being based on a large body of evidence not readily
available to earlier workers.
The following chapter, on ecology and geography, includes sections
on climate, soils and habitat, on fire phenology, on pollination biology,
on geography and on centres of diversity. It is interesting that variations
in floral structure and colour can be directly correlated to different
pollinators, as has been borne out by observations by the author and others
in the field. Thus the long-tubed red-flowered species are generally
bird-pollinated, and the short-tubed pink-flowered species are generally
bee-pollinated.
As in Moraea, Watsonia occurs in both the winter and summer
rainfall areas of southern Africa, with the northernmost record being
from Woodbush near Pietersburg. In the winter rainfall area the highest
concentration occurs in the south-western Cape, with no less than 16
species in the Worcester quarter degree square. There is a secondary
centre of diversity in the southern Cape, with 11 species recorded in the
Oudtshoorn grid. In the summer rainfall area a centre of high diversity
is found in the Lydenburg and Pilgrim’s Rest grids, and a second one
in the Port Edward and Port St. Johns grids. Interestingly, a viviparous
Bothalia 20,1 (1990)
cultivar of W. meriana (L.) Mill, has become naturalised on Mauritius
and Reunion and in Australia.
The final chapter in this part has comprehensive notes on cultivation.
The horticultural value of the genus has long been recognized, as many
of the species are spectacular in massed display, and more importantly
are easy to grow, provided that their requirements regarding watering
(this depends on the particular rainfall area to which they are native)
and frequent lifting and division of the corms, are met.
The second and major part of this volume is devoted to a systematic
treatment, with a description of the genus, a key to the species (which
works!) and then an enumeration, references, synonymy, description and
discussion of each taxon, with distribution maps, specimen citation and
black-and-white illustrations. Finally there is a discussion of hybrids,
doubtful or unknown species, and excluded species, plus an extensive
bibliography and a taxonomic index to the species.
Turning now to a discussion of the plates and figures: twenty-seven
watercolour plates are included, of which 14 have appeared in early
volumes of The Flowering Plants of Africa. Except for plate 10, Watsonia
pulchra, originally published as Watsonia densiflora in The Flowering
Plants of Africa t. 1293, it is the first time that these original paintings
have been reproduced. (The reason for this is that up until Volume 24,
The Flowering Plants of Africa was printed in England. These early plates
were reproduced in black-and-white by the lithographic process, then
sent to competent hand-colour artists. Mainly due to the scarcity of these
artists, especially during the 1939—45 war, it was decided in 1944 to
change over entirely to colour printing.)
It is difficult to make constructive comment about the colour reproduc-
tion on the early plates, as most are housed in Bolus Herbarium, Cape
Town, and therefore a comparison to the originals cannot be made.
However, the pencil habit and dissection drawings have been printed far
too dark, making them appear very crude. They distract and spoil
otherwise very attractive plates (note also, style drawings on Fay
Anderson’s plates).
The seven illustrations commissioned from Fay Anderson and one from
John Manning have been beautifully executed and are refreshingly crisp.
Unfortunately, in a number of them the colour printing is poor, leaving
them very washed out (compare the cover illustration of W. dubia with
that of plate 6). This problem which is also encountered in the Moraea
revision, could have easily been avoided by better supervision of colour
proofing and printing.
Unfortunately the illustration facing the title page is unsigned and the
artist is not acknowledged.
The text figures are very disappointing. They are generally poorly
arranged, and do not employ the allocated space to best advantage, e.g.
figures 10 and 20. The use of small areas of solid shading is, if anything,
distracting, and serves no scientific or artistic purpose. The corms of
Watsonia provide important diagnostic characters (especially useful to
the horticulturist and herbarium worker), yet in the majority of figures
they have been omitted.
It is a pity that greater effort was not made to select more plates of
high quality, and that more originals were not commissioned.
There are a couple of minor errors in the acknowledgements: e.g.
Cythna Letty — not Cynthia — and seven of the plates — not six— are
by Fay Anderson, and one is by John Manning.
A pleasing touch for the benefit of the non-botanist is the brief
explanation of the specific name for each species.
In conclusion, Watsonia has long been a troublesome genus for
the herbarium worker and the present publication will surely solve
many of the problems experienced. It will also prove valuable to the
horticulturist and interested non-botanist.
C. REID and G. CONDY
Bothalia 20,1: 133-140 (1990)
Guide for authors to Bothalia
This guide is updated when necessary and includes an
index. The latest version 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 National Botanical
Institute, Pretoria, is devoted to the furtherance of botanical
science. The main fields covered are taxonomy, ecology,
anatomy and cytology. Two parts of the journal and an in-
dex to contents, authors and subjects are published
annually.
1 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 En-
glish or Afrikaans.
Articles are assessed by referees, both local and
overseas. Authors are welcome to suggest possible referees
to judge their work. Authors are responsible for the factual
correctness of their contributions. Bothalia maintains an
editorial board (see title page) to ensure that international
standards are upheld.
Articles should preferably be submitted on PC diskettes
or stiffies but the format of all articles should conform
to paragraphs 3.2 to 3.5. Articles not submitted in
electronic form should be arranged according to section 3.
2 Requirements for a diskette
2.1 data must be IBM compatible and written in ASCII.
2.2 a printout of the diskette should be supplied to indicate
(in pencil) the necessary underlining, paragraphs etc.
2.3 tables need not be placed on the diskette — a typed
version is adequate.
2.4 the diskette must have single line spacing, the printout
with markings must be in double line spacing.
2.5 do not justify lines.
2.6 do not break words, except hyphenated words.
2.7 all lines, headings, keys, etc., should start flush at the
margin, therefore no indentations of any kind.
2.8 no italics, bold or underlined words.
2.9 paragraphs and headings are delineated by an extra
line spacing (carriage return) and no indentation.
2.10 a hyphen is designated as one dash, with no space
between the letter and the dash, e.g. ovate-lanceolate. See
also 17.6.
2.11 an N-dash is typed as two hyphens with no space
between the letter and the hyphen, e.g. 2-5 mm (typeset,
it looks like this, 2—5 mm).
2.12 an M-dash is typed as three hyphens with no space
between the letter and the hyphen, e.g. computers — what
a blessing! (typeset, it looks like this, computers — what).
2.13 do not use a double space between words, after
commas, full stops, colons, semicolons or exclamation
marks.
2.14 use lower case x as a times sign, with one space on
either side of the x, e.g. 2x3 mm.
2.15 use single (not double) opening and closing quotes.
2.16 keys — put only three leader dots before number and
name of taxon (with a space before and a space after the
first and last dot), regardless of how far or near the word
is from the right margin, e.g. ... 1. R. ovata.
3 Requirements for a manuscript
3.1 Manuscripts should be typewritten on one side of
good quality A4-size paper, double-spaced throughout
(including abstract, tables, captions to figures, literature
references, etc.) and have a margin of at least 30 mm all
round. 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.
3.2 Papers should conform to the general style and layout
of recent issues of Bothalia (from volume 17 onwards).
3.3 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.
3.4 The sequence continues with Introduction and
aims. Material and methods. Results, Interpretation
(Discussion), Acknowledgements, Specimens examined (in
revisions and monographs), References, Index of names
(recommended for revisions dealing with more than about
15 species), Tables, Captions for figures and figures. In
the case of short notes and book reviews, keywords and
abstract are superfluous.
3.5 All pages must be numbered consecutively beginning
with the title page to those with references, tables and
captions to figures.
3.6 For notes on the use of hyphens and dashes see 2.10
to 2.12.
4 Author(s)
When there are several authors the covering letter
should indicate clearly which of them is responsible for
correspondence and, if possible, telephonically available
while the article is being processed. The contact address
and telephone number should be mentioned if they differ
from those given on the letterhead.
5 Title
The title should be as concise and as informative as
possible. In articles dealing with taxonomy or closely
134
Bothalia 20,1 (1990)
related subjects the family of the taxon under discussion
(see also 13.2) should be mentioned in brackets but author
citations should be omitted from plant names.
6 Keywords
Up to 10 keywords (or index terms) should be provided
in English in alphabetical sequence. The following points
should be borne in mind when selecting keywords:
6.1 Keywords should be unambiguous, internationally
acceptable words and not recently-coined little-known
words.
6.2 they should be in a noun form and verbs should be
avoided.
6.3 they should not consist of an adjective alone; adjec-
tives should be combined with nouns.
6.4 they should not contain prepositions.
6.5 the singular form should be used for processes and
properties, e.g. evaporation.
6.6 the plural form should be used for physical objects,
e.g. augers.
6.7 location (province and/or country); taxa (species,
genus, family) and vegetation type (community, veld type,
biome) should be used as keywords.
6.8 keywords should be selected hierarchically where
possible, e.g. both family and species should be included.
6.9 they should include terms used in the title.
6.10 they should answer the following questions:
6.10.1 what is the active concept in the document (activity,
operation or process).
6.10.2, what is the passive concept or object of the active
process (item on which the activity, operation or process
takes place).
6.10.3, what is the means of accomplishment or how is the
active concept achieved (technique, method, apparatus,
operation or process).
6.10.4 what is the environment in which the active concept
takes place (medium, location).
6.10.5 what are the independent (controlled) and dependent
variables?
6.11 questions 6.10.1 to 6.10.3 should preferably also be
answered in the title.
7 Abstract
7.1 Abstracts of no more than 200 words should be
provided in English and Afrikaans. Abstracts are of great
importance and should convey the essence of the article.
7.2 They should refer to the geographical area concerned
and, in taxonomic articles, mention the number of taxa
treated. They should not contain information not appearing
in the article.
7.3 In articles dealing with taxonomy or closely related
subjects all taxa from the rank of genus downwards should
be accompanied by their author citations.
7.4 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.
8 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.
9 Acknowledgements
Acknowledgements should be kept to the minimum
compatible with the requirements of courtesy. Please give
all the initials of the person(s) you are thanking.
10 Literature references
10.1 Literature references in the text should be cited as
follows: ‘Jones & Smith (1986) stated...’, or ‘...(Jones &
Smith 1986)’ or (Ellis 1988: 67) when giving a reference
simply as authority for a statement. For treatment of
literature references in taxomonic papers see 14.
10.2 When more than two authors are involved in the paper
use the name of the first author followed by et al.
10.3 When referring to more than one literature reference,
they should be arranged alphabetically according to author
and separated by a semicolon, e.g. (Anon. 1981, 1984;
Davis 1976; Nixon 1940).
10.4 Titles of books and names of journals should prefer-
ably not be mentioned in the text. If there is good reason
for doing so, they should be treated as described in 10.17.
10.5 Personal communications are given only in the text,
not in the list of references. Please add the person’s full
initials to identify the person more positively, e.g. C.
Boucher pers. comm.
10.6 References of the same author are arranged in
chronological sequence.
10.7 Where two or more references by the same author
are listed in succession, the author’s name is repeated with
every reference.
10.8 All publications referred to in the text, including those
mentioned in full in the treatment of correct names in
taxonomic papers, but no others, and no personal
communications, are listed at the end of the manuscript
under the heading References.
10.9 The references are arranged alphabetically accord-
ing to authors and chronologically under each author, with
a, b, c, etc. added to the year, if the author has published
more than one work in a year.
10.10 If an author has published both on his own and as
a senior author with others, the solo publications are listed
first and after that, in strict alphabetical sequence, those
published with one or more other authors.
10.11 Author names are typed in capitals.
10.12 Titles of journals and of books are written out in full
and are underlined as follows: Transactions of the Linnean
Society of London 5: 171-217, or Biology and ecology of
weeds : 24.
Bothalia 20,1 (1990)
135
10.13 Titles of books should be given as in Taxonomic
literature , edn 2 by Stafleu & Cowan and names of journals
as in the latest edition of World list of scientific periodicals.
10.14 If the same author is mentioned more than once, the
name is written out in full and not replaced by a line.
10.15 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, Lon-
don.
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 ofRooiberg
State Forest. Department of Forestry, Technical Bulletin.
VOGEL, J.C. 1982. The age of the the Kuiseb river silt terrace at Homeb.
Palaeoecology of Africa 15. In press.
WEISSER, P.J., GARLAND, J.F. & DREWS, B.K. in prep. Dune
advancement 1937—1977 and preliminary vegetation succession
chronology at Mlalazi Nature Reserve, Natal, South Africa.
Bothalia.
Thesis
KRUGER, F.J. 1974. The physiography and plant communities of the
Jakkalsrivier Catchment. M.Sc. (Forestry) thesis. University of
Stellenbosch.
Miscellaneous paper, report, unpublished article, technical
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 Institute.
In A.E.F. Heydorn, Proceedings of workshop research in Cape
estuaries: 105-107. National Research Institute for Oceanology,
CSIR, Stellenbosch.
NATIONAL BUILDING RESEARCH INSTITUTE 1959. Report of the
committee on the protection of building timbers in South Africa
against termites, woodboring beetles and fungi, 2nd edn, CSIR
Research Report No. 169.
11 Tables
11.1 Each table should be presented on a separate sheet
and be assigned an Arabic numeral, i.e. the first table
mentioned in the text is marked ‘Table 1’.
11.2 In the captions of tables the word ‘table’ is written
in capital letters. See recent numbers of Bothalia for the
format required.
11.3 Avoid vertical lines, if at all possible. Tables can often
be reduced in width by interchanging primary horizontal
and vertical heads.
12 Figures
12.1 Figures should be planned to fit, after reduction, into
a width of either 80, 118 or 165 mm, with a maximum
vertical length of 230 mm. Allow space for the caption
in the case of figures that will occupy a whole page.
12.2 Line drawings, including graphs and diagrams, should
be in jet-black Indian ink, preferably on fine Felix
Schoeller parole or similar board, 200 gsm, or tracing
film. Lines should be bold enough to stand reduction.
12.3 It is recommended that drawings should be twice the
size of the final reproduction.
12.4 Photographs should be of excellent quality on glossy
paper with clear detail and moderate contrast, and they
should be the same size as required in the journal.
12.5 Photograph mosaics should be submitted complete,
the component photographs mounted neatly on a white
flexible card base leaving a narrow gap of uniform width
(2 mm) between each print. Note that grouping photo-
graphs of markedly divergent contrast results in poor
reproductions.
12.6 Lettering and numbering on all figures should be done
in letraset, stencilling or a comparable method. If symbols
are to be placed on a dark background it is recommended
that black symbols are used on a small white disk ± 7
mm in diameter and placed in the lower left hand corner
of the relevant photo.
12.7 If several illustrations are treated as components of
a single composite figure they should be designated by
capital letters.
12.8 Note that the word ‘figure’ should be written out in
full, both in the text and the captions.
12.9 In the text the figure reference is then written as in
the following example: ‘The stamens (Figure 4A, B, C)
are. . .’
12.10 In captions, ‘figure’ is written in capital letters.
Magnification of figures should be given for the size as
submitted.
12.11 It is recommended that scale bars or lines be used
on figures.
12.12 In figures accompanying taxonomic papers, voucher
specimens should be given in the relevant caption.
12.13 Figures are numbered consecutively with Arabic
numerals in the order they are referred to in the text. These
numbers, as well as the author’s name and an indication
of the top of the figure, must be written in soft pencil on
the back of all figures.
12.14 Captions of figures must not be pasted under the
photograph or drawing.
12.15 Authors should indicate in pencil in the text where
they would like the figures to appear.
12.16 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’.
136
Bothalia 20,1 (1990)
12.17 Authors wishing to use illustrations already published
must obtain written permission before submitting the
manuscript and inform the editor of this fact.
12.18 Captions for figures should be collected together and
typed on a separate sheet headed Captions for figures.
12.19 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 reduction
to 80 mm (recommended size: letraset 5 mm diameter).
12.20 Blank maps are available from the editor.
13 Text
13.1 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, 51 and 56.
13.2 Names of genera and infrageneric taxa are usually
underlined, with the author citation (where relevant) not
underlined. Exceptions include names of new taxa in the
abstracts, correct names given in the synopsis or in para-
graphs on species excluded from a given supraspecific
group in taxonomic articles, in checklists and in indices,
where the position is reversed, correct names not being
underlined and synonyms underlined.
13.3 Names above generic level are not underlined.
13.4 In articles dealing with taxonomy and closely related
subjects the complete scientific name of a plant (with
author citation) should be given at the first mention in the
text. The generic name should be abbreviated to the initial
thereafter, except where intervening references to other
genera with the same initial could cause confusion.
13.5 In normal text, Latin words are italicized, but in the
synopsis of a species, Latin words such as nom. nud. are
not italicized.
13.6 Names of authors of plant names 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.
13.7 Modern authors not included in the list should use
their full name and initials when publishing new plant
names. Other author names not in the list should be in
agreement with the recommendations of the Code.
13.8 Names of authors of publications are written out in
full except in the synonymy in taxonomic articles where
they are treated like names of authors of plant names.
13.9 Names of plant collectors are underlined whenever
they are linked to the number of a specimen. The collec-
tion number is also underlined, e.g. Acocks 14407.
13.10 Surnames beginning with ‘De’, ‘Du’ or ‘Van’ begin
with a capial letter unless preceded by an initial.
13.11 For measurements use only units of the International
System of Units (SI). Cm should not be used, only mm
and/or m.
13.12 The use of ‘±’ is preferred to c. or ca.
13.13 Numbers ‘one’ to ‘nine’ are spelled out in normal
text, and from 10 onwards they are written in Arabic
numerals.
13.14 In descriptions of plants, numerals are used through-
out. Write 2,0— 4,5 (not 2—4,5). When counting mem-
bers write 2 or 3 (not 2-3) but 2—4.
13.15 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.
13.16 Apart from multi-access keys, indented keys should
be used with couplets numbered la— lb, 2a— 2b, etc.
(without full stops thereafter).
13.17 Keys consisting of a single couplet have no number-
ing.
13.18 Manuscripts of keys should be presented as in the
following example:
la Leaves closely arranged on an elongated stem; a sub-
merged aquatic with only the capitula exserted ... lb. E.
setaceum var. pumilum
lb Leaves in basal rosettes; stems suppressed; small marsh
plants, ruderals or rarely aquatics:
2a Annuals, small, fast-growing pioneers, dying when the
habitat dries up; capitula without coarse white setae;
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:
13.19 Herbarium voucher specimens should be referred to
wherever possible, not only in taxonomic articles.
14 Species treatment in taxonomic papers
14.1 The procedure to be followed is illustrated in the
example (17, 17.8), which should be referred to, because
not all steps are described in full detail.
14.2 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 (not underlined).
14.3 Thereafter all literature references, including those
of the synonyms, should only reflect author, page and year
of publication, e.g. C.E. Hubb. in Kew Bulletin 15: 307
(1960); Boris et al. : 14 (1966); Boris: 89 (1967); Sims:
t. 38 (1977); Sims: 67 (1980).
14.4 The description and the discussion, which should
consist of paragraphs commencing, where possible, with
italicized leader words such as flowering time, diagnostic
characters, distribution and habitat.
14.5 When more than one species of a given genus is dealt
with in a paper, the correct name of each species should
be prefixed by a sequential number followed by a full stop,
the first line of the paragraph to be indented. Infraspecific
taxa are marked with small letters, e.g. lb., 12c., etc.
14.6 Names of authors are written in the same way (see
13.1, 13.6), irrespective of whether the person in question
is cited as the author of a plant name or of a publication.
Bothalia 20,1 (1990)
137
14.7 The word ‘figure’ is written as ‘fig.’, and ‘t.’ is used
for both ‘plate’ and ‘tablet’.
14.8 Literature references providing good illustrations of
the species in question may be cited in a paragraph
commencing with the word leones followed by a colon.
This paragraph is given after the last paragraph of the
synonymy, see 17.8.
15 Citation of specimens
15.1 Type specimen in synopsis: the following should be
given (if available): country (if not in RSA), province, grid
reference (at least for new taxa), locality as given by
original collector, modem equivalent of collecting locality
in square brackets (if relevant), date of collection
(optional), collector’s name and collecting number (both
underlined).
15.2 The abbreviation s.n. ( sine numero) is given after the
name of a collector who usually assigned numbers to his
collections but did not do so in the specimen in question.
The herbaria in which the relevant type(s) are housed are
indicated by means of the abbreviations given in the latest
edition of Index Herbariorum.
15.3 The holotype (holo.) and its location are mentioned
first, followed by a semicolon, the other herbaria are
arranged alphabetically, separated by commas.
15.4 Authors should indicate by means of an exclama-
tion mark (!) which of the types have been personally
examined.
15.5 If only a photograph or microfiche was seen, write
as follows: Anon. 422 (X, holo.-BOL, photo.!).
15.6 Lectotypes or neotypes should be chosen for correct
names without a holotype. It is not necessary to lectotypify
synonyms.
15.7 When a lectotype or a neotype are newly chosen this
should be indicated by using the phrase ‘here designated’.
If reference is made to a previously selected lectotype
or neotype, the name of the designating author and
the literature reference should be given. In cases
where no type was cited, and none has subsequently
been nominated, this may be stated as ‘not designated’.
15.8 In brief papers mentioning only a few species and a
few cited specimens the specimens should be 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.
15.9 Grid references should be cited in numerical sequence.
15.10 Locality records for specimens should preferably be
given to within a quarter-degree square. Records from the
same one-degree square are given in alphabetical order,
i.e (—AC) precedes (—AD), etc. Records from the same
quarter-degree square are arranged alphabetically
according to the collectors’ names; the quarter-degree
references must be repeated for each specimen cited.
15.11 The relevant international code of the 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\ Koffiefontein
(-AB), Taylor 720 (PRE); Cathedral Peak Forest Station (— CC), Marriot
74 (KMG); Wilgerfontein, Roux 426. Grid ref. unknown: Sterkstroom,
Strydom 12 (NBG).
15.12 For records from outside southern Africa authors
should use degree squares without names, e.g.:
KENYA. — 0136: Nairobi plains beyond race course, Napier 485.
15.13 Monographs and revisions: in the case of all major
works of this nature it is assumed that the author has
investigated the relevant material in all major herbaria and
that he has provided the specimens seen with determinavit
labels. It is assumed further that the author has submitted
distribution maps for all relevant taxa and that the
distribution has been described briefly in words in the text.
Under the heading ‘Vouchers’ no more than five specimens
should be cited, indicating merely the collector and the
collector’s number (both 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 semi-
colon. The purpose of the cited specimens is not to indicate
distribution but to convey the author’s concept of the taxon
in question.
15.14 The herbaria in which the specimens are housed are
indicated by means of the abbreviation given in the latest
edition of Index Herbariorum. They are given between
brackets, arranged alphabetically and separated by commas
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 4451 (BM, BOL, GRA, K, PRE).
15.15 If long lists of specimens are given, they should be
listed together at the end of the article under the heading
Specimens 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 mentioned is indicated as 1.1. This is followed by
the international herbarium designation. Note that the
name of the collector and the collection number are
underlined:
Acocks 12497 (2. lb) 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.
16 Synonyms
16.1 In a monograph or a revision covering all of southern
Africa, all synonyms based on types of southern African
origin, or used in southern African literature, should be
included.
16.2 Illegitimate names are designated by nom. illeg. after
the reference, followed by non with the author and date,
if there is an earlier homonym.
16.3 Nomina nuda {nom. nud.) and invalid names are
excluded unless there is a special reason to cite them.
138
Bothalia 20,1 (1990)
for example if they have been used in prominent publi-
cations.
16.4 In normal text Latin words are italicized, but in the
synopsis of a species Latin words such as nom. nud. are
not italicized.
16.5 Synonyms should be arranged chronologically into
groups of nomenclatural synonyms, i.e. synonyms based
on the same type, and the groups should be arranged
chronologically by basionyms, except for the basionym of
the correct name which is dealt with in the paragraph
directly after that of the correct name.
16.6 When a generic name is repeated in a given synonymy
it should be abbreviated to the initial except where inter-
vening references to other genera with the same initial
could cause confusion.
17 Description and example of species treatment
17.1 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.
17.2 As a rule shape should be given before measurements.
17.3 In general, if an organ has more than one of the parts
being described, use the plural, otherwise use the singular,
for example, petals of a flower but blade of a leaf.
17.4 Language must be as concise as possible, using
participles instead of verbs.
17.5 Dimension ranges should be cited as in the example
below.
17.6 Care must be exercised in the use of dashes and
hyphens: a hyphen is a short stroke joining two syllables
of a word, e.g. ovate-lanceolate or sea-green, with no space
between the letter and the stroke; an N-dash (en) is a longer
stroke commonly used instead of the word ‘to’ between
numerals, ‘2—5 mm long’ (do not use it between words
but rather use the word ‘to’, e.g. ‘ovate to lanceolate’); it
is produced by typing 2 hyphens 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 in-
stead of the ftill species name; it is produced by typing
3 hyphens next to one another.
17.7 The use of ‘±’ is preferred to c. or ca when describing
shape, measurements, dimensions, etc.
17.8 Example:
1. Bequaertiodendron magalismontanum (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.) Aubr6v. & Pelegr.: 105 (1958); Justin: (1973).
Chrysophyllum argyrophyllum Hiem: 721 (1850); Engl.: 43 (1904).
Boivinella argyrophylla (Hiem) Aubr6v. & Pellegr.: 37 (1958); Justin:
98 (1973). Types: Angola, Welwitsch 4828 (BM!, lecto., here designated;
PRE!); Angola, Welwitsch 4872 (BM!).
Chrysophyllum wilmsii Engl.: 4, t. 16 (1904); Masonet: 77 (1923);
Woodson: 244 (1937). Boivinella wilmsii (Engl.) Aubrdv. & Pellegr.: 39
(1958); Justin: 99 (1973). Type: Transvaal, Magoebaskloof, Wilms 1812
(B, holo.; K!, P!, lecto., designated by Aubr6v. & 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); Pelmer: 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 oblanceolate,
obtuse, base broad, half-clasping. Heads heterogamous,
campanulate, 7—8x5 mm, solitary, sessile at tip of
axillary shoots; involucral bracts in 5 or 6 series, inner
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: 2n = 22. Figure 23B.
18 New taxa
18.1 The name of a new taxon must be accompanied by
at least a Latin diagnosis. Authors should not provide
full-length Latin descriptions unless they have the required
expertise in Latin at their disposal.
18.2 It is recommended that descriptions of new taxa be
accompanied by a good illustration (line drawing or
photograph) and a distribution map.
18.3 Example:
109. Helichrysum jubilatum Hilliard , sp. nov. H.
alsinoidei DC. affmis, sed foliis ellipticis (nec spatulatis),
inflorescentiis compositis a foliis non circumcinctis,
floribus femineis numero quasi dimidium hermaphrodi-
torium aequantibus (nec capitulis homogamis vel floribus
femineis 1-3 tantum) distinguitur.
Herba annua e basi ramosa; caules erecti vel decum-
bentes, 100-250 mm longi, tenuiter albo-lanati, remote
foliati. Folia plerumque 8—30 x 5—15 mm, sub capitulis
minora, elliptica vel oblanceolata, obtusa vel acuta,
mucronata, basi semi-amplexicauli, utrinque cano-lanato-
arachnoidea. Capitula heterogama, campanulata, 3,5— 4,0
X 2,5 mm, pro parte maxima in paniculas cymosas
terminales aggregata; capitula subterminalia interdum
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 subaequalibus 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, apicibus scabridis, basibus non
cohaerentibus.
Bothalia 20,1 (1990)
139
TYPE. — Cape, 2817 (Vioolsdrif): (— CC), Richters-
veld, ± 5 miles E of Lekkersing on road to Stinkfontein,
kloof in hill south of road, annual, disc whitish, 7.11.1962,
Nordenstam 1823 (S, holo. ; E, NH, PRE).
19 Proofs
Only page proofs are normally sent to authors. They
should be corrected in red ink and be returned to the editor
as soon as possible.
20 Reprints
Authors receive 100 reprints free. If there is more than
one author, this number will have to be shared between
them.
21 Documents consulted
Guides to authors of the following publications were
made use of in the compilation of the present guide: Annals
of the Missouri Botanic Garden, Botanical Journal of the
Linnean Society, Flora of Australia, Smithsonian Con-
tributions to Botany, South African Journal of Botany
(including instructions to authors of taxonomic papers),
South African Journal of Science.
22 Address of editor
Manuscripts should be submitted to: The Editor,
Bothalia, National Botanical Institute, Private Bag X101,
Pretoria 0001.
INDEX
abbreviation, 13.4, 13.5, 13.12, 13.15, 14.7, 15.2, 15.14, 16.2, 16.3, 16.4, 16.6
abstract (uittreksel), 3.2, 7, 13.2
acknowledgements, 9
address of
authors, 3.3, 4
editor, 22
alphabetical, 6, 10.3, 10.9, 10.10, 15.3, 15.10, 15.13, 15.14, 15.15
Arabic numerals, 11.1, 12.13, 13.3
ASCII, 2.1
author(s), 1, 3.1, 4, 10.15, 12.15
address, 3.3, 4
citation, 5, 7.3, 13.2, 13.4, 14.2
first, 10.2
names, 3.3, 10.3, 10.7, 10.9, 10.11, 10.14, 12.13, 13.7, 13.8, 14.3, 14.6, 15.7,
16.2
names of plant names, 13.6, 13.7, 13.8
senior, 10.10
book reviews, 1, 3.4
books, 10.4, 10.12, 10.13, 10.15
Bothalia, 1, 3.2, 11.2, 22
brief taxonomic articles, 15.8
c., 13.2, 17.7
ca, 13.2, 17.7
capitals, 11.2, 12.7, 12.10, 14.2, 15.8
captions, 3.1, 3.4, 3.5, 11.2, 12.8, 12.10, 12.12, 12.14, 12.18
checklist, 13.2
chromosome number, 17.1, 17.8
chronological sequence, 10.6, 10.9, 16.5
citation
author, 5, 7.3, 13.2, 13.4, 14.2
literature, 14.4
of specimens, 15
cm, 13.11
collection
date, 15.1
number, 13.9, 15.1, 15.2, 15.11, 15.13, 15;15
collective book, 10.15
collector, 13.9, 15.1, 15.2, 15.10, 15.13, 15.15
colon, 2.13
comma, 2.13
compass directions, 13.15
composite figure, 12.7
congress proceedings, 10.15
contents, 8
correspondence, 4
countries, 6.7, 15.8
description and example of species treatment, 17
diagrams, 12.2
discussion, 3.4, 14.4
diskette, 1, 2.4
distribution maps, 12.19, 12.20, 15.13, 18.2
documents consulted, 21
dot maps, 12.19, 12.20, 15.13, 18.2
double
line spacing, 2.4
space, 3.1, 2.13
drawing paper, 12.2
drawings, 12.2
edition, 13.15
editor, 13.15, 22
editorial
board, 1
policy, 1
et al., 10.2, 13.6, 14.3
example of
new taxa, 18.3
species treatment, 17.8
exclamation mark, 2.13, 15.4
family name, 5, 6.7
fig., 14.7
figure(s), 12, 14.7, 17.1
reduction of, 12.1, 12.2, 12.19
returned, 12.16
first author, 10.2
flora, 1, 10.15
footnote, 3.3
full stop, 2.13, 13.15, 13.16, 14.5
genera, 13.2
generic name, 13.3, 13.4, 16.6
geographical area, 7.2
GIBBS RUSSELL, G.E. et al. List of species of southern African plants.
Memoirs of the Botanical Survey of South Africa Nos 48, 51 &
56, 10.15, 13.1, 13.6
granting agencies, 3.3
graphs, 12.2
grid reference system, 15.1, 15.8, 15.9, 15.11
headings, 2.7, 2.9
sequence of, 3.3, 3.4
herbaria, 15.2, 15.3, 15.11, 15.13, 15.14
herbarium
code, 15.11
designations, 13.15, 15.15
voucher specimens, 12.12, 13.19
holo., 15.5, 17.8, 18.3
holotype, 15.3, 15.6
homonym, 16.2
hyphenated words, 2.6
hyphen, 2.10-2.12, 17.6
IBM compatible, 2.1
icones, 10.2, 17.8
illegitimate names (nom. illeg.), 16.2
illustrations, 12.3, 12.7, 12.17, 14.8
previously published, 12.17
Index Herbariorum, 15.2, 15.14
index of names, 3.4
infrageneric taxa, 13.2
initials, 9, 10.5, 13.7
in prep., 10.15
in preparation, 10.15
in press, 10.15
International
Code of Botanical Nomenclature, 13.7
System of Units (SI), 13.11
invalid names, 16.3
italics/underlining, 7.4, 10.12, 13.2, 13.3, 13.5, 13.9, 14.2, 15.1, 15.13, 15.15
journals, 10.4, 10.12, 10.15
names of, 10.1, 10.13
140
Bothalia 20,1 (1990)
justify, 2.5
keys, 2.7, 2.16, 13.16, 13.17, 13.18
keywords, 3.3, 3.4, 6
Latin, 13.5, 15.2, 16.2, 16.3, 16.4
descriptions, 18.1
layout, 3.2
lecto., 15.6, 15.7, 17.8
lectotype, 15.6, 15.7, 17.8
letraset, 12.6, 12.19
lettering, 12.6
line
drawings, 12.2, 18.2
spacing, 2.4, 2.9
literature
citations, 14.4
references, 3.1, 10, 10.7
within synonymy, 10.7, 14.8
localities outside southern Africa, 15.12
locality, 15.1, 15.10
m, 13.11
magnification of figures, 12.3, 12.10
manuscript
language, 1
requirements, 3
map, distribution, dot, 12.19, 12.20, 15.13, 18.2
M-dash, 2.12, 17.6
mm, 13.11
margin, 2.7, 2.16, 3.1, 17.1
material, 3.3, 3.4
measurements, 13.11, 17.2, 17.7
methods, 3.4, 6.10.3
microfiche, 15.5
miscellaneous paper, 10.15
monograph, 3.4, 15.13, 16.1
name
collector’s, 15.10
illegitimate, 16.2
invalid, 16.3
name(s) of
author(s), 3.3, 10.7, 10.9, 10.11, 10.14, 13.7, 13.8, 14.6
of plant names (TN TAX2/1), 5, 13.1, 13.6, 14.6
of publications, 13.8
plant collectors, 13.9
publication, 14.2
taxa, 2.16, 5, 7.4, 10.8, 13.2, 13.3
N-dash, 2.11, 17.6
neotype, 15.6, 15.7
new
combinations, 7.4
taxa, 7.4, 13.2, 13.7, 15.7, 18
nom. illeg. , 16.2
nom. nud., 13.5, 16.3, 16.4
notes, 1, 3.4, 10.15
technical, 10.15
number, chromosome, 17.1, 17.8
numbering, 13.13
figures, 12.6, 12.13, 17.1
keys, 13.16, 13.17
pages, 3.5, 13.4
taxa, 2.16, 7.2, 13.4, 14.5, 15.15
numerals, Arabic, 11.1, 12.13, 13.3
PC diskettes, 2
pers. comm., 10.5
personal communications (pers. comm.), 10.5, 10.8
photocopies, 3.1
photograph, 12.4, 12.14, 15.5, 18.2
mosaic, 12.5
plant name, 5, 13.4, 13.6, 13.7, 13.8, 14.6
plate (t.), 14.7
prepositions, 6.4
proceedings, 10.15
proofs, 19
provinces, 6.7, 15.1, 15.8
publications, 10.8, 13.8, 14.3
name of, 14.2
solo, 10.10
year of, 10.9, 14.3
quarter-degree squares, 15.10
quotes, 2.15
reduction of figures, 12.1, 12.2, 12.19
referees, 1
reference, 3.4, 10.6, 10.7, 10.9, 10.15
figure, 12.9
grid, 15.1, 15.8, 15.9, 15.11
list, 10.5, 10.8, 10.9
literature, 3.1, 10, 10.7
report, 10.15
reprints, 20
requirements for
diskette, 2
manuscript, 3
results, 3.4
revision, 3.4, 8, 15.13, 16.1
scale bar, 12.11
semicolon, 2.13, 10.3, 15.3, 15.13
senior author, 10.10
sequence of headings, 3.3, 3.4
short notes, 1, 3.4
single line spacing, 2.4
species treatment in taxonomic papers, 14
specimens examined, 3.4, 15.5
STAFLEU, F. A. & COWAN, R.S. 1976-1988. Taxonomic literature. Vols
1-7, 10.13
stiffies, 1
surnames, 13.10
symbols, 12.6
synopsis, 13.2, 13.5, 15.1, 16.4
synonymy, 10.7, 13.8, 14.4, 14.8, 16.6
t., 14.3, 14.7, 17.8
table, 2.3, 3.1, 3.4, 3.5, 11
of contents, 8
tablet (t.), 14.7
name of, 2.16, 5, 7.4, 10.8, 13.2, 13.3
new, 7.4, 13.2, 13.7, 15.7, 18
numbering of, 2.16, 7.2, 13.4, 14.5, 15.15
taxonomic
articles/papers, 7.2, 10.8, 12.12, 12.19, 13.2, 13.8, 14
revision, 8
taxonomy, 5, 7.3, 13.4, 15.8
technical note, 10.15
text, 3.1, 10.1, 10.4, 10.5, 10.8, 11.1, 12.8, 12.9, 12.13, 12.15, 13, 15.13, 15.15,
16.4
thesis, 10.15
times sign, 2.14
title, 3.3, 5, 6.9, 6.11
of books, 10.4, 10.12, 10.13, 10.15
of journals, 10.4, 10.12, 10.13, 10.15
page, 1, 3.3, 3.5
type, 15.2, 15.4, 15.7, 16.1, 16.6, 17.8
here designated, 15.7, 17.8
not designated, 15.7
specimen, 15.1
underlining/italics, 7.4, 10.12, 13.2, 13.3, 13.5, 13.9, 14.2, 15.1, 15.13, 15.15
uittreksel (abstract), 7.1
units of measure, 13.11, 13.15
unpublished article, 10.15
vouchers, 15.13, 15.14
voucher specimens, 12.12, 13.19
World list of scientific periodicals , 10. 13
year of publication, 10.9, 14.3
BOTHALIA
Volume 20,1
May/Mei 1990
CONTENTS — INHOUD
1. Studies in the genus Lotononis (Crotalarieae, Fabaceae). 9. Four new species of the L. pentaphylla group,
section Lipozygis. B-E. VAN WYK 1
2. Studies in the genus Lotononis (Crotalarieae, Fabaceae). 12. Four new species of the L. falcata group,
section Leptis. B-E. VAN WYK 9
3. Studies in the genus Lotononis (Crotalarieae, Fabaceae). 13. Two new species and notes on the occur-
rence of cleistogamy in the section Leptis. B-E. VAN WYK 17
4. Studies in the genus Riccia (Marchantiales) from southern Africa. 15. R. hirsuta and R. tomentosa,
sp. nov., two distinct species previously treated as one. O.H. VOLK and S.M. PEROLD .... 23
5. Studies in the genus Riccia (Marchantiales) from southern Africa. 16. R. albomarginata and R. simii,
sp. nov. S.M. PEROLD 31
6. New species of Erica (Ericaceae) from the Cape Province. E.G.H. OLIVER 41
7. Studies in the southern African species of Justicia and Siphonoglossa (Acanthaceae): seeds. K.L.
IMMELMAN 49
8. Studies in the southern African species of Justicia and Siphonoglossa (Acanthaceae): indumentum.
K.L. IMMELMAN 61
9. Notes on African plants:
Asclepiadaceae. Corona lobe variation and the generic position of Asclepias macra. A. NICHOLAS
and D.J. GOYDER 87
Asteraceae. A new species of Pterothrix (Gnaphalieae) from the northern Cape. F. BRUSSE ... 67
Fabaceae. Studies in the genus Lotononis (Crotalarieae. 10. L. esterhuyseniana, a new species
from the south-western Cape. B-E. VAN WYK 70
Fabaceae. Studies in the genus Lotononis (Crotalarieae). 11. A new species of the section
Leobordea from north-western Namibia. B-E. VAN WYK 73
Fabaceae. Studies in the genus Lotononis (Crotalarieae). 14. Three new species of the sections
Telina and Polylobium. B-E. VAN WYK 75
Liliaceae/Asphodelaceae. The correct author citations of Aloe bowiea and A. myriacantha
(Alooideae). G.F. SMITH 80
Poaceae. Two new species of Stipagrostis (Aristideae) from the dune-Namib Desert, Namibia.
B. DE WINTER 82
10. Vegetative morphology and interfire survival strategies in the Cape Fynbos grasses. H.P. LINDER and
R.P. ELLIS 91
11. The ecology of the False Bay estuarine environments, Cape, South Africa. 1. The coastal vegetation.
M. O’CALLAGHAN 105
12. The ecology of the False Bay estuarine environments. Cape, South Africa. 2. Changes during the last
fifty years. M. O’CALLAGHAN 113
13. Miscellaneous note:
Notes on the distribution and habitat of Aloe bowiea (Liliaceae/Asphodelaceae: Alooideae) — an
endangered and little known species from the eastern Cape. G.F. SMITH and A.E. VAN
WYK 123
14. Obituary: Mary Davidson Gunn (1899—1989). D.M.C. FOURIE 127
15. Book reviews 131
16. Guide for authors to Bothalia 133
Abstracted, indexed or listed in AGRICOLA, Biological Abstracts, Current Advances in Plant Science, Current Contents, Field Crop Abstracts,
Forestry Abstracts, Herbage Abstracts, Excerpta Botanica, Revue of Plant Pathology, Revue of Medical and Veterinary Mycology and The Kew
Record of Taxonomic Literature.
ISSN 0006 8241
O anti published by the National Botanic Gardens/Botanical Research Institute, Private Bag X101, Pretoria 0001, South Africa. Printed by Gutenberg
Book Printers (Pty) Ltd, 141 Industrial Rd, Pretoria West 0183. Tel. (012) 386-1133/4/5/6/7/8. Obtainable from the National Botanical Institute,
Private Bag X101, Pretoria 0001, South Africa.