ISSN 0006 8241 = Bothalia

Bothalia

’N TYDSKRIF VIR PLANTKUNDIGE NAVORSING

A JOURNAL OF BOTANICAL RESEARCH

Vol. 24,1

May/Mei 1994

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BOTHALIA

Bothalia is named in honour of General Louis Botha, first Premier and

Mi nister of Agriculture of the Union of South Africa. This house journal

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’n Palaeoflora met ’n uitleg vergelykbaar met die van die Flora van

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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. Obtain-

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ANNALS OF KIRSTENBOSCH BOTANIC GARDENS

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on southern African flora. Published: Vol. 14-19 (earlier volumes

published as Supplementary volumes to the Journal of South African

Botany )

-n Reeks gewy aan die publikasie van monografiee en belangrike werke oor

flora van suidelike Afrika. Gepubliseer: vol. 14-19 (vroeere volumes

gepubliseer as Supplementary volumes van die Journal of South African

Botany).

BOTHALIA

’N TYDSKRIF VIR PLANTKUNDIGE NAVORSING

A JOURNAL OF BOTANICAL RESEARCH

Volume 24,1

Scientific Editor/Wetenskaplike Redakteur: O.A. Leistner

Technical Editor/Tegniese Redakteur: B.A. Momberg

NATIONAL

Botanical

INSTITUTE

NASIONALE BOTANIESE

INSTITUUT 3

Private Bag X 101 PRETORIA 0001

1994 -05- 0'

Prlvaatsak X 101 PRETORIA 0001

NATIONAL BOTANICAL

INSTITUTE

2 Cussonia Avenue, Brummeria, Pretoria

Private Bag XI01, Pretoria 0001

ISSN 0006 8241

May/Mei 1994

Editorial Board/Redaksieraad

D.F. Cutler

B.J. Huntley

P.H. Raven

J.P. Rourke

M.J. Werger

Royal Botanic Gardens, Kew, UK

National Botanical Institute, Cape Town, RSA

Missouri Botanical Garden, St Louis, USA

Compton Herbarium, NBI, Cape Town, RSA

University of Utrecht, Utrecht, Netherlands

CONTENTS— INHOUD

Volume 24,1

1. Three new species of Serruria (Proteaceae) from the southwestern Cape. J.P. ROURKE 1

2. Studies in the Marchantiales (Hepaticae) from southern Africa. 4. Mannia capensis, section and subgenus

Xeromannia (Aytoniaceae). S.M. PEROLD 9

3. Studies in the Marchantiales (Hepaticae) from southern Africa. 5. The genus Exormotheca, E. pustulosa

and E. holstii. S.M. PEROLD 15

4. Studies in the Ericoideae (Ericaceae). XIV. Notes on the genus Erica. E.G.H. OLIVER and INGE M.

OLIVER 25

5. Notes on African plants:

Asphodelaceae/Aloaceae. Was Gasteria nitida var. armstrongii validly published? G.F. SMITH,

B-E. VAN WYK & E.J. VAN JAARSVELD 34

Asphodelaceae/Aloaceae. Aloe barberae to replace A. bainesii. G.F. SMITH, B-E. VAN WYK &

H.F. GLEN 34

Boraginaceae. The correct author citation for Lobostemon montanus. M.H. BUYS and J.J.A. VAN

DER WALT 35

Rosaceae. Cliffortia longifolia, a 'good' species or should it be a variety under C. strobiliferal . A.C.

FELLINGHAM.V 31

6. Ferns and flowering plants of Klaserie Private Nature Reserve, eastern Transvaal: an annotated checklist.

N. ZAMBATIS 37

7. External fruit morphology of southern African Arundineae (Arundinoideae: Poaceae). N.P BARKER . 55

8. Names of the southern African species of Rhus (Anacardiaceae) and their etymology. R.O. MOFFETT . 67

9. Flowering phenology in the arid winter rainfall region of southern Africa. M. STRUCK 77

10. Miscellaneous notes:

Chromosome counts for seven species of Cineraria (Senecioneae, Asteraceae). G.V. CRON, B-E.

VAN WYK and P.L.D. VINCENT 91

Acytotaxonomic study of some representatives of the tribe Cynodonteae (Chloridoideae, Poaceae).

A. STRYDOM and J.J. SPIES 92

Systematic studies in the genus Mohria (Anemiaceae). V. Karyology (Pteridophyta). J.P. ROUX . 97

1 1. Embryo sac development in some representatives of the tribe Cynodonteae (Poaceae). A. STRYDOM and

J.J. SPIES 101

12. Breeding systems in some representatives of the genus Lycium (Solanaceae). L. MINNE, J.J. SPIES, H.J.T.

VENTER and A.M. VENTER 107

13. A life dedicated to lichens and literature, Ove Almbom (30-07-1914— 06-03-1992). E.I. KARNEFELT . Ill

14. Book reviews 117

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Bothalia 24,1: 1-7(1994)

Three new species of Serruria (Proteaceae) from the southwestern Cape

J.P. ROURKE*

Keywords: new species, Proteaceae, Serruria, southwestern Cape

ABSTRACT

Three new species of Serruria are described, namely, S. altiscapa Rourke from the Villiersdorp and Hottentots Holland

Mountains, characterised by its unusually tall paniculate inflorescences; S. gremialis Rourke, a multistemmed lignotuberous

species with adaxially cygneous, basally pubescent styles and cyiindric pollen presenters from the Villiersdorp and Houhoek

Mountains as well as the north slopes of the Riviersonderend Range, and S. balauocephala Rourke, a single-stemmed re-seeding

species from the north slopes of the Langeberg, distinguished by its adaxially cygneous, basally pubescent styles and conic-acute

pollen presenters.

UITTREKSEL

Drie nuwe Serruria- spesies word beskryf, naamlik S. altiscapa Rourke van die Villiersdorp- en Hottentots-Hollandberge,

gekenmerk deur buitengewoon lang, pluimvormige bloeiwyses; S. gremialis Rourke, 'n veelstammige spesie met verhoute

knolle, adaksiaal swaannekvormige, basaal sagharige style en silindriese stuifmeelaanbieders, van die Villiersdorp- en

Houhoekberge en die noordelike hange van die Riviersonderendreeks, en S. balauocephala Rourke, ’n enkelstammige

hersaaiende spesie van die noordelike hange van die Langeberg, gekenmerk deur adaksiaal swaannekvormige, basaal

saagharige style en konies-skerppuntige stuifmeelaanbieders.

INTRODUCTION

Serruria Salisb., is the largest genus of the South Afri-

can Proteaceae which is endemic to the southwestern and

southern Cape. The most recent comprehensive account

of the genus recognised 48 species and listed a further 1 8

as ‘imperfectly known species’ (Phillips & Hutchinson

1912). Exploration of mountainous areas of the Cape dur-

ing the present century has brought a number of new taxa

to light, several of which have already been described

(Rourke 1982, 1990, 1991). This paper enumerates an-

other three new species of Serruria prior to the preparation

of a complete account of the genus.

Serruria altiscapa Rourke , sp. nov., a 5. williamsii

Rourke habitu humili compacto, inflorescentiis pani-

culatis peraltis 700-1 200 mm, bracteis glabris transverso-

ovatis manifeste alatis, perianthioque proximaliter

utriculoso differt.

Frutex compactus, rotundatus, 200-300 mm altus,

monocaulis. Folia 130-230 x 65-140 mm, glabra,

tripinnatifida; petioli 60-75 mm longi, pulvinis pro-

minentibus; foliola 2 mm lata, canaliculata, apices obtusi.

Inflorescentia panicula crassa capitulis composita 200-

1 240 mm longa. Capitula ovoidea 35 — 40 mm in diam.

Bracteae involucrales glabrae, valde transverso-ovatae, 5-

6 x 8-10 mm, manifeste alatae, alis hyalinis. Perianthium

rectum, 15-16 mm longum; tubus inflatus 5 mm longus,

proximaliter glaber, distaliter sparse sericeus; ungues gra-

ciles, adpresso-sericei, recurvati post anthesin; limbi

* Compton Herbarium, National Botanical Institute, Private Bag X7,

Claremont 7735.

MS. received: 1993-02-15.

lanceolato-acuti. dense adpresso-sericei. Stylus rectus, gla-

ber, 12-14 mm longus. Stigma cylindrico-clavatum 1.5

mm longum, proximaliter leviter annulatum. Ovarium

ovoideum, 1 mm longum, dense villosum. Squamae

hypogynae lineari-subulatae, 1.5 mm longae. Fructus

achenium, late cylindricum vel parum obovoideum.

villosum, 7-8 x 3 mm, ad apicem rostratum, basis truncata

et pedicellata, fimbriata trichomatibus crassis (Figurae 1,

2 & 3).

TYPE. — Cape Province, 3319 (Worcester): Blokkop.

above Villiersdorp on southeast-facing slopes, (-CD), 2-

7-1992, Rourke 1971 , (NBG, holo.; BOL, E, K, MO,

NSW, PRE, S, STE).

Compact rounded shrub 200-300 mm high with single

stout main stem, 10-15 mm in diam., branching near

ground level, initially with fine juvenile leaves at base.

Mature leaves 130-230 x 65-140 mm. glabrous, usually

tripinnatifid, petiolate; petiole 60-75 mm long with prom-

inent pulvinus; leaflets canaliculate on upper surface 2

mm broad, apices obtuse. Inflorescence a stout panicle of

capitula opening basipetally, 700-1 240 mm long. Pedun-

cle up to 900 mm long, terete, glabrous, 4-6 mm in diam.

Capitula ovoid, 35—40 mm in diam. Involucral bracts gla-

brous, very broadly transverse-ovate, 5-6 x 8-10 mm,

prominently winged with hyaline wings. Receptacle very

narrowly conic-oblong, 15 x 3 mm. Floral bracts very

broadly transverse-ovate, glabrous, with hyaline wings.

Perianth straight in bud. 15-16 mm long: tube region in-

flated, 5 mm long, glabrous basally but sparsely sericeous

above; claws slender, adpressed sericeous, recurved at an-

thesis; limbs lanceolate-acute, densely adpressed seri-

ceous. Style straight, glabrous, 12-14 mm long. Pollen

presenter clavate-cylindric, slightly annulately thickened

at junction with style, 1.5 mm long; stigmatic groove ter-

Bothalia24,] (1994)

FIGURE 1. — Sen hi in ulusiupa Rourke, inflorescence and llowcrs. A, end of scape showing panicululely arranged eapiiula opening in basipeial

succession; B, longitudinal section through capitulum; C, perianth and floral bract prior to opening; D, perianth and floral bract after opening;

E, involucral bract; F, single perianth segment; G, gynoecium; H, mature fruit. A-G, Rourke 1971', H, Rourke 2005.

minal. Ovary ovoid, 1 mm long, sharply differentiated

from style, densely villous. Hypogynous scales linear-su-

bulate, 1 .5 mm long. Fruit broadly cylindric to slightly

obovoid, villous achene, 7-8 x 3 mm, beaked terminally,

truncate and pedicellate at base with basal fringe of stout

trichomes.

Diagnostic characters: Serruria altiscapa is related to

S. williamsii Rourke but is readily distinguished by its

short, compact habit, its exceptionally tall 700-1 200 mm

long inflorescences (hence the specific epithet), by the gla-

brous, prominently winged transverse-ovate involucral

and floral bracts and by the very utriculose perianth tube.

Bothalia 24,1 ( 1994)

3

sal shoot with emergent inflorescence scape (left) and the

strongly lignified persistent remains of previous years’ scape

(right). Type material, Rourke 1971 .

In S. williamsii the involucral bracts are clasping and cym-

biform with villously fimbriate margins and the inflores-

cences are usually shorter, from 100-400 mm. In addition,

the leaflet apices of S. altiscapa are obtuse, unlike the

sharply cuspidate leaflet apices in S. williamsii.

Serruria altiscapa is also related to S. elongata (P.J.

Bergius) R. Br. On Blokkop above Villiersdorp they occur

sympatrically and flower simultaneously, yet there is no

evidence of hybridisation, suggesting that there is a strong

genetic barrier between the two species.

The annual vegetative growth increments in this spe-

cies are rather short resulting in a low, compact, rounded

form in mature plants. In early winter certain vegetative

shoots produce a terminal peduncle 700-1 200 mm long

which develops into a large panicle of capitula with sec-

ond order branching. By early June the capitula begin

opening, in centrifugal succession, continuing until matu-

rity in November, by which time a new, short, vegetative

growth flush commences. An axillary bud below the fruit-

ing inflorescence develops into a new, short vegetative

shoot which then terminates in a further inflorescence the

following winter. As the peduncles are highly lignified

they persist for several years on the adult plants before

decaying. In some plants these lignified peduncles may

persist for from three to four years before disintegrating.

Distribution and habitat : at present only two occur-

rences of this species are known from nearly adjacent

mountains in the southwestern Cape. Several populations,

FIGURE 3. — Serruria altiscapa Rourke. Complete plant showing ha-

bit. Current season's inflorescence and persistent lignified re-

mains of inflorescences from three previous flowering seasons

are evident. Type material, Rourke 1971.

4

Bothalia 24,1 ( 1994)

each consisting of approximately 300 plants, occur on the

middle south slopes of Blokkop above Villiersdorp and

about 15 km to the west; there are a few smaller popu-

lations on the lower south slopes of the Hottentots Holland

Mountains in the Viljoen’s Pass-Nuweberg area (Figure

4A). Serruria altiscapa grows in mesic mountain fynbos

on south-facing slopes at elevations between 700 and

1 100 m. Flowering commences in early June, reaches a

peak in August and continues until October. Fruits are

shed during November and December.

It is regrettable that at the time of collecting the type

material (1992) the main populations of S. altiscapa on

Blokkop above Villiersdorp were in imminent danger of

being choked by Pinus pinaster saplings. Unless immedi-

ate attention is paid to removing this massive infestation

of alien vegetation, S. altiscapa has little chance of sur-

vival at this locality.

The oldest specimen of this species is a single leafless

inflorescence in the Burmann collection of the Delessert

herbarium in Geneva (sheet 005770/000143 in G-DEL).

The locality given is merely ‘Caput bonae Spei’ and al-

though no collector’s name is mentioned it was probably

gathered by H.B. Oldenland (1663-1697). Some 250

years elapsed before T.P. Stokoe obtained further herbar-

ium material in May 1935 since which time only seven

subsequent collections have been made, which is surpris-

ing considering dramatic growth form of the species.

CAPE. — 3319 (Worcester): Blokkop above Villiersdorp, (-CD), Nov.,

Paterson -Jones s.n. ( NBG 122855 ); Blokkop above Villiersdorp, May,

Rourke 1 7/0(NBG); Blokkop, above Villiersdoip, July. Rourke 1971 (BOL,

FIGURE 4. — A, distribution of Serruria altiscapa Rourke; B, distribu-

tion of 5. gremialis Rourke. •; S. balanocepMala Rourke, O .

E, K, MO, NBG, NSW, PRE, S, STE); Blokkop above Villiersdorp, Nov.,

Esterhuysen 35578 (BOL); Drakenstein Mts near Villiersdorp, June, Van

Breda 1875 (PRE); Blokkop above Villiersdorp Wild Flower Reserve, in

fruit, Pan., Rourke 2005 (NBG). 3419 (Caledon): Nuweberg Forest Reserve,

(-AA), Aug., Rourke 827 (NBG); between Viljoen's Pass and Somerset

Sneeuwkop, Nov. 1942, Stokoe 8609 (BOL); between Viljoen’s Pass and

Somerset Sneeuwkop, 28-5-1935, Stokoe 3003 (BOL); Nuweberg, towards

Boesmanskloof, May, H.C. Taylor 10301 (PRE, STE).

Serruria gremialis Rourke sp. nov„ a habitu multi-

cauli patulo, capitulis manifeste pedunculatis, stylis

arcuatis adaxiali in dimidio inferiore pubescentibus, et

stigmate cylindrico-clavato, distinguitur.

Fmtex multicaulis e lignotubere, suberectus vel patu-

lus, 300-500 mm altus, ad 1 m in diam. Rami erecti ad

patentes, puberuli. Folia ascendentia vel subpatentia,

puberula sed glabrescentia; bipinnatifida ad tripinnatifida,

10-30 x 10-25 mm; petioli 2-12 mm longi; foliola can-

aliculata, acuta. Inflorescentia capitulum globosum 25-30

mm in diam., solitaria et terminalis vel terminalis aliquot

capitulis axillaribus; pedunculi 10-60 mm longi. Bracteae

involucrales ovato-acutae, dense velutinae. Perianthium

adaxialiter valde arcuatum, 12-15 mm longum, sericeum

vel villosum. Stylus cygneus, adaxialiter valde curvatus,

12-15 mm longus, puberulus in dimidio inferiore, glaber

et decrescens apicem versus. Stigma cylindraceo-clava-

tum, 2 mm longum. Ovarium ovoideum 2 mm longum,

lanatum. Squamae hypogynae subulato-acuminatae, 2 mm

longae. Fructus achenium ellipsoideum 7-8 mm longum,

ad apicem rostratum, basis truncata et pedicellata, fim-

briata (Figura 5).

TYPE. — Cape Province, 3319 (Worcester): Jonaskop,

Riviersonderend Range, lower north slopes below

Jonasplaats, (-CD), 30-7-1987, Rourke 1877 (NBG, holo.;

BOL, E, K, MO, NSW, PRE, S, STE).

An erect, suberect to slightly sprawling shrublet, 300-

500 mm in height, up to 1 m in diam., with multiple stems

produced from the crown of a persistent lignotuber.

Branches erect to sprawling, 2 mm in diam., puberulous.

Leaves , ascending to subpatent, puberulous initially, later

tending to glabrescent or glabrous, bipinnate to tripinnate,

10-30 x 10-25 mm, shortly petiolate; petiole 2-12 mm

long; leaflets terete, upper surface canaliculate, apices

acute. Inflorescence a pedunculate globose capitulum 25-

30 mm in diam., solitary and tenninal or terminal with

up to 3 axillary inflorescences; peduncle 10-60 mm long.

Involucral bracts ovate-acute, closely adpressed, cartilag-

inous, densely velutinous. Involucral receptacle oblong,

10-12 x 3-4 mm. Floral bracts broadly ovate, 4-6 x 4—5

mm, apex cuspidate; outer surface lanate proximally, se-

riceous distally. Perianth very strongly adaxially curved,

12-15 mm long; tube glabrous, 2 mm long; claws and

limbs sericeous to villous; limbs elliptic, 2 mm long, very

strongly adaxially curved. Style cygneous, strongly adaxi-

ally curved, 12-15 mm long, basal half puberulous, upper

half glabrous, carmine, tapering terminally. Pollen presen-

ter cylindric-clavate, 2 mm long; stigmatic groove termi-

nal. Ovary ovoid, sharply differentiated from style, 2 mm

long, lanate. Hypogynous scales subulate-acuminate, 2

mm long. Fruit an ellipsoid achene 7-8 x 3 mm, ad-

pressed sericeous, beaked terminally, truncate and pedi-

cellate basally, with basal fringe of trichomes.

Bothalia 24,! (1994)

5

FIGURE 5. — Serruria gremialis Rourke. A, flowering shoot; B. leaf; C, longitudinal section through inflorescence; D, open flower and subtending

floral bract; E, involucral bract; F, single perianth segment; G. gynoecium; H. habit; I, mature fruit. Type material, Rourke 1877.

Diagnostic characters'. Serruria gremialis is a multi-

stemmed lignotuberous species with a suberect to sprawl-

ing habit, prominently pedunculate capitula (peduncles

10-60 mm long), strongly adaxially curved perianths, cyg-

neous, strongly adaxially curved styles, puberulous in the

lower half and cylindric-clavate pollen presenters 2 mm

long.

Distribution and habitat', this species occupies a variety

of habitats in the Breede River valley from Stettynsberg

along the southern foothills of the Villiersdorp Mountains

to Viljoen's Pass, the northern foothills of Groenlandberg,

Houhoek and eastwards along Ezeljacht to the Caledon

Swartberg. However, its main area of distribution occurs

on the northern slopes of the Riviersonderend Mountains

from Donkerhoek and Boskloof in the west, eastwards to

McGregor reaching Bushman’s River at its most easterly

limits. Isolated populations also occur on Sandberg near

Robertson (Figure 4B). Most populations occur in mesic

mountain fynbos at elevations between 270 and 760 nt.

6

Bothalia24, 1 (1994)

Flowering extends over a period of almost eight months,

from July to February but is dependant on local factors

such as site, aspect and elevation.

Serruria gremialis is a relatively common and wide-

spread species which was collected as early as the 1 830’s

by Zeyher at Houhoek but was not subsequently recog-

nised as distinct. Meisner (1856) misidentified Zeyher’s

collection from Houhoek and cited it under S. elevata R.

Br. (= S. fucifolia Salisb. ex Knight). Later, in the Flora

capensis , Phillips & Hutchinson (1912) cited material be-

longing to no less than four different species under their

concept of S. acrocarpa R. Br., including what is here

described as S. gremialis. It is therefore not surprising that

since then herbarium material of the species now known

as S. gremialis has been identified under a number of dif-

ferent names, particularly as S. incrassata Meisn. and S.

acrocarpa R. Br.

The distribution ranges of S. gremialis and S. acro-

carpa are contiguous and almost mutually exclusive ex-

cept at one site, namely Silverstream near Boskloof in the

western Riviersonderend Mountains where they grow

sympatrically (Rourke 2000 , 2002).

The epithet gremialis (= growing in a cluster from a

stump) refers to the species’ habit of sprouting multiple

shoots from a lignotuber after burning.

CAPE. — 3319 (Worcester): French Hoek Pass, Villiersdorp bridge,

(-CC), Jan., Compton 8172 (NBG); on mountain slopes between

Villiersdorp and Franschhoek, Nov., Bolus 5240 (BOL); near Genadendal,

(-CD), Burchell 7852 (SAM); Boschjeveld Mountains, Worcester side,

(-CD), Oct., Stokoe s.n. ( SAM 67852)-, Stettynsberg Mountains, (-CD),

June, Stokoe s.n. (SAM 61894)-, east slopes of Stettynsberg, (-CD), July,

Esterhuysen 15581 (BOL); Blokkop above Villiersdorp, (-CD), July,

Rourke 1 970 (NBG ); Jonaskop, Ri vier-sonderend Range, lower north slopes

below Jonasplaats, (-CD), July, Rourke 1877 ( BOL, E, K. MO, NBG, NSW,

PRE, S, STE); Riviersonderend Mts, Boschkloof, overlooking Farm Re-

treat, Poejenels Valley, (-DC), Nov., McDonald 1725 (PRE); Robertson

Dist., road between Agterkliphoogte and Robertson, (-DD), Sept., Simpson

95 (NBG); Sandberg, near Robertson lower slopes, (-DD), Sept., Rourke

1987 (NBG). 3419 (Caledon): Viljoen’s Pass, (-AA), Nov., Compton 8172

(NBG); Houhoek, (-AA), April, Guthrie s.n. ( NBG 105028)-, Groot

Houhoek, (-AA), Dec., Zeyher 3707 (SAM); Houhoek, (-AA), Feb.,

Schlechter 7330 (PRE); Caledon Baths, (-AB), Dec., Guthrie 3588 (NBG);

Silverstream near Boskloof between Villiersdorp and Genadendal, (-AB),

Nov., Esterhuysen 35297 (BOL, NBG); Silver-stream, upper reaches of

Waterkloof stream, western Riviersonderend Mts, (-AB), Nov., Rourke

2000 (NBG); Eseljagt Mountain, (-AB), July, Rourke 1969 (NBG);

Boesmanskloof Pass near McGregor, (-BA), Sept., Lewis 6069 (NBG);

Donkerhoek between Genadendal and Villiersdorp. (-B A), Dec., Pappe s.n.

(SAM); Bushmans River, (-BB), Sept., Compton 11914 (NBG).

Serruria balanocephala Rourke, sp. nov., a habitu

monocaule, capitulis pedunculatis terminalibus, stylis

basaliter puberulis adaxiale cygneis, stigmatibus conico-

acutis et perianthiis villosis, distinguitur.

Frutex erectus, monocaulis, 0.5- 1.0 m altus. Rami

erecti, puberuli demum glabrescentes. Folia ascendentia,

profunde bipinnatifida ad tripinnatifida 40-60 x 15-30

mm; petioli 10-20 mm longi; foliola canaliculata, sparse

puberula demum glabra. Inflorescentia capitulum

globosum pcdunculatum et aliquot capitula axillaria. Ca-

pitula ovoidea 25-30 mm in diam. Pedunculi 20-50 mm

longi, puberuli, graciles. Bracteae late ovatae, 5-6 x 2-3

mm, dense lanatae sed apices recurvi, glabri. Perianthium

adaxialiter valde cygneum, 12-14 mm longum villosum.

Stylus cygneus, adaxialiter valde curvatus, 12-14 mm

longus, basaliter puberulus. Stigma deorsum curvata,

ovoideo-acuta, stylo juncta collo angusto. Ovarium

ovoideum, 1 mm longum dense lanatum. Squamae

hypogynae subulatae, 1.5 mm longae. Fructus achenium

anguste ellipsoideum 8-10 x 2-3 mm, dense adpresso-

sericeum, ad apicem rostratum; basis truncata et pedi-

cellata, manifeste fimbriata (Figura 6).

TYPE. — Cape Province, 3320 (Montagu):, Muurrivier

Kloof, north slopes of the Langeberg in the Boosmansbos

Wilderness Area, (-DD), 7-9-1990, Rourke 1933 (NBG,

holo.; B, BOL, E, K, MO, NSW, PRE, S, STE).

An erect rounded shrub 0.5-1. 0 m in height, up to 1.5

m in diam., with single main stem up to 20 mm in diam.,

branching near ground level. Branches erect, puberulous,

tending to glabrescent later, 1. 5-2.0 mm in diam. Leaves

ascending, deeply bipinnatifid to tripinnatifid, 40-60 x

15-30 mm, petiolate; petiole 10-20 mm long. Leaflets

terete, canaliculate on upper surface, sparsely puberulous,

later glabrescent to glabrous, apices obtuse. Inflorescence

a single terminal pedunculate capitulum with several ax-

illary pedunculate capitula clustered towards apex of flow-

ering shoot, opening in basipetal succession. Capitula

ovoid, 25-30 mm in diam., peduncles 20-50 mm long,

crisped-puberulous, slender, peduncular bracts ovate-acu-

minate, glabrescent. Receptacle narrowly obconic, 10-12

x 3 mm. Floral bracts broadly ovate, 5-6 x 2-3 mm,

densely lanate, terminating in thickened slightly recurved

glabrous apex. Perianth strongly cygneous and adaxially

curved, 12-14 mm long; tube laterally compressed, gla-

brous, 2 mm long; claws strongly incurved, sericeous;

limbs ovate-acute, 1 mm long, 3 abaxial limbs villous,

adaxial limb adpressed-sericeous. Style strongly adaxially

cygneous, 12-14 mm long, lower third puberulous, car-

mine becoming reddish black towards apex. Pollen pre-

senter downwardly curved, ovoid-acute, with angular

facets, separated from style by narrow neck, dark reddish

black; stigmatic groove terminal. Ovary ovoid 1 mm long,

densely lanate. Hypogynous scales subulate, yellow, 1.5

mm long. Fruit a narrowly ellipsoid achene, 8-10 x 2-3

mm, densely adpressed-sericeous, beaked terminally, trun-

cate and pedicellate basally with prominent fringe of tri-

chomes.

Diagnostic characters'. Serruria balanocephala is a

single-stemmed re-seeder, with an erect, upright habit at-

taining 1 m in height at maturity with terminal peduncu-

late capitula. It is distinguished by its adaxially cygneous

style, puberulous in the lower half, the conic-acute,

slightly faceted pollen presenter and villous perianth.

Distribution and habitat: this species has a rather re-

stricted distribution along the northern slopes of the

Langeberg Range. It occurs from a few kilometres south-

east of Montagu, eastwards to Barrydale and thence to

Brandrivier just west of Garcia’s Pass, a distance of ap-

proximately 80 kilometres, but has not been recorded east

of Garcia’s Pass (Figure 4B). Serruria balanocephala ap-

pears to be confined to a narrow belt of dry mountain

fynbos, at elevations of approximately 600 to 650 m in

the lower northern foothills of the Langeberg, fringing the

Little Karoo. Flowering takes place between late August

Bothalia 24. 1 (1994)

FIGURE 6. — Serruria balanocephala

Rourke. A, flowering shoot,

bud and post pollination

phase; B, longitudinal section

through inflorescence; C, open

flower and subtending floral

bract; D, gynoecium; E, invo-

lucral bract; F, perianth limb;

G, leaf; H, lateral view of

conic-acute, slightly faceted

pollen presenter; I, mature

fruit. Type material, Rourke

1933.

and November. The inflorescences are pinkish silvery and

have no perceptible scent.

The epithet balanocephala is a compound from the

Greek balanos , an acom, and cephale, a head, alluding to

the distinctive ovoid-acute form of the pollen presenter.

CAPE. — 3320 (Montagu): north base of Langeberg Mts, 5-6 miles ESE

of Montagu towards Barrydale, (-DC), Aug. 1965, Dahlgren & Strid 2268

(NBG); Langeberg above Witbooisrivier, (-DD), Jan., Burger 97 (PRE,

STE); Grootvadersbos State Forest, north side of Langeberg above

Witbooisrivier, (-DD), Sept., McDonald & Morley 1013 (STE); Muurrivier

Kloof, north slopes of Langeberg near Barrydale, (-DD), Aug., Rourke 1881

(NBG ); Langeberg Mountain catchment area, Barrydale, municipal area,

lower slopes and next to Huis River, (-DD), Oct., T.J. van der Merwe 157

(PRE, STE); Muurrivier Kloof, north slopes of Langeberg in Boosmansbos

Wilderness Area, (-DD), Sept., Rourke 1933 (NBG. B, BOL, E, K, MO,

NSW, PRE, S, STE). 3321 (Ladismith): Brandrivier. between Barrydale and

Muiskraal, (-CC), 7-9-1967, Williams 1054 (NBG).

ACKNOWLEDGEMENTS

I am most grateful to Mrs Ellaphie Ward-Hilhorst for

the care with which she prepared the line drawings and

to Mrs Jeanette Loedolff for the photograph in Figure 3.

REFERENCES

MEISNER, C.F. 1856. Proteaceae. In A.P. De Candolle, Prodromus sys-

tematic universalis regni vegetabilis 15: 283-299.

PHILLIPS, E.P. & HUTCHINSON, J. 1912. Proteaceae. In W.T.

Thiselton-Dyer, Flora capensis 5: 654-685.

ROURKE, J.P 1982. Two new species of Serruria Salisb. (Proteaceae)

from the southwestern Cape. Journal of South African Botany 48:

285-293.

ROURKE, J.P. 1 990. Four new species of Serruria (Proteaceae) from the

southwestern Cape. South African Journal of Botany 56: 497-

505.

ROURKE, J.P. 1991. 51 Serruria stellata Rourke. The Flowering Plants

of Africa: t. 2032.

'

Y

1

Bothalia 24,1: 9-14(1994)

Studies in the Marchantiales (Hepaticae) from southern Africa. 4. Mannia

capensis , section and subgenus Xeromannia (Aytoniaceae)

S.M. PEROLD*

Keywords: Aytoniaceae, Hepaticae, Mannia capensis , M. capensis var. pallida, Marchantiales, southern Africa

ABSTRACT

A taxonomic account of the genus, Mannia, and its only local representative, M. capensis, together with its distribution in

southern Africa as currently known, is given. Mannia capensis var. pallida S.W. Arnell is not recognized here as a distinct

variety, since the 'sulphuric yellow' spores by which Arnell (1963) segregated it, can be ascribed to immaturity.

UITTREKSEL

'n Taksonomiese verslag oor die genus Mannia en die enigste plaaslike verteenwoordiger, M. capensis, saam met die

verspreiding daarvan in Suider-Afrika soos tans bekend, word gegee. Mannia capensis var. pallida S.W. Arnell word nie hier

as 'n egte varieteit erken nie, aangesien die ‘swawel-geel’ spore, waarvolgens Arnell (1963) dit afgesonder het, bloot aan

onvolwassenheid toegeskryf kan word.

Mannia Opiz in Corda, Genera hepaticarum.

Beitrage zur Naturgeschichte 12: 646 (1829); Frye & L.

Clark: 60 (1937); R. Schust.: 60 1 ( 1 953); D. Shimizu & S.

Halt.: 60 (1953); S.W. Arnell: 267 (1956); S.W. Arnell: 71

( 1963). Type species: M. raddii Opiz = M. triandra (Scop.)

Grolle.

Grimaldia Raddi: 356 (1818); Steph.: 792 (1898);

Howe: 40 (1899); Evans: 43 (1923); Sim: 21 ( 1926). Type

species: G. dichotoma Raddi, nom. illeg.

Synonymy partly after Grolle: 56 (1983a).

Thalloid, smallish to medium-sized, firm, compact and

somewhat leathery to lacunose dorsally, linear or strap-

shaped, bright green fresh, turning grey in older, dead

parts, margins black; in crowded patches, on shallow soil

covering rocky outcrops in exposed or somewhat sheltered

niches, xerophytic.

Branches simple, or once, rarely twice pseudo-dichoto-

mously branched, occasionally with latero-ventral or api-

cal innovations; thickened over midrib, gradually thinning

laterally into wings; apex entire, black tips of scale ap-

pendages reflexed over it; not grooved. Dorsal epidermis

hyaline, cell walls thin to thickened, trigones present; air

pores simple, inconspicuous, slightly raised, with 3 con-

centric rings of cells, their walls thin, leading below into

small air chambers, these storied, mostly not containing

chlorophyllose filaments; oil cells present; storage tissue

up to twice (or more times) thicker than assimilation tissue

above; ventrally reddish black; rhizoids some smooth, oth-

ers pegged; ventral scales in one row on either side of

midrib, red-black, large, obliquely triangular or semilunate

with 1 or 2 linear-lanceolate appendages, not constricted

at base, with a few scattered, colourless oil cells.

* National Botanical Institute, Private Bag X101, Pretoria 0001.

MS. received: 1993-03-26.

Dioicous , rarely monoicous, when gametangia on dif-

ferent branches. Antheridia immersed in rows along centre

of branches with projecting conical papillae turning pur-

ple. Carpocephala terminal, arising from apical notch and

raised on stalk with single rhizoidal furrow, disk apically

domed to low conical, not lobed, continuous with cam-

panulate involucres, each with protruding spherical cap-

sule, dehiscing by discrete apical lid, the wall lacking

annular thickenings. Spores with convex distal face bear-

ing conspicuous hemispherical bullae, wing thick and un-

dulating, proximal face on raised platform, with low

triradiate mark; elaters tapering, 2- or 3-spiral.

Mannia capensis (Steph.) S. W. Arnell, in Mit-

teilungen der Botanischen Staatssammlung, Miinchen 16:

263; S.W. Arnell 72: (1963); O.H. Volk: 233 (1979). Type:

Bloemfontein, leg. Rehmann (G, holo.!).

Thallus smallish to medium-sized, linear or strap-

shaped, occasionally jointed, firm and compact, older

parts rarely somewhat lacunose, bright green fresh, some-

times on exposure to strong sunlight, becoming reddish

or bronzed, especially toward margins, speckled with nu-

merous small, slightly raised stomata, boundaries of nar-

row air chambers below faintly visible from above, not

grooved, flat to slightly concave dorsally, margins with

black borders narrow distally, but gradually widening

proximally, somewhat scalloped, apically with cluster of

shiny, reddish black, lanceolate scale appendages reflexed

over edge when wet (Figure 1 A); Hanks black and shiny,

transversely slightly wrinkled, tightly incurved and meet-

ing along middle above dorsal face, becoming almost tu-

bular, and partly exposing closely imbricate ventral scales

with forwardly directed, oblique appendages, when dry;

in crowded patches, simple or once, rarely twice, pseudo-

dichotomously branched, sometimes with ventral innova-

tions emerging laterally or from apical V-shaped sinus.

Branches with total length up to 12 mm. terminal seg-

ments 3-7 mm long, closely to widely divergent, 2-4 mm

10

Bothalia 24.1 (1094)

FIGURE I .—Mannia capensis. A, dorsal aspect of male thallus with rows of antheridia; B, apical portion of female thallus with stalk arising rom

apical sinus and interrupted further up, at its tip bearing unlobed archegoniophore, involucres partly covering 2 capsules; C, ventral aspect o

thallus showing scales; D, transverse section of thallus; E, transverse section of male thallus through antheridia; F, transverse section of air

pore, dorsal cells, and assimilation tissue; G, air pore and dorsal cells from above; H, scales from foot of stalk; I, transverse section of stalk,

.1, ventral scale with one appendage; K, ventral scale with two appendages. A-D, G-I, K, Glen 2117; E, F-J, Koekemoer 637. Scale bars.

A_C, I mm; D, E, 500 pm; F, G, 50 pm; H, 1, 250 pm; J, K, 500 pm. Drawings by A. Pienaar.

Bothalia 24,1 (1994)

wide, 525-775 |im thick over midrib, laterally thinning

out into wings (Figure ID); apex entire, hidden by re-

flexed appendages of distalmost ventral scales; margins

acute, dorsally black or bronze-black along edge, ventrally

entirely black, flanks sloping obliquely upward and out-

ward; ventral face medianly broadly keeled, black, on ei-

ther side of midrib a row of imbricate, obliquely directed

reddish black scales with appendages (Figure 1C). Dorsal

epidermal cells unistratose, hyaline, from above somewhat

polygonal, 27-30 x 15-25 pm. covered with minute spic-

ules, trigones conspicuous but not bulging, in transverse

section closely packed, rectangular, 30.0-37.5 x 15.0 pm;

marginal cells purple-black, isodiametric, 20 x 20 pm, or

rectangular, 27 x 15 pm, 2 or 3 rows wide, row at edge

with outer wall convex, trigones lacking; air pores slightly

raised, simple, oval or rounded, up to 25 x 15 pm. (88-)

100-200 pm distant from each other, bounded by 3 con-

centric rings of cells, the radial walls not thickened: in-

nermost row, the remains of a collapsed cell ring, usually

numbering 7 flattened cells, trapezoidal in shape, 10 pm

long, inner wall 5 pm wide, outer wall 10 pm wide, next

row generally composed of 8 cells, transversely ovate, ±

12.5 x 15.0 pm. partly overlying outer row of somewhat

curved, larger cells, ± 30 x 32 pm (Figure 1G); assimi-

lation tissue 200-250 pm thick in transverse section, air

chambers in several layers, medianly vertically storeyed,

toward margins obliquely sloping, densely crowded,

small, especially the lower ones, incompletely bounded

by uniseriate walls consisting of cells mostly 17.5 x 12.5

pm, and in uppermost layer, below air pore, with a few

erect chlorophyllose filaments, the top cell of which con-

ical, up to 37 x 15 pm, those below rounded, 22 x 25

pm, containing numerous chloroplasts, otherwise almost

impossible to distinguish between bounding cells and

those forming filaments (Figure IF); storage tissue 300-

500 pm thick, cells angular, 6- or 7-sided, 40-62 pm

wide; oil bodies solitary, scattered throughout thallus, al-

most entirely filling cell, ± 67 x 37 pm; rhizoids arising

from ventral epidennis, some smooth, up to 32 pm wide,

others pegged, only 10 pm wide. Scales ventral, in 2 lon-

gitudinal rows, one on either side of midrib, black to red-

dish black, obliquely triangular or semilunate, with

concave base, imbricate, large, body of scale ± 600 x 600

pm, cells rectangular, (35— )52— 62 x 17 pm, walls straight

or slightly sinuous, bearing 1 (Figure 1 J) or 2 (Figure IK)

(rarely 3), forwardly directed, linear- lanceolate append-

ages, not reaching thallus margins, except at apex, up to

± 110 pm long, 17.5 pm or 5 cells wide below, tapering

to single cell width toward apex, cells 32^f0 x 15-17

pm, remains of oil cells scattered, clear, round, 20 x 30

pm, margins entire, occasionally with a few mucilage pa-

pillae.

Dioicous (rarely monoicous, but gametangia on differ-

ent branches). Androecia consisting of up to 4 rows of

conical papillae, 100 pm high, acropetally arranged along

centre of branches (Figure 1A), green when young, turn-

ing purple with age, each with opening leading into im-

mersed antheridial cavity below (Figure IE). Gynoecia

terminal, emerging knob-like from V-shaped, apical sinus

and surrounded by numerous, arched, dark purple lanceo-

late scales up to 1150 pm long, narrowing from 4-6 cells

wide at base to one cell wide at apex (Figure 1H), stalk

eventually 5-25 mm long, cylindrical, 400 pm in diame-

ter, irregularly ovate in transverse section, with single rhi-

zoidal furrow (Figure II), only a few scales carried along

and also present, but obscured at top of stalk, where joined

to carpocephalum. the latter 2 mm across, straw-coloured,

domed to low conical, papillose centrally (with compound

pores), not or scarcely lobed, continuous with ( 1— )2 — 4- in-

FIGURE2. — Mannia capensis. SEM micrographs of spores and elaler. A, distal lace of spore; B, distal face ol spore slighil) tilted; C. distal lace ol

‘yellow’ spore; D, spore seen from side; E, proximal face of spore; F, elater. A, E, Eyles CH 1179; B, Bosnian 199; C. F, Volk 00828; D,

Burrows 2521. A, E, x 540; B. D. x 480; C. x 570; F , x 740.

12

Bothalia 24,1 (1994)

volucres, these membranous, wide-mouthed and bell-

shaped, each with single, protruding, globular, dark brown

capsule (Figure IB), the wall with 4-6-sided cells, 27-62

x 37 pm, lacking annular thickenings and dehiscing along

row of smaller cells, 37 x 27 pm, forming a well-defined

lid. Spores 75-80 pm in diameter, polar, roughly triangu-

lar-globular, light brown, semitransparent, with distal face

convex (Figure 2A, B), bearing 10-13 crowded and very

conspicuous, hemispherical bullae, ± 22 pm wide, mostly

well separated from each other, occasionally 1 or 2 col-

lapsed; wing lighter, straw-coloured, 7.5 pm wide, porate

at each of 3 angles (or in between as well), wavy and

undulating, inflated and twisted in side view (Figure 2D);

proximal face with raised base, triradiate mark present but

weak (Figure 2E), slightly raised or not, arms not extend-

ing to edge, with shallow, crowded and overlying areolae

in centre, both faces generally covered with granules. EI-

aters reddish brown, tapering toward ends, 212-275 pm

long. 12.5 pm wide in centre, 7.5 pm wide at tips, 2-3-

spiral (Figure 2F). Chromosome number, n = 9 (Bomefeld

1987).

Mannia is said by Schuster (1992) to be a relatively

large genus with ± 18 species worldwide, but Engel

(1990) says that there are only six species. All of them,

except M. capensis , are from the northern hemisphere.

The species are generally distinguished from each other

by the size of the plants, the compactness of the assimi-

lation tissue, the colour and size of the scales and their

appendages and by the dorsal cells of the thallus, whether

they are thin- or thick-walled, whether the epidermis re-

mains intact and leathery, or regularly becomes lacunose

on ageing and whether the plants are aromatic or not.

Mannia capensis has spores which, in morphology, are

closely similar to those of the American M. califomica,

but in the latter species they are purple in colour. Two

subfamilies are recognized in the Aytoniaceae, Aytoni-

oideae and Reboulioideae. In southern Africa the latter

comprises the genera Asterella, Cryptomitrium and

Mannia.

The genus was originally known as Grimaldia Raddi,

but the specimen on which it was based, contained two

elements. Evans (1938) showed that the name, Grimaldia,

is illegitimate and recently Grolle (1983b) submitted a

proposal to conserve the name, Mannia Opiz (1829),

against Cyathophora S.F. Gray (1821). Two subgenera are

recognized by Grolle (1976): Mannia and Arnelliella, with

the former having two sections, Mannia and Sindonisce\

M. capensis was placed by him in section Mannia. Later

on, after the introduction of the new version of Article

10, ICBN (Voss et al. 1983), Grolle (1983c) changed this

grouping as follows:

subgenus Mannia (subgenus) Arnelliella (Mass.) Grolle,

with section Neesiella (Schiffn.) D. Shimizu & S. Hatt.

subgenus Xeromannia Grolle with sections Xeromannia

and Sindonisce.

The two species, M. fragrans and M. androgyna (the

latter selected as the type of subgenus Xeromannia ), are

classified in section Xeromannia. They are closest to M.

capensis and therefore, it too is placed in section Xero-

mannia, subgenus Xeromannia.

Schuster (1956, 1992) is, however, convinced that the

division of Mannia into subgenera, based largely on the

form of the aerenchyma tissue ‘compact’ versus ‘loose’,

is artificial. He concluded that it ‘is perhaps an artifact

based on environmentally induced distinctions’ (Schuster

1956). Furthermore. Schuster (1992) (and others) regard

the air chambers as lacking chlorophyllose filaments, stat-

ing that ‘secondary walls are largely incomplete, their free

margins are usually irregular, or even produced as short

teeth, which may give the superficial impression of rep-

resenting ‘free photosynthetic filaments’, as illustrated in

Figure IF of this study.

Grolle (pers. comm.) suspects that M. capensis could

be conspecific with M. androgyna. From the few authen-

tically named specimens of M. androgyna examined for

this study, it is difficult to come to a definite decision

about this, although its spore ornamentation is very similar

to that of M. capensis. The dorsal epidermal cells in M.

androgyna, described by Muller (1951-1958) as having

delicate walls, appear to be somewhat thicker-walled and

yellowish, the width of the pigmented margins of the thalli

is greater, it is generally a larger plant and the scale ap-

pendages extend beyond the thallus margins, particularly

the distal ones. Live plants of M. androgyna were unfor-

tunately not available for cultivation and comparative

study with M. capensis.

Fertile plants of M. capensis can be distinguished quite

readily from other members of the Marchantiales by the

stalked, domed and unlobed carpocephala, lacking a

pseudoperianth. Sterile specimens are more difficult to

distinguish, particularly from Targionia hypophylla and

small Asterella plants. In T. hypophylla, however, the dor-

sal air pores are larger, and, when dry, white-encircled; in

Asterella species, with the exception of A. muscicola, the

air pores and air chambers are generally difficult to detect

from above, wet or dry.

Although not common, Mannia capensis has been col-

lected over a wide area in southern Africa. It is known

from Namibia, western, central and eastern Transvaal, Or-

ange Free State, northern, central, western and southern

Cape Province, as well as from Zimbabwe (Best 1990)

(Figure 3). Bizot & Poes (1982) reported its occurrence

in East Africa on the Uluguru Mountains, as disjunct. Here

Bothalia 24,1 (1994)

13

too, it grows in open, exposed sites. Otherwise it is found

in rocky clefts or on shallow soil covering granite, quartz-

ite, sandstone or sometimes dolomite, occasionally in as-

sociation with other liverworts such as Riccia spp. and

Plagiochasma spp. At Kogmanskloof, near Montagu in

the Cape, it grew together with Exormotheca pustulosa.

It is a pronounced xerophyte, with the storage tissue thick-

er than the assimilation tissue.

Arnell (1963) described a new variety from Kog-

manskloof, M. capensis var. pallida , on the basis of its

sulphuric yellow spores. These spores, as remarked by

Garside (note found with specimen Arnell 798 , (BOL

54675)). were, however, only immature. Garside further

noted that 'the bullae of the spore coat were also open at

the apex — perhaps by mounting in glycerine, they had

burst. I saw Dr Arnell’s preparation (at Sea Point where

he stayed)’. Signed 'S.G.' In a specimen, Volk 00828 , col-

lected at Okahandja. Namibia, two capsules were borne

in the same carpocephalum: one contained bright yellow

spores whereas the other had the usual brownish spores.

The ornamentation of the yellow spores was less well-

formed, with the papillae low or collapsed (Figure 2C)

and sometimes confluent; that of the brown spores was

as described above, thus furnishing clear proof for

Garside’s statement, that the yellow colour of the spores

must be ascribed to immaturity and does not justify vari-

etal status. Mannia capensis var. pallida S.W. Arnell is

accordingly not recognized here as a distinct variety.

SPECIMENS EXAMINED

NAMIBIA. — 1918 (Grootfontein): Ossa, around shaly base of rocks,

(-CA), Volk 81/147 (M, PRE). 2016 (Otjiwarongo): 7 km S of Otji-

warongo, prominent granite hill near road, on soil in shade of boulders,

(-DA), Brusse 4206 (PRE). 2017 (Waterberg): ETO 98 Oros, Fels,

schattig, (-AA), Volk 00452 (M). 2116 (Okahandja): Okahandja OK 206,

auf Granit, (-DD), Volk 00828 (M). 2216 (Otjimbingwe): L WIN 329,

Matchless Mine, Glimmerfelstrip, (-DB), Volk 006771 (M). 2217

(Windhoek): Neudamm, Glimmerschiefer. Uberhiinge am Rivier, (-AD),

Volk 00685. 00686 (M); 10 km ostlich Kl. Windhoek, Glimmerschiefer,

Uberhang, (-CA), Volk 00905 (M); Bellerode, Glimmerschiefer,

Uberhang, (-CA), Volk 86-879a. 86-880 (M); WIN 63 Glimmerschiefer,

Oberhange schattig, in Rasen, (-CB). Volk 00948 (BOL. M); Voigtland.

Klipp-Kamp, an Felsen, (-CB), Volk 11400 (BOL, M). 2317 (Rehoboth):

REH 20, Giillschau, auf Granit, (-AC), Volk 6104 (M).

TRANSVAAL. — 2427 (Thabazimbi): Sterkstroom 250 KO farm, on

top of koppie between farmhouse and R5 17 road between Vaal water and

Ellisras, on soil overlying Waterberg sandstone, (-BD), Glen 2114. 2117

(PRE). 2527 (Rustenburg): Pilanesberg Nat. Park, along new road be-

tween main road and Mankwe loop, 1.0-1 .5 km from main road, on

humus-rich soil over white felsite, (-AC), Glen 2461. 2466 (PRE);

Rustenburg, 14 km N of, on road to Northam. on soil over granite

outcrops, (-CA), S.M. Perold 227 (PRE); Magaliesberg, Rietfontein

Farm, Hamerkop Kloof, on streambank on humus-rich soil overlying

quartzite, (-CD), Glen 3176 (PRE). 2528 (Pretoria): Pretoria, Won-

derboom, (-CA), Bosman CH 199 (PRE), Bonomley BOL 54667 ( BOL);

Hennops River Valley, (-CA), Meeuse 9621 (BOL, PRE); Magaliesberg

Range, 18 km W from Lanseria Airport. Plover’s Lake Farm (owner

Everard Read), up the hill, (-DD), Condy 46 (PRE). 2530 (Lydenburg):

15 miles NE of Lydenburg (-B A), Schelpe 5906 (BOL); Sudwala Caves,

between boulders on path between Dinosaur Park and cave entrance,

(-BC), Koekemoer 637a (PRE); Sudwala Caves, ± 30 km NW of

Nelspruit, on earth bank next to path leading to Dinosaur Park, (-BC),

S.M. Perold 402 (PRE). 2626 (Klerksdorp): Klerksdorp, (-DC), Sister

Victoria 273 (PRE). 2628 (Johannesburg): Kuilfontein Farm, 10 miles

SE of Heidelberg, quartzite rock, acid, sandy soil, (-CB), Mogg 37590

(PRE). 2630 (Carolina): Kangwane, Songimuelo Game Res., Farm

Kortbegin 168 IT, on soil along flat granite outcrop, (-BB), Smook 8236a

(PRE).

O.F.S. — 2729 (Volksrust): 42 km SE of Vrede on road to Ver-

kykerskop, Farm Drie Kop at Aasvoelkop, on ground between boulders

on steep slope above stream, (-CB). Smook 6422a (PRE). 2926

(Bloemfontein): Bloemfontein, Wilde Als Kloof, (-AA), Potts CH 1178

(PRE); Bloemfontein, Eagle’s Nest, (-AA), Potts CH 1266 (PRE);

Bloemfontein Botanical Garden, uncultivated part on hill, (-AA), Volk

81/062 (BOL, M, PRE). 3025 (Colesberg): 4 miles S of Trompsburg.

around base of boulders, S aspect, (-BB), Schelpe 5282 (BOL). 3026

(Aliwal North): 12 km SE of Bethulie, Cliftonvale Farm. Albert Dist..

foot of rocky cliffs at seepage zone, (-CA), H.H. Burrows 2521 (PRE).

CAPE. — 2723 (Kuruman): just east of Bretby Mine, in damp rock

crevices and under overhanging rocks in kloof, (-CA), Oliver 1450

(BOL). 2820(Kakamas): Augrabies, Granit Felssockel Uberhang, (-CB)

Volk 00554 (M). 2823 (Griekwastad): 32 miles east from Griquatown, in

seasonally wet limestone crevices, (-DC), Schelpe 5884 (BOL). 3123

(Victoria West): Three Sisters, (-CC), Schelpe 5830 (BOL). 3219

(Wuppertal): Clanwilliam Dist., Farm Mertenhof, Bidouw Valley, on

slope facing east below a Bushman Cave, (-AA), Oliver 1465 (BOL).

3224 (Graaff-Reinet): Naudesberg Pass, 40 km N of Graaff-Reinet, at

seepage at large vertical rocks at roadside, on soil, (-BA), S.M. Perold

947 { PRE). 3320 (Montagu): Kogmanskloof, (-CC), 5. Arnell 791. 795.

798 (BOL). 3322 (Oudtshoorn): Swartberge, First River, Nordseite,

Uferboschung, (-AC), Liibenau-Nestle SA 122 (PRE).

ACKNOWLEDGEMENTS

I wish to thank the following for the loan of specimens:

Prof. O.H. Volk. Wurzburg, the curators of BOL and G,

as well as Dr Liibenau-Nestle, Kempten, Germany, for the

loan of specimens. Dr R. Grolle is cordially thanked for

refereeing this paper and for drawing my attention to the

close similarity between Mannia capensis and M. an-

drogyna and their possible conspecificity. My thanks also

to my colleagues at NB1 for collecting specimens, to the

artist, Ms A. Pienaar, the typist, Mrs J. Mulvenna, and the

photographer. Mrs A. Romanowski. for their valued con-

tributions.

REFERENCES

ARNELL, S.W. 1956. Moss Flora of Fennoscandia. 1. Hepaticae.

Gleerup, Lund.

ARNELL, S.W. 1957. Hepaticae collected in South West Africa by Prof.

Dr O.H. Volk. Mitteilungen der Botanischen Staatssammlung,

Miinchen 16: 262-272.

ARNELL, S.W. 1963. Hepaticae of South Africa. Swedish Natural Sci-

ence Council, Stockholm.

BEST, E.B. 1 990. The Bryophyta of Zimbabwe — an annotated checklist.

Kirkia 13: 293-318.

BIZOT, M. & POCS, T. 1982. East African bryophytes V. Acta Botanica

Academiae Scientiarum Hungaricae 28: 15-64.

BORNEFELD, T. 1987, The natural system of the Marchantiales based

upon cytogenical and morphological evidence. Nova Hedwigia

45: 41-52.

CORD A, A.J.C. 1829. Genera hepaticarum. Beitrage zurNaturgeschichte

23: 643-655.

ENGEL, J.J. 1990. Falkland Islands (Islas Malvinas) Hepaticae and An-

thocerotophyta: a taxonomic and phytogeographic study. Fieldi-

ana 25: 1-209.

EVANS, A.W. 1923. Rebouliaceae. North American Flora 14: 39-56.

EVANS, A.W. 1938. The invalidity of the genus Grimaldia of Raddi.

Chronica Botanica IV: 223-225.

FRYE, T.C. & CLARK, L. 1937. Hepaticae of North America. University

of Washington Publications in Biology 6: 8-104.

GRAY, S.F. 1821. A natural arrangement of British plants 1: 1-824.

GROLLE, R. 1976. Verzeichnis der Lebermoose Europas und benachbar-

ter Gebiete. Feddes Repertorium 87: 171-279.

GROLLE, R. 1983a. Nomina generica hepaticarum, references, types and

synonymies. Acta Botanica Fennica 121: 1-62.

GROLLE, R. 1983b. Proposals to conserve Mannia and Pellia (Hepatic-

opsida). Taxon 32: 135-137.

GROLLE. R. 1983c. Hepatics of Europe including the Azores. Journal

of Bryology 12: 403^-59.

HOWE, M.A. 1899. The Hepaticae and Anthocerotes of California. Mem-

oirs of the Torre y Botanical Club 7: 1-208.

MULLER, K. 1951-1958, Die Lebermoose Europas. Dr L. Rabenhorst’s

Kryptogamen-Flora 6: 416-471.

RADDI, G. 1818, Novarum vel rariorum ex Cryptogamia stirpium in agro

Florentino collectarum decades duae. Opuscoli scientifici di Bolo-

gna 2: 349-361.

14

Bothalia 24,1 ( 1994)

SHIMIZU, D. & HATTORI. S. 1953. Marchantiales of Japan 2. Journal

of the Hattori Botanical Laboratory 10: 49-55.

SIM, T.R. 1926. The Bryophyta of South Africa. Transactions of the

Royal Society of South Africa 15: 1^475.

SCHUSTER. R.M. 1953. Boreal hepaticae. A manual of the liverworts of

Minnesota and adjacent regions. The American Midland Natural-

ist 49: 257-684.

SCHUSTER, R.M. 1956. Review: Marchantiales of Japan I-IV in Journal

of the Hattori Botanical Laboratory 9: 32-44; 10: 49-55; 12:

53-75; 14: 91-107, by D. Shimizu & S. Hattori.

SCHUSTER, R.M. 1992. The Hepaticae and Anthocerotae of North

America 6: 1-937.

STEPHANI, F. 1898. Species hepaticarum. Bulletin de THerbier Boissier

6: 792-796.

VOLK, O.H. 1979. Beitriige zur Kenntnis der Lebermoose (Marchan-

tiales) aus Siidwest-Afrika (Namibia) 1. Mitteilungen der

Botanischen Staatssanvnlung, Miinchen 15: 223-242.

VOSS, E.G., BURDET, H.M., CHALONER, W.G., DEMOULIN, V„

HIEPKA, P, McNEIL, J.. MEIKLE, R.D.. NICOLSON, D.H.,

ROLLINS. R.C., SILVA. PC. & GREUTER, W. (eds) 1983.

International Code of Botanical Nomenclature adopted by The

Thirteenth International Botanical Congress. Sydney, August

1981. Regnum Vegetabile 111. Bohn, Scheltema & Holkema,

Utrecht, Antwerpen.

Bothalia 24, 1 : 15-23(1994)

Studies in the Marchantiales (Hepaticae) from southern Africa. 5. The genus

Exormotheca , E. pustulosa and E. holstii

S.M. PEROLD *

Keywords: Exormothecaceae, Exormotheca holstii, E. megastomata, E. pustulosa, E. youngii , Hepaticae, Marchantiales, southern Africa

ABSTRACT

A taxonomic account of the genus, Exormotheca, and its local representatives, E. pustulosa and E. holstii, together with their

distribution in southern Africa as currently known, is given. Exormotheca megastomata is here treated as a synonym pf E. holstii

since no distinct morphological differences could be found between them.

UITTREKSEL

’n Taksonomiese verslag oor die genus Exormotheca en die plaaslike verteenwoordigers, E. pustulosa en E. holstii, saam

met hul verspreiding in Suider-Afrika, soos tans bekend, word gegee. Exonnotheca megastomata word hier as a sinoniem van

E. holstii beskou, aangesien geen duidelike verskille tussen hulle gevind kon word nie.

Exormotheca (Godm.) Mitten in Natural history of

the Azores or Western Islands: 325 (1870); Schiffn.: 29

(1893-1895); Solms: 2 (1897); Steph.: 218 (1899); K.

Mull.: 292 (1905-1916); Schiffn.: 40 (1942); K. Mull.:

398 ( 1951-1958); Hiissel de Menendez: 193 (1962) Type

species: E. pustulosa.

Myriorrhynchus Lindb. & H.W. Amell: 8 (1884). Type

species: M. fimbriatus.

Corbierella Douin & Trab.: 321 (1919). Type species:

C. algeriensis.

Thalloid, small to medium-sized to quite large, silvery

glaucous green, dorsally with numerous conical evagi-

nations of the epidermis; in crowded patches, in damp to

rather dry, exposed areas on gravelly or sandy soil or on

soil overlying rocky outcrops, xerophytic. Branches sim-

ple, or once, sometimes twice pseudo-dichotomously fur-

cate, apex entire to shallowly notched, dorsally not

grooved, slightly concave. Dorsal epidermis raised as pus-

tular or conical evaginations over low or tall air chambers,

opening above via simple air pores, encircled by thin-

walled cells and basally occupied by chlorophyllose,

sometimes branched, cell filaments, the lateral walls ver-

tical, hyaline; storage tissue % or ± Vl the thickness of

thallus, a rather loose mesh of open, round spaces or so-

called ‘Schleintzellen’ surrounded by smaller angular

cells, single oil bodies in scattered upper and lower cells;

ventrally keeled to rounded, green to purple, flanks slop-

ing obliquely upward and outward; rhizoids some smooth,

others tuberculate. Scales smallish to medium-sized, pur-

ple, oblong or rounded, occasionally with 1 or 2 append-

ages. otherwise large, hyaline with purple base, obliquely

triangular, with long, filiform appendages from apex.

Monoicous or ?dioicous. Antheridia in rows along mid-

dle of thallus, in shallow groove where development of

* National Botanical Institute, Private Bag X101 , Pretoria 0001 .

MS. received: 1993-03-26.

air chambers temporarily suppressed, sunken, necks pro-

truding conspicuously. Gynoecia raised on stalk with sin-

gle rhizoidal furrow, receptacle erect or horizontal, when

hammer-like, the parenchymatous centre covered by air

chambers that open via simple pores, above or laterally,

with 1 or 2 capsules, each supported by a seta, exserted

from bilabiate involucre and dehiscing by 4 or 5 valves

after shedding operculum; otherwise subsessile, lacking

stalk, central dome with air chambers above, epidermis

extended laterally and turning purple, forming involucres

that cover capsules on either side; capsule wall with cells

containing semi-annular thickenings. Spores 70-75 pm or

almost twice as large, ± 140 pm. triangular-globular, distal

face with large hollow, conical papillae or convoluted

areas covered with granules, proximal face without trira-

diate mark, heavily encrusted with granules; elaters taper-

ing, up to 150 pm long and trispiral or blunt at one end,

much shorter and unispiral or ringed.

Two genera are classified in the family Exormothe-

caceae K. Mull, ex Grolle (1983): Exormotheca Mitten

and the Indian Stephensoniella Kashyap. Three subgenera

are recognized in the genus Exormotheca: Exormotheca

(for E. pustulosa and E. tuberifera), Corbierella (Douin

& Trab.) Schiffn. (for E. welwitschii , E. algeriensis and

E. holstii) and Myrriorhynchus Lindb. & H.W. Amell (for

E. fimbriata) (Schiffner 1942).

Exormotheca pustulosa Mitten in Natural history

of the Azores or Western Islands: 326 (1870); Solms: 2

(1897); Steph.: 218 (1899); K.I. Goebel: 244 (1905); K.

Mull.: 292 (1906-1912); Schiffn.: 46 (1942); K. Mull:

399 (195 1-1958); S.W. Amell; 74 (1963); O.H. Volk: 232

(1979). Type: Madeira, Pico de Barcellos, leg. Johnson

(NY, holo.!).

E. africana Steph.: 18 (1917). Type: Transvaal, in Wa-

terfall gorge near Belfast, leg. Pole Evans (G).

16

Bothalia 24.1 (1994)

FIGURE 1 . — Exormolheca puslulosa. Anatomy of thallus. A, dorsal 1’acc of monoieous thallus seen from abo\ e. with row s ol anihernlia m groos es

toward apices and with young gynoecium at furcation; B. ventral face of thallus, showing scales; C, stalk (cut off) emerging just proximal to

furcation of thallus; D, 'hammer-like' head of carpocephalum with 2 dehisced capsules; E, air chamber with assimilation tissue filaments and

some storage tissue cells; F, transverse section of thallus showing two antheridia; G, air pore and surrounding cells from above; H, scale; I,

capsule wall in transverse section; .1, capsule wall from above; K, transverse section of stalk with one rhizoidal furrow. A, B, F, H,S.A/. Perold

2604 ; C, D. I-K, S'. W. Arnell 79E, G, Cl. Reid 1107a. Scale bars: A-D, 1 mm; E, G, I, J, 50 |im; F, H, 500 pin; K, 100 pm. Drawings by A.

Pienaar.

Botha] ia 24,1 (1994)

17

Thallus rather small, linear to ligulate, glaucous green,

with numerous conspicuous, partly adjoined conical

evaginations of the air chambers forming pustules (Figures

1A; 2E), opening above via air pores, dorsally toward

apex slightly concave medianly, otherwise Hat and not

grooved along midline, margins with scales extending

above or hardly so, when wet; flanks covered by imbric-

ate, purple or partly hyaline scales, generally tightly in-

curved over whitish, dorsal face when dry; in crowded

patches, simple, or once, occasionally twice (-3 times)

symmetrically or asymmetrically pseudo-dichotomously

furcate. Branches with total length up to 8 or 9 mm, seg-

ments 3-5 mm long, variously divergent, 2(-3) mm wide,

± 1000 pm thick; apex slightly tapering to rather blunt,

shallowly notched or entire; margins somewhat obtuse,

becoming more acute proximally; ventrally quite strongly

keeled, green, flanks sloping obliquely upward and out-

ward, covered by scales (Figure IB). Dorsal epidermal

cells unistratose. hyaline, rectangular to 5-sided or poly-

gonal. 30-75 x 22-25 pm, thin-walled and completely

lacking trigones, raised into conical protuberances (Fig-

ures IE; 2F); ± 300 pm high and 150-180 pm wide, in

(8)9-11 irregular rows across width of thallus; air pores

(Figure 1G) at or near top of cone, simple, round or oval,

up to 62 pm wide, bounded by 2 concentric rings of

smaller, thin-walled cells, inner cells mostly 8, ± 20.0 x

1 2.5 pm, outer cells ± 45 x 25 pm; assimilation tissue ±

200 pm thick, occupying lower ± 2/3 of air chambers,

and consisting of densely crowded filaments, sometimes

branching from close to base, top 3-5 cells free, apical

cell conical, 25-50 x 15 pm, those below rectangular,

27-32 x 22-25 pm, filled with numerous chloroplasts;

storage tissue ± 700 pm thick, loosely composed, with

larger empty spaces or so-called ‘slime’ cells, up to 80

pm wide, encircled by smaller cells ± 50 pm wide, in

between a number of cells, also ± 50 pm wide, each con-

taining a brown, granular oil body. 37.5 x 30.0 pm; rhi-

zoids arising from ventral epidermis, some smooth, 1 2.5—

20.0 pm wide, but mostly tuberculate, ± 12.5 pm wide.

Scales imbricate, apical ones oblong to rounded (Figure

IH), proximally shorter than wide, 500-1000 x 540-850

pm, occasionally with 1 or 2 appendages at margin, pur-

ple, generally with 1 or 2 cell rows at apex and base

hyaline, rarely entirely hyaline, projecting above thallus

margins or not, cells in body of scale 5- or 6-sided, up to

137.0 x 37.5^12.5 pm, smaller at margin and often brick-

shaped, 17.5 x 50.0 pm, lacking oil cells.

Monoicous (or rarely ?dioicous). Androecia in 1-3

rows along middle of thallus, close to female receptacle

(distal or proximal to it), antheridia sunken, their necks ±

175 pm long, protruding above surface between air cham-

bers, which are suppressed here to form a shallow groove

(Figure 1A). Gynoecia just proximal to bifurcation of 2

terminal branches, emerging as a central, round green

cushion, at maturity raised on a cylindrical stalk (Figure

1C), basally purple-streaked, the remainder yellowish,

length variable, up to 10 mm, diameter 350 pm, with a

single rhizoidal furrow, in transverse section (Figure IK)

cortical cells similar to medullary ones, average size 25.0

x 17.5 pm, top of stalk loosely sheathed in a short col-

lar-like outgrowth of the head, the latter internally filled

with parenchymatous tissue and covered by a row of fila-

ment-containing air chambers that open via hardly raised

simple air pores, laterally with a capsule exserted from

bilabiate involucre on either side, 3 mm across and ham-

mer-like in appearance (Figure ID), frequently, however,

bearing only 1 capsule above, when erect and oblong.

Capsule sheathed in thin calyptra, ± spherical, wall brown,

unistratose, upper part forming an operculum, cells some-

what smaller, otherwise similar to the rest. ± 57.5 x 25.0

pm, spindle- or irregularly shaped with semi-annular

thickenings (Figure 1J). in transverse section 30 pm thick,

with ‘rods’ projecting inwardly (Figure II). dehiscing by

FIGURE 2. — Exonnotheca puslulosa. Spores and thallus. A, distal lace of spore; B, outer wall of papilla broken down, show ing minute internal

granular ‘pillars’ ; C, proximal face of spore; D, granules on proximal face; E, thallus; F, conical air chamber opening above via air pore. A-D.

5.VE A me II 79 1\ E, F, S.M. Perold2604. A, C,x 515; B, D, x 3980; E, x 16; F, x 265.

18

Bothalia 24. 1 (1994)

4 or 5 irregularly shaped valves and folding back, petal-

like, seta up to 1625 x 500 pm, with ± 30 rows of cortical

cells in transverse section, foot rounded. Spores 70-75

pm in diameter, polar, triangular-globular, bright honey-

brown, distal face (Figure 2A) rounded, with up to ± 50

crowded, hollow, conical papillae, 10 pm high and 10 pm

wide, walls of papillae composed of numerous adjoining

granules stacked into tiny pillars in some areas and only

exposed where wall has broken down (Figure 2B); prox-

imal face with vestigial triradiate mark or part of it occa-

sionally present (Figure 2C), entirely encrusted with fine

granules (Figure 2D); wing absent, margin scalloped by

protruding papillae on distal face. Elaters honey-brown,

not tapering toward ends, up to 150 x 10 pm, trispiral.

Chromosome number, n = 16 (Bischler 1976).

DISCUSSION

As a member of this rather rare genus Exormotheca,

E. pustulosa is quite widespread, although the single,

highly disjunct record from Mexico (Bischler 1976) may

be an introduction (Gradstein et al. 1983). The latter au-

thors regard it as an Afro-American disjunct with a Sub-

tropical-Mediterranean range. It is known from the

following Mediterranean c mntries: Portugal, Spain,

France (with only one locality (Bischler & Jovet-Ast

1981)) and Italy, as well as from the Atlantic islands:

Azores, Madeira, the Canaries, Cape Verde and St Helena;

also from two island groups (or islands) in the Indian

Ocean: the Comores and Reunion (Bischler 1976). It is

further known from Saudi Arabia, United Arab Emirates

and Oman (Frey & Kurschner 1988) as well as from the

following African countries: Morocco, Chad, Ethiopia,

and also from Kenya, Tanzania (Bizot & Poes 1979), An-

gola, Zimbabwe and southern Africa. In southern Africa

(Figure 3) E. pustulosa has been quite rarely collected in

Namibia (O.H. Volk pers. comm.), as well as in western,

central, southern and eastern Transvaal, Orange Free State,

Lesotho and southwestern Cape.

Frey & Kurschner (1988) regard E. pustulosa as a xero-

thermic Pangaean taxon. They define the xerothermic

Pangaean element as comprising representatives with a

present pattern of distribution corresponding to the Per-

motriassic continental Pangaea region. Exormotheca

pustulosa grows in association with other liverworts, such

as Riccia species and Mannia capensis on soil around

granite or sandstone outcrops, which are generally only

temporarily wet. Under rather wetter conditions the ven-

tral scales are less conspicuous and almost entirely hya-

line. Vegetative reproduction is by ventral tubers

(Knollchen) which were spherical, scale-clad, 500 pm

wide structures in S.W. Arnell 791, one of only three

southern African specimens that had mature sporophytes.

Schiffner ( 1 942) had already placed E. africana Steph.

in synonymy under E. pustulosa, when Arnell (1953a)

stated that he could not find any real differences between

E. pustulosa and E. africana. The Indian species, E.

tuberifera, described by Kashyap (1914), seems very

closely related to E. pustulosa and so does E. ceylonensis

Meijer (1956). Exormotheca pustulosa can be distin-

guished by its low conical air chambers ± % filled with

chlorophyllose cell filaments, by its small size, oblong or

rounded purple scales, by its stalked carpocephala, by its

spore ornamentation and by its spherical tubers.

FIGURE 3. — Distribution of Exormotheca pustulosa , U, and E. holstii ,

•, in southern Africa.

SPECIMENS EXAMINED

TRANSVAAL. — 2427 (Thabazimbi): Kransberg, Farm Geelhout-

bos, NE of Thabazimbi, at T-junction of upper 2 kloofs, (-BC), S.M.

Perold 2993 (PRE); Waterberg, Welgevonden Estate, drift on Sterk-

stroom above farmhouse on cliffs, (-BD), Glen 2140. 2145 (PRE). 2527

(Rustenburg): Rustenburg Golf Course, (-CA), Heart CH 3664 (PRE).

2528 (Pretoria): Baviaanspoort, (-CB), Bosnian CH 1533 (PRE); 18 km

NE of Cullinan, on road to Sybrandskraal, near turnoff to De Tweede-

spruit, Konsensusgrotte above Malanspruit, on soil under overhanging

rock, (-DA), S.M. Perold 2604 (PRE). 2529 (Witbank): Loskop Dam

Nature Reserve, Rhenosterhoek, in gully under vegetation on edge of

exposed rock sheet, on loamy soil, small local population with Riccia

natalensis, (-AD), Cl. Reid 1107(a) (PRE); Krantz, 3 miles N of Middel-

burg, high above river, on moist soil, (-CD), Van der Merwe CH 229

(PRE). 2627 (Potchefstroom): 6 km N of Carltonville, on the road to

Pretoria, at rock outcrop immediately N of Wonderfontein, with Riccia

spp., (-AD), S.M. Perold 1207 (PRE).

O.F.S. — 2925 (Jagersfontein): Fauresmith, on Reserve koppie, (-

CB), Liebenberg CH 3659 (PRE). 3025 (Colesberg): 4 miles south of

Trompsburg, at round boulder bases on S aspect of outcrop, (-BB),

Schelpe 5284 (BOL).

LESOTHO. — 2828 (Bethlehem): ± 35 km before Oxbow on road

from Butha-Buthe, on vertical soilbank of small stream, (-CB/CD),

Perold & Koekemoer 2941 (PRE); ± 1 8 km before Oxbow on road from

Butha-Buthe, at seepage area above road, (-DC), Perold & Koekemoer

2952 (PRE).

CAPE. — 3219 (Wuppertal): Olifants River Valley, between Klawer

and Citrusdal, on moist soil by wayside, (-AC?), Wilman 664 (BOL,

PRE). 3320 (Montagu): Kogmanskloof, (-CC), 5.VT Antell 791, 798

(BOL).

Exormotheca holstii Steph. in Bulletin de FHerbier

Boissier 7: 145 (1899); Schiffn.: 66 (1942); S.W. Arnell:

76 (1963); O.H. Volk: 231 (1979). Type: Deutsch-Ost-

Afrika (Tanzania), Muse, plains, in moist sandy spots

[regio campestris, in locis arenosis humidis], leg. Holst

3107 (G 024591, holo.!).

E. youngii S.W. Arnell: 283 (1953b). Type: Transvaal,

Pilgrim’s Rest, Hendriksdal, on dry rocky veld, leg. E.M.

Young ( Duthie 5211) (BOL 54651, holo.!).

E. megastomata C. Marquand: 237 (1930). Type:

Transvaal, Krantz, 3 miles N of Middelburg, leg. F. Van

der Merwe CH 214 (BM, holo.), CH 214 (PRE, iso.!);

BOL 54643 (Duthie 5042), iso.!, synon. nov.

Bothalia 24,1 (1994)

19

Thallus medium-sized to quite large, broadly linear to

± ovate (Figure 4A), silvery green, with numerous con-

spicuous, conical evaginations of the air chambers (Figure

5B, C), medianly narrow and somewhat lower, laterally

mostly wider and taller, each opening above via an air

pore (Figure 5A), dorsally flat to centrally slightly con-

cave or shallowly grooved toward apex, scales at margins,

except at apex, mostly hidden when wet; concave to ap-

parently slightly grooved along midline of dorsal face,

white, with hyaline, apical and distal scales incurved over

margins or erect, when dry; in crowded patches, simple

or once, rarely twice or 3 times pseudo-dichotomously

furcate, symmetrical or not. Branches with total length

8—1 5(— 22) mm, terminal segments generally 4— 7(— 10) mm

long, moderately divergent, (2.5-)3.0-5.0(-7.0) mm wide,

2125-2900 pm thick; apex slightly tapering, with shallow

notch, margins obscured by air chambers; ventrally

rounded to flattish, green, or with transverse extensions

of purple scale bases or entirely purple, flanks basally

tinged with purple, sloping slightly obliquely upward and

outward (Figure 4C) covered with large hyaline scales.

Dorsal epidermal cells unistratose, hyaline, raised into 4—

6-sided air chambers, centrally 1000 x 150 pm and mar-

ginally 1500 x up to 500 pm. in 8-12 irregular rows

across width of thallus, attached to each other at sides,

but narrower apical part free (Figure 4E) for ± 250 pm,

at margins of thallus at least 500 pm at top free; cells

mostly 5- or 6-sided (Figure 4E) near top of elevated

cones 65-75 x 42-50 pm. lower down elongated, 1 25—

1 75(— 260) x 30-50 pm, thin-walled, air pores at or near

tip of cone (Figures 4D; 5D), simple, rounded or elon-

gated, ( 80—) 1 25—1 37 x 50— 75(— 85 ) pm, bounded by

smaller cells, 75-112 x 27-30 pm, sometimes a single

cell, ± 37 x 37 pm. or part of a larger cell jutting into

lumen of air pore; assimilation tissue 350—100 pm thick,

occupying basal Vs-l A of air chambers, and composed of

densely crowded filaments (Figure 4F), 7 or 8 cells long,

top cell with conical tip, 62.5 x 17.5 pm, others 67.5 x

32.5 pm, some filaments branching from near base, filled

with chloroplasts; storage tissue 1000-1150 pm thick,

cells forming a rather open mesh with ‘rounded spaces’,

150 x 100 pm, surrounded by smaller, mostly angular

cells, ± 62 x 50 pm, toward the top and base of storage

tissue and quite numerous, cells containing oil bodies,

these transversely oval. 67.5 x 55.5 pm, yellow, not en-

tirely filling cells; ventral epidermal cells isodiametric or

not, giving rise to rhizoids, some smooth, 17.5 pm wide,

others tuberculate, 12.5 pm wide. Scales imbricate (Figure

5E, F). large, 1250-1625 pm long, 900-1500 pm wide at

base, hyaline but base purple, somewhat obliquely trian-

gular with vertical side ± entire, facing toward apex of

thallus, diagonal side often toothed, cells in body of scale

long-hexagonal. 225-250 x 37-62 pm, smaller at base,

lacking oil bodies, apex mostly with branched or un-

branched, filiform appendages up to 5 (Figure 4G), 700-

1000 pm long, cells 150 x 20 pm.

Monoicous or dioicous or ?protandrous, rarely produc-

ing gametangia of both sexes simultaneously. Androecia

in 2^1 irregular rows along midline of thallus (Figure 4B),

in 1 or 2 successive linear groups, antheridia sunken,

necks protruding conspicuously, ± 1000 pm, between cen-

tral air chambers, development of the latter here tempo-

rarily suppressed. Gynoecia developing near apex of

thallus (which continues growth), causing widening and

hollowing of latter, somewhat sunken, sessile, mostly in

single groups (Figure 4A) ± 4 mm wide, occasionally with

a second one behind the first, supported on central core

of dense parenchymatous tissue, 1500 pm wide, on either

side I (or 2) slightly obliquely held capsules, 1500 pm

wide, wall with cells containing semi-annular thickenings

(Figure 41, J), on short seta, 500 x 550 pm, ± oval in

shape, cortical cells hardly differentiated from medullary

ones (Figure 4K), sheathed in thin calyptra, capsules sep-

arated by taller central dome, ± 2250 pm wide, containing

elongated air chambers (Figure 4H), basally with cellular

filaments and opening above via simple air pores, sur-

rounded by thin-walled cells, and otherwise covered by

larger cells with thicker, purple-stained walls; covering

layer of dome forming a groove laterally and continuing

on both sides as deeply purple-stained extensions; the in-

volucres, containing numerous cells with oil bodies, partly

covering capsules and at scalloped margins with smaller,

thinner-walled cells. Spores 117.5-142.5 pm in diameter,

polar, triangular-globular, dark red, distal face (Figure 6A)

rounded, with 6-8 highly convoluted, raised areas across,

22.5-27.5 pm wide, bordered by superimposed layers of

granules and hollowed in the centre, distinctly or poorly

separated by deep furrows (Figure 6B); proximal face with

triradiate mark absent (Figure 6D), but with slight flatten-

ing of the 3 facets, entirely encrusted with numerous, tiny

granules; wing absent (Figure 6C, E), margin scalloped

by projecting convoluted areas from distal face. Elaters

brownish red, tapering slightly at one end, blunt and

thicker at the other, 70-90 x 20 pm, unispiral or ringed

(Figure 6F). Chromosome number, n = 18, 32 (T.

Bomefeld pers. comm, via O.H. Volk).

DISCUSSION

Exormotheca holstii and E. pustulosa are the only two

species belonging to this rather rare genus, that also occur

in southern Africa. Exormotheca holstii has been reported

in southern Africa (Figure 3) from Namibia [several col-

lections recorded by Volk (1979)]. Botswana, northern,

central, eastern and southern Transvaal, Natal, Orange

Free State and northern Cape Province. Most gatherings

are from the Transvaal, as in recent times it has become

a more thoroughly collected area, as far as bryophytes are

concerned. Exormotheca holstii has also been found else-

where in Africa, namely Tanzania (locus classicus ) and

Zimbabwe (Best 1990). It appears to prefer quite dry,

somewhat sandy or gravelly soil, sometimes between

grass or in exposed areas that are only occasionally wet

and that overlie sandstone or quartzitic or granitic rock.

It sometimes grows in association with Riccia species,

such as R. volkii, R. rosea and R. albovestita.

Marquand (1930) described a new species of Exor-

motheca. E. megastomata, based on a sterile specimen

from Middelburg, Transvaal. This plant and its scales are

somewhat larger than in E. holstii Steph.. but the ratio of

the surface area of an apical scale to that of a section of

the thallus, works out to the same for both; no other sig-

nificant differences between E. megastomata and E. holstii

could be found after careful study of the type specimens

of both species. I. accordingly, regard them as conspecific

and place E. megastomata in synonymy under E. holstii.

Marquand stated that E. megastomata ‘is distinct from all

previously described species of that genus in the very tall

20

Bothalia 24,1 ( 1994)

I l( JURE 4. — Exormotlieculiolsiii. Anatomy of thallus. A, tliallus with sessile caipoeephalum IVom above: B, thallus with antheridial necks emerging

between air chambers in midline; C, transverse section of thallus showing tall air chambers, chlorophyllose cell filaments and storage tissue;

D, air pore and surrounding cells from above; E, top part of two partly adjoining air chambers; F, assimilation tissue with cell filaments; G,

scale with branched filiform appendages; H, caipoeephalum, upper half three dimensional, lower half in transverse section; I, capsule wall

from above; J, capsule wall in transverse section; K, transverse section of seta. A, H, K, Gennishuizen 2839; C, Perold & Koekemoer 2872;

D, S.M. Perold 2702; E, F, Glen 2190 ; G, I, J, Holst 3107. Scale bars: A-C, H, 1 mm; D-F, I— K, 100 pm; G, 500 pm. Drawings by A.

Pienaar.

Bothalia 24. 1 (1994)

21

FIGURE 5. — Exonnotheca Iwl.siii. Thai 1 us, A, thallus from abo\c, with conical air chambers; B, air chambers seen from side; C. air pores seen from

above; D, air pore at apex of cone; E, scales inflexed over apex of thallus; F, scales seen from side. A-F, R. Smit s.n. A, x 8; B, x 26; C, E x

34; D, x 150; F, x 35.

stomata’, for which he named it, and so he made the same

error as Stephani ( 1899) in referring to an entire air cham-

ber as a stoma — Schiffner (1942) had already drawn at-

tention to this. In his protologue. Marquand compared

only E. africana Steph. (a synonym of E. pustulosa) to

his new species, which is a much larger plant and differs

in several other aspects as well, such as the tall air cham-

bers and the conspicuous, triangular, mostly hyaline scales

with filiform apical appendages. Curiously, Marquand re-

ported the marginal scales of E. megastomata as ‘minute’

and was followed in this by Arnell (1963), who also re-

ferred here, but not elsewhere, to the air chambers as sto-

mata. In his description of a new species, Exonnotheca

youngii , Arnell (1953b) reported that it differed from E.

megastomata by the large marginal scales and by the free

air chambers (‘free from one another down to the base’).

FIGURE 6. — Exonnotheca hoist ii. Spores. A, distal luce; B, granules on distal face much enlarged; C. side view; D. proximal face; E. boundary

between proximal and distal face; F, elater. A, D, Volk 011670 ; C, Volk 85-766\ B. E, F, Holst 3107. A, x 293: B. E, x 794; C, x 324; D, x

300; F, x 700.

22

Bothalia 24,1 (1994)

Arnell (1963) later placed E. youngii in synonymy under

E. holstii. In his key to three species of Exormotheca,

Arnell (1963) also referred to the air chambers in E. holstii

as being free almost to the base. This is contrary to my

findings (see above). Stephani (1899) also described the

stomata (sic) of E. holstii as ‘densissima altissima, ad

% coalita, tertio supero libera’, and Schiffner (1942) re-

ferred to the air chambers as ‘seitlich bis drei Viertel

verwachsen'. AmeH’s observations must, therefore, have

been based on an error. In Schiffner’s (1942) monographic

study of the genus Exormotheca , he makes no reference

to E. megastomata. In this work he refers to the contro-

versy between himself and Muller (1940, 1941) concern-

ing the application of the epithet, E. bullosa , which he

suggested ‘als Art einzuziehen ... isf . I discussed this in

greater detail in Perold ( 1991 ), urging a return to its earlier

epithet, E. welwitschii , and furnishing reasons for it.

Fruiting collections of those Exormotheca species that

are characterized by sessile carpocephala, are extremely

rare and have only been illustrated once before by Douin

& Trabut (1919). They described Corbierella algeriensis,

a synonym of E. algeriensis , as dioicous. Of the speci-

mens studied in the present work, Giess 15383 is defi-

nitely monoicous; of the remaining 53 specimens, only

the type specimen of E. holstii, Holst 3107, Volk 01160

and Germishuizen 2839 have mature sporangia and are

dioicous or possibly protandrous.

Volk (in litt.) identified his collection Volk 81/015, from

Middelburg (the type locality of E. megastomata ), as E.

megastomata', Bomefeld (in litt.) found its chromosome

number to be n = 18, whereas specimens from Namibia

have n = 32 chromosomes. Bomefeld and Volk are of the

opinion that the material from Middelburg and from

Namibia represents two different species. However, two

different chromosome numbers manifested here are not

reflected in morphological differences. The same phenom-

enon was observed in some Riccia species, e.g. R.

argenteolimbata (Volk et al. 1988), where as many as five

chromosome numbers were found in the same species.

Exormotheca holstii can be distinguished from other

species in the genus by its large size, by its tall air cham-

bers, basal V5-V3 occupied by chlorophyllose filaments,

and laterally adjoined except for the apical 250 pm toward

the centre of the thallus and marginally for the apical ±

500 pm; by the large, hyaline, triangularly shaped scales

with long filiform appendages and by its subsessile

gynoecia. Tubers were not found.

SPECIMENS EXAMINED

NAMIBIA. — 1918 (Grootfontein): Gaikos, on quartzite sand, (-AD),

Volk 81/124 (M, PRE). 2116 (Okahandja): Erichsfelde, (-DA), Volk 11925

(M, PRE); Fort Garden Park, Okahandja, (-DD), Giess 15383 (PRE). 2217

(Windhoek): WIN 85, Farm Rietfontein, Granitzersatz, flach, durch

Sickerwasser zeitweise feucht bis nass, (-CA), Volk 01160 (BOL, PRE).

BOTSWANA. — 2127 (Francistown): Shashe Dam, 30 km S of Francis-

town, damp soil in sandy wash, ( — DA), Cl. Reid & Barker 14 (PRE).

TRANSVAAL. — 2229 (Waterpoort): Farm Driehoek, on top of sand-

stone koppie, on sandy soil, (-DC), Fourie 31 (PRE); Lokovhela 793 Farm,

campsite near Zwarthoek boundary, near edge of abandoned Eucalyptus

plantation, on well-drained sand/gravel overlying Waterberg sandstone,

(-DD), Glen 2650 (PRE). 2230 (Messina): Maname, on soil, (-CD), A.E.

van Wyk 5424 (PRE). 2329 (Pietersburg): Soutpansberg, Koedoesvlei 47

Farm, Lejuma, foot of krantzes, stony soil overlying Waterberg quartzite,

(-AB), Glen 2190 (PRE); Pietersburg, Farm Bloedrivier, 696LR, on ground

in open patches, (-CD), F. Venter 12229 (PRE). 2330 (Tzaneen): Mountain

Home Farm, Woodbush, in exposed positions on southern slopes, (-CC),

Mogg CH 3172 (PRE). 2331 (Phalaborwa): Silonque, in large colonies, in

open areas on loamy soil, weakly drained, (-CC), I.M. Retief246 (PRE).

2428 (Nylstroom): Loubad, on road between Nylstroom and Alma, near

bridge over Sand River, on soil between flat rock outcrops above river,

(-CA), S.M. Perold 820 (PRE). 2429 (Zebediela): Chuniespoort, (-BC), F.

van der Merwe CH 3666 (PRE). 2430 (Pilgrim's Rest): Bourke's Luck

Potholes, 27 km N of Graskop on R532 road, (-DB ), S.M. Perold 412 (PRE);

on R532 road to Bourke’s Luck Potholes, ± 1 km before coming to SADF

Dog Training Centre, on dry sandy slope above stream, between grass,

(-DB), S.M. Perold 2702 (PRE). 2528 (Pretoria): Soutpan road, on sandy

soil between grass, (-CA), Odendaal CH 12611 (PRE); Pretoria, (-CA),

Wager CH 3819 (PRE); Zwavelpoort, near Willows road. (-CA). Watson

PRE 56943 ; Magaliesberg, summit Boekenhoutskloof, sandy flats, (-CB),

Mogg CH 853 (PRE); Pienaarspoort, (-CB), Wager CH 852 (PRE);

Donkerpoort, (-CD), Bottomley CH 3566 (PRE). Bottomley & Doidge CH

3590 (PRE); Donkerhoek, 22.5 km E of Pretoria, along Pretoria- Witbank

Freeway, just beyond road cutting, seepage area, (-CD), M. Crosby 464

(PRE), Germishuizen 5624 (PRE), S.M. Perold 330. 2015. 2795. 2796. 2884

(PRE); open veld east of Premier Mine, (-DA), Pole Evans s.n. (T.R. Sim

Herb. 1 826) (PRE); BronkhorstspruitDist., 19kmNoftownonGroblersdal

road, at left hand side of road on soil overlying base of granite, (-DC), S.M.

Perold 144 (PRE); 33 km E of Pretoria, on R5 1 5 road to Cullinan/Bapsfont-

ein, rock outcrop next to road, at small streamlet on damp soil beneath grass,

(-DC), S.M. Perold 1371 (PRE); between Cullinan and Bapsfontein on

R515 road, just after T-junction from Rhenosterfontein, rock outcrop next

to road, (-DC ), S.M. Perold 2010 (PRE); near Bronkhorstspruit, (-DC), leg.

not known, CH 3566a (PRE). 2529 (Witbank): Triegaardts Poort, (-CA),

Verdoom CH 3759 (PRE); Middelburg, along extension of Noordkant

Street, W of town, on hill next to road, (-CB), Van Rooy & Perold 644. 645

(PRE); on road from Middelburg to township, sandstone, on slope opposite

Olifants River, (-CB), Volk 81/015 (BOL, M, PRE); Witbank, on soil,

(-CD). Reinet Smit s.n. (PRE); Middelburg, Gravel Krantz, (-CD), Van der

Merwe CH 214 (type of E. megastomata), CH 3657. CH 3665 (PRE). 2530

(Lydenburg): halfway between Dullstroom and Lydenburg, Farm Klipsteen,

seepage area, on gravel overlying quartzite, near path, (-AB), H. Anderson

1244 (PRE); 29 km from Dullstroom on road R540 from Lydenburg, at dirt

road turnoff to Boschhoek, near Marmerkop Station, on soil on hillside,

(-AB), S.M. Perold 422 (PRE); south of Lydenburg, Spitskop, on

Coromandel Farm (opposite side to homesteads), on soil on dry ledge above

waterfall, (-AB), Perold & Koekemoer 2872 (PRE); Hendriksdal, on dry

rocky veld, (-BB), Young (type of E. youngii). 2628 (Johannesburg):

Benoni, Farrarmere, on soil at rocky outcrop near Homestead Lake, (-AB ),

S.M. Perold 461 (PRE); 5 km NE of Kriel on road to Vanwyksdrift, beyond

old bridge, coarse rock outcrop, (-BB), S.M. Perold 2477 (PRE). 2629

(Bethal): 5 km W of Kriel on road to Vandyksdrift, on left side of road, on

soil in depression at flat rocks, (-AB), S.M. Perold 348 (PRE). 2630

(Carolina): Carolina, (-AA), Van der Merwe CH 3583 (PRE); Knock Dhu

Farm, 1 3 km SE of Lake Chrissie on Lothair road, common on rich black

loamy soil, in grassland, (-AD), Germishuizen 2839. 2887 (PRE).

NATAL. — 2732 (Ubombo): Zululand, Muzi swamp near Phelandaba

Cash Store, south of foot-and-mouth barrier, sedge meadow, in bare

patches overlain by coarse sand, (-BA), Cl. Reid 1024 (PRE). 3030 (Port

Shepstone): Lynton Farm, N of east end of Oribi Gorge, overlooking

Umzimkulu River. ± 2 km W of ‘Rock of Gibraltar’, on stony soil

overlying Natal Mountain sandstone, (-CB), Glen 2243 (PRE).

O.F.S. — 2627 (Potchefstroom): Sasolburg, on soil, (-DD), Kroon s.n.

(PRE). 2828 (Bethlehem): 14 km E of Paul Roux on road to Bethlehem,

at flat weathered sandstone rocks, on soil, (-AC), S.M. Perold 1349

(PRE). 2925 (Jagersfontein): Fauresmith, Pasture Research Station,

(-CB), Henrici CH 11226 (PRE). 2926 (Bloemfontein): grassveld in

University grounds, Bloemfontein, (-AA), Van Zinderen Bakker 7494

(BOL).

CAPE. — 2724 (Taung): Farm Zoetvlei, ± 50 km W of Vryburg and 6

km from turnoff on road to Louwna, 50 m W of pan, on higher ground,

with Riccia volkii, (-AA), M. Crosby 1115 (PRE); Vryburg, Heuningvlei

Salt Pan, west of town, in shallow damp hollow, occasional, (-AA),

Oliver 1446 (BOL). (A Wager bequest, PRE CH 3839, is said to be from

Stellenbosch, but this is surely a mistaken locality, as Exormotheca holstii

has not otherwise been recorded in the southwestern Cape).

ACKNOWLEDGEMENTS

I wish to thank Professors S. Jovet-Ast and T. Poes for

refereeing this article and for their helpful suggestions;

also Prof. Dr O.H. Volk, Wurzburg, for the loan of his

specimens and for fruitful discussions and Dr T. Bomefeld

Bothalia 24,1 (1994)

23

for chromosome counts. The curators of BOL, G and NY

are thanked for the loan of type specimens. My thanks

also to my colleagues at NBI for collecting specimens, to

the artist, Ms A. Pienaar, the typist, Mrs J. Mulvenna and

the photographer, Mrs A. Romanowski, for their valued

contributions.

REFERENCES

ARNELL. S.W. 1953a. Notes on South African Elepaticae. Revue

Bryologique et Lichenologique XXII, fasc. 1-2.

ARNELL, S.W. 1953b. A new South African species of Exormotheca.

Transactions of the British Bryological Society 2: 283, 284.

ARNELL, S.W. 1963. Hepaticae of South Africa. Swedish Natural Sci-

ence Council, Stockholm.

BEST, E.B. 1990. Bryophyta of Zimbabwe — an annotated checklist.

Kirkia 13:293-318/

BISCHLER, H. 1976. Exormotheca pustulosa Mitten. Revue Bryo-

logique et Lichenologique 42: 769-783.

BISCHLER. H. & JOVET-AST, S. 1981. The biological significance of

morphological characters in Marchantiales (Hepaticae). The

Bryologist 84: 208-215.

BIZOT, M. & POCS. T. 1979. East African Bryophytes III. Acta Botanica

Academiae Scientiarum Hungaricae 25: 223-261.

DOUIN, C. & TRABUT, L. 1919. Deux Hepatiques peu connues. Revue

generate de Botanique 31: 321-328.

FREY, W. & KURSCHNER. H. 1988. Bryophytes of the Arabian Penin-

sula and Socotra. Nova Hedwigia 46: 37-120.

GOEBEL, K.I. 1905. Zur Kenntnis der Verbreitung und der Lebensweise

der Marchantiaceen-Gattung Exormotheca. Flora 95: 244—250.

GRADSTEIN, S.R.. POCS, T. & VANA, J. 1983. Disjunct hepaticae in

tropical America and Africa. Acta Botanica Hungarica 29: 127-

171.

GROLLE, R. 1983. Nomina generica Hepaticarum. references, types and

synonymies. Acta Botanica Fennica 121: 1-62.

HASSEL DE MENENDEZ, G.G. 1962. Estudio de las Anthocerotales y

Marchantiales de la Argentina. Opera Lilloana 7: 1-297.

KASHYAP, S.R. 1914. Morphological and biological notes on new and

little-known West Himalayan liverworts. II. New Phytologist 13:

308-323.

LINDBERG, S.O. & ARNELL, H.W. 1884. Sandea et Myriorrhynchus,

nova hepaticarum genera. Acta Soc. pro fauna et flora Fennica

2,5: 1-9.

MARQUAND, C.V.B. 1930. XXVI. A new species of Exormotheca from

South Africa. Kew Bulletin 1930: 237-239.

MEIJER, W. 1956. A new species of Exormotheca from Ceylon. Journal

of the Hattori Botanical Laboratory 16: 12-1 A.

MITTEN, W. 1870. Hepaticae. In F.C. Godman. Natural history of the

Azores or Western Islands'. 316-328. Van Voorst, London.

MULLER, K. 1906-1912. Die Lebermoose. Dr L. Rabenhorst's Krypto-

gamen-Flora 6: 1-870.

MULLER, K. 1940. Exormothecaceae. Die Lebermoose Europas. Dr L.

Rabenhorst’s Kryptogamen-Flora 6: 275-280.

MULLER. K. 1941. Beitrage zur Systematik der Lebermoose II.

Hedwigia 80: 90-118.

MULLER, K. 1951-1958. Die Lebermoose Europas. Dr L. Rabenhorst’s

Kryptogamen-Flora 6: 417^171.

PEROLD, S.M. 1991. Studies in the genus Riccia (Marchantiales) from

southern Africa. 23. R. bullosa : typification and a full description.

Bothalia 21: 129-135.

SCHIFFNER. V.F. 1893-1895. Hepaticae. Die Nattirlichen Pflanzen-

f ami lien 1: 3-141.

SCHIFFNER, V.F. 1942. Monographic der Gattung Exormotheca.

Hedwigia 8 1 : 40-74.

SOLMS-LAUBACH, H. 1897. liber Exormotheca Mitten, eine wenig

bekannte Marchantiaceengattung. Botanische Zeitung 55: 1-16.

STEPHANI. F. 1899. Species hepaticarum. Bulletin de THerbier Boissier

7: 198-225.

STEPHANI, F. 1917. Species hepaticarum. Bulletin de THerbier Boissier

6: 18.

VOLK. O.H. 1979. Beitrage zur Kenntnis der Lebermoose (Marchan-

tiales) aus Siidwestafrika (Namibia). 1. Mitteilungen der Bota-

nischen Staatssammlung, Miinchen 15: 223-242.

VOLK. O.H., PEROLD, S.M. & BORNEFELD, T. 1988. Studies in the

genus Riccia (Marchantiales) from southern Africa. 10. Two new

white-scaled species of the group ‘Squamatae’: R. argenteo-

limbata and R. albomata. Bothalia 18: 155-163.

*

Bothalia 24,1: 25-30(1994)

Studies in the Ericoideae (Ericaceae). XIV. Notes on the genus Erica

E.G.H. OLIVER* and INGE M. OLIVER*

Keywords: Erica, nomenclature, southern Africa, taxonomy

ABSTRACT

Recent investigations of all the recorded southern African species of Erica have been undertaken to establish the characters

of importance in generic delimitation in the subfamily Ericoideae. As a result several anomalies have been found in the

circumscription and identification of certain species and in the nomenclature of others. The corrections and alterations are

published here.

U1TTREKSEL

Ondersoeke van al die bekende Suider-Afrikaanse Erica-spesies is onlangs onderneem om die kenmerke wat belangrik is

vir generiese afbakening in die subfamilie Ericoideae te bepaal. Verskeie ongerymdhede het in die omskrywing en identifikasie

van sekere spesies en in die nomenklatuur van ander aan die lig gekorn. Die aanpassings en veranderinge word hier gepubliseer.

INTRODUCTION

The last full revision of the southern African members

of the genus Erica was provided in Flora capensis by

Guthrie & Bolus ( 1905). Subsequently numerous new spe-

cies have been added by many workers with a major con-

tribution being the abbreviated revision of the genus by

Duller in 1965. In this work Duller did much to sort out

the nomenclature and references to our species in the lit-

erature. Unfortunately he did not examine the complete

collections in any of the major herbaria housing material

from southern Africa, namely BM, BOL, K, NBG, PRE

and STE. His revision also lacked descriptions of the spe-

cies.

Since the revision by Guthrie & Bolus (1905) some

180 additional species have been added which has made

it increasingly difficult to gain an overall view of the

genus. Within recent years we have come across anoma-

lies in the circumscription of certain species, in the appli-

cation of names and in the identification of certain

collections. These records and observations are rational-

ized and published in this paper.

OVARY COMPLEMENT

The genus Erica is generally characterized by having

a 4-locular, many ovuled ovary that forms a dehiscent

capsule. Guthrie & Bolus (1905) recorded that the ovary

was mostly 4-celled, very rarely 8-celled. They noted

under the species treatments that the 8-celled condition

occurred in only E. perspicua Wendl. and E. verticillata

Berg, and 4—8 in E. propendens Andrews.

During our detailed examination of all 657 species of

Erica, several additions to this list of species having non-

standard ovaries have been recorded:

* Stellenbosch Herbarium, National Botanical Institute. P.O. Box 471,

Stellenbosch 7599.

MS. received: 1993-03-31.

E. macowanii Cufino

This species is very closely related to E. perspicua. but

was distinguished by Guthrie & Bolus as having a 4-loc-

ular ovary. Examination of an isotype in BOL and numer-

ous recent collections has shown that the ovary is always

8-locular. The species is, in fact, very difficult to distin-

guish from E. perspicua.

E. colorans Andrews

This species is also closely related to E. perspicua and

has been found to have 4— 6-locular ovaries.

E. annectens Guthrie & Bolus

This species, which is endemic to the Cape Peninsula,

was not recorded by either Guthrie & Bolus (1905) or

Salter (1950) as having an 8-locular ovary.

E. pinea Thunb.

The ovary complement of this species was never men-

tioned by Guthrie & Bolus so we presume they regarded

it as having the standard 4-locular ovary. We have found

the species has 8-locular ovaries, but some collections

from the Hermanus area exhibit 6-locular ovaries.

E. trichophora Benth./E. octonaria L. Bolus/E. dulcis L.

Bolus

This group of species which forms a complex with E.

perspicua Wendl. and E. propendens Andrews, all possess

8-locular ovaries. Esterhuysen (1963) placed E. dulcis in

synonymy under E propendens and Dulfer (1965) dis-

cussed most of the above species as very closely related

under E. perspicua.

26

Bothalia 24,1 (1994)

CORRECTIONS AND ALTERATIONS OF NAMES

Erica alexandri

Until recently this species was known from only three

collections from sandy flats in the Paarl area: the type

collection by Alexander- Prior of 1847, a second collection

of his in K, not cited in Guthrie & Bolus (1905) and a

third one by F.H. Cooper in 1922. The spread of farms

and pine plantations has destroyed almost all natural veg-

etation on sandy flats in the area. Searches for the species

in likely places in the Paarl area have not met with suc-

cess.

The species was, however, recently found growing in

a wetland near the Wemmershoek sawmill east of Groot

Drakenstein towards Fransch Hoek. These plants match

those of Alexander-Prior and Cooper exactly and are the

only known surviving population of the species.

E. alexandri is strikingly similar to E. acockii , which

is known only from a few collections made in the 1930’s

from the Brackenfell/Kraaifontein area and is listed as ex-

tinct in the South African Red Data Book (Hall & Veld-

huis 1988). It has not been found since, in spite of

searches of the area, including the type locality, pointed

out to one of us (EGHO) by Acocks in 1965. It must be

assumed that the species has been destroyed by housing

development and alien acacias.

E. acockii grew in very similar conditions to E. al-

exandri, namely sandy flats with some indication of water

seepage just below the surface. This factor may be rele-

vant to the fact that both species flower(ed) in the driest

months of the year, February through to early May. The

localities of the current and former populations of the two

species are only about 40 km apart.

Morphologically the two species are very similar de-

spite the fact that they were placed in two different sec-

tions in the genus, E. alexandri in section Pachysa and

E. acockii in Section Pseuderemia. Both have heads of

mauvish pink sticky flowers with manifest dark anthers,

long gland-tipped hairs on the leaves, stems, pedicels,

bracts, bracteoles and sepals. There are only some slight

differences between the collections from the two areas.

The only clear difference lies in the leaves of E. acockii

which are shorter and more rounded than those of E. al-

exandri. Surprisingly the type collection of E. acockii has

obovoid-obconical corollas with more erect lobes as op-

posed to the urceolate corollas with spreading to reflexed

lobes in the other collections of E. acockii and of E. al-

exandri.

With only one measurable difference between the two

species coupled to the spatial separation of the popula-

tions, we believe that recognition of E. acockii at subspe-

cific level is warranted.

Erica alexandri Guthrie & Bolus in Flora capensis

4: 195 (1905); Duller: 98 (1965).

subsp. alexandri. Type: Paarl Div.; sandy flats

below Paarl Mountain, Alexander 17 (K, holo.!; BOL

fragm.!).

subsp. acockii (Compton) E.G.H. Oliv., stat. et

comb. nov.

E. acockii Compton: 43 (1934); Dulfer: 111 (1965). Type:

Stellenbosch Dist.; Brackenfell, Acocks [Acock] 1319 (BOL, holo.!; K!,

S!, STE!).

Erica auriculata/E. greyi

Both these species were based on a single specimen

and published in the same work (Guthrie & Bolus 1905).

The type of E. auriculata had only a few flowers which

were still in bud, and according to the authors, anther

appendages unlike any other species. On account of its

puberulous flowers Guthrie & Bolus (1905) placed the

species in Section Ephebus. The type of E. greyi had nu-

merous flowers in 6-8-flowered heads, and the authors

made no reference to any distinctive feature of the anthers.

They placed it in Section Pseuderemia.

Comparison of the two types showed that their anthers

are identical. Other characters of flowers and vegetative

parts also matched, and there is no doubt that the two

collections belong to the same species.

No other collections of these two species have been

made and a search for them in the more exact locality

given by Schlechter has not produced any plants. Unfor-

tunately the whole area was burnt in 1991 and thus it will

be several years before further searches will be worth un-

dertaking as it would appear that the plants are reseeders.

Erica greyi Guthrie & Bolus in Flora capensis 4:

231 (1905); Dulfer: 110 (1965). Type: Ceres Div.; Cold

Bokkeveld, Grey 658 (K, holo.!; BOL, fragm.!).

E. auriculata Guthrie & Bolus: 1 19 ( 1905); Dulfer: 68 (1965). Type:

Ceres Div.; on the Skurfde Bergen near Klein Valley, 5800 ft, Schlechter

10207 (BOL, holo.!).

Erica cerviciflora/E. sphenanthera

Both of these species have been known only from the

type collections. They were both placed in Section

Didymanthera on their possession of long tubular flowers

with exserted anthers. An examination of these collections

has shown characters not previously noted in assessing

their true relationships, namely the shape of the anthers

with minute decurrent appendages and fringed corolla

lobes. These characters allied the two species to E.

grandiflora which is highly variable in the position of its

anthers which may be included to exserted even further

than other species of Section Didymanthera.

There are clearly no characters to separate the above

two species from the older E. grandiflora and so they are

here reduced to synonymy under the older epithet.

Erica grandiflora L. /., Supplementum plantarum

: 223 (1782); Benth.: 628 (1839); Guthrie & Bolus: 57

(1905). E. exsurgens Andr. var. grandiflora (L. f.) Dulfer:

35 (1965). Type: Cape, Thunberg s.n. (UPS!).

E. cerviciflora Salish. : 362 (1802); Benth.: 664 (1839); Guthrie &

Bolus: 53 (1905); Dulfer: 33 (1965). Type: Hottentots Holland, Mulder

s.n. (K, holo.!).

E. sphenanthera Tausch: 626 (1834); Guthrie & Bolus: 52 (1905);

Dulfer: 33 (1965). Type: without locality or collector (PRAG, holo.!).

Bothalia 24,1 (1994)

27

Erica esterhuyseniae

When Compton described this species he provided a

Latin diagnosis and full English description, but did not

cite a type. However, he stated that there were clearly two

distinct varieties, var. a tetramera and var. b trimera, for

which he provided the diagnoses and holotypes. Accord-

ing to the current Code this would automatically imply

that there was a typical variety var. esterhuyseniae with

the same type as the species name.

Compton’s application of the term ‘Var. a’ would sug-

gest that he intended this variety to be the typical variety

but that he supplied the incorrect epithet for it. We are

therefore replacing Compton’s name with the correct au-

tonym and regarding the holotype of his var. tetramera as

the type of the species name.

Erica esterhuyseniae Compton in Journal of South

African Botany 7: 193 (1941); Dulfer: 75 (1965). Type:

Uniondale Div., Kamanassieberg (Mannetjieberg), 1 700

m, 1 Feb. 1941, Compton 10557 (NBG!).

var. esterhuyseniae [as var. a tetramera ]

var. trimera Compton: 193 (1941); Dulfer: 75 (1965). Type: Oudts-

hoorn Div., Swartberg Pass, 2 000 m, 28 Jan. 1941, Bond 866 (NBG!).

Erica filiformis

Guthrie & Bolus (1905) recorded the species as having

a variable number of stamens, namely usually 8 but often

7-4. They noted that 4-stamened specimens were 'tech-

nically Blaeria and not Erica'. In a recent paper (Oliver

1993) this species was discussed in detail in connection

with the relationship between Erica and Blaeria. An ex-

amination of all the available collections of E. filiformis

showed that some collections have 4-stamened flowers,

others 8-stamened flowers, with only one collection show-

ing any intermediate numbers.

In the same work Bolus (1905) described the variety,

var. maritima Bolus, from a single collection made in the

Agulhas area. An examination of the type showed that the

4-stamened flowers look superficially very similar to those

of E. filiformis, but do not possess the 4-locular. multi-

ovuled ovary typical of Erica. Instead the flowers have

2-, rarely 1 -locular ovaries with a single ovule per locule,

a feature characteristic of certain minor genera. The ma-

terial was in fact an isosyntype of Acrostemon schlechteri

N.E. Br.!

The synonymy is therefore as follows:

Acrostemon schlechteri N.E. Br. in Flora capensis

4: 353 (1906). Types: Bredasdorp Div.; Rhenosterkop, 50

ft., Schlechter 10576 (BM!, BOL!, G!, GRA!, K!, Z!);

Cape Agulhas, 250 ft., Schlechter 10559 (BM!. BOL!, G!,

GRA!, K!, MO!, P!, PRE!, S!, W!, Z!); lectotype still to

be chosen.

Erica filifonnis Salisb. var. maritima Bolus: 150 (1905). Type:

Bredasdorp Div.; hills near Cape Agulhas, 250 ft, Schlechter 10559

(BOL, holo.!).

Erica gillii

Until recently the type at Kew was the only known

collection of this species with none apparently occurring

in southern African herbaria. Various recent collections,

Oliver 4127, Schumann 793 and Vlok 2499, made in the

Attaquas Kloof area have turned out to be this species

when compared with the type. Also an investigation of

the type sheet of Erica rhodantha Guthrie & Bolus in

Section Polycodon showed that two collections were

mounted on the sheet, one being the type, Galpin 3706,

from Garcia’s Pass and the other being a collection of E.

gillii , Bolus 11339, also from Garcia’s Pass. A duplicate

of the Bolus collection turned out to be included with E.

rhodantha in STE. The two species are superficially very

similar, but may be distinguished on their anthers, black

with a large apical ridge in E. gillii. typical of the Section

Melastemon, versus brown and not ridged in E. rhodantha.

This latter species was also poorly represented in herbaria

with, surprisingly, only a few old collections from the

Garcia's Pass, an area which has been very well collected.

It has also recently been turned up on the lowest northern

slopes of the Langeberg west of Garcia’s Pass (Oliver

10242).

Erica gillii Benth. in De Candolle, Prodromus 7:

684 (1839); Guthrie & Bolus: 302 (1905); Dulfer: 136

(1965). Type: Attaquaskloof, Gill s.n. (K, holo.!).

Erica leptostachya

The type and only authentic material attributable to this

name came from the cultivated collections of William

MacNab. Many of MacNab's collections of heaths culti-

vated in Edinburgh in the early 1800's were known to

have originated from material collected at the Cape, par-

ticularly through the efforts of James Niven. So when de-

scribing this species Guthrie & Bolus noted ‘though its

origin is somewhat uncertain, it is most probably South

African’. A close examination of the fragment of the type

in BOL showed that the material identified in herbaria as

E. leptostachya did not match the type which was recog-

nized as being identical to specimens in STE of the Eu-

ropean species, Erica scoparia L. This species is charac-

terized by small, creamish green, wind-pollinated flowers

with large red stigmas. The flowers are borne on absolute

brachyblasts in the axils of foliage leaves up the main

branches, thus giving the appearance of a pseudoraceme.

Dulfer (1965) identified two sheets of unlocalized ma-

terial in W as clearly belonging to this species, but also

did not realise the European connection.

Therefore E. leptostachya Guthrie & Bolus becomes a

taxonomic synonym of E. scoparia L.

Erica scoparia L., Species plantarum edn 1, 1: 353

(1753); Benth.: 692 (1839); Dulfer: 149 (1965); D.A.

Webb & Rix: 8 ( 1972); D.C. McClint.: 286 (1989) et auct.

mult. Type: species still to be lectotypified.

E. leptostachya Guthrie & Bolus: 217 (1905); Dulfer: 105 (1965).

Type: South Africa?, without note of origin. Herb. MacNab 405a. 109

(K, holo.!).

28

Bothalia 24,1 (1994)

Dulfer (1963) described a variety of Guthrie & Bolus’

species as var. glabra Dulfer, based on a recent collection

from Mariepskop in the Transvaal. He noted that his type

material had originally been identified as E. leucopelta

Tausch var. ephebioides Bolus. Bolus’ type also came

from Mariepskop, as do the type and paratype of another

species described by Dulfer, E. merxmuelleri Dulfer. The

specimens of the latter species appear to us to be E.

natalitia Bolus.

Dulfer’s taxon can clearly not be part of the European

E. scoparia. Until such time as we can sort out the tax-

onomic problems of the E. leucopelta/natalitia/ merx-

muelleri complex this variety must be left unplaced, if

indeed it is a sound taxon.

Erica longisepala ^

When describing E. longisepala , Guthrie & Bolus

( 1905) placed it in the long-tubed Section Pleurocallis and

allied it to E. grandiflora. They commented that the spe-

cies was a link between the Sections Pleurocallis and

Hermes. Both of these sections are characterised by a

pseudospicate inflorescence and are separated only on the

size of the flowers. Guthrie & Bolus noted that their spe-

cies was related to E. parilis Salisb. in Section Hermes.

Since their publication much material of E. parilis and

E. longisepala has been collected which shows a consid-

erable amount of variability in the size of the flowers from

as short as 5 mm up to 25 mm. There is no clear demar-

cation of a boundary between the two species. Es-

terhuysen (1963) noted that these two species were

synonymous, but Dulfer (1965) did not follow this. We

confirm Esterhuysen’s opinion with the following synon-

ymy.

Erica parilis Salisb. in Transactions of the Linnean

Society 6: 371 (1802); Benth.: 664 (1839); Guthrie &

Bolus: 209 (1905); Dulfer: 102 (1965); Esterhuysen: 57

(1963). Type: Hottentots Holland, Masson s.n. (BM,

holo.!).

E. longisepala Guthrie & Bolus: 57 ( 1905); Dulfer: 36 ( 1965). Type:

Clanwilliam Div.; without precise locality, Maders.n. (SAM, holo.!).

Erica minutissima

An examination of a portion of the type which Bolus

fortunately acquired from Berlin has revealed that the

specimen is a monstrosity having a double calyx, no co-

rolla and no stamens. Bolus noted on his sheet that there

were two sheets of this 'species’ in Berlin, labelled as

locality 83, ‘Stellenbosch Umgebung Somerset West’. In

BOL there is a collection of Zeyher (?Ecklon) which

matches the Berlin fragment exactly, but is labelled as

moist places at Bloemendal. The material in our opinion

is referrable to E. quadrangularis which occurs sporadi-

cally in the area and which must have been very common

before the spread of housing and farms.

Erica quadrangularis Salisb., Prodromus 297

(1796); Guthrie & Bolus: 182 (1905); Dulfer: 93 (1965).

E. quadraejlora Salisb.: 375 (1802); Benth.: 679 (1839),

nom. illegit. Type: in Promontorio Cap, Jac. Mulder s.n.

(K, holo.!).

E. minutissima Klotzsch ex Benth.: 691 (1839);

Guthrie & Bolus: 223 (1905); Dulfer: 107 (1965). Type:

in montibus Hottentots Holland, Ecklon & Zeyher s.n. (B,

holo.f; BOL, fragm.!).

Erica monadelphia

The name of this species is problematic in having no

relevance to the species. The anthers of the species are

not joined together into one unit as they so clearly are in

the anomalous E. embothriifolia Salisb.

There has been much confusion with the publication

dates of Andrews’ species, particularly those in the first

volume of his folio edition. Coloured engravings of

heaths, which appeared in the four bound volumes pub-

lished from 1802 onwards. This led Dulfer (1965) to

change the authorship, as given by Bentham (1839) and

Guthrie & Bolus (1905), from Andrews (1802) to

Willdenow (1799).

Andrews produced his first drawings in separate parts

of three loose plates each, before he had them bound as

the first volume. Some, but unfortunately not all, of the

plates in the first volume of his work are dated. Recently

an excellent set of the first 22 parts of his Coloured en-

gravings was found in the library of the Farm Vergelegen,

Somerset West. From this set the publication date of E.

monadelphia was June 1, 1797 in part 8. A note on the

Andrews’ publications on heaths is being prepared for

publication.

Erica monadelphia Andrews, Coloured engravings

of heaths, part 8 (1797); Willd.: 396 (1799); Benth.: 622

(1839); Guthrie & Bolus: 51 (1905); Dulfer: 32 (1965).

Type: Andrews: t. 38 (1797).

Erica newdigatei

Due to an earlier homonym, the name of this species

had to be changed. Dulfer chose to commemorate one of

the collectors mentioned by Guthrie & Bolus ( 1905), Miss

Caroline Newdigate (1857-1937) of Forest Hall,

Plettenberg Bay. Thus Dulfer’s use of the masculine end-

ing ‘newdigatei’ must be corrected.

Erica newdigateae Dulfer in Annalen des Natur-

historischen Museums Wien 68: 138 (1965). E. longipes

Klotzsch ex Benth.: 684 (1839), non Bartl. (1832) [= E.

coarctata Wendl. var. longipes (Bartl.) Bolus]; Guthrie &

Bolus: 306 (1905). Type: in hills at Grasrugg [Grasrug?],

Uitenhage, Ecklon & Zeyher s.n. (B, holo.f; ?iso.).

Erica priorii

Until recently this species was known only from the

type collection in Kew, collected by Alexander-Prior near

George in 1847. Some recent collections, Vlok 1120 &

Schumann 590 from Karatara Forest Reserve, were com-

pared with the type and found to be this species. Subse-

quently a collection made by Keet in 1920 from Spitzkop,

Knysna and placed under incertae and one by Taylor from

the Outeniqua Pass in 1962 placed under E. coarctata

Wendl., have been found to be this species.

Bothalia 24, 1 (1994)

29

The fresh material from Vlok has shown us that the

species is a distinct one favouring moist, humic, condi-

tions on steep, southern slopes of the Outeniqua Moun-

tains. The shrubs grow to 1.2 m tall and bear bright pink

flowers in dense pseudospikes towards the ends of the

branches.

Erica priorii Guthrie & Bolus in Flora capensis 4:

216 (1905); Dulfer: 105 (1965). Type: George Div.; near

George, Alexander s.n. (K, holo.!).

E. recurvata

This name has been maintained in all major revisions

of the family with the citation of no collections from the

wild, only material cultivated in Europe during the early

1800’s. From the scraps of cultivated material available

for us to examine we find that the material is clearly a

lush cultivated form of E. cumuliflora Salisb. with its dis-

tinctive corolla lobes. These lobes are relatively long, sub-

spathulate and suberect forming windows around the base

of the corolla interstices and edged with short hairs in the

window areas. Only two species of Erica possess these

peculiar lobes, E. cumuliflora and E. genistifolia Salisb.

The former has 4-nate leaves and denser heads of 5-12

flowers whereas the latter has 3-nate leaves and fewer

flowers (3-4) per inflorescence.

Erica cumuliflora Salisb. in Transactions of the

Linnean Society 6: 336 (1802); Benth.: 657 (1839);

Guthrie & Bolus: 237 (1905); Dulfer: 112 (1965). Type:

Hottentots Holland. Mulder s.n. (K, holo.!).

E. recurvata Andrews: t. 262 ( 1 809): Lodd. : t. 1093 ( 1 825 ): Bot. Mag.

t. 3427 (1835): Benth.: 657 (1 839); Guthrie & Bolus: 236 (1905); Dulfer:

1 1 2 ( 1 965). Type: Andrews: t. 262 ( 1 809 ).

Erica revoluta/austrovema

When going through the collections of incertae in

BOL, PRE and STE, several specimens were found that

matched two recently described species, the Transvaal ma-

terial being E. revoluta in Section Arsace and the Natal

material being E. austrovema in Section Pyronium. An

additional specimen was found filed under E. woodii

Bolus in Section Chlorocodon. This led to an investigation

of the two species.

The material cited by Davidson (1985) under E. revo-

luta (Bolus) L.E. Davidson and Hilliard & Burtt (1985)

under E. austrovema Hilliard and the additional material

clearly belong to one, variable, widespread species. There

is much variation in the degree of revoluteness of the

leaves and of hairiness on the branches and pedicels and

in the shape of the sepals.

The collections from Natal and Swaziland tend to have

the leaves more revolute and therefore narrower, whereas

most of the collections from the eastern Transvaal have

broad open-backed leaves. However, the type of E.

revoluta. Wilms 908. and Keet STE 16476 have the narrow

leaves of the Natal material and both come from near

Lydenburg.

Erica revoluta (Bolus) L.E. Davidson in Journal of

South African Botany 51: 71 (1985). E. subverticillaris

Diels ex Guthrie & Bolus var. revoluta Bolus: 224 (1905);

Dulfer: 108 (1965). Type: Spitz Kop, near Lydenburg,

Wilms 908 (K, holo.!: BOL!, WU).

E. austrovema Hilliard in Hilliard & B.L. Burtt: 243 (1985). Type:

Natal, Vryheid Dist., Hlobane, 10-09-1950 , Johnstone 433 (NU, holo.!;

E. STE fragm.!).

Erica solandra/setulosa

In 1963 Dulfer described the variety mollis under E.

solandra Andrews based on a collection from Seven

Weeks Poort. We subsequently found material growing on

the Rooiberg in the Little Karoo and realized that the

growth form and habitat were quite unlike those of E.

solandra which is known only from the Outeniqua Moun-

tains near George. Several other collections from the

Seven Weeks Poort area matched Dulfer’s material.

During an investigation of all the species in the Section

Pseuderemia, it was found that a distinct new species

could be described, E. ingecina E.G.H. Oliv. (Oliver &

Oliver 1991) and that Dulfer’s variety was not part of E.

solandra and also constituted a distinct separate species.

Later during our survey of all Erica species the material

attributed to var. mollis was found to match a collection

by Elsie Esterhuysen from the extreme western end of the

Langeberg and which she had tentatively placed under E.

setulosa Benth.

E. setulosa is known only from Bentham’s type in Kew,

a specimen collected without precise locality by Niven in

the 1790’s and fragments of which were given to H. Bolus

and are now housed in BOL. A detailed examination of

these fragments showed that they match the Esterhuysen

collection in most respects. They also indicate that

Dulfer’s variety is indeed E. setulosa.

The placing of E. setulosa under the Section Ephebus

must have been based solely on the possession of the hairy

corolla which fact overlooked the head of flowers similar

to those found in Pseuderemia which also has some spe-

cies in which the corolla is hairy. This clearly indicates a

rather tenuous distinction between the two sections.

Erica setulosa Benth. in De Candolle, Prodromus

7: 682 (1839); Guthrie & Bolus: 123 (1905); Dulfer: 69

(1965). Type: Cape Colony, Niven s.n. (K, holo.; BOL,

fragm.!).

E. solandra Andrews var. mollis Dulfer: 33 (1963); Dulfer: 69 (1965).

Type: Prince Albert Div., Seven Weeks Poort Berg, rocky S. slopes, 5000

ft, 13-10-1955, Esterhuysen 24820 (W, holo.; BOL!, STE!).

Erica turmalis

Salisbury based his E. turmalis on one of the many

collections supposedly made by the enigmatic I. Mulder.

Salisbury is the only botanist to have cited this collector,

as Jac. Mulder (Salisbury 1796) and I. Mulder (Salisbury

1802), but nowhere is there a collection of his labelled as

such. There is no Salisbury material bearing the name

‘ turmalis ’ in either K or BM. Salisbury described his spe-

cies as 4-5-anthered and noted that it had the facies of

his E. bruniifolia. A collection by Mund in K with a frag-

ment in BOL, determined as this species by Bentham. is

clearly part of the E. cordata Andrews complex. Until

such time as an authentic specimen can be located we are

30

Bothalia24.1 (1994)

removing this name from the list of currently accepted

southern African species and placing it in the list of "spe-

cies non satis cognitae .

Erica turmalis Salisb. in Transactions of the Linnean Society 6: 342

(1802); Benth.: 616 (1839); Guthrie & Bolus: 234 (1905); Dulfer: 111

(1965). Type: in Hottentots-Holland, Mulder s.n. (?, holo.).

Erica umbelliflora

This species has remained uncollected since it was de-

scribed in 1839. An examination of the material cited by

Bentham revealed that E. umbelliflora is, in fact, a distinct

and well-known species in the southern Cape and that all

the collections assignable to this species were filed under

two recently described species.

The current sectional subdivision of the genus Erica is

a complex and unfortunately unnatural system which has

led to a number of irrelevant new species being described

(see several cases "cited above). The circumscription of

some of the sections has been rather vague and with the

arbitrary placing of species in these sections true relation-

ships have inevitably been overlooked in the describing

of new taxa. This has been the case with two species, E.

manifesta Compton and E. ionii H.A. Baker.

*

E. umbelliflora was placed by Guthrie & Bolus in the

Section Pachysa because they assumed that the flowers

were viscid. Compton looked in the Section Gypsocallis

for species allied to his E. manifesta on the grounds of

slightly exserted stamens and a pseudospicate inflores-

cence. Baker placed his E. ionii in Section Pyronium on

account of the exserted stamens and terminal flowers. In

all three cases the true relationship of the taxa was not

possible to ascertain because of their diverse placement

in the genus.

The variability of the flowers is such that they may be

viscid because of the glands on the calyx, may have in-

cluded to exserted stamens and have the flowers arranged

from terminal on short lateral branchlets to aggregated in

a pseudospicate arrangement.

Later Baker (1970) recognized that his species was

synonymous with Compton’s species, but overlooked the

relationship with E. umbelliflora. The synonymy of the

species is thus:

Erica umbelliflora Klotzsch ex Benth. in De Can-

dolle, Prodromus 7: 659 (1839); Guthrie & Bolus: 197

( 1905); Dulfer: 98 ( 1965). Types: in monte Zwartberg ad

Attaquaskloof et flumen Calutso, Masson s.n. et Drege

s.n. (lectotype still to be chosen).

E. manifesta Compton: 37 (1935); Dulfer: 84 (1965). Type: Uni-

ondale Dist.; hills northeast of Avontuur, 3100 ft. Sept. 1932, Fourcade

4606 (BOL, holo.!).

E. ionii H.A. Baker: 148 ( 1965); Dulfer: 86 (1965). Type: Uniondale

Div. On sandy slopes on Potjieskloof Pass beside the National Road from

George to Uniondale at about 900 m ( 3000 ft). Baker 2330 (BOL, holo. ! ).

REFERENCES

ANDREWS, H.C. 1794— 71830. Coloured engravings of heaths. London.

BAKER. H.A. 1965. Notes on some species of Erica, with descriptions of

three new species and two new species and two new varieties.

Journal of South African Botany 31: 145-161 .

BAKER, H.A. 1970. Taxonomic notes on Erica. Journal of South African

Botany 36: 267-270.

BARTLING. T.F. 1832. Plantae Ecklonianae, Ericaceae. Linnaea 7: 627-

652.

BENTHAM, G. 1839. Ericaceae. In A.P De Candolle, Prodromus sys-

tematis naturalis regni vegetabilis 7: 580-733. Paris.

BOLUS, H. 1905. Erica. Flora capensis 4,1: 4—315. Reeve, London.

BROWN, N.E. 1906. Ericaceae. Flora capensis 4,1: 315-336. Reeve,

London.

COMPTON, R.H. 1934. Erica acockii. Journal of Botany 72: 43.

COMPTON, R.H. 1935. Plantae novae africanae. Series IV. Journal of

South African Botany 1: 37-39.

COMPTON, R.H. 1941 . Plantae novae africanae. Series XVII. Journal of

South African Botany 7: 193-207.

DAVIDSON, L.E. 1985. A change in status for Erica subverticillaris var.

revoluta from the eastern Transvaal. South African Journal of

Botany 51:71-73.

DULFER. H. 1963. Neue Arten und Varietaten der Gattung Erica L. aus

Siidafrika. Annalen des Naturhistorischen Museums Wien 66:

19-33.

DULFER. H. 1965. Revision der siidafrikanischen Arten der Gattung

Erica L. 2. Annalen des Naturhistorischen Museums Wien 68:

25-177.

ESTERHUYSEN, E. 1963. Notes on South African species of Erica.

Journal of South African Botany 29: 51-58.

GUTHRIE, F. & BOLUS, H. 1905. Erica. Flora capensis 4,1: 4-315.

Reeve, London.

HALL, A.V. & VELDHUIS, H.A. 1988. South African Red Data Book:

plants — Fynbos and Karoo Biomes. South African National Sci-

entific Programmes Report No. 117. CSIR. Pretoria.

HILLIARD, O.M. & BURTT, B.L. 1985. Notes on some plants of south-

ern Africa chiefly from Natal: XL Notes from the Royal Botanic

Gardens Edinburgh 42,2: 227-260.

LINNAEUS, C. 1753. Species plantarum edn 1, 1:352-356.

LINNAEUS, C. fil. 1782. Supplementum plantarum. Braunschweig.

LODD1GES, C. 1825. The botanical cabinet: t. 1093.

MCCLINTOCK, D. 1989. The heathers of Europe and adjacent areas.

Botanical Journal of the Linnean Society 101: 279-289.

OLIVER, E.G.H. & OLIVER. I.M. 1991. Studies in the Ericoideae

(Ericaceae). VIII. New species in Erica, section Pseuderemia,

from southern Africa. Botlialia 21: 137-142.

OLIVER. E.G.H. 1993. Studies in the Ericoideae (Ericaceae). XI. The

generic relationship between Erica and Blaeria. Kew Bulletin 48:

771-780.

SALISBURY, R.A. 1796. Erica. Prodromus : 292-298.

SALISBURY, R.A. 1802. Species of Erica. Transactions of the Linnean

Society’ 6: 316-388.

SALTER, T.M. 1950. Ericaceae. In R.S. Adamson & T.M. Salter, Flora of

the Cape Peninsula: 627-657. Juta, Cape Town.

TAUSCH, J.F. 1834. Bemerkungen tiber Erica. Flora 17: 615-622.

WEBB, D.A. & RIX, E.M. 1972. Erica. In V.H. Heywood, Flora eu-

ropaea 3: 5-8.

WILLDENOW. C.L. 1799. Caroli a Linne species plantarum 2: 396.

Berlin.

Bothalia 24,1: 31-35(1994)

Notes on African plants

VARIOUS AUTHORS

ROSACEAE

CLIFFORTIA LONG1FOL1A. A ‘GOOD' SPECIES OR SHOULD IT BE A VARIETY UNDER C. STROBIL1FERA ?

Ecklon & Zeyher (1836) described a variety of

Cliffortici strobilifera L. and named it var. longifolia , but

pointed out that it might be a separate species. Their type,

Ecklon & Zeyher 1 753B, was observed by the authors as

‘wanting flowers' whereas Weimarck (1933) noted that it

had ‘an abundance of male flowers'.

As the female flowers were not seen by Weimarck

(1933), his decision to raise the taxon to specific level

was based on the differences in the male bracteoles and

the brachyblasts. Observed inconsistencies in Weimarck's

(1933, 1934) descriptions and figures depicting the respec-

tive sizes of the leaf stipules and male bracteoles in C.

longifolia and the indumentum of the male bracteoles in

C. strobilifera , added to the doubt concerning the validity

of such a decision. In fact, the investigation of the male

bracteoles and pedicels in C. strobilifera , revealed a very

similar structure to those in C. longifolia but on a some-

what smaller scale. (Figure 1M & N compared to 1A &

B).

Further confusion was caused by the observed variation

in leaf size and sheath length in C. strobilifera, the larger

forms being difficult to distinguish from C. longifolia on

vegetative characters only.

The investigation of female flowers seemed to be the

only way to settle this question. If the female flowers

should show the same degree of similarity as the male

flowers, differing in size only, the case for raising the sta-

tus of the taxon would be very flimsy. If however, the

female flower and fruit should be markedly different, there

could no longer be room for doubt.

In the search for female flowers, short shoots with ob-

vious male flowers but apparently totally lacking in female

flowers, were removed from herbarium specimens and

softened by boiling and soaking. Successive leaf sheaths

were removed from each short shoot and the axils exam-

ined for flowers.

Each axil contained a single flower and even though

the intact short shoots showed no evidence of the presence

of female flowers, these were in fact found. A single fe-

male flower occurred in the axil of the leaf halfway up

the short shoot, preceded by 3(4) old male flower remains

(pedicels and bracteoles) and followed by a similar group

of young male flowers (buds). Fully matured fruits were

also totally obscured by the leaf sheaths.

With the discovery of the female flowers and fruits of

C. longifolia, the necessity for amendments to the descrip-

tions of both taxa, became obvious.

The specimens of C. strobilifera selected for examina-

tion included the extremes from the small-leafed form nor-

mally regarded as the typical C. strobilifera, to the large-

leafed form that is almost indistinguishable from C. longi-

folia on the vegetative characters only.

Cliffortia longifolia ( Eckl. & Zeyh.) Weim., Mono-

graph of the genus Cliffortia : 144 (1934).

C. strobilifera L. var. longifolia Eckl. & Zeyh. : 271 (1836). Type:

Swellendam: ad llumen Breederivier, Murid s.n. sub Ecklon s.n. ( B t, S!.

lectotype here designated).

Weimark (1934) quoted a Mund collection under

Ecklon & Zeyher 1753B as the type in B and S. The Berlin

specimen was destroyed in the Second World War and the

Stockholm specimen bears no collector’s number. A hand-

written label (not in Weimarck’s handwriting) bearing the

name ‘ Cliffort strobilifera B longifol' and a reprint of

Ecklon & Zeyher’s description of the variety giving the

locality as the Breede River and the collector as Mund.

are attached to the Stockholm specimen. Weimarck seems

to have found these two labels sufficient evidence that the

Stockholm specimen is a duplicate of the Berlin one in

spite of the absence from the Stockholm specimen of the

Ecklon & Zeyher collector’s number.

Leaves : stipules up to 20 mm long, densely shortly

hairy; leaflets 15.0-65.0 x 2. 0-6.0 mm. Male flowers:

pedicel 6.75 mm long with 2.4 mm above upper bracteole

and 2.9 mm below lower bracteole, densely hairy; upper

bracteole 63-6.5 mm long, lower bracteole 9.25-9.50 mm

long, situated 1.45 mm apart on pedicel; receptacle 1.0-

1.25 x 0.65 mm, glabrous; sepals 3, 10.0 x 2.0 mm, (Fig-

ure 1A & B). Female flowers: pedicel 1.6 mm long with

half of its length above upper bracteole, densely hairy ex-

cept for glabrous apical cone (Figure IF & K); bracteoles

7. 0-8.4 mm long, originating one above the other on ped-

icel, basally ovate abruptly narrowing to linear, keel of

basal ovate part and entire linear portion densely hairy,

margin of basal ovate portion glabrous but for a sparsely

ciliate edge (Figure ID & F); sepals 3, 5.0-6. Ox 1 .0—1.2

mm, linear-lanceolate, glabrous (Figure 1C); ovary 2.6 x

1.5 mm. ovate, truncate at both ends, glabrous, irregularly

longitudinally ribbed with 19 (20) low ridges apically

joined to form roughly flattened top bearing staminodes,

ridges basally lost in a somewhat swollen and hollow part

that fits over cone of pedicel, adaxially irregularly grooved

32

Bothalia 24. 1 (1994)

FIGURE 1 . — Cliffortia longijolia. A, ubaxiul \ lew ol male flower (stamens removed): Aa, reeeplacle; Ab & Ad, densely hairy pedicel. B. adaxial

view of male flower, receptacle and calyx removed: Bb. Be, Bd, parts of pedicel. C, female calyx lobes; D, lower female bracteole; E, style

and stigma; F, upper female bracteole attached to pedicel; G, ovary; H, female flower. (D-G in situ). 1, female bracteoles, fruit removed: Ie,

part of pedicel. J, fruit viewed from top. K, top half of densely hairy pedicel: Kf, conical apex. L, fruit: Lg, swollen part, with basal hollow

that fits over cone on pedicel. Cliffortia strobilifera. M, adaxial view of male flower (stamens removed): Ma, glabrous receptacle; Mb, Md.

hairy pedicel. N. abaxial view of male flower, receptacle and calyx removed: Nb. Nc & Nd, parts of pedicel. O, female calyx lobes; P, style

and stigma; Q, fruit, irregularly rugose, top truncate; R, intact female flower and bracteoles. A, B, Burgers 1668 (STE); C-H, Hugo 1800

(STE); I-L, Oliver 8494 (STE); M, Rudatis 999 (STE); N, Thompson 3691 (STE); O-R, Oliver 10053 (STE). Scale bar: 1 mm.

by edges of the bracteoles (Figure 1 G); style solitary, 3. 1

mm long, stigma 3.3 mm long sparsely fimbriate on one

side only (Figure IE). Fruit: 4.0 x 1.5 mm, elliptic, trun-

cate at both ends, longitudinally ridged, without irregular

adaxial groove as seen on ovary, the fruit having devel-

oped beyond bracteoles (Figure 1H, J & L).

Cliffortia strobilifera L., Systema vegetabilium

edn 13 : 749 (1774). Weim. : 140 (1934); auct. mult. Type:

Pluk.: t. 275, fig. 2 (1694).

Although a Plukenet specimen associated with the cited

figure exists under Flerb. Sloane 99: 179, upper right (BM-

SL), this specimen cannot be designated lectotype as it

was never seen by Linnaeus, according to Dr F.R. Barrie

of the Linnaean Plant Name Typification Project (pers.

comm.). The only possible type referred to by Linnaeus

(1774) in his description of the species, is the Plukenet

figure.

Change of author citation

The author of C. strobilifera has always been cited as

J. A. Murray in Systema vegetabilium edn 13: 749 (1774).

Stafleu & Cowan (1981) however, in referring to this doc-

ument, added the following: ‘N.B. The new taxa in this

work must be attributed to Linnaeus; Murray was simply

editor.’

Leaves : vagina 3-9 mm long; stipules 2^1 mm long,

ciliate; leaflets glabrous to hairy all over or on veins only,

10-70 x 1.5-8 mm, linear-lanceolate, base somewhat nar-

rowed to channelled and petiolate, apex acute to tridentate

or with main vein bisecting lamina obliquely. Male flow-

ers: pedicel 1.0-1. 3 mm long, hairy; bracteoles 4.0-4.75

mm long, linear subulate, the lower implanted 0.4 mm

below the upper, keel ciliate; sepals ± 4.0 x 1.0 mm,

linear-lanceolate apiculate, glabrous (Figure 1M & N). Fe-

male flowers: sessile; bracteoles 2.5-3 .5 mm long, lan-

ceolate-linear acuminate, apex sometimes spirally twisted;

sepals 3, 3.0-3. 3 x 0.6-0.8 mm, linear lanceolate, glabrous

Bothalia 24,1 (1994)

33

(Figure 10); ovary linear, apex truncate, base cuneate, gla-

brous, irregularly rugose; style solitary, 1 .5 mm long,

stigma 1.5 mm long, fimbriate mainly on one side (Figure

IP) . Fruit: 23-2.4 mm x 0.75-1.0 mm. cuneate linear

truncate, mgose to very obscurely longitudinally ridged,

apex concave giving the appearance of an opening (Figure

IQ) .

Distribution and habitat: C. longifolia occurs in wet-

lands, half-wetlands, swamps, vleis and on streamsides

near the coast, at low altitudes, from Langebaan Lagoon

in the west to the Cape Peninsula, then along the southern

coast to Ystervarkpunt (Gouriqua) near Albertinia ( Figure

2).

C. strobilifera shares the habitat with C. longifolia. but

has a much wider distribution and altitudinal range; the

localities near the Cape southwestern coast are at lower

altitudes than those inland and northerly, which occur as

high as 1 600 m.

DISCUSSION

Some herbarium specimens of C. strobilifera have both

small and large leaves. These are often borne on separate

branches, or the large ones are on long shoots, the small

ones on short shoots, apparently indicating different

growth phases. Field observations confirmed this: the new

season’s growth is represented by a much more robust and

larger long shoot with a thicker stem and larger leaves

than the existing stems and leaves. These leaves also have

the typically soft texture and hairiness of new growth.

Where the larger leaves occur on herbarium specimens of

C. strobilifera , these are always together with smaller

ones, either on the same branch and then with the large

leaves on the long shoot sheathing the short shoots, or on

different branches where the robust branch represents the

new growth referred to above. In contrast, the leaves of

C. longifolia appear homogeneous or at least far less vari-

able in size. The leaves are also of a more robust texture

and the margins thicker.

Whereas most other differences between C. longifolia

and C. strobilifera could very well be dismissed as merely

of degree, the differences in fruit morphology establish C.

longifolia as a separate entity beyond all doubt. C.

longifolia has the clearly marked ribs typical of section

FIGURE 2. — Known distribution of Cliffurtici longifolia.

Costatae, whereas C. strobilifera not only has the

smoother fruit less often encountered in this section, but

the shape of the fruit is also unusual and the concave apex

even more so.

Weimarck (1934) gave the distribution of C. longifolia

as the type locality only. Six years later, Weimarck (1940)

quoted one other collection and extended the range to in-

clude the Cape Peninsula. Later collections filled in the

gap between these two localities and extended the distri-

bution to the west coast.

SPECIMENS EXAMINED

C. longifolia

CAPE. — 3318 (Cape Town): Farm Geelbek, eastern shores,

Langebaan Lagoon, marsh occasionally saline, ± 5 m, 14-08-1975,

(-AA), Boucher 2813 (STE). 3318/3418 (Cape Town/Simonstown):

Cape Peninsula without precise locality, banks of streams, ± 160-320 m,

1887, Thode 9183 (STE). 3419 (Caledon): Ratel River Mouth, 50 m,

07-06-1984, (-DC), O'Callaghan et al. 483 (PRE. STE). 3420

(Bredasdorp): De Hoop-Potberg Nature Reserve, Windhoek, Rietkloof,

damp soil along stream, 10 m, 16-01-1979, (-AD). Burgers 1668 (PRE);

between Noetzie and Elandspad, streamside vegetation, 60 m, 11-04-

1979. (-BC), Hugo 1800 (STE); Potberg area, Hamerkop Resort,

streamside in kloof, 60 m, 19-06-1984, (-BC). Oliver 8494 (PRE. STE).

3421 (Riversdale): Rransfontein Farm, Still Bay, river bank, 30 m,

07-10-1980, (-AD), Bohnen 7730 (PRE, STE); Albertinia, near

Ystervarkpunt, on Gouriqua land, in a marsh, on gentle south-facing

slope, in coastal fynbos, 10 m, 18-02-1989, (-BC), Vlok 2098 (PRE);

Albertinia, Ystervarkpunt (Gouriqua), along stream, 25 m, 18-03-1987,

(-BD), Willemse 150 (PRE. STE).

C. strobilifera

Robust form

NATAL. — 2930 (Pietermaritzburg): Inanda. 06-1880. (-DB), Wood

795 (STE). 2931 (Stanger): Chakaskraal, by streams, 100 m, 02-1916,

(-AC), Thode 4407 (STE). 3030 (Port Shepstone): Fairfield, Dumisa, 750

m, 14-01-191 1, (-AD). Rudatis 1303 (STE); Oribi side river, in riverbed

between boulders, 23-01-1973, (-CB), Slrey 11061 (PRE); Izotsha, riv-

erbank, mud, 16-11-1969. (-CD), Slrey 9283 (PRE).

CAPE. — 3227 (Stutterheim): Komgha River, 330 m. 11-1892,

(-DB), Flanagan 1452 (STE). 3318 (Cape Town): Cape Peninsula, Table

Mountain, Orange Kloof, 08-1903, (-CD), Marloth 3431 (STE);

Jonkershoek, Bosboukloof, 320 m, 05-1967, (-DD), Kerfoot 5805 (STE).

3319 (Worcester): near Karoo Botanic Garden, beside dam, 330 m,

11-10-1969, (-CB), Rycroft 3055 (NBG. STE). 3323 (Willowmore):

Avontuur Poort, 4 miles from Uniondale, 810 m, 12-1930, (-CA), Four-

cade 4520 (STE). 3326 (Grahamstown): Grahamstown, 01-04-1922,

(-BC), Wilson SA142 (STE). 3418 (Simonstown): lower Eerste River

below Krammat near mouth on Zandvliet allotment, riverbank, 7 m.

06-01-1978, (-BB), Boucher 3475 (STE); Sir Lowry’s Pass, Steenbras

side, 04-1948, (-BB), Stokoe SAM 61497 (PRE, SAM); Palmiet River

Mouth near Kleinmond, 5 m, 17-03-1983, (-BD), Van Wyk 1183 (STE).

3419 (Caledon): Palmiet River, 25-04-1948, (-AA), De Vos 895 (STE).

3423 (Knysna): Concordia, along streams and on moist ground. 230 m,

03-1921, (-AA).A'eet 705(STE); Keurboo ms River, 160 m, 1969, (-AB),

Heinecken K58 (STE). 3424 (Humansdorp): Kromme River at Assegai

Bosch, 223 m, 03-1931, (-BA), Fourcade 4565 (STE).

Smaller form

CAPE. — 3318 (Cape Town): Blaauwklip Stellenbosch. 01-1926,

(-DD), Gilletts. n. (STE). 3319 (Worcester): Wemmershoek, ProteaState

Forest Reserve, ENE of Wemmershoek Station, 800 m. 29-02-1992,

(-CC), Oliver 10053 (STE). 3320 (Montagu): Langeberg East.

Lemoenshoek. 550 m, 25-02-1988, (-DD), Van der Merwe 226 (STE).

3321 (Ladismith): Elandskloof between Vleiland and Seweweekspoort,

1 300 m, 20-02-1986. (-AD). Moffett & Steensma 3824 (STE); Garcia’s

Pass near Tollhouse, 19-04-1983, (-CC). Fellingham 445 (STE). 3322

(Oudtshoorn): Boomplaas, Cango Valley on banks of Grobbelaars River,

24-06-1974, (-AC), Moffett 62 (STE). 3418 (Simonstown): Cape of

Good Hope Nature Reserve, 330 m, 09-03-1973, (-AB), Taylor 8352

34

Bothalia 24,1 ( 1994)

(STE); Gordon’s Bay, March. (-BB), Duthie 762 (STE); Rooiels, River

Mouth, near Hangklip Floodplain, 11-03-1981, (-BD), Parsons 59

(STE). 3419 (Caledon): Oudebos, Riviersonderend, 04-1930, (-AB),

Stokoe s.n. (STE); between Avoka and Goedvertrou, 83 m, 22-05-1971,

(-DA), Thompson 1207 (STE). 3420 (Bredasdorp): Breede River at

Malgas, 16 m, 07-03-1971. (-BC). Heard 3 (STE). 3421 (Riversdale):

Korente River Dam, 200 m, 26-09-1979, (-AA), Bohnen 6621 (STE).

REFERENCES

ECKLON, C. F. & ZEYHER, C. 1836. Enumeratio plantarum africae

australis extra tropicae. part 2.

LINNAEUS, C. 1774. Systema vegetabilium...editio decima tertia , edn

13. J.A. Murray (ed.). Gotha, Gottingen.

PLUKENET, L. 1694. Alrnagestum botanicum: t. 275, fig. 2. London.

STAFLEU, F. A. & COWAN, R. S. 1981. Taxonomic Literature 3: 670.

Bohn, Scheltema & Holkema, Utrecht; Junk, The Hague.

WEIMARCK, H. 1933. New species in the genus Cliffortia. Botaniska

Notiser 1933: 158-160.

WEIMARCK. H. 1934. Monograph of the genus Cliffortia. Gleerupska

Universitets Bokhandeln. Lund.

WEIMARCK, H. 1940. Some notes on the genus Cliffortia. Botaniska

Notiser 1 940: 400.

A C. FELLINGHAM*

*Stellenbosch Herbarium, National Botanical Institute, P.O. Box 471,

Stellenbosch 7599.

MS. received: 1993-04-01.

ASPHODELACEAE/ALOACEAE

WAS GASTER1A N1TIDA VAR. ARMSTRONG!! VALIDLY PUBLISHED?

In the recently published synopsis of the genus Gas-

teria Duval (Van Jaarsveld 1992: 12), the full page refer-

ence to the basionym of Gasteria nitida var. armstrongii

(Schonland) Van Jaarsv. was not given where the new

combination was made. The combination therefore ap-

pears not to have been validly published (Eggli & Taylor

1993). Under Art. 33.2 of the Botanical Code (Greuter et

al. 1988), on or after 1 January 1953, a new combination

is not validly published unless its basionym is clearly in-

dicated and a full and direct reference -.given to its author

and place of valid publication with page or plate reference

and date.

However, the name has been validly published because

the page reference required for validation is given in the

list of references of the same paper (Van Jaarsveld 1992).

The entry 'Schonl. in Rec. Albany Mus. 1912', on p. 12,

is a perfectly acceptable, unambiguous citation of the item

fully listed under 'References’ on p. 28. This point of view

is in accordance with the interpretation of Brummitt

(1969: 45), who determined that the combination Erysi-

mum arbuscula (Lowe) Snogerup was validly published

by citation of the basionym reference in detail in the bib-

liography, but indirectly only in the text (Snogerup 1967:

9). Furthermore, the policy of the herbarium of the Royal

Botanic Gardens, Kew, regarding validly published names

included in the Kew Record of Taxonomic Literature was

formulated as follows: ‘A reference to the basionym is

accepted if it is cited adjacent to, or clearly associated

with, a new combination, or in a bibliography at the end

of the paper, but not if it is elsewhere in the same paper

without any clear connection’ (Royal Botanic Gardens,

Kew 1974: 383).

The full citation to the name therefore is:

Gasteria nitida var. armstrongii (Schonland) Van

Jaarsv. Basionym: Gasteria armstrongii Schonland in Re-

cords of the Albany Museum 2,4: 258 (1912).

ACKNOWLEDGEMENTS

Dr Donald B.J. Killick, formerly of the Botanical Re-

search Institute, Pretoria, and Prof. W. Greuter and Dr B.

Zimmer of the Botanisches Garten und Botanisches Mu-

seum, Berlin-Dahlem, are thanked for their comments on

an earlier draft of this paper.

REFERENCES

BRUMMITT, R.K. 1969. On some requirements for valid publication of

names. Regnum Vegetabile 60: 42-57.

EGGLI, U. & TAYLOR, N.R (eds) 1993. Aloaceae. In Repertorium

Plantarum Succulentarum ( 1992) 43: 6.

GREUTER, W. et al. 1988. International Code of Botanical Nomencla-

ture. Regnum Vegetabile 1 18.

ROYAL BOTANIC GARDENS, KEW 1974. The Kew Record, a new

annual bibliography. Taxon 23: 381-386.

SCHoNLAND, S. 1912. A new species of Gasteria. Records of the

Albany Museum 2: 258.

SNOGERUP, S. 1967. Studies in the Aegean flora. VIII. Erysimum sect.

Cheiranthus. A. Taxonomy. Opera Botanica 13: 3-70.

VAN JAARSVELD, E.J. 1992. The genus Gasteria, a synoptic review.

Aloe 29: 5-32.

G.F. SMITH*, B-E. VAN WYK** and E.J. VAN JAARSVELD***

* National Botanical Institute, Private Bag X101, Pretoria 0001.

** Department of Botany, Rand Afrikaans University, P.O. Box 524,

Auckland Park 2006.

*** Kirstenbosch National Botanical Garden, National Botanical Insti-

tute, Private Bag XI, Claremont 7735.

MS. received: 1993-11-24.

ALOE BARBERAE TO REPLACE A. BAINESII

In an enumeration of the tree aloes of southern Africa,

Dyer (1874a) described as new two tree aloes from the

subtropical eastern coastal areas: A. bainesii from northern

Natal and A. barberae from ‘Caffraria’ (Transkei). These

two ‘species’ were described as differing mainly in their

leaf characters, A. barberae having longer, less glaucous,

and more widely spaced leaves.

A few months later Dyer (1874b), in a short note in

the same journal, decided to combine the latter two spe-

Bothalia 24,1 (1994)

35

cies and expressly chose the name A. barberae for the

southeastern African tree aloe. Unfortunately this publica-

tion was overlooked by all subsequent research workers

on Aloe, notably Baker (1880) and Reynolds (1950). How-

ever, Baker (1885) regarded A. bainesii and A. barberae

as conspecific, but incorrectly chose as the correct name

A. bainesii. An attempt by Baker (1896) to taxonomically

reinstate A. barberae as a variety of A. bainesii has re-

ceived little support.

Therefore, although the name A. bainesii has been

widely adopted by most southern African 11 ora writers and

succulentists, it has to be rejected in favour of the name

A. barberae , which was chosen as the one to be accepted

by the original author.

Aloe barberae Dyer in The Gardeners' Chronicle

(n.s.) 1: 568 ( 1874a);’ Dyer: 720 (1874b); Dyer: 90

(1875a); Dyer: 49 (1875b). Type: Hort ..Anon. s.n. (K!).

A. bainesii Dyer: 568 (1874a). Baker: t. 6848 (1885); Baker: 326

(1896); Berger: 319 ( 1908); Marloth: 92 (1915); Sim: 152 ( 1919); Reyn-

olds: 498 (1950); Jeppe: 59 (1969); Bomman & Hardy: 281 (1972);

Palmer & Pitman: 339 (1972); Compton: 98 (1976). Type: Natal. Grey-

town Dist., T. Baines s.n. (K!).

A. bainesii Dyer var. barberae (Dyer) Baker: 326 (1896).

ACKNOWLEDGEMENTS

Dr Colin C. Walker, Open University, Milton Keynes,

U.K. and Mrs Estelle Potgieter, Chief Librarian, Mary

Gunn Library, are thanked for locating and supplying cop-

ies of scarce publications and illustrations. Dr Donald J.

B. Killick is thanked for his comments on an earlier draft

of this paper.

REFERENCES

BAKER, J.G. 1880. A synopsis of Aloineae and Yuccoideae. Journal of

the Linnean Society, Botany 18: 148-241.

BAKER, J.G. 1885. Aloe bainesii. Curtis’ Botanical Magazine 111: t.

6848.

BAKER, J.G. 1896. Aloe. In W.T. Thiselton-Dyer, Flora capensis 6,2:

302-320. Reeve, London.

BERGER, A. 1908. Liliaceae-Asphodeloideae-Aloineae. Aloe. In A.

Engler, Das Pflanzenreich 4.38.3.2 (Heft 33): 150-326. Engel-

mann, Leipzig.

BORNMAN, H. & HARDY. D.S. 1972. Aloes of the South African veld.

Voortrekkerpers, Johannesburg.

COMPTON, R.H. 1976. Flora of Swaziland. Journal of South African

Botany supplementary vol. 11: 1-684.

DYER. W.T. THISELTON 1874a. The tree aloes of South Africa. The

Gardeners' Chronicle (n.s.) 1: 566-569. 571.

DYER. W.T. THISELTON 1874b. Tree aloes of South Africa. The

Gardeners ' Chronicle (n.s.) 2: 720.

DYER, W.T. THISELTON 1875a. The tree aloes of South Africa. Nature

11: 89-91.

DYER. W.T. THISELTON 1875b. Short notes and queries. Tree aloes.

Journal of Botany, London 13: 49, 50.

JEPPE. B. 1969. South African aloes. Purnell. Cape Town.

MARLOTH, H.W.R. 1915. The flora of South Africa. Vol. 4. Monocoty-

ledons. Darter, Cape Town.

PALMER, E. & PITMAN, N. 1972. Trees of southern Africa. Balkema,

Cape Town.

REYNOLDS, G.W. 1950. The aloes of South Africa. Aloes of South

Africa Book Fund, Johannesburg.

SIM. T.R. 1919. Flowering trees and shrubs. Specialty, Cape Town.

G.F. SMITH*. B-E. VAN WYK** and H.F. GLEN*

* National Botanical Institute. Private Bag X101, Pretoria 0001.

** Department of Botany, Rand Afrikaans University, P.O. Box 524,

Auckland Park 2006.

MS. received: 1993-11-24.

BORAGINACEAE

THE CORRECT AUTHOR CITATION FOR LOBOSTEMON MONTANUS

Herman (1993) cited Lobostemon montanus (DC.) H.

Buek; however, Buek described L. montanus in 1837,

whereas De Candolle published the combination Echium

montanum (H. Buek) DC. in 1846. The correct author

citation is therefore L. montanus H. Buek.

REFERENCES

BUEK, H.W. 1837. Echia capensia. Linnaea 11: 132.

DE CANDOLLE. A.P. 1 846. Prodromus systematis naturalis regni vege-

tabilis ... 10: 15. Treuttel & Wiirtz, London.

HERMAN, P. 1993. Boraginaceae. In T.H. Arnold & B.C. de Wet. Plants

of southern Africa: names and distribution. Memoirs of the Bota-

nical Sur\’ey of South Africa No. 62.

M.H. BUYS and J.J.A. VAN DER WALT*

* Department of Botany, University of Stellenbosch, Private Bag X5018,

Stellenbosch 7599.

MS. received: 1993-11-09.

'

Bothalia 24,1: 37-53 (1994)

Ferns and flowering plants of Klaserie Private Nature Reserve, eastern

Transvaal: an annotated checklist

N. ZAMBATIS*

Keywords: checklist, eastern Transvaal, Klaserie, Savanna Biome

ABSTRACT

An annotated checklist of the plant taxa of the Klaserie Private Nature Reserve, eastern Transvaal Lowveld, is presented. Of

the 6 1 8 infrageneric taxa recorded, six are pteridophytes and the remainder angiosperms. Of these, 161 are monocotyledons and

451 dicotyledons. Five of the latter are currently listed in the Red Data List of the Transvaal, two of which are first records for

the Transvaal Lowveld. The vegetation of the reserve shows strong affinities with the Savanna Biome, and to a lesser degree,

with the Grassland Biome.

UITTREKSEL

'n Geannoteerde kontrolelys van die planttaksons van die Klaserie Privaatnatuurreservaat, Oos-Transvaalse Lae veld, word

gegee. Van die 618 taksons wat aangeteken is, is ses pteridofiete en die res angiosperme, waarvan 161 monokotiele en 451

dikotiele is. Vyf van laasgenoemde taksons word tans in die Rooidatalys van Transvaal ingesluit. Twee hiervan verteenwoordig

eerste rekords vir die Transvaalse Laeveld. Die plantegroei van die reservaat toon 'n sterk affiniteit tot die Savanne Bioom en,

tot ’n mindere mate, met die Grasveld Bioom.

INTRODUCTION

No comprehensive plant list of Klaserie Private Nature

Reserve (KPNR) has previously been produced. Wit-

kowski (1983) recorded a total of 124 taxa, whereas this

list comprises a total of 618 taxa. Additions to this list

will no doubt be made in the future. Apart from free-rang-

ing exotic taxa, no other exotic (or indigenous) taxa

planted in gardens were included. No algae, mosses, or

fungi have been collected or recorded by the author nor,

as far as is known, by any other collectors.

STUDY AREA

The Klaserie Private Nature Reserve (KPNR) is situ-

ated between the town of Hoedspruit and the Kruger Na-

tional Park (KNP), with which it shares a common

boundary of some seven kilometres. To the east lie the_

Timbavati, Umbabat. and Ntsiri Private Nature Reserves

(Figure 1 ).

The reserve extends from 24°02' to 24°16'S, and

31°03' to 31°19'E and thus falls within the following quar-

ter degree square grids: 2431 AA, 2431AB, 243 1AC, and

2431 AD.

The area was proclaimed as a nature reserve in 1972

and is jointly owned by 107 members. Prior to its proc-

lamation, the area was partly utilized for cattle farming.

Diseases and predators, however, led to the abandonment

of this form of land use.

* National Parks Board, Kruger National Park, Private Bag X402,

Skukuza 1350.

MS. received: 1993-02-04.

With a surface area of 62 818 ha, the KPNR is the

largest privately owned nature reserve in the Transvaal

Lowveld. and probably the largest in the country. Together

with the other two large private nature reserves, Timbavati

to the east and Sabi-Sand in the south, as well as a number

of smaller private nature reserves in the area, it comprises

a portion of an important conservation, tourism, and hunt-

ing area of over 180 000 ha.

In terms of a recent Contractual Park Agreement be-

tween the National Parks Board and a number of privately

owned nature reserves, including the KPNR, bordering

the Kruger National Park (KNP), common fences between

these reserves and the KNP have been removed.

All the more common larger mammals, including the

full spectrum of carnivores occurring in the KNP are also

found in the KPNR.

Plant specimens were collected or recorded routinely

during the course of other work undertaken in KPNR.

This formed part of an overall objective of compiling a

herbarium collection of the central Transvaal Lowveld re-

gion outside the KNP.

Geologically, KPNR consists primarily of granitoid

rocks of the Swazian Period, i.e. Makhutswi Gneiss. The

gneiss is intruded by a few relatively small, scattered out-

crops of Harmony Granite and the Phalaborwa complex.

Other formations which occur in the area include Milky

Quartz Pegmatite and a very small outcrop of Dark Green-

ish Black Clinopyroxene Hornblende of the Rubbervale

Formation (Geological Survey 1986).

The geomorphology of KPNR can generally be de-

scribed as gently rolling or undulating over most of its

area, the southern part being almost flat. The northern

38

Bothalia 24,1 (1994)

FIGURE 1.— The locality of the

Klaserie Private Nature Re-

serve. Broken lines denote

quarter-degree squares. Inset

shows the reserve’s locality

within the Transvaal. Biomes

after Rutherford & Westfall

(1986), figure taken from

Huntley (1989).

third is somewhat more rugged, however, with many rel-

atively short drainage lines. Boulder or rock outcrops are

common here, in places forming low koppies or ridges.

Soils of this region generally comprise shallow sandy

loam or gravel, and loose surface stones and rocks are

common. Elsewhere, soils are generally deeper and pri-

marily sandy to loamy or loamy in texture. Along lower-

lying areas, clay-loam or clay predominates, particularly

along watercourses. Alluvial sand ts frequently encoun-

tered along the banks of the rivers and larger seasonal

streams.

Drainage is provided primarily by the Olifants,

Klaserie, and Ntsiri Rivers. The Olifants and Klaserie are

normally perennial, though in recent times, particularly

during periods of drought, the Klaserie ceases to How,

and the Olifants decreases to a small stream.

Mean annual rainfall recorded in the reserve over the

past 16 years totals 424.7 mm. During this period, a mean

annual rainfall of 452.8 mm has been recorded at Ncheni

Gate, 408.5 mm at Xanatseni Gate, and 432.0 mm at

Xiplapaleni Gate. A mean of 407.8 mm over a 33-year

period has been recorded at the reserve’s headquarters

(Warden’s records, unpublished).

The reserve's mean annual rainfall is well below 530

mm, the approximate upper limit for the arid subdivision

of the Savanna Biome, and is close to 400 mm, the lower

limit given for the dry subdivision of the Grassland Biome

(Rutherford & Westfall 1986).

Altitude ranges from 303 m in the northeast at the con-

fluence of the Klaserie and Olifants Rivers, to 535 m in

the southwestern comer — a difference of 232 m over a

distance of 33 km.

The vegetation of KPNR falls within the Savanna

Biome (Gibbs Russell 1987; Rutherford & Westfall 1986).

According to Acocks (1988), the only Veld Type of the

reserve is Arid Lowveld of the Tropical Bush and Savanna

Types (Bushveld). In the northeastern and eastern sectors

of the reserve however, mopani veld ( Colophospermum

mopane) is present and is dominant over large areas, de-

creasing in extent southwards, where it occurs in the form

of small, scattered pockets. The vegetation of KPNR has

been mapped in greater detail at the reconnaissance level.

Bothalia 24, 1 (1994)

39

and 15 major associations were identified (Zambatis

1983).

METHODS

The great majority of taxa given in the list which fol-

lows were collected or recorded over the period 1973 to

1987 by the author whilst in the service of the Transvaal

Directorate of Nature and Environmental Conservation

(TDNEC). Apart from duplicate specimens donated to the

National Botanical Institute's National Herbarium in Pre-

toria (which undertook the identification of all the speci-

mens collected), the collection is housed in the herba-

rium of the Hans Hoheisen Wildlife Research Station

(HHWRS), bordering the Kruger National Park at Orpen

Gate.

Taxa recorded but not collected by the author were

listed after being identified by reference to herbarium ma-

terial collected elsewhere in the region. Where no doubt

existed as to their identity, these were recorded without

any reference to herbarium material.

Nomenclature, taxonomic sequence of arrangement

and spelling used throughout this list follows that of the

National Botanical Institute’s PRECIS system as detailed

in Gibbs Russell et al. (1985, 1987, 1988; De Wet et al.

1989, 1990, 1991; Arnold & De Wet 1993). Taxonomic

numbering has however been omitted in order to enhance

clarity and brevity.

Synonyms are given only in those cases where taxo-

nomic revision and subsequent name changes occurred

after specimens were originally named.

With few exceptions, the taxa have been annotated

with summarized field notes made at the time of collec-

tion. A standard sequence of annotation has been followed

whereby the first category represents the life form, fol-

lowed by the height range, vegetation type, habitat, and

soil texture. These categories are separated by a comma.

Where a category was not recorded, this is indicated by

a dash.

The vegetation and habitat types given for each taxon

refer only to the specimens collected or recorded. This

does not imply that the taxon in general is restricted to

this vegetation or habitat type. Many of the taxa occur in

a variety of vegetation and habitat types.

Life form classes are according to the Raunkiaer sys-

tem as used by Miiller-Dombois & Ellenberg (1974) and

Rutherford & Westfall (1986). Height classes of phanero-

phytes are those of Miiller-Dombois & Ellenberg (1974).

The following classes are used:

Phanerophytes (P): perennial plants, usually woody,

mean height of the renewal buds > 0.7 m above ground.

Mesophanerophytes (Me): mean height of the renewal

buds 5-50 m. Microphanerophytes (Mi): mean height of

the renewal buds 2-5 m. Nanophanerophytes (N): mean

height of the renewal buds < 2 m. Chamaephytes (Ch):

perennial plants, generally woody or partly woody, mean

height of the renewal buds < 0.7 m. Hemicryptophytes

(H): perennial plants, generally herbaceous, renewal buds

at, or usually, close to ground level and seldom over 0. 1

m. Geophytes (G): perennial plants, usually herbaceous,

renewal buds below ground level. Therophytes (T):

ephemeral plants (annuals). Lianas (L): plants that grow

by supporting themselves on others. Epiphytes (E): plants

that germinate and root on other plants, including dead

standing plants. Parasites (Pa): green plants growing at-

tached to other living autotrophic plants.

The other annotations are explained below:

Height: given in metres. In most cases, a height range

is given, though where only one specimen was collected

or recorded, only one height is given. Vegetation type: the

dominant (i.e. most abundant) woody species, such as

Combretum, Acacia, Colophospermum, etc. 'Mixed wood-

land' is used in cases where dominance by one or more

species is not clearly evident and is thus a mixture of

various species. The term 'woodland' is used in a broad

context and does not necessarily imply a dominance by

trees. A common feature in fact is the presence of both

trees and shrubs, in varying degrees of dominance, though

in no case where 'woodland' is used, is the tree stratum

absent. 'Riverine' refers to the woodland and shrubveld

along the two perennial rivers, the Olifants and Klaserie,

whereas ‘seasonal stream' refers to the vegetation occur-

ring along the banks of the numerous watercourses of a

seasonal nature, varying from substantial rivers such as

the Ntsiri, to the many minor streams, gullies and other

tributaries of the perennial rivers. In both these forms of

riverine vegetation, physiognomic variations exist and

range from poorly developed, to closed tall woodland,

with or without a dense (or more open) thicket under-

storey. Although many taxa are common to both of these

riverine forms, sufficient physiognomic and floristic dif-

ferences in these (and in the water regime) exist to war-

rant a separation between them. 'Koppie' is applied to a

range of vegetation types associated with rocky ridges,

koppies, or isolated boulder or rock outcrops of varying

extent and height above the surrounding terrain. 'Open

parkland' refers to the vegetation structure of the seepline

complex where trees are scattered or absent, with a scat-

tered shrub stratum, whereas 'grassland' refers only to sea-

sonally or perennially waterlogged marshy areas (vleis),

the presence of woody plants being a rare exception. Hab-

itat: the term 'dryland' refers to the broadly homogeneous

area occurring between one drainage line and another, but

excludes distinctly different or specialized habitats which

are floristically and physiognomically atypical of the sur-

rounding area and which generally occupy the same to-

pographic position. Examples of these exceptions are

koppies and rock outcrops, seasonal seepage areas, vleis,

and riverine habitats. An overriding feature of dryland is

the presence of a woody component in the form of one

or two strata, namely tree and shrub layers, each occurring

in varying frequency or dominance, ranging from sparse

to dense. 'Dryland' is thus synonymous with 'veld', in a

broad context, but within the confines of the Savanna

Biome of Rutherford & Westfall (1986) and Gibbs Russell

(1987).

A 'seepline complex' represents a specialized habitat

for a variety of plant types, a number of which appear to

be restricted to this type of habitat. These areas are com-

monly located on slopes on a catenary sequence of soil

40

Bothalia 24,1 ( 1994)

types. Sandy soils occur on the upper side of the seepage

area, grading into sandy loam and finally into clayey loam

or clay at the bottom of the slope. During the rainy season,

and particularly after several days of heavy rain, water

seeps to the surface at the sand/clay interface, creating

damp or even waterlogged conditions for prolonged peri-

ods. A gradient in the distribution of plant types along

these areas is also evident, with Tenninalia sericea char-

acteristically demarcating the upper edge. These areas are

vulnerable to erosion when heavily utilized by herbivores

due to the concentration of salts which lead to defloccula-

tion and collapse of the clay structure. When in this state,

they are known as sodic areas (Scholes 1985). In this ar-

ticle, no distinction has been made between the two, as

certain plant taxa occur on both types of habitats, hence

the term 'complex’.

A 'rock outcrop' refers to the habitat structure of the

koppie vegetation type and can comprise large and exten-

sive boulders, or isolated clumps of rocks. In practically

all rock outcrop habitats, soils are shallow to very shallow,

and sandy or gravelly, witli smaller, loose stones being

common.

RESULTS AND DISCUSSION

A total of 618 infrageneric, or 'lower’, taxa have been

recorded in the KPNR. This does not include specimens

identified to the genus level only, except those which are

the only members recorded in a particular family or genus.

Taxa of the same species but with different subspecies or

varieties were regarded as different taxa. These are

summarised in Table 1 according to the number of fam-

ilies, genera, and taxa in the pteridophytes, monocotyle-

dons, and dicotyledons.

Twenty-seven (26.2%) of the total number of families

contribute 1% or more of the total number of taxa (Table

2). Genera with six or more taxa (> I % of the total number

of taxa) are listed in Table 3.

Free-ranging exotics are represented by 39 taxa.

Five taxa, listed as 'rare' in the Red Data List for the

Transvaal (Fourie 1986; revised by the Flora Division of

TDNEC, unpublished), have been recorded for KPNR.

These were all collected by the author (collector’s number

in brackets): Aloe vandennerwei (880); Nymphaea lotus

(1092); Phyllanthus pinnatus (891, 1214); Ceropegia

mafekingensis (1536); Orbea maculata (1602).

TABLE 2. — Families with 1% or more of the total number of infrageneric

taxa

Two of these, N. lotus and C. mafekingensis , are first re-

cords for the Lowveld (E. van Hoepen pers. comm.). Nei-

ther of these taxa have yet been collected in the KNP,

though N. lotus has apparently been recorded by S.P.

Fourie of the TDNEC. In the case of A. vandennerwei,

the only specimen collected was from a small group which

appears to have been planted at Xanatseni Gate. A search

for wild-growing specimens in the vicinity of this gate

proved fruitless. It is possible that these specimens were

introduced into the reserve from beyond its boundaries.

This plant thus remains to be confirmed in its natural hab-

itat within the reserve.

Despite its relatively arid climate, and the fact that the

surface area of KPNR is a mere 0.1% of the total surface

area of the Savanna Biome of 632 034 km, the reserve’s

These represent 2% of the total number of 25 1 taxa

currently listed in the Red Data List for the Transvaal.

TABLE 1. — Number of families, genera and infrageneric taxa recorded

in the Klaserie Private Nature Reserve

Pterido- Monocoty- Dicoty-

phyta ledonae ledonae

TABLE 3. — Genera with 1% or more of the total number of infrageneric

taxa

Bothalia 24, 1 (1994)

41

618 infrageneric taxa represent 10.7% of the total number

of taxa (5 788) recorded in this biome (Gibbs Russell

1987). The riverine habitats along the two perennial rivers

of the reserve partly account for this relatively high num-

ber of taxa. If the frequency and duration of desiccation

of these rivers, however, increases in the future (as it is

likely to do, given the ever-increasing human demands on

all water resources), a decline of these plant communities,

or even the disappearance of certain taxa is likely to occur.

The quarter-degree squares of the KPNR were not in-

cluded in the PRECIS search undertaken by Gibbs Russell

(1987) in her analysis of the southern African biomes, and

in the determination of the core areas of these biomes. In

this search, she recorded 21 families, comprising 1% or

more of the taxa, which together account for 55-60% of

the total number of taxa recorded in the Savanna Biome.

These families, all of which have been recorded in KPNR

as well, account for 459 (74.2%) of the reserve’s total

number of taxa. These relationships consequently reflect

a strong affinity of the flora of KPNR with that of the

Savanna Biome in general.

A total of 22 families and 36 genera with 10 taxa or

more, have their centres of diversity in the Savanna and

Grassland Biomes (Gibbs Russell 1987). All except one

of these (Aspleniaceae) and six genera, have also been

recorded in the KPNR, indicating a strong relationship

with the Grassland Biome as well. It is significant to note

though, that the Orchidaceae, represented by a total of 53

taxa common to the Savanna and Grassland Biomes, and

which, together with the Lamiaceae, distinguishes the

Grassland Biome (Gibbs Russell 1987), is very poorly

represented in KPNR, with only two taxa being recorded.

This poor representation of the Orchidaceae can most

probably be ascribed to the relatively arid climate of

KPNR.

In the Umfolozi Game Reserve, on the other hand, this

family is absent, in spite of a mean annual rainfall of 625

mm. Downing & Gibbs Russell (1981) suggest that wild

herbivores could have severely depleted the rhizomes of

any ground orchids present, while extensive and intensive

spraying of insecticides during the tse-tse fly eradication

campaign could have inhibited sexual reproduction by

eliminating insect pollinators of both ground and epiphytic

orchids. A similar situation is evident in the Manyeleti

Game Reserve, bordering the KNP on the western side of

the central region of the Park. Here, Bredenkamp (1982)

only recorded one taxon of this family, even though this

reserve falls within the mesic subdivision of the Savanna

Biome (Rutherford & Westfall 1986). No insecticide

spraying has been undertaken in KPNR.

The generally arid nature of a major part of the

Lowveld consequently appears to be the reason for the

relatively poor representation of this family in this part of

the Savanna Biome, which therefore weakens the link be-

tween this biome and the Grassland Biome somewhat, at

least in this region.

The aridity of KPNR is most probably also the reason

for the absence of the Aspleniaceae. This large family ap-

pears to be confined to the higher rainfall regions of the

country, as shown on the distribution maps of Jacobsen

(1983) and Burrows (1990). This is further reflected by

the fact that the only two members of this family recorded

in the KNP are restricted to the Pretoriuskop area of the

Park, which receives the highest mean annual rainfall for

the Park of some 722 mm.

Table 4 summarises the life forms of the KPNR flora.

This shows that woody plants — phanerophytes and cham-

aephytes, together comprise 31.7% of all taxa, whereas

herbaceous plants, the hemicryptophytes and therophytes,

comprise 51.7%. The high frequency of woody and her-

baceous taxa is further evidence that the vegetation of the

reserve comprises a woody layer above an herbaceous

layer.

TABLE 4. — Number of infrageneric taxa and percentage of total per life

form category

Mes, mesophanerophyte; Mi, microphanerophyte; N. nanophanero-

phyte; Ch, chamaephyte; H, hemicryptophyte; G, geophyte; T, thero-

phyte; L, liana; E. epiphyte; Pa. parasite; P. phanerophyte.

CONCLUSION

The primary purpose of this article is to provide a sys-

tematic list of the flora of KPNR, which can be used in

a variety of more detailed studies. Nevertheless, this rather

brief analysis of the reserve’s flora shows that very strong

affinities exist with the Savanna Biome, and to a lesser

degree, with the Grassland Biome as well. The vegetation

of the Klaserie Private Nature Reserve can therefore be

regarded as comprising part of the core area of the Sa-

vanna Biome.

ACKNOWLEDGEMENTS

The many landowners of KPNR are thanked for al-

lowing me to collect plant specimens on their properties,

as is the Warden, Mr E.P Leibnitz, for his interest and

support.

The identification of the large number of specimens

collected over the years was undertaken by the National

Botanical Institute, Pretoria. Without the assistance of the

many persons involved, it would not have been possible

to compile this list. In particular, the assistance of Mrs E.

van Hoepen, Ms E. Retief, and Mrs B.C. de Wet is grate-

fully acknowledged.

The Directorate of Nature and Environmental Conser-

vation (Transvaal) is thanked for providing the support

which made this work possible.

REFERENCES

ACOCKS. J.RH. 1988. Veld types of South Africa, 3rd edn. Memoirs of

the Botanical Survey of South Africa No. 57.

42

Bothalia 24,1 (1994)

ARNOLD, T.H. & DE WET, B.C. 1993. Plants of southern Africa: names

and distribution. Memoirs of the Botanical Survey of South Africa

No. 62.

BREDENKAMP, G.J. 1982. 'n Plantekologiese studie van die Manyeleti-

wildtuin. D.Sc. thesis. University of Pretoria, Pretoria.

BURROWS, J.E. 1990. Southern African ferns and fern allies. Frandsen

Publishers, Sandton.

DE WET, B.C., GIBBS RUSSELL, G.E., GERMISHUIZEN, G„

SHRIRE, B.D.. JORDAAN, M„ PIENAAR, B.J., WELMAN,

W.G., REID, C., VAN WYK, C.M., FISH. L„ IMMELMAN,

K.L., VAN ROOY, I, GLEN, H.F. & BARKER, N.P. 1989. New

taxa, new records and name changes for southern African plants.

Bothalia 19: 275-294.

DE WET, B.C., GERMISHUIZEN, G„ SHRIRE, B.D., JORDAAN, M„

PIENAAR, B.J., WELMAN, W.G., REID, C., VAN WYK, C.M.,

FISH, L„ IMMELMAN, K.L., VAN ROOY, J„ PEROLD, S.M.,

TAUSSIG, J„ BARKER, N.P. & GLEN, H.F. 1990. New taxa,

new records and name changes for southern African plants.

Bothalia 20: 249-266.

DE WET, B.C., ARCHER, R„ FISH, L„ GERMISHUZEN, G„ HER-

MAN, P.P., JORDAAN, M„ PEROLD, S.M., REID, C„ VAN

ROOY, J„ WELMAN, W.G. & GLEN, H.F. 1991. New taxa, new

records and name changes for southern African plants. Bothalia

21: 191-213.

DOWNING, B.H. & GIBBS RUSSELL, G.E. 1981. Phytogeographic

and biotic relationships of a savanna in southern Africa: analysis

of an angiosperm checklist from Acacia woodland in Zululand.

Journal of South African Botany 47: 721-742.

FOURIE, S.P. 1986. The Transvaal, South Africa, Threatened Plants

Programme. Biological Conservation 37: 23^-2.

GEOLOGICAL SURVEY 1986. 1:250 000 Geological Series. 2430

Pilgrim’s Rest. Map published by Dept, of Mineral and Energy

Affairs, Pretoria. Government Printer, Pretoria.

GIBBS RUSSELL, G.E. 1 987. Preliminary floristic analysis of the major

biomes in southern Africa. Bothalia 17: 231-227.

GIBBS RUSSELL, G.E., REID, C., VAN ROOY, J. & SMOOK, L. 1985.

List of species of southern African plants, edn 2, part 1 . Memoirs

of the Botanical Survey of South Africa No. 5 1 .

GIBBS RUSSELL, G.E., REID, C„ FISH, L„ GERMISHUIZEN, G„

VAN WYK, M„ VAN ROOY, J. & STAFF 1987. New taxa, new

records and name changes for southern African plants. Bothalia

17: 269-275.

GIBBS RUSSELL, G.E., WELMAN, W.G., GERMISHUIZEN, G„

RETIEF, E„ PIENAAR, B.J., REID, C., FISH, L„ VAN ROOY,

J., VAN WYK, C.M., KALAKE, E. & STAFF 1988. New taxa,

new records and name changes for southern African plants.

Bothalia 1 8: 293-304.

HUNTLEY, B.J. 1989. Biotic diversity in southern Africa: concepts and

conservation. Oxford University Press, Cape Town.

JACOBSEN, W.B.G. 1983. The fern and fern allies of southern Africa.

Butterworths, Pretoria.

MULLER-DOMBOIS , D. & ELLENBERG, H. 1974. Aims and methods

of vegetation ecology. John Wiley, New York.

RUTHERFORD, M.C. & WESTFALL, R.H. 1986. Biomes of southern

Africa — an objective categorization. Memoirs of the Botanical

Survey of South Africa No. 54.

SCHOLES, R.J. 1985. A guide to bush clearing in the eastern Transvaal

Lowveld. Resource Ecology Group, Dept, of Botany, University

of the Witwatersrand, Johannesburg.

WITKOWSKI, E.T.F. 1983. Ecology of the Klaserie Private Nature

Resen’e. M. Sc. thesis. University of the Witwatersrand. Johan-

nesburg.

ZAMBATIS, N. 1983. The major vegetation associations of the Klaserie

Private Nature Resen’e. Unpublished map, Transvaal Nature

Conservation Division.

SYSTEMATIC CHECKLIST

After the name of the author(s) of the species name, the sequence of annotation is as follows:

collection number : without initials — author; with initials — RPE = R.P. Ellis, JdK = J.P.S. de Kock, P.J.M. = P.J.M. Muller.

§ taxa recorded, but not collected by author.

+ recorded by Witkowski ( 1983).

* exotic taxa, including naturalized taxa.

life form: Mes, mesophanerophyte; Mi, microphanerophyte; N, nanophanerophyte; Ch, chamaephyte; H, hemicryptophyte; G, geophyte; T, thero-

phyte; L, liana; E, epiphyte; Pa, parasite,

height range in metres,

vegetation type,

habitat,

soil texture.

- not recorded.

PTERIDOPHYTA

SELAGINELLACEAE

Sellaginella

dregei (C. Presl) Hieron., 1118 , H, 0.02, open parkland, seepline com-

plex, shallow sandy loam on flat sheetrock

MARSILEACEAE

Marsilea

ephippiocarpa Alston, RPE 3464 , §, T, 0.1-0.15, Colophospermum

woodland, seasonal pans & still pools, sandy, or black clay

ADIANTACEAE

Actiniopteris

radiata (J. Kiinig ex Sw. ) Link, 1591 , H, 0.05-0. 1 , boulder bases, steep

south slopes, shade or sunlight, calcrete

Cheilanthes

involuta (Sw.) Schelpe & N. C. Anthony var. involuta, 1594, H, 0. 1, - ,

boulder bases, steep south slopes, shade, shallow sandy

Pellaea

calomelanos (Sw.) Link var. calomelanos, 984, 1 176, 1595, H, 0.2-0. 3,

Combretum woodland, steep south or east slopes, shade or sun-

light, sandy

THELYPTERIDACEAE

Ampelopteris

prolifera (Retz.) Cope!.. 983, H, 0.4-0. 6, riverine, reed bed, damp or

saturated river sand

ANGIOSPERMAE-MONOCOTYLEDONAE

TYPHACEAE

Typha

capensis (Rohrb.) N.E. Be, 970, H, 2.0, aquatic, dam shores, mud

POACEAE

Sorghum

bicolor (L.) Moench subsp. arundinaceum (Desv.) De Wet <£ Harlan

(= S. verticilliflorum (Steud.) Stapf), §, T, 1.0-1. 5, -, seasonally

wet areas, loam, sandy loam

versicolor Andersson, 913, 1121, H, 1.0, grassland, vlei edge, clayey

loam, stony to sandy loam

Bothriochloa

radicans (Lehm.)A. Camus, 307, 349, 624, 760, 1030, H, 0.4-0.9, Com-

bretum woodland, riverine, dryland, termitarium, river bank,

alluvial sand, sandy loam, loam

Dichanthium

annulatum (Forssk.) Stapf var. papillosum (A. Rich.) De Wet & Harlan

(= D. papillosum (Hochst.) Stapf), 606, H, 1 .7, riverine, river bed,

river sand

Schizachyrium

exile (Hochst.) Pilg.. RPE 3462, T, -, Colophospermum veld, -, gravelly

Andropogon

chinensis (Nees) Merr.

(= A. schinzii Hack.), 311, 759, 909, H, 1.0- 1.5, Combretum &

mixed woodland, -, sandy loam

gayanus Kunth var. polycladus (Hack.) Clayton

(= var. squamulatus (Hochst.) Stapf), 527, 908, 968, H, 2.0,

Bothalia 24,1 (1994)

43

riverine, riverbed, rock outcrops, river sand, gravel, stony or

sandy loam

Cymbopogon

plurinodis (Stapf) Stapf ex Burn Davy, 352, H, 0.9, Combretum wood-

land, dryland, sandy loam

Hyperthelia

dissoluta (Nees ex Steud .) Clayton, 963, 990, 1736, H, 2. 0-2. 5, Colo-

phosperrnum woodland, disturbed (de-bushed), seasonal stream

bank, rock outcrop, stony or sandy loam

Heteropogon

contortus (L.) Roem. & Schult., 910, H, 1.0, mixed woodland, dryland,

sandy loam

Diheteropogon

amplectens(Ateey) Clayton, 966, 967, H, 0.9, Combretum woodland, rock

outcrop, sandy loam

Themeda

triandra Forssk., §, +, H, 0.6, -

Digitaria

eriantha Steud, 592

(= subsp. eriantha), 877, 929, 971

(= subsp .pentzii (Stent) Kok), 931

(= subsp. stolonifera (Stapf) Kok), 876, 878, 930, 939

(= subsp. transvaalensis Kok), 1366

All specimens combined: H, 0.4— 1.2, Combretum woodland,

mixed woodland, riverine, dryland, seepline complex, river bed,

river sand, sandy or stony loam, clay

seriata Stapf, 996, H, 0.5, riverine, river bank, alluvial sand

Eriochloa

meyeriana (Nees) Pilg. subsp. grandiglumis (Stent & J.M. Rattray ) Gibbs

Russell, 322, H, 0.4, riverine, seasonal stream, river sand

meyeriana (Nees) Pilg. subsp. meyeriana, 1029, RPE 3460, H, -, riverine,

river bank, alluvial sand

Brachiaria

deflexa ( Schumach .) C.E. Hubb. ex Robyns

(= Pseudobrachiaria deflexa (Schumach.) Launert), 593, 602, T,

0.7-0.9, Combretum woodland, dryland, disturbed (old lands),

loam

nigropedata (Ficalho & Hiem) Stapf, 616. 945, H, 0.6, Combretum &

mixed woodland, dryland, sandy loam

serrata (Thunb.) Stapf 528, H. 0.5, Combretum woodland, dryland, rock

outcrop, -

xantholeuca (Schinz) Stapf, 601, T. 0.7, disturbed (old lands), loam

Paspalum

urvillei Steud. * 1025, H, 2.0, aquatic, shores of dam, sandy loam

Urochloa

mosambicensis (Hack.) Dandy, §, +, H, 0.4, Combretum woodland,

dryland, sandy loam

trichopus (Hochst.) Stapf, 331, T, 0.3, riverine, -, sandy loam

Echinochloa

jubata Stapf, §, H, -

Panicum

coloratum L. var. coloratum. 611, H, 0.4—0.5, Combretum woodland,

dryland, sandy loam

deustum Thunb., 361, 926, H, 1.3, riverine, woodland, river and stream

bank shade, rock outcrop, river sand, stony

maximum Jacg., 313, 360, 589, 591. 949, H, 0. 7-2.0, woodland, riverine,

dryland shade, termitarium, river banks, sandy loam, river sand

Setaria

incrassata (Hochst.) Hack.

(= S. woodii Hack. var. woodii), 604, H, 1 .5, grassland, vlei, clay,

clayey loam

sagittifolia (A. Rich.) Walp.

(= Cymbosetaria sagittifolia (A. Rich. ) Schweick., 1096), T, 0.5-

1 .0, -, rock outcrop, loam

ustilata De Wit, 915, 1646, T, 0.4-0. 6, mixed woodland, seasonal stream.

shade under Combretum, river sand, sandy loam

verticillata (L.) P. Beauv., 603, T, 0.7, disturbed (old lands), loam

Melinis

repens (Willd.) Z.izka subsp. repens

(= Rhynchelytrum repens (Willd.) C.E. Hubb.), 387, T, 0.6,

disturbed (roadside, de-bushed), sandy loam

Tricholaena

monachne (Trin.) Stapf & C.E. Hubb. 363, H, 0.6, Combretum woodland,

dryland, sandy loam

Pennisetum

macrourum Trin., 1676, H, 2.0-3. 0, riverine, river bed, river sand

setaceum (Forssk. ) Chiov. * 987, H, 0.9, -, abandoned garden, stony loam

Cenchrus

ciliaris L„ 324, 334, 526, H, 0.3-0. 5, riverine, termitarium, stream bank,

sandy stony loam, loam

Phragmites

mauritianus Kunth, 1022, H, 5.0, riverine, river bed, river sand

Stipagrostis

hirtigluma (Trin. & Rupr.) De Winter subsp. patula (Hack.) De Winter,

1674, H, 0.4, Combretum woodland, dryland, shallow gravel

Aristida

adscensionis L.

(= subsp. guineensis (Trin. & Rupr.) Henrard), 761. 916, 948. T.

0.5-0.9, mixed woodland, disturbed, roadside, stony or sandy

loam

(= subsp. adscensionis, 989), T, 0.8, Combretum-Commiphora

woodland, rock outcrop, shallow stony

canescens Henrard subsp. ramosa De Winter, 351, 398, 594, H, 0.2-0. 6,

Combretum woodland, dryland, sandy loam, loam

congesta Roem. & Schult. subsp. barbicollis (Trin. & Rupr.) De Winter,

398, 594, 997, 998, T, 0.4— 0.6, Combretum woodland, dryland,

disturbed (roadside, overgrazed), stony or sandy loam, loam

meridionalis Henrard, 610, H, 0.9, Combretum woodland, dryland, rock

outcrop, stony gravel

scabrivalvis Hack, subsp. scabrivalvis, 944, T, 0.5, -, roadside, sandy

loam

stipitata Hack, subsp. graciliflora (Pilg.) Melderis, 391, T, 0.5, Com-

bretum woodland, dryland, sandy loam

Tragus

berteronianus Schult., 325, T, 0.3, Combretum woodland, disturbed

(trampled), dryland, sandy loam

Perotis

patens Gand., §, T, 0.3, Combretum woodland, dryland, sandy

Sporobolus

festivus A. Rich.

(= var. fibrosus Stent), RPE 3464D, H, -, Colophospermum

woodland, -, shallow gravelly

fimbriatus (Trin.) Nees, 933, 935, 937, H, 1.0, mixed woodland, riverine,

dryland, river bed, stony or sandy loam

nitens Stent, 350, 927, 1010, H, 0.2, Combretum woodland, dryland,

sandy

panicoides A. Rich., 936, T, 0.7, mixed woodland, dryland, sandy loam

stapfianus Gand., §, H, 0.2, -, -, -

virginicus (L. ) Kunth, RPE 3463, H. -, -, -

Eragrostis

ciliaris (L.) R. Br„ §, T. 0.3, -, disturbed (garden), sandy

cylindriflora Hochst., 390, 977, T, 0.2-0.4, -, disturbed (roadside), sandy

loam

gummiflua Nees, §, H. 0.3-0. 5, grassland, open parkland, seepline com-

plex, sandy clay, clayey loam

heteromera Stapf, 917, H. 1.5, riverine, grassland, seasonal stream, vlei,

sandy, clay loam, clay

inamoena K. Schum., 946, H, -, grassland, open parkland, seepline

complex, sandy clay

lehmanniana Nees var. lehmanniana, RPE 3464, H, -, -, -

pilosa (L.) P Beauv. *, 892, 1009, T, 0. 2-0.3, -, garden lawn, sandy loam

rigidior Pilg., 934, H, 1.0, mixed woodland, dryland, sandy loam

rotifer Rendle, 321, 607, 912, H, 0.9-1 .5, riverine, river bed, river sand,

coarse river sand

superba Peyr., 316, 938, H, 0.4— 0.5, Combretum & Colophospennum

woodland, dryland, sandy loam

trichophora Coss. & Durieu, 932, H, 0.3, -, disturbed (roadside, old

lands), sandy loam

sp., 613, 947, -, 0.6-1. 5, riverine, river bed, seasonally waterlogged river

sand, sandy loam

Cynodon

dactylon (L. ) Pers., §, +. H, 0.2, riverine, river bank, sand bank, alluvial

sand

Enteropogon

macrostachyus (A. Rich.) Benth., 525, 911, H, 1.0, mixed woodland,

dryland, sandy loam, loam

Chloris

gayana Kunth, 358, H, 0.9-1. 0, riverine, riverbed, river sand

mossambicensis K. Schum., 928, H. 0.4, riverine, seasonal stream, river

& stream bank, clayey loam

roxburghiana Schult., 353, H, 0.7, -, termitarium, sandy clay

Oropetium

capense Stapf, RPE 3464E, H, -, Colophospermum veld, -, shallow gravel

Dactyloctenium

aegyptium (L.) Willd., 561, T, 0.6, open parkland, seepline complex,

garden, loam

geminatum Hack., 1410, H, 0.7, riverine, reed bed, river sand

giganteum Fisher & Schweick., 965, 1645, T. 0. 5-1.0, riverine, outer

edge of river bed, alluvial sand

44

Bothalia24,l (1994)

Pogonarthria

squarrosa (Roem. & Schult.) Pilg ., §, H, 0.2-0.4, Combretum & mixed

woodland, dryland, overgrazed areas, sandy

Diplachne

fusca (L.) P. Beauv. ex Roem. & Schult.. RPE 3464A, H, Colophospermum

veld, -, heavy black turf

Trichoneura

grandiglumis (Nees) Ekman var. grandiglumis, §, +, H, 0. 2-0.3, Com-

bretum woodland, dryland, sandy

Enneapogon

cenchroides {Roem. & Schult. ) C.E. Hubb.. 355, 988, T, 0.3- 0.6, Acacia

woodland, dryland, rock outcrop, shallow stony loam

Schmidtia

pappophoroides Steud., 309. 969 , H, 0.2-0. 5, Combretum woodland,

riverine, dryland, riverbank, sandy, alluvial sand

Fingerhuthia

africanaLe/zm., 312, H, 0.5, Colophospermum woodland, dryland, sandy

Megastachya

mucronata (Poir. ) P. Beauv., +, T, -

CYPERACEAE

Cyperus

articulatus L., 585, H, 1.2, riverine, river bed, river sand

compressusL., 918, 1647, H, 0.5, seasonal stream, stream bed, river sand

difformis L., 595, 612, H, 0.3-0. 7, grassland, riverine, vlei, river bed,

river sand

distans L. f, 1586, H, 1.4, riverine, reed bed, river sand

fastigiatus Rottb., 587, H, 1 .8, riverine, reed bed, river sand

immensus C.B. Clarke, 588, H, 2.7, riverine, reed bed, river sand

iria L., 596, H, 0. 3-0.4, grassland, vlei, sandy

margaritaceus Vahl, +, H, -, -, -

obtusiflorus Vahl var. obtusiflorus, 346B, 920, H, 0.2-0.4, Combretum

and mixed woodland, tree shade in dryland, sandy loam

rupestris Kunth var. rupestris

(= C. rupestris Kunth), 258, H, 0.1, grassland, vlei, black clayey

loam

sexangularis Nees, 320, H, 0.9, riverine, river bed, river sand

sphaerospermus Schrad., §, H. -, -

tenax Boeck., 586, 599, H, 1.4, riverine, river bed, river sand

thomcroftii McClean, 1633, H, 0.3, rock outcrop, edge of small seasonal

rock pool, granite rock

Pycreus

macrostachyos (Lam.) J. Raynal, 597, H, 0.6-0.7, grassland, vlei, sandy

pumilus (L.) Nees

(= subsp. patens (Vahl) Podlech), 609, T, 0. 1, riverine, river bed,

river sand

Mariscus

indecorus (Kunth) Podlech, §, H, -, -, -

rehmannianus C.B. Clarke, 919, T. 0.4, mixed woodland, dryland, sandy

loam

squarrosus (L.) C.B. Clarke, §, H, -, -, -

Kyllinga

alba Nees, 257, H, 0. 1 , grassland, vlei, black clayey loam

Fuirena

ciliaris (L.) Roxb. var. ciliaris, 1117, H, 0.4, open parkland, seepline

complex, waterlogged sandy clayey loam

pubescens (Poir.) Kunth, 625, H, 0.4, grassland, vlei, sandy

Fimbristylis

bisumbellata (Forssk.) Bubani, 1347, H, 0.2-0. 3, riverine, river bed,

alluvial & river sand

ferruginea (L.) Vahl, §, H, 0.5, -, -, -

microcarya F. Mueli, 608, H, 0. 1 -0.2, riverine, river bed, river sand

Bulbostylis

burchellii (Ficalho & Hiem ) C.B. Clarke, 986, H, 0.4, rock outcrop,

seepage area amongst boulders, sandy

hispidula (Vahl) R.W. Haines, 397, H, 0.3, -, garden, sandy loam

Cladium

mariscus (L.)Pohl subsp. jamaicensef Crantz) Kiik., 890. H, 2.0, riverine,

dam shore, alluvial sand, clay

ARECACEAE

Phoenix

reclinata Jacq., $, Mi, 2,0. riverine, seasonal stream, river & stream bank

or bed, alluvial sand, river sand

Hyphaene

coriacea Gaertn.

(= H. natalensis Kuntze), §, Mi, 1.3, -, -

ARACEAE

Sty loch iton

natalensis Schott, 1365, G, 0.5, mixed woodland, Combretum-Conuni-

phora woodland, dryland, sandy or stony loam

COMMELINACEAE

Commelina

africanaL. var. lancispatha C.B. Clarke, 1058, H, 0.3-0. 7, mixed wood-

land, dryland, sandy loam

diffusa Burnt, f. subsp. scandens (C.B. Clarke ) Oberm., 1368, H, 0.6,

riverine, reed bed, river sand

subulata Roth, 1095, T, 0.3, aquatic, seasonal pan, waterlogged clay

Cyanotis

speciosa (L. f) Hassk., 1088, H, 0. 1-0.2, Colophospermum woodland,

bare soil, sandy clay

COLCHICACEAE

Gloriosa

superba L., §, G, 1.3, mixed woodland, dryland, sandy loam

Camptorrhiza

strumosa ( Baker ) Oberm., 1537, G, 0.2, Colophospermum woodland/riv-

erine ecotone, dryland, sandy clayey loam

ASPHODELACEAE (PART A)

Trachyandra

sal tii (Baker) Oberm. var. saltii, 253, 1533, G, 0. 1 , Combretum woodland,

dryland, sandy loam

Anthericum

cooperi Baker, 254, G, 0. 1, Combretum woodland, dryland, sandy loam

galpinii Baker var. galpinii, 373, 1052, 1350, G, 0. 2-0.3, Acacia/Com-

bretum woodland, open parkland, dryland, seepline complex,

stony sandy loam

longistylum Baker, 1216, G, 0.3, open parkland, seepline complex, sandy

clayey loam

HYACINTHACEAE (PART A)

Schizobasis

intricata (Baker) Baker, 1505, 1623, G, 0.3-0. 5, Combretum-Commi-

pltora woodland, dryland, shallow gravel, sandy loam

ERIOSPERMACEAE

Eriospermum

burchellii Baker, 314, G, 0.3, Colophospermum woodland, dryland,

gravel

galpinii Schinz, 617, 1056, G, 0.2, grassland, open parkland, vlei, seepline

complex, loam, sandy clay

luteo-rubrum Baker, §, G, -, -, -, -

ASPHODELACEAE (PART B)

Aloe

lutescens Groenew., §, N, -, -, -, -

marlothii A. Berger subsp. marlothii, §, N, -, -, -, -

vandermerwei Reynolds, 880, N, 1.2, -, yard of entrance gate (planted ?),

sandy, stony

HYACINTHACEAE (PART B)

Albuca

angolensis Welw., 1617, 1635, G, 0.6-2.0, koppie, rock outcrop, shallow

loam, humus-rich loam

Urginea

epigea R.A. Dyer, 1049, G, 1.0-1. 4, mixed woodland, dryland, sandy

loam

sanguineaSc/zhzz, 1624, G,-, Combretum woodland, dryland, sandy loam

Drimia

sp. cf. D.elata7flci7., 1618, G, 0.3, open mixed woodland, dryland, clayey

loam

Dipcadi

glaucum (Ker Gaxvl.) Baker, 1587, G, 0.4, Combretum woodland, dry-

land, shallow sandy or stony loam

gracillimum Baker, 359, G, 0.4— 0.5, Combretum woodland/riverine eco-

tone, river bank, dryland, alluvial sand, stony loam

viride (L.) Moench, 1586, G, 1.0, Combretum woodland, dryland, sandy

loam

Ornithogalum

seineri (Engl. & K. Krause) Oberm., 326, 855, G, 0.2-0.3, Combretum

woodland, dryland, old lands, gravel, sandy loam

Drimiopsis

burkei Baker, §, G, 0.2, Combretum woodland, koppie, dryland shade,

rock outcrop, sandy, shallow

Bothalia 24,1 (1994)

45

Ledebouria

apertiflora (Baker) Jessop, 1305, G, 0.2, open parkland, seepline com-

plex, sandy clay

cooperi (Hook, f.) Jessop, §, G, -, -, -, -

marginata (Baker) Jessop, 1254, G, 0.1, Combretum woodland, dryland,

clayey loam

sp., 1509, -, 0.3, mixed woodland, dryland, sandy loam

DRACAENACEAE

Sansevieria

hyacinthoides (L.) Druce, 1372, 1373, H, 0. 3-0.5, koppie, mixed wood-

land, open parkland, rock outcrop, seepline complex (upper

edge), shallow, sandy, sandy clayey loam

pearsonii N.E. Br., 1611, H, 0.3-0. 7, koppie, rock outcrop, sandy loam

ASPARAGACEAE

Protasparagus

africanus (Lam.) Oberm.

(= Asparagus africanus L.), §, G, 0.6, Combretum woodland,

dryland, sandy loam

buchananii (Baker) Oberm.

(= Asparagus buchananii Baker), §, G, 2.0-3. 0, koppie, rock

outcrop, sandy loam

exuvialis (Burch.) Oberm.

(= Asparagus exuvialis Burch.), 376, G, 0.6, Combretum wood-

land, dryland, sandy loam

natalensis (Baker) Oberm.

(= Asparagus falcatus L. var. ternifolius (Baker) Jessop), 1067,

G, 1 .0, Colophospermum woodland, erosion area, gravelly clay

AMARYLLIDACEAE

Scadoxus

multiflorus (Martyn) Raf. subsp. multiflorus, 1257, G, 0.3, mixed wood-

land, termitarium, sandy loam, sandy clayey loam

Boophane

disticha (L.f.) Herb., §, G, 0.5, mixed woodland, Combretum woodland,

dryland, sandy loam

Crinum

buphanoides Welw. ex Baker, §, G, 0.4, open parkland, seepline complex,

clayey loam, clay

graminicola L Verd., §, G, 0.6. Combretum woodland, dryland, sandy

loam

minimum Milne-Redh., 1236, G, 0.2, Combretum woodland, dryland,

sandy loam

Ammocharis

coranica (Ker Gawl.) Herb., 1504, G, 0.3, riverine, river bank, alluvial

sand

Pancratium

tenuifolium Hochst. ex A. Rich., §, G, 0.2, Combretum woodland, dry-

land. sandy loam

HYPOXIDACEAE

Hypoxis

hemerocallidea Fisch. & C.A. Mey.

(= H. rooperi S. Moore), §, G, 0.3, -, -

VELLOZIACEAE

Xerophyta

retinervis Baker, §, Ch, 0.4-0.7, Combretum woodland, dryland, shallow

sandy or gravelly loam

DIOSCOREACEAE

Dioscorea

sylvatica ( Kunth ) Eckl. var. sylvatica, 1287, Ch, 1.5, seasonal stream,

stream bank, shallow, stony

Eulophia

petersii Rchb.f, 1516, H, 1.2, koppie, rock outcrop, shallow leaf litter

and humus

ANGIOSPERMAE-DICOTYLEDONAE

SALICACEAE

Salix

mucronata Thunb. subsp. capensis (Thunb.) Immelman

(= S. capensis Thunb.), 1028, Mi, 3.0, riverine, river bank,

alluvial sand

MORACEAE

Madura

africana (Bureau) Corner, 1035. Mi, 3.0. riverine, river bank, alluvial

sand

Ficus

abutilifolia (Miq.) Miq.

(= F. soldanella Warb. ), 992, Me, 7.0-10.0, koppie, rock outcrop,

stony loam

glumosa (Miq. ) Delile

(= F. sonderi Miq., §, +. Me, -, -, -, -

ingens (Miq.) Miq. var. ingens, 1517, Me, 5.0, riverine cliff, rock outcrop,

shallow, gravelly

sycomorus L„ 1401, Me, 15.0-20.0, riverine, river bank, alluvial sand

URTICACEAE

Pouzolzia

mixta Solms

(= P. hypoleuca Wedd.), 1596, N, 1.5, koppie, rock outcrop,

humus-rich sandy loam

LORANTHACEAE

Plicosepalus

kalachariensis (Schinz) Danser, 1162, Pa, -, woodland. Acacia nigres-

cens, -

SANTALACEAE

Osyridicarpos

schimperianus (Hochst. ex A. Rich.) A. DC., 1612, H, 0.5, seasonal

stream, vertical stream bank, alluvial sand

OLACACEAE

Ximenia

americana L. var. americana, 577, N, 1.5, Combretum woodland, dryland,

loam

caffra Sond. var. caffra, 1742 , N, 1. 2-2.0, Combretum-Commiphora

woodland, seasonal stream, dryland, stream bank thicket, stony

sandy loam, alluvial sand

POLYGONACEAE

Polygonum

aviculare L. *, §, H, -, -, -, -

Persicaria

serrulata (Lag.) Webb & Moq., 600, 1027, 1393, Ch, 0.6-1. 0, riverine,

river bed, river sand, alluvial sand

Oxygonum

sinuatum (Hochst. & Steud. ex Meisn.) Dammer, 368, 1309, 1642, H,

0.2-0.3, -, disturbed (old lands), de-bushed area, sandy loam

CHENOPODIACEAE

Chenopodium

ambrosioides L. *, 943, T, 0.6, riverine, river bed, river sand

schraderianum Roem. & Schult. *, 1418, T, 0.7, riverine, river bed, river

sand

IRIDACEAE

Lapeirousia

masukuensis Vaupel & Schltr. 1345, G, 0.4, open parkland, seepline

complex, sandy clay

sandersonii Baker, §, G, 0.4— 0.5, open parkland, seepline complex, sandy

clay

ORCHIDACEAE

Ansellia

africana Lindl.

(= A. gigantea Rchb. f. var. nilotica (Baker) Summerh.), 1288,

1311, E, 0.6, riverine. Acacia woodland, Diospyros mespili-

formis, Acacia nigrescens, -

AMARANTHACEAE

Hermbstaedtia

odorata (Burch.) T. Cooke var. odorata, 562, Ch, 0.6, Combretum wood-

land, dryland, loam

Sericorema

remotiflora (Hook, f.) Lopr., §, Ch, 0.6, Colophospermum woodland,

dryland, clayey loam

Kyphocarpa

angustifolia (Moq.) Lopr., 1007, H, 1.0, mixed woodland, dryland, sandy

loam

cruciata (Schinz) Schinz, 1187, T, 0.3-0.4, mixed woodland, trampled

veld, clayey loam

Cyathula

cylindricaMoc/., 590, T, 1.0, Combretum woodland, dryland, sandy loam

46

Bothalia 24,1 (1994)

lanceolata Schinz

(= C. hereroensis Schinz), 777, Ch, 0.3, -, disturbed (de- bushed),

sandy

Pupalia

lappacea (L.) A. Juss. var. lappacea *

(= P. atropurpurea (Lam.) Moq.), 1160, 1342 , H, 0.4-1. 5, river-

ine, river bank, disturbed (roadside), gravel, sandy loam

Achvranthes

asperaL. var. aspera *, 1159, 1477 , Ch, 0.4-1 .5, riverine, shade of riverine

woodland, alluvial sand

Alternanthera

pungens Htiinb., Bonpl. & Kunth *, 1330. H, 0.03-0.05, disturbed areas,

gravel, sandy loam

Gomphrena

celosioides Mart. *, 1319. H, 0.3, aquatic, seepage below dam, sandy,

stony

NYCTAGINACEAE

Commicarpus

fallacissimus (Heimerl) Heimerl ex Oberm., Schweick. & 1. Verd., 1306,

Ch, 0.2, Combretum woodland, disturbed (trampled area next to

waterhole), sandy clay loam

plumbagineus (Cav.) Standley, §, Ch, 1.2, riverine, river bank, sandy

loam

Boerhavia

diffusa L.*, 283, 285, 1253, Ch. 0.2-0.6, grassland, vlei, disturbed

(garden), sandy clayey loam, loam, clayey loam

AIZOACEAE (PART A)

Limeum

dinteri G. Schellenb., §, H, -, -, -, -

fenestratum (Fenzl) Heimerl var. fenestratum, 942, H, 0.4, -, disturbed

(roadside), gravel

myosotis H. Walter var. confusum Friedrich, §, T, -, -, -, -

pterocarpum (J. Gay) Heimerl var. pterocarpum, 1581 , T, 0.5, Com-

bretum woodland, heavily trampled and overgrazed areas, sandy

loam

sulcatum (Klotzsch) Hutch, var. sulcatum. 280, 622, 1474, H. 0. 1-0.5,

Combretum woodland, dryland, disturbed (garden), roadside,

sandy, sandy loam

AIZOACEAE (PART B )

Gisekia

africana (Lour.) Kuntze var. africana, 941, 1580, T, 0.02-0.05, Com-

bretum woodland, dryland, roadside, sandy loam, gravel

africana (Lour.) Kuntze var. cymosa Adamson, 335, T, 0.3, Combretum

woodland, dryland, sandy

Mollugo

cerviana (L.) Ser. ex DC., 389, T, 0.06-0.1, -, disturbed (garden), sandy

loam

Glinus

bainesii (Oliv.) Pax, 1319, T, 0.3, aquatic, seepage below dam, sandy,

stony

Hvpertelis

salsoloides (Burch.) Adamson var. salsoloides, 364, H, 0.2, Combretum

woodland, dryland, sandy loam

Corbichonia

decumbens (Forssk.) Exell, 377, 1069, Ch, 0.03-0.05, Colophospermum

woodland, dryland, disturbed (roadside), sandy gravel, clay

PORTULACACEAE

Talinum

caffrum ( Thunb .) Eckl. & Zeyh., 1252, G, 0.2, open parkland, seepline

complex, clayey loam

Portulacaria

afra Jacq., §, N, 1 .7, koppie, rock outcrop, sandy, gravelly

Portulaca

oleracea L. *, 1259, Ch, 0.06, -, disturbed (garden), sandy loam

pilosa L, 338, 1583. H, 0.04-0.08, Combretum woodland, koppie, rock

outcrop, disturbed (roadside), shallow, stony, sandy loam

quadrifida L., 1078, T, 0.1, open parkland, seepline complex, sandy clay

MENISPERMACEAE

Cocculus

hirsutus (L. ) Diels *, 1620, L, 2.0, riverine, river bank, shallow loam on

quartzite

Tinospora

fragosaf/. Verd.) I. Verd. & Troupin

(= Desmonemafragosum I. Verd.), §, L, 5.0, koppie, rock outcrop,

sandy

ANNONACEAE

Hexalobus

monopetalus (A. Rich.) Engl. & Diels var. monopetalus, 960, 1627, 1682,

Mi, 2. 0-4.0, koppie, rock outcrop, shallow gravel, sandy loam

PAPAVERACEAE

Argemone

mexicana L.*, 1217, T, 0.6, -, disturbed (old lands, roadside ), sandy loam

ochroleuca Sweet subsp. ochroleuca *

(=A. subfusiformis G. B. Ownbey),iS6,T,0.6, riverine, disturbed

(old lands, roadside), river sand, sandy loam

BRASSICACEAE

Lepidium

africanum (Burnt, f.) DC. subsp. africanum, 1080, T, 0.3, -, disturbed

(garden), sandy loam

CAPPARACEAE

Cleome

angustifolia Forssk. subsp. petersiana (Klotzsch ex Sond.) Kers, 336, T,

0.7, Combretum woodland, dryland, gravelly loam

gynandra L., §, T, 0.5, -, disturbed (old lands), sandy loam

hirta (Klotzsch) Oliv., 1019, T, 0.6-1 .0, riverine, river bank, alluvial sand

maculata (Sond.) Szyszyl.. 1592, T, 0.2-0.3, mixed woodland, dryland,

trampled and overgrazed areas, shallow, sandy

monophylla L., 392, T, 0.2-0. 3, Combretum woodland, dryland, sandy

loam

Capparis

tomentosa Lam., §, +, L, 1.5-10.0, riverine, seasonal stream, river &

stream banks, alluvial sand, loam, clay loam

Boscia

albitrunca (Burch.) Gilg & Gilg-Ben. var. albitrunca

(= Boscia albitrunca (Burch.) Gilg & Gilg-Ben.), 976, Me,

4. 0-6.0, koppie, rock outcrop, stony loam

Maerua

juncea Pax subsp. crustata (Wild) Wild, 1687, 1716, N, 1. 5-2.0, open

parkland, dryland, alluvial sand, sandy loam

parvifolia Pax, 339, 1003, N, 0.6-0.8, Combretum woodland, dryland,

seasonal stream bank, sandy loam, loam

CRASSULACEAE

Cotyledon

barbeyi Schweinf. ex Baker, 1175, H, 0.6, riverine, quartzite cliffs, shal-

low sandy gravel

Kalanchoe

lanceolata (Forssk.) Pers., §, H, 0.6, open parkland, seepline complex,

clayey loam

longiflora Schltr.exJ.M. Wood, 1033, 1173, H, 1.0, koppie, riverine, rock

outcrop, quartzite cliffs, sandy gravel

paniculata Harv., 1038, 1174, 1194, H, 1. 5-2.0, koppie, open parkland,

mixed woodland, rock outcrop, seepline complex, erosion areas,

shallow gravel, sandy loam, clayey loam

rotundifolia (Haw.) Haw., 1480, H, 0.3-0. 5, open Acacia woodland,

seepline complex, sandy loam

Crassula

expansa Dryand. subsp. fragilis (Baker) Toelken, 1475, H, 0.3, open

parkland, seepline complex, sandy loam

VAHLIACEAE

Vahlia

capensis (L.f.) Tlumb. subsp. capensis, 365, H, 0.2, Combretum wood-

land, dryland, sandy loam

capensis (L.f.) Thunb. subsp. vulgaris Bridson var. longiflora (Gand.)

Bridson, 1263, H, 0.2, -, disturbed (de-bushed), sandy loam

NYMPHAEACEAE

Nymphaea

lotus L., 1092, G, -, aquatic, seasonal pan, clay

nouchali Burm.f. var. caerulea (Savigny) Verde.

(= N. caerulea Savigny), 1108, G, -, aquatic, seasonal pan, clay

(= TV. capensis L.), 1107, G, -, aquatic, seasonal pan, clay

FABACEAE

Albizia

anthelmintica (A. Rich.) Brongn., 1619, Mi, 3.0-4.0, riverine, river bank,

sandy loam

brevifolia Schinz, 1412, 1590, Mi, 5.0, koppie, rock outcrop, -

forbesii Benth., 1171, Me, 20.0, open parkland, riverine, seasonal stream,

seepline complex, river bank, sandy clayey loam

Bothalia 24,1 (1994)

47

harveyi E. Fourn., 261, 904, Me, 5,0-8. 0, Acacia & mixed woodland,

dryland, seepline complex, sandy loam, loam

Acacia

erubescens Wei w. ex Oliv., 1589, 1622, Me, 5.0-6.0, Combretum-Com-

miphora woodland, riverine, open parkland, dryland, river bank,

seepline complex, shallow gravel, shallow stony loam

exuvialis /. Vercl., 1396, Mi, 1. 5-3.0, mixed woodland, dryland, shallow

stony loam

gerrardii Benth. var. gerrardii, 1352, 1353, 1679, Mi, 3. 0-4.0, mixed

woodland, open parkland, riverine, dryland, seepline complex,

stream bank, sandy loam, sandy clayey loam

grandicornuta Gerstner, +, Me, -, -, -

mellifera ( Vahl) Benth. subsp. detinens (Burch. ) Brenan, 1184, 1715, Me,

6.0-10.0, riverine, mixed woodland, dryland, river bank, river

bed, alluvial sand, clayey loam

nigrescens Oliv., §, +, Me, -, -, -

nilotica(L. I Willd. exDelile subsp. kraussiana (Benth.) Brenan, §, +, Me,

robusta Burch, subsp. clavigera (E. Mey.) Brenan, 1195, Me, 10.0,

riverine, river bank, alluvial sand

schweinfurthii Brenan & Exell var. schweinfurthii, §, L, 10.0, riverine,

river bank, alluvial sand

Senegal (L.) Willd. var. leiorhachis Brenan, §, Me, 6.0, Combretum

woodland, dryland, stony sandy loam

Senegal (L.) Willd. var. rostrata Breitan, 598, Mi, 1 .8—4.5, Combretum -

Acacia woodland, open parkland, dryland, seepline complex,

loam, clayey loam

tortilis (Forssk.) Hayne subsp. heteracantha (Burch.) Brenan, 914, Me,

7.0, riverine. Acacia woodland, river bank, dryland, disturbed

(old lands & cattle kraals), alluvial sand, stony loam, loam

Dichrostachys

cinerea (L.) Wight & Ant. subsp. africana Brenan & Brummitt var.

africana, 388, Mi, 1.8, woodland, shrubveld, dryland, sandy loam

Colophospermum

mopane (J. Kirk, ex Benth.) J. Kirk ex J. Leonard, §, +, Me, 2.0-15.0,

woodland & shrubveld dominant, -, sandy loam, clayey loam

Schotia

brachypetala Sond., §, +, Me. 10.0, riverine, termitarium, river bank,

dryland, sandy clayey loam

capitata Bolle, §, N, 0.5-1 .5, open parkland, seepline complex, clayey

loam

Bauhinia

galpinii N.E. Br., §, +, Mi, 5.0, riverine, river bank, alluvial sand, loam

Cassia

abbreviata Oliv. subsp. beareana (Holmes) Brenan , §, +, Me, 3.0-6.0,

Acacia & mixed woodland, dryland, sandy loam

Chamaecrista

mimosoides (L.) Greene

(= Cassia mimosoides L.), §, Ch, 0.3, Combretum & mixed

woodland, dryland, sandy loam

stricta E. Me v.

(= Cassia quarrei (Ghesq.) Steyaert), 1104, Ch, 0.3, Combretum

woodland, dryland, sandy loam

Senna

italica subsp. arachoides (Burch.) Lock

(= Cassia italica (Mill.) Lam. ex F.W. Andrews subsp. arachoides

(Burch.) Brenan), 366, H, 0.2, Colophospermum-Combretum

woodland, dryland, sandy loam

occidentals (L.) Link *

(= Cassia occidentalis L. ), §, Ch, 1.1, riverine, river bank, sandy

loam

petersiana (Bolle) Lock

(= Cassia petersiana Bolle), §, N, 1.5, roadside, sandy loam

Pterolobium

stellatum (Forssk.) Brenan, 1181 , Mi, 3.0, riverine, river bank, alluvial

sand

Peltophorum

africanum Sond., §, +, Me, 6.0, mixed woodland, dryland, sandy loam

Bolusanthus

speciosus (Bolus) Harms §, +, Me, 8.0, Acacia & mixed woodland,

riverine, seasonal stream, dryland, river & stream bank, sandy

loam

Crotalaria

burkeana Benth., 1089, Ch, 0.4, disturbed (old lands, roadside), sandy

loam

labumifolia L. subsp. australis (Baker f.) Polhill, 1314, Ch, 0.6, -,

disturbed (roadside), sandy loam

monteiroi Taub. ex Baker f. var. galpinii Burtt Davy ex 1. Verd.

(- C. rigidula Baker f.), §, Ch, 1 .0. seasonal stream, stream bank,

sandy loam

schinzii Baker f, 277, 308, 1134, H, 0. 2-1.0, Combretum woodland,

dryland, disturbed (roadside), sandy loam

virgulata Klotzsch subsp. grantiana (Harv. ) Polhill, 575, 1105, H, 0.2-0.3,

Combretum woodland, dryland, disturbed (roadside), sandy

loam, loam

Indigofera

bainesii Baker, 964, H, 0.2, -, disturbed (roadside), stony loam

costata Gud I . & Perr. subsp. macra (E. Mey.) J.B. Gillett

(= /. macra E. Mey.), §, H, 0.5, seasonal stream, stream bank,

sandy loam

fdipes Benth. ex Harv., 1017, 1321, T, 0.5, Combretum woodland,

dryland, disturbed (roadside), gravel, sandy or stony loam

galpinii N.E. Br., 1511, T, 0.3, -, disturbed (garden), stony or sandy loam

lydenburgensis N.E. Br., 1473, Ch, 0.5-0. 6, mixed woodland, open

parkland, seepline complex, sandy loam

Tephrosia

longipes Meisn. subsp. longipes, 881, 1071 , Ch, 0.6-1 .0, riverine, river

bed, disturbed (roadside), river sand, stony loam

polystachya E. Mey. var. latifoiia Harv., §, Ch, 0.4, -, -, -

rhodesica Baker f. var. rhodesica, 982, Ch, 0.4, -, disturbed (de- bushed),

sandy clayey loam

villosa (L.) Pers. subsp. ehrenbergiana(5c/twe/n/ ) Bmmmittv ar. daviesii

Brummitt, 1130, Ch, 1.0, -, disturbed (roadside), gravel, sandy

clay

Mundulea

sericea (Willd.) A. Chev., §, +, Mi, 2.0. open parkland, upper edge of

seepline complex, sandy clayey loam

Sesbania

bispinosa (Jacq.) W. Wight var. bispinosa *, 921, 1135, Mi, 2.0-2. 5,

seasonal stream, stream bed, river sand, silt

sesban (L. ) Merr. subsp. sesban var. nubica Chiov., 1493, H, 2.0, riverine,

river bank, alluvial sand

Ormocarpum

trichocarpum (Taub.) Engl., 524, Mi, 1. 6-2.4. mixed woodland, open

parkland, dryland, seepline complex, stony loam, sandy clayey

loam

Aeschynomene

indicaL., 1166, T, 1.0, seasonal stream, stream bank, alluvial sand

Stylosanthes

fruticosa (Retz.) Alston, 1147, H, 0.3, open parkland, seepline complex,

sandy clayey loam

Zornia

glochidiata DC.. 1124, T, 0.3, -, disturbed (roadside), sandy loam

Dalbergia

melanoxylon Guill. & Perr., 1402, Mi, 3.0, open parkland, riverine, upper

edge of seepline complex, alluvial sand, sandy clayey loam

Lonchocarpus

capassa Rolfe, 341 , Me, 10.0, riverine, woodland, river bank, dryland,

alluvial sand, loam

Xanthocercis

zambesiaca (Baker) Dumaz-le-Grand, 579, Me, 12.0-15.0, riverine,

Combretum-Commiphora woodland, river bank, termitarium,

alluvial sand, clayey loam

Abrus

precatorius L. subsp. africanus Verde., 1115, L, 2.0. riverine, seasonal

stream, river & stream bank, alluvial sand, sandy loam

Erythrina

humeana Spreng., §, N, 1 .5-2.0, riverine, seasonal stream, river & stream

bank, alluvial sand, sandy loam

Rhynchosia

caribaea (Jacq.) DC., 886, L, 1.0, Colophospermum woodland, dryland,

sandy clay

minima (L.) DC. var. prostrata (Harv.) Meikle. 382, H, 0.6, Combretum

woodland, dryland, disturbed (roadside), gravel

sp., 276, -, 0.6, Combretum woodland, dryland, sandy

Eriosema

psoraleoides (Lam.) G. Don, 884, Ch, 2.0, riverine, riverbed, river sand

Vigna

unguiculata (L.) Walp. subsp. unguiculata, 1112, 1643, L. 1. 0-2.0. river-

ine, mixed woodland, river bank, dryland, alluvial sand, sandy

loam

Dolichos

trilobus L. subsp. transvaalicus Verde., 1320, L, 0.6, aquatic, seepage

below dam, deep shade, clayey loam

Macrotyloma

axillare (E. Mey.) Verde, var. axillare, 1123, L, 0.5, Combretum woodland,

dryland, sandy or stony loam

Decorsea

schlechteri (Harms) Verde., 1355, L, 2.0, seasonal stream, woodland,

stream bank, dryland, alluvial & river sand, sandy loam

48

Bothalia 24,1 (1994)

GERANIACEAE

Monsonia

angustifoliaE. Mey. ex A. Rich.. 1077. Ch, 1.0, mixed woodland, dryland,

sandy loam

burkeana Planch, ex Harv., 1059. T, 0.3, Combretum woodland, dryland,

sandy loam

glaucaR. Knuth. 1020, 1053, H, 0.2-0.4, Combretum woodland, koppie,

dryland, rock outcrop, stony loam

OXALIDACEAE

Oxalis

corniculata L. *, 1207. H, 0.08, -, disturbed (garden lawn), sandy loam

latifolia Humb.. Bonpl. & Kunth *, 1021, T, 0.2, mixed woodland, deep

shade in dryland, loam

ZYGOPHYLLACEAE

Tribulus

terrestris L., 328, T, 0.2, riverine, river bank, disturbed (roadside, de-

bushed, trampled), sandy loam, loam

zeyheri Sond. subsp. zeyheri, 1004, T, 0.2, -, disturbed (roadside), sandy

loam

BALANITACEAE

Balanites

maughamii Sprague, 1405, Me, 12.0, open woodland, dryland, alluvial

sand, sandy loam

RUTACEAE

Zanthoxylum

capense (Tlmnb.) Harv., §, Mi, 1.2, mixed woodland, dryland thicket,

stony loam

Teclea

pilosa (Engl.) 1. Verd., 1048, N, 1.0, riverine, river bank, sandy clayey

loam

SIMAROUBACEAE

Kirkia

wilmsii Engl., 1413, Me, 6.0, koppie, rock outcrop, shallow sandy

BURSERACEAE

Commiphora

africana (A. Rich.) Engl., 340, 576, 974, Mi, 1. 2-3.0, koppie, Combretum

woodland, rock outcrop, dryland, stony loam, sandy loam

glandulosa Schinz, §, Mi, 4.0, Combretum woodland, dryland, shallow

gravelly

merkeri Engl., 342, Me, 1.2, riverine, river bank, stony loam

mollis (Oliv.) Engl., 957, Me, 5.0, Combretum-Commiphora woodland,

mixed woodland, dryland, shallow stony

pyracanthoides Engl., 993, JdK 10, N, 1 .5, koppie, rock outcrop, sandy

PTAEROXYLACEAE

Ptaeroxylon

obliquum (Thunb.) Radik., §, Mi, 3.0, koppie, rock outcrop, sandy,

gravelly

MELIACEAE

Turraea

obtusifolia Hochst., 887, 922, 1086, Ch, 0.7- 1.0, Colophospermwn,

Combretum, & mixed woodland, dryland, stony, sandy loam,

sandy clay loam

Trichilia

emetica Vahl, §, Me, 10.0, riverine, river bank, alluvial sand

MALPIGHI ACEAE

Triaspis

hypericoides (DC.) Burch, subsp. nelsonii (Oliv.) Immelman, 1349

(= T. nelsonii Oliv.), 773, N, 0.5-1. 5, Combretum & mixed

woodland, dryland, rocky, stony

Sphedamnocarpus

pruriens (Juss.) Szyszyl. subsp. pruriens, 885, 1110, L, 1. 5-2.0, Col-

ophospermum, Combretum, and mixed woodland, dryland, sandy

loam, sandy clayey loam

EUPHORBI ACEAE

Securinega

virosa (Roxb. ex Willd.) Pax & K. Hoffm., 333, N, 1. 5-2.0, riverine,

seasonal stream, roadside, open parkland, river bank, stream

bank, seepline complex, sandy loam, loam, clay loam

Phyllanthus

maderaspatensis L., 347, Ch, 0.5, Combretum woodland, dryland, sandy

pinnatus (Wight) G.L. Webster

(= P. kirkianus Miill.Arg.), 891, 1214, N, 1.5, Colophospermwn

woodland, dryland, stony loam

reticulatus Poir., 1675, Mi, 1. 0-3.0, riverine, seasonal stream, river bank,

stream bank, alluvial sand, sandy loam

Bridelia

cathartica Bertol.f, 1129, 1403, Mi, 2.0-3. 0, riverine, river bank, alluvial

sand

micrantha (Hochst.) Baill., §, +. Mi, 5.0. riverine, river bank, alluvial

sand

mollis Hutch., §, +, Me, 6.0, koppie, rock outcrop, sandy

Croton

megalobotrysM;7//.Arg., 1404, Me, 4.0, riverine, river bank, alluvial sand

Acalypha

indica L.. 940, 1005, 1429, 1585, T, 0.4— 0.5, -, disturbed (roadside,

garden), gravel, sandy loam

villicaulis Hochst.

(=A. petiolaris Hochst.), §, H. 0.3-0. 5, riverine, seasonal stream,

river & stream bank, alluvial sand, sandy loam

Tragia

dioica Sond., 772, H, 0.3, Combretum & mixed woodland, dryland,

disturbed (overgrazed, trampled), sandy

rupestris Sond., 1068, 1113, L, 0.6-1. 5, riverine, seasonal stream, river

& stream bank, alluvial sand, sandy loam

Dalechampia

galpinii Pax, 1116, L, 1.0, mixed woodland, dryland, stony loam

Jatropha

curcas L. *, 1610, Ch, 0.7, koppie, rock outcrop, sandy loam

spicata Pax

(= J. messinica E. A. Bruce), 958, Ch, 0.7, koppie, rock outcrop,

sandy or stony loam

zeyheri Sond. var. zeyheri, 259, 621, 1262, H, 0.2-0.3, Combretum

woodland, dryland, sandy loam

Spirostachys

africana Sond., §, +, Me, 4.0-10.0, riverine, seasonal stream, open

parkland, river & stream bank, seepline complex, clayey loam,

clay

Euphorbia

cooperi N.E. Br. ex A. Berger var. cooperi, §, Mi, 3.0, Combretum-Com-

miphora woodland, dryland, stony sandy loam

guerichiana Pax, 925, N, 1.3, mixed woodland, dryland, stony

schinzii Pax, §, Ch, 0.5, Combretum-Commiphora woodland, dryland,

shallow stony sandy loam

tirucalli L., §, L, 6.0, riverine, rocky river bank (cliff), shallow stony

Chamaesyce

hirta (L. ) Millsp.

(= Euphorbia hirta L.), 1008, H, 0. 1-0.3, -, disturbed (garden,

roadside), sandy loam

neopolycnemoides (Pax & K. Hofftn.) Koutnik

(= Euphorbia neopolycnemoides Pax & K. Hoffm. ), 1094, H, 0.3,

koppie, rock outcrop, loam

ANACARDI ACEAE

Selerocarya

birrea (A. Rich.) Hochst. subsp. caffra (Sond.) Kokwaro

(= S. caffra Sond.), §, +, Me, 3.0-15.0, Acacia & Combretum

woodland, dryland, sandy loam

Lannea

schweinfurthii (Engl.) Engl. var. stuhlmanni (Engl.) Kokwaro

(= L. stuhlmannii (Engl. ) Engl.), §, Me, 8.0, Combretum & mixed

woodland, dryland, sandy loam

Ozoroa

paniculosa (Sond.) R. Fern. <£ A. Fern. var. paniculosa, 895, 1332, JdK

8, Mi, 2. 0-4.0, mixed woodland, dryland, stony loam

Rhus

gueinzii Sond, 980, Mi, 2.0-4. 0, riverine, seasonal stream, river & stream

bank, alluvial sand, sandy loam

pentheri Zahlbr., 1140, Mi, 1.0, -, termitarium, sandy clay

POLYGALACEAE

Polygala

sphenoptera F resen., 1146, H, 0.5, Combretum woodland, dryland, sandy

loam

CELASTRACEAE

Maytenus

heterophylla (Eckl. & Zeyh.) N. Robson, 893, 1057, Mi, 2.0-3 .0, riverine,

seasonal stream, open parkland, mixed woodland, river & stream

Bothalia 24,1 (1994)

49

bank, seepline complex, dryland, alluvial sand, sandy loam,

sandy clay

senegalensis (Lam.) Exell, §, +, JdK 21. Mi, 1. 0-2.5, riverine, seasonal

stream, river & stream bank, alluvial sand, sandy loam

Cassine

aethiopica Thunb. §, +, Mi, -

burkeana (Sond.) Kuntze. 1044, Mi, 2.0-3. 0, riverine, seasonal stream,

river & stream bank, sandy loam

transvaalensis (Burn Davy) Codd, 1186, Me, 6. 0-8.0, mixed woodland,

koppie, dryland, rock outcrop, sandy loam

Hippocratea

longipetiolata Oliv., §, +, L, 3.0, koppie, seasonal stream, rock outcrop,

stream bank, termitarium, shallow stony & sandy, sandy clay

SAPINDACEAE

Cardiospermum

corindum L., 1015, 1064, L, 1. 0-2.0, Combretum & mixed woodland,

dryland, stony loam

Pappea

capensis Eckl. & Zeyh., §, +, Mi, 4.0, koppie, open parkland, rock

outcrop, seepline complex, sandy, gravelly, clayey loam

Dodonaea

angustifolia L.f.

(= D. viscosa Jacq. var. angustifolia Benth.), 1628, Mi, 3. 0^1.0,

riverine, river bed, river sand

RHAMNACEAE

Ziziphus

mucronata Willd. subsp. mucronata, §, +, Me, 6.0, mixed woodland,

dryland, sandy loam

Berchemia

discolor (Klotzsch) Hemsl., 961. 1036, Me, 3. 0-6.0, koppie, riverine,

rock outcrop, river bank, alluvial sand, sandy

zeyheri (Sond.) Grubov, +, Me, -, -, -, -

VITACEAE

Rhoicissus

tridentata (L.f.) Wild & R.B. Drumm. subsp. cuneifolia (Eckl. & Zeyh.)

N.R. Urton, 1158, L, 3.0, riverine, river bank, alluvial sand

Cissus

comifolia (Baker) Planch.

(= C. lonicerifolia C.A. Sm.), §, N, 1.2, Combretum & mixed

woodland, dryland, sandy loam

quadrangularis L., §, L, 4.0. Combretum & mixed woodland, dryland,

sandy loam

rotundifolia (Forssk.) Vahl. 1032. L, 3.0, koppie, rock outcrop, sandy

Cyphostemma

puberulum (C.A. Sm.) Wild & R.B. Drumm., 1111. 1114, L. 1.5, mixed

woodland, riverine, dryland, river bank, alluvial sand, sandy

loam, clayey loam

schlechteri (Gilg & M. Brandt) Desc. ex Wild & R.B. Drumm.. 1258, L,

- , -, termitarium, red sandy loam

TILIACEAE

Corchorus

asplenifolius Burch., 1639, H, 0.2, Combretum woodland, dryland, sandy

loam

confusus Wild, 281, H, 0.2, Combretum woodland, dryland, sandy

Grewia

bi color Juss., 310, 501, 1398. PJM 02, 03. 04, JdK 28, 29, Mi, 1. 2-3.0,

Combretum & Combretum-Commiphora woodland, dryland,

shallow gravel, stony loam, sandy loam

flavaDC., 279, Mi, 1.5, Combretum woodland, dryland, sandy

flavescens Juss. var. flavescens, JdK 25, 26, §, +. Mi, 2.0- 3.0, Com-

bretum-Commiphora woodland, dryland, shallow gravel

hexamita Burret, 252, Mi, 1. 5-3.0, Combretum woodland, dryland, stony

loam

monticola Sond., +, Mi, -, -, -, -

subspathulata N.E. Br.. 1397, PJM 01, JdK 27, Mi. 2.0, Combretum &

Combretum-Commiphora woodland. -, shallow gravel, sandy

loam

sulcata Mast., 1026, 1414, Mi. 1.0-1. 5, riverine, river bank, alluvial sand

villosa Willd., 999, N, 0.7, Acacia & mixed woodland, dryland, termitar-

ium, sandy clayey loam

Triumfetta

rhomboideayao/., 1640. Ch, 1.5, riverine, river bank, alluvial sand

austro-africanum Hochr., 275, 1055, H, 0.3-0.4, Combretum & mixed

woodland, dryland, sandy loam

englerianum Ulbr., 374, H, 0.5, -, termitarium, loam

ramosum (Cav.) Guill. &Perr., 1641, H, 1.5, riverine, river bank, alluvial

sand

Sida

chrysantha Ulbr., 282. 1054, 1133, 1265, H, 0.3-0.5, Combretum &

mixed woodland, dryland, disturbed (roadside), gravelly loam,

sandy loam, sandy clayey loam

cordifolia L., 951 , 1144, H, 0.5-0.6, koppie, rock outcrop, disturbed (old

lands), course gravel, sandy loam

rhombifolia L., 1334, H, 0.4, riverine, river bed, river sand

Pavonia

burchellii (DC.) R.A. Dyer, 619, 906. 1051. 1083, H, 0.3-0.6, Acacia,

Combretum, & mixed woodland, dryland, termitarium, sandy

loam, loam, clayey loam

Hibiscus

calyphyllus Cav., 1081 , H, 0.4, open parkland, seepline complex, sandy

clay

cannabinusL., 1018, 1163, H, 1.7, seasonal stream, stream bed, river sand

coddii Exell, 1614. H, 1.0, koppie, rock outcrop, shallow sandy

engleri K. Schum., 354, 1367, H, 0. 7-1.0, koppie, riverine, rock outcrop,

river bank, shallow sandy loam

meyeri Harv. subsp. meyeri, 1261, H. 1.0, Combretum woodland, dry-

land, sandy loam

micranthus Lf, 251 . H, 0.9, Combretum woodland, dryland, stony loam

praeteritus R.A. Dyer, 385, 874, 1099, H, 1. 0-2.0, riverine, seasonal

stream, river & stream bank, sandy loam, loam

pusillus Thunb., 1087, H. 0.3, Combretum & mixed woodland, dryland,

sandy clayey loam

schinzii Giirke, 1149, H, 0.1, -, disturbed (old lands, roadside), sandy

loam

sidiformis BailL, 1340, H, 0.3, -, disturbed (roadside), gravel, sandy loam

upingtoniae Giirke, 1085, L, 0.4, mixed woodland, dryland, sandy clayey

loam

Gossypium

herbaceum L. subsp. africanum ( Watt) Vollesen

(= var. africanum (Watt) Hutch. & R.L.M. Ghose), 873, N, 1.5,

mixed woodland, dryland, sandy loam

BOMBACACEAE

Adansonia

digitataZ.., §, Me, 15.0, Acacia woodland, dryland, sandy loam

STERCULIACEAE

Melhania

acuminata Mast. var. acuminata, §, Ch, 0.5-0. 6, Colophospermum wood-

land, disturbed (overgrazed & trampled), sandy, gravel

forbesii Planch, ex Mast., §, +. Ch, 0.6. Combretum, Acacia, & mixed

woodland, riverine, dryland, disturbed (overgrazed & trampled),

river bank, sandy loam

prostrata DC., 379, H, 0.3, -, disturbed (roadside), gravel

rehmannii Szyszyl. 1714, Ch, 0.3-0.7, Combretum woodland, dryland,

shallow gravel

Dombeya

rotundifolia (Hochst.) Planch, var. rotundifolia, §, Mi, 3.0, koppie,

riverine, seasonal stream, rock outcrop, river & stream bank,

sandy loam

Hermannia

glanduligera K. Schum.

(= H. viscosa sensu Burtt Davy non Hiem), 1400, H, 0.6, -,

disturbed (roadside), dryland, sandy loam

modesta (Ehrenb.) Mast., 278. 1125, T. 0. 2-0.5, Combretum woodland,

dryland, disturbed (roadside), stony, sandy loam

Waltheria

indica L.. 605, 1126, H, 0.4— 0.5, Combretum woodland, dryland, dis-

turbed (roadside), stony, sandy loam

Sterculia

rogersii N.E. Br., 955, Mi, 3.0, koppie. rock outcrop, shallow stony loam

OCHNACEAE

Ochna

inermis (Forssk.) Schweinf, 991, 1045, Mi, 2.0, koppie, rock outcrop,

stony loam

pretoriensis E. Phillips, +, Mi, -, -, -, -

MALVACEAE

Abutilon

angulatum ( Guill. & Perr. ) Mast. var. angulatum, §, T, 1 .2, riverine, river

bank, sandy, loam

CLUSIACEAE

Garcinia

livingstonei T. Anderson, 1415, Me, 15.0, riverine, river bank, alluvial

sand

50

Bothalia 24,1 (1994)

ELATINACEAE

Bergia

decumbens Planch, ex Harv., 1615 , H, 0. 1-0.2, aquatic, dam shores, clay

PLUMBAGINACEAE

Plumbago

zeylanica L., §, H, 1.2, riverine, river bank, sandy loam

VIOLACEAE

Hybanthus

capensis (Thunb.) Engl., 255, H, 0.3, Combretum woodland, dryland,

loam

enneaspermus (L.) F. Muell. *, 1082, 1255, T, 0. 1 , Combretum woodland,

disturbed (de-bushed), dryland, sandy loam, sandy clayey loam

TURNERACEAE

Tricliceras

glanduliferunt (Klotzsch) R. Fern.

(= Wormskioldia glandulifera Klotzsch ), 1339, T, 0.3, -, disturbed

(roadside), sandy loam

laceratum (Oberm.) Oberm.

(= Wormskioldia lacerata Oberm.), 620, T, 0.2, Combretum

woodland, -, sandy, stony

longipedunculatum ( Mast .) R. Fern. var. longipedunculatum

(= Wormskioldia longipedunculata Mast.), §, T, 0.3, -, disturbed

(roadside), gravel, sandy loam

SAPOTACEAE

Manilkara

mochisia (Baker) Dubard, 362, Mi, 2. 5-5.0, riverine, open parkland,

river bank, seepline complex, sandy loam, clayey loam

EBENACEAE

Euclea

divinorum Hiern, 1042, Mi, 2.0-3. 0, seasonal stream, open parkland,

stream bank, seepline complex, sandy-loam, sandy clayey loam

natalensis A. DC. subsp. natalensis, 1046, Mi, 2.0, riverine, seasonal

stream, river & stream bank, alluvial sand, sandy loam

undulata Thunb. var. undulata, 375, 1331, N, 1. 0-2.0, Combretum &

mixed woodland, dryland, sandy loam, loam

Diospyros

mespiliformis Hochst. ex A. DC., 399, Me, 15.0-20.0, riverine, seasonal

stream, river & stream bank, termitarium, sandy loam, clayey

loam

PASSIFLORACEAE

Adenia

digitata (Harv.) Engl., 1507, L, 1.5, Combretum woodland, dryland,

shallow stony loam

hastata (Harv.) Schinz var. hastata, 1354, L, 2.0, Combretum woodland,

dryland, red sandy loam

OLEACEAE

Jasminum

fluminense Veil., 1063, L, 0.6, seasonal stream, stream bank, sandy loam

multipartitum Hochst., 344, L, 4.0, Combretum woodland, dryland,

gravel

sp. cf. J. stenolobum Rolfe, 1534, N, -, koppie, rock outcrop, gravel

THYMELAEACEAE

Gnidia

rubescens B. Peterson. 319, H, 0.4, Combretum woodland, dryland,

disturbed (roadside), gravel

LYTHRACEAE

Ammannia

senegalensis Lam. ex Poir, 1164, T, 0.6, seasonal stream, stream bed,

damp river sand

COMBRETACEAE

Combretum

apiculatum Solid, subsp. apiculatum, §, +, Mi, 3.0-5. 0, woodland domi-

nant, dryland, sandy loam

erythrophyllum (Burch.) Sond., 1417, Me, 5.0, riverine, river bank, river

bed, alluvial & river sand

hereroense Schinz, 343, Mi, 3.0-5.0, Combretum woodland, riverine,

seasonal stream, dryland, river & stream bank, gravel, alluvial

sand, sandy loam

imberbe Wawra, §, +, Me, 10.0, mixed woodland, riverine & seasonal

stream, dryland, river & stream bank, clayey loam, clay

microphyllum Klotzsch

(= C. paniculatum Vent, subsp. microphyllum (Klotzsch)

Wickens), §, L, 10.0, riverine, river bank, alluvial sand

mossambicense (Klotzsch) Engl., 323, Mi, 1.5-2. 5, riverine, Combretum

woodland, river bank, stony or sandy loam

zeyheri Sond., 262, Me, 4. 5-6.0, mixed woodland, dryland, loam

Terminalia

prunioides M.A. Lawson. 317, Me, 6.0, Combretum woodland, dryland,

shallow stony loam

sericea Burch, ex DC., §, +, Me, 6.0, Combretum woodland, open

parkland, upper edge of seepline complex, sandy loam

MYRTACEAE

Syzygium

guineense (Willd.) DC., §, Me. 10.0, riverine, river bed, river sand

ONAGRACEAE

Ludwigia

octovalvis ( Jacq.) P.H. Raven subsp. octovalvis, 1128, 1338, H, 1.0,

riverine, river bed, wet river sand

polycarpa Short & Peter ex Ton: & A. Gray, 1165, T, 0.9, seasonal stream,

stream bed, river sand

stolonifera (Guill. & Perr.) P.H. Raven, 1535, H, 0.1-0. 2, aquatic, river

bed, mud

AP1ACEAE

Steganotaenia

araliacea Hochst., 1613, Mi, 4.0, koppie, rock outcrop, loam

LOGANIACEAE

Strychnos

madagascariensis Poir., 981, JdK 5, Mi, 1 .5-3.5, Combretum woodland,

koppie, dryland, rock outcrop, gravel, stony loam, sandy

spinosa Lam., 1406, Mi, 3.0, open woodland, dryland, alluvial sand

Nuxia

oppositifolia (Hochst.) Benth., 1336, 1409, Mi, 3. 0-4.0, riverine, seasonal

stream, river & stream bed, river sand, sandy loam

GENTIANACEAE

Enicostema

hyssopil'olium (Willd.) I. Verd., 1079, H, 0.1, open parkland, seepline

complex, sandy clay

APOCYNACEAE

Carissa

bispinosafLJDey/i exBrenan subsp. bispinosa, §, +, N, 1.0-1 .7, riverine,

seasonal stream, river & stream bank, sandy loam, loam

Adenium

multiflorum Klotzsch

(= A. obesum (Forssk.) Roem. & Schult. var. multiflorum

(Klotzsch) Codd), §, +, N, 0.5-1. 2, koppie, open parkland, rock

outcrop, seepline complex, sandy, sandy clay loam

Pachypodium

saundersii N.E. Br., 962, N, 0.7-1. 2, koppie, rock outcrop, sandy loam

Strophanthus

gerrardii Stapf 1303, L, 7.0, riverine, seasonal stream, river & stream

bank, alluvial sand

PERIPLOCACEAE

Cryptolepis

obtusa N.E. Br., §, L, 1.3- 1.8, riverine, river bed (reed bed), river sand

Stomatostemma

monteiroae (Oliv.) N.E. Br, 1329, Ch, 0.4, koppie, rock outcrop, seepline

complex, loam, clay

Raphionacme

burkei N.E. Br., 1625, 1626, G, 0.2-0.3, Combretum woodland, dryland,

sandy loam

galpinii Schltr., 1593, G, 0. 1-0.2, Combretum woodland, dryland, sandy

loam

monteiroae (Oliv.) N.E. Br., §, G, -, -

sp., 1579, 1598, -, 1 .0, koppie, rock outcrop (dolerite), loam, humus-rich

loam

ASCLEPIADACEAE

Schizoglossum

sp., 1264, H, 0.2, Combretum-Acacia woodland, dryland, clay loam

Kanahia

laniflora (Forssk.) R. Br, 1377, H, 1.0, riverine, river bed river sand

Bothalia 24,1 (1994)

51

Stenostelma

capense Schltr.

(= Schizoglossum capense (Schltr.) H. Huber), 337, G.0.1, Com-

bretum woodland, dryland, shallow sandy

Asclepias

fruticosa L., §, H, 1.0, riverine, river bed. river sand

Sarcostemma

viminale (L.) R. Br., §, L, 4.0, mixed woodland, dryland, sandy loam

Secamone

parvifolia (Oliv.) Bullock, 1308 , L, 0.05, riverine, river bank, alluvial

sand

Ceropegia

mafekingensis (N.E. Br.) R.A. Dyer, 1536, G, 0.3, Colophospermum

shrubveld, dryland, sandy clayey loam

rendallii N.E. Br., 1170, G, 1.5, seasonal stream, stream bank thicket,

sandy gravelly loam

Stapelia

gettliffei Pott-Leend.. 1599, Ch, 0.2, Combretum-Commiphora wood-

land, dryland, shallow sandy & stony

gigantea N.E. Br., §, Ch, 0.3, open riverine shrubveld, riverbank, sandy

loam

Orbea

maculata (N.E. Br.) L.C. Leach, 1602, Ch, 0. 1-0.2, Combretum-Com-

miphora woodland, dryland, shallow sandy & stony

Pachycymbium

rogersii (L. Bolus) M.G. Gilbert

(= Caralluma rogersii (L. Bolus) E.A. Druce & R.A. Dyer), §,

Ch, 0.2-0.3, open parkland, seepline complex, clayey loam

Huernia

kirkii N.E. Br., §, Ch, 0.1, Colophospermum woodland, dryland, red

sandy loam

Pergularia

daemia (Forssk.) Chiov. var. daemia, 1161, L, 2.0, riverine, disturbed

(roadside), stony sandy loam

Fockea

angustifolia K. Schum., 1427, 1600, G, 0. 2-0.4, open Combretum &

Colophospermum woodland, dryland, shallow stony, clay loam,

clay

CONVOLVULACEAE

Evolvulus

alsinoides (L.) L. var. linifolius (L.) Baker, 348, H. 0.1, Combretum

woodland, dryland, sandy

Seddera

suffruticosafSc/n'nzj Hallierf. §, H, 0.4, mixed woodland, dryland, sandy

loam

Merremia

kentrocaulos (C.B. Clarke) Rendle, §, L, 2.0, Combretum woodland,

dryland, gravel, loam

palmata Hallierf., §, L, 1 .0, Colophospermum woodland, dryland, sandy

loam

tridentata (L.) Hallier f. subsp. angustifolia (Jacq.) Ooststr. var. an-

gustifolia, 979, H, 0.06, -. disturbed (roadside), sandy loam

Ipomoea

albivenia (Lindl.) Sweet, §, L, 6.0, riverine, seasonal stream, open park-

land, river & stream bank, seepline complex, clayey loam

arachnosperma Welw., §, L. 1.2, Combretum woodland, dryland, sandy

loam

bolusiana Schinz subsp. bolusiana, §, H, 0.1, Combretum & mixed

woodland, dryland, sandy loam

coptica (L.) Roth ex Roem. & Schult. var. coptica, 1139, H, 0.05, Com-

bretum woodland, disturbed (roadside), sandy

crassipes Hook., 346A. 723, 1154, H, 0.05-0.2, Combretum & mixed

woodland, dryland, disturbed (roadside), sandy, stony sandy loam

hochstetteri House, 1114A, L, 2.0, riverine, mixed woodland, riverbank,

dryland, alluvial sand, sandy loam

magnusiana Schinz var. eenii (Rendle) A. Meeuse, 1013, L, 0.1, open

mixed woodland, dryland, stony loam

magnusiana Schinz var. magnusiana, 1114, L, 2.0, riverine, mixed wood-

land, river bank, dryland, alluvial sand, sandy loam

obscura (L.) Ker Gawl. var. fragilis (Choisy) A. Meeuse, 1351, L, 0.1,

Combretum woodland, dryland, shallow stony

sinensis (Desr.) Choisy subsp. blepharosepala (Hochst. ex A. Rich.)

Verde., §, L, 1.3, Combretum woodland, dryland, sandy clayey

loam

BORAGINACEAE

Cordia

monoica Roxb., 1597

(= C. ovalis R. Br. ex DC.), 1119, 1152, JdK 16, Mi, 2.0-4.5, open

parkland, koppie, closed riverine woodland, seepline, termitar-

ium, rock outcrop, sandy humus-rich loam, sandy clay loam,

clayey loam

Ehretia

amoena Klotzsch, 722, 894, Mi, 1. 3-2.0, open woodland, seasonal

stream, dryland, stream bank, stony, sandy loam

obtusifolia Hochst. ex DC., 1233, N, 2.0, Combretum-Terminalia wood-

land, dryland, shallow stony

rigida (Thunb.) Druce, §, N, 0.2-0. 6, open parkland. Acacia woodland,

seepline complex, dryland, sandy clayey loam

Heliotropium

lineare (A. DC.) Giirke, §, T, 0.3, -, -, -

ovalifolium Forssk., §, H, -, -, -

steudneri Vatke, 274, H, 0. 2-0.4, Acacia woodland, disturbed (over-

grazed & trampled), sandy loam

strigosum Willd., 907 , T, 0.3, -, disturbed (roadside), gravel

VERBENACEAE

Verbena

bonariensis L. *, H, 1.4, grassland, vlei margin, clay loam

Lantana

rugosa Thunb., 1070, Ch, 0.6-0. 8, mixed woodland, dryland, sandy loam

Lippia

javanica (Bunn, f.) Spreng., §, N, 1.0-1. 3, riverine, river bank, alluvial

sand

Phyla

nodi fl ora (L.) Greene var. nodiflora, 1343, H, 0.06, aquatic, dam seepage,

sandy clay

Plexipus

hederaceus (Sond.) R. Fern. var. hederaceus

(= Chascanum hederaceum (Sond.) Moldenke var. hederaceum ),

770, H. 0.3, Combretum woodland, dryland, stony, sandy

incisus (H. Pearson) R. Fern.

(= Chascanum incisum (H. Pearson) Moldenke), §, H, 0.3, Col-

ophospermum woodland, dryland, sandy loam

pinnatifidus(L./) R. Fern. var. racemosus (Schinz ex Moldenke) R. Fern.

(= Chascanum pinnatifidum (L. f.) E. Mey. var. racemosum

Schinz ex Moldenke), 520, 1289, H, 0.4— 0.5, Colophospennum

woodland, disturbed (de-bushed), shallow stony loam, loam

Priva

meyenJaub. & Spach var. meyeri, §, H, 1.0, riverine, river bank, alluvial

sand

Duranta

erecta L. *

(= D. repens L.), §, Ch, 3.0, -, -, -

Clerodendrum

tematum Schinz var. tematum

(= C. ternatum Schinz var. lanceolatum (Giirke) Moldenke), §,

Ch, 0.3, koppie, rock outcrop, sandy, gravelly

LAMIACEAE

Leonotis

nepetifolia (L.) R. Br., 995. T, 1.5, seasonal stream, stream bank, clayey

loam

Leucas

neuflizeana Courbon, §, H. 0.3-0.4, Combretum woodland, dryland,

sandy loam

sexdentata Skan, 1000, 1106, H, 0.2-0.4, Combretum & open woodland,

dryland, disturbed (roadside), stony sandy loam

Tetradenia

riparia (Hochst.) Codd

(= Iboza riparia (Hochst.) N.E. Br.), §, N. 1. 2-2.0, riverine, river

bank, rocky sandy loam

Endostemon

tereticaulis (Poir.) M.R. Ashby, 1584, H, 0.3, Combretum woodland,

dryland, sandy loam

Plectranthus

tetensis (Baker) Agnew, 953, 1472, 1603. H, 0.05-0.12, koppie, Acacia

woodland, rock outcrop, seepline complex, sandy loam, loam,

clayey loam

xerophilus Codd, §, H, 2.0, koppie, rock outcrop, stony loam

Hoslundia

opposita Vahl, 1337, H, 2.0, seasonal stream, stream bank, sandy loam

Hemizvgia

elliottii (Baker) M. Ashby, 306, 623, 1327. 1335, H. 0.4-0. 5, Combretum

woodland, koppie, dryland, rock outcrop, gravelly, stony, sandy

petrensis (Hiem) M. Ashby, §, H, 0.3-0.4, Combretum woodland, dry-

land, gravelly, stony

Ocimum

canum Sims, §, H, 0.4, Combretum & Acacia woodland, dryland, sandy

loam

52

Bothalia 24,1 (1994)

Becium

filamentosum (Forssk.) Chiov.

(= B. knyanum (Vatke) N.E. Br. ex Broun & R.E. Massey), §, H,

0.5, Combretum & Colophospermum woodland, dryland, sandy

loam

Orthosiphon

suffrutescens (Thonn.) J.K. Morton, §, H, 0.5, Colophospermum wood-

land, riverine, dryland, river bank, alluvial sand, gravelly, sandy

loam

SOLANACEAE

Solarium

coccineum Jcicq., 1084, Ch, 0.4-0. 5, open parkland, seepline complex,

disturbed (overgrazed & trampled), sandy clay

incanum L„ 371 , Ch, 1.3, -, disturbed (old lands), sandy loam

panduriforme E. Mey., 250, Ch, 0.5, -, disturbed (overgrazed, trampled,

& roadside), gravel, sandy loam

Datura

stramonium L. *, §, T, 1.2, riverine, river bed, river sand

SCROPHULARIACEAE (PART A)

Aptosimum

lineare Marloth & Engl., 614, H, 0.2, Combretum woodland, dryland,

sandy loam

Peliostomum

leucorrhizum E. Mey. ex Benth. var. leucorrhizum

(= P. leucorrhizum E. Mey. ex Benth.), 523, H, 0.4, Combretum

woodland, dryland, stony loam

SCROPHULARIACEAE (PART B)

Buchnera

reducta Hiern, 1132, 1148, H, 0.3-0.4, open parkland, seepline complex,

sandy clayey loam, waterlogged clayey loam

Cycnium

adonense E. Mey. ex Benth. subsp. adonense, §, H, 0.1, Combretum

woodland, dryland, sandy loam

Striga

asiatica (L.)Kuntze, 1145, T, 0.2, open parkland, seepline complex, sandy

clayey loam

gesnerioides (Willd.) Vatke ex Engl., 1120, T, 0.4, Combretum woodland,

dryland, stony sandy loam

BIGNONIACEAE

Rhigozum

zambesiacum Baker, §, +, Mi, 1 .5-2.5, Acacia, Combretum, & mixed

woodland, dryland, rock outcrop, shallow loam

PEDALIACEAE

Pterodiscus

aurantiacus Welw., 372, H, 0.2, Combretum woodland, disturbed (old

lands), dryland, sandy loam

luridus Hook, f, 1061, H, 0.3, open parkland, seepline complex, sandy

clayey loam

Har pagophytu m

zeyheri Decne. subsp. zeyheri, 367, H, 0.1, Combretum woodland, dry-

land, sandy loam

Holubia

saccata Oliv., 883, 1582, T, 0. 3-0.4, Combretum woodland, disturbed

(overgrazed, roadside), stony sandy loam

Sesamum

alatum Thonn., 1016, T, 1.0, Combretum woodland, disturbed (over-

grazed, roadside), stony

Ceratotheca

triloba (Bernh.) Hook.f, 882, 1006, T, 1.0, Combretum & mixed wood-

land, disturbed (overgrazed & roadside), stony sandy loam

Dicerocaryum

eriocarpum (Decne.) Abels

(= D. zanguebarium (Lour.) Merr. subsp. zanguebarium), §, H,

0.1,-, disturbed (roadside, trampled river bank), sandy, gravelly

LENTIBULARIACEAE

Utricularia

stellaris L.f, 1024, T, 0.05, aquatic, seasonal pan, clay

ACANTHACEAE

Thunbergia

negiecta Solid. , 356, 1060, H, 0.2-0.6, riverine. Acacia & mixed wood-

land, dryland, sandy loam, loam

Dischoriste

rogersii S. Moore, 1155, Ch, 0.3, Acacia, Combretum, & mixed wood-

land, dryland, rock outcrop, disturbed (roadside), shallow sandy

loam, sandy clayey loam

Ruellia

patula Jacq., 345, 1050, 1256, 1260, H, 0. 1-0.3, Combretum woodland,

dryland, disturbed (overgrazed & de-bushed), stony loam, sandy

loam, sandy clayey loam

Crabbea

velutina S. Moore, 1065, H, 0.2, mixed woodland, dryland, sandy loam

Barleria

affinis C.B. Clarke, 972, H, 0.6, koppie, rock outcrop, stony

crossandriformis C.B. Clarke. 1023, H, 0.6, koppie, rock outcrop

elegans S. Moore ex C.B. Clarke, 1141, H, 0.6, seasonal stream, stream

bank, sandy loam

lancifolia T. Anderson, 956, 1093, 1131, H, 0.4-0. 7, koppie, mixed

woodland, rock outcrop, dryland, stony sandy loam, loam

senensis Klotzsch, 923, H, 0.7, mixed woodland, dryland, stony, shallow

Blepharis

aspera Oberm., 973, H, 0.3, koppie, rock outcrop, stony

Crossandra

mucronata Lindau, §, H, 0.3, open parkland, seepline complex, clayey

loam, clay

Hypoestes

aristata ( Vahl ) Sol. ex Roem. & Schult. var. aristata

(= H. verticillaris (L. f.) Sol. ex Roem. & Schult.), 1395, 1478,

H, 0.3, riverine, deep shade on river bank, alluvial sand, loam

Rhinacanthus

xerophilus A. Meeuse, 1333, H, 0.4— 0.5, Combretum woodland, shade in

dryland, sandy loam

Justicia

flava (Vahl) Vahl, 357, H, 0.3, Acacia woodland, dryland, loam

protracta (Nees) T. Anderson subsp. protracta, 1476, H, 0.3, riverine,

shade on river bank, alluvial sand

Monechma

divaricatum (Nees) C. B. Clarke, 952, H, 0.4, koppie, rock outcrop, loam

RUBIACEAE

Kohautia

caespitosa Schnizl ■ subsp. brachyloba (Sond.) D. Mantell

(= K. caespitosa Schnizl. var. delagoensis (Schinz) Bremek.),

394, T, 0.5, Combretum woodland, dryland, sandy loam

cynanchia DC.

(= K. omahekensis ( K. Krause ) Bremek. ), 395, T, 0.4, Combretum

& Colophospermum woodland, dryland, loam

virgata (Willd.) Bremek., 393, H, 0.4, Combretum woodland, dryland,

sandy loam

Agathisanthemum

bojeri Klotzsch subsp. bojeri

(= subsp. australe Bremek. var. australe ), 1001, H, 0.3, -, dis-

turbed (roadside), sandy loam

sp., 578, -, 0.3, Combretum woodland, dryland, sandy

Pentodon

pentandrus (Schumach. & Thonn.) Vatke var. minor Bremek., 1348, T,

0.3, riverine, river bed, damp river sand

Breonadia

salicina (Vahl) Hepper & J.R.I. Wood

(= Adina microcephala (Delile) Hiern var. galpinii (Oliv.) Hiern,

§, + , Me, 1 5.0-20.0, riverine, river bed, river sand amongst rocks

Gardenia

volkensii K. Schum. subsp. spatulifolia (Stapf & Hutch.) Verde.

(= G. spatulifolia Stapf & Hutch.), §, +, Mi, 3. 0-4.0, Acacia,

Combretum, & mixed woodland, dryland, shallow gravelly &

sandy

Tricalysia

junodii (Schinz) Brenan var. junodii

(= T. allenii (Stapf) Brenan var. australis (Schweick.) Brenan),

959, 994, N, 1.0-1. 5, koppie, Combretum-Commiphora wood-

land, rock outcrop, stony sandy loam

Sericanthe

andongensis (Hiern) Robbr. var. andongensis

(= Tricalysia andongensis Hiern), 1506, N, 1 .2-1.5, Combretum-

Commiphora woodland, dryland, shallow sandy loam

Vangueria

infausta Burch, subsp. infausta, §, +, Mi, 3.0, koppie, rock outcrop, sandy

Plectroniella

armata (K. Schum.) Robyns, 1189, Mi, 3.0-4.0, riverine, seasonal stream,

river bank, stream thicket, clayey loam

Bothalia 24,1 ( 1994)

53

Pyrostria

hystrix (Bremek.) Bridson

(= Dinocanthium hystrix Bremek.), 1191 , Mi, 2.0-3. 0, riverine,

seasonal stream, river & stream bank, sandy loam

Pavetta

catophylla K. Schum., 975, 1047 , N, 0.8-1 .0, koppie, Combretum wood-

land, rock outcrop, partial shade in dryland, stony loam, sandy

clayey loam

gardeniifoliaA. Rich. var. gardeniifolia, 1638, 1677, Mi,2.0-4.0, koppie,

seasonal stream, rock outcrop (dolerite), stream bank, sandy

loam, loam

Richardia

brasiliensis Gomes *, 1215 , T, 0.05, -, disturbed (garden), sandy loam

scabra L. *, 1127, T, 0.05, -, disturbed (roadside), sandy loam

CUCURBITACEAE

Kedrostis

hirtella (Naudin) Cogn., §, L, 2.0. koppie, rock outcrop, gravelly, sandy

Momordica

balsamina L., 1090, L, 3.0, seasonal stream, stream bank, alluvial sand,

sandy loam

boivinii Baill., 1304, L, 0.2, riverine, seasonal stream, river & stream

bank shade, humus-rich alluvial sand

Cucumis

anguria L.

(= var. longipes (Hook, f.) A. Meeuse), §, L, 1.0, Acacia &

Combretum woodland, dryland, gravelly, sandy

hirsutus Sond., §, L. 1 .6, Acacia & Combretum woodland, dryland, sandy

metuliferus Naudin, 1157, L, 3.0, riverine, seasonal stream, river &

stream bank, alluvial sand

zeyheri Sond., 370,

(= C. prophetarum L. subsp. zeyheri (Sond.) C. Jeffrey), 1150, L.

0.05-0.1, -, disturbed (old lands), sandy loam

Lagenaria

siceraria ( Molina ) Standi, 1644, L. 5.0, riverine, seasonal stream, river

& stream bank, alluvial sand

Coccinia

rehmannii Cogn., 327,

(= var. rehmannii ), 1091, 1341. 1399, L. 0. 1-2.0, riverine, sea-

sonal stream, open parkland, Combretum woodland, river &

stream bank, seepline complex, dryland, disturbed (roadside),

gravelly, stony loam, sandy loam, loam

CAMPANULACEAE

Wahlenbergia

undulata (L.f.) A. DC., §, H, 0.3, riverine, river bed, river sand

ASTERACEAE

Ethulia

conyzoides L.f. subsp. conyzoides, 1378, 1419, T, 0.6-0. 7. riverine, river

bed, reed bed. waterlogged river sand

Vemonia

crataegifolia Hutch., §, N, 1.5, riverine, river bank, sandy loam

fastigiata Oliv. & Hiem, 1098, T, 0.4, -. disturbed (roadside), sandy loam

glabra (Steetz) Vatke var. glabra, §, H, 0.6, -, disturbed (roadside), gravel,

stony loam

oligocephala (DC.) Sch.Bip. ex Walp., §, H. 0.3, Acacia woodland,

dryland, sandy loam

steetziana Oliv. & Hiem

(= V. poskeana Vatke & Hildebr. var. chlorolepis (Steetz) O.

Hoffm.), 1109, T. 1.0, Combretum woodland, disturbed (over-

grazed and trampled), sandy loam

Aster

squamatus (Spreng.) Hieron. *, 1172, T, 1 .0, aquatic, dam seepage, sandy

gravel

Nidorella

resedifolia DC. subsp. resedifolia, 384, T, 1.5, riverine, river bank, sandy

loam

Blumea

cafra (DC.) O. Hoffm., 1318, T, 0.5, aquatic, dam seepage, stony, sandy,

clayey loam

gariepina DC., 1218, T, 1.0, -, disturbed (de-bushed, roadside), shallow

gravel

Pluchea

dioscoridis (L.) DC., 1031, N, 0.6, riverine, Combretum woodland, river

bed, alluvial & river sand

Pechuel-Loeschea

leubnitziae (Kuntze) O. Hoffm., §, Ch, 1.2, Acacia, Combretum, &

Colophospermum woodland, riverine, seasonal stream, river &

stream bank, alluvial sand, clayey loam

Denekia

capensis Thunb., §, T, 0. 1-0.2, aquatic, edge of seasonal pan, mud

Epaltes

gariepina (DC.) Steetz, 1156, 1371 , Ch. 0.3-0.4, riverine, open parkland,

river bed, dam shores, seepline complex, disturbed (roadside),

alluvial sand, silt

Sphaeranthus

incisus Robyns, 1192, T, 0.3, aquatic, dam shore, clay

Helichrysum

candolleanum H. Buek, 1002, 1151 . H, 0. 2-0.3, -, disturbed (old lands,

roadside), sandy loam

Calostephane

divaricata Benth., 924, 1616, 1713, H, 0.3-0.6, Combretum & mixed

woodland, dryland, disturbed areas (overgrazed & trampled),

shallow sandy gravel

Pegolettia

senegalensis Cass., 1122, T, 0.4, -, disturbed (roadside), gravel

Geigeria

burkei Harv. subsp. fruticulosa Merxm., §, H, 0.4, riverine, river bed.

sand bank, river sand

omativa O. Hoffm., §, Ch, 0.2-0.3, Acacia woodland, dryland, gravelly,

sandy

Acanthospermum

hispidum DC. *, 1479, T, 0.5-0.6, riverine, river bank (trampled), stony

or sandy loam

Xanthium

strumarium L. *, 1446, T, 0.4, aquatic, dam seepage, sandy gravel

Zinnia

peruviana (L.) L. *, §, T, 0. 2-0.3, -, disturbed (overgrazed, trampled, &

roadside), sandy gravel

Eclipta

prostrata (L.) L. *, 1344, T, 0.4, aquatic, riverine, dam seepage, river &

stream bed sandy, river sand

Aspilia

mossambicensis (Oliv.) Wild. 1153, 1328, Ch, 0.6-1. 0, mixed woodland,

riverine, seasonal stream, shade in dryland & river & stream

banks, shallow stony sandy loam

Melanthera

scandens (Schumach. <£ Thonn.) Roberty subsp. dregei (DC.) Wild, 383,

T, 1.5, riverine, river bank, loam

triternata ( Klatt) Wild

(= M. marlothiana O. Hoffm.), 1346, T, 1.5, riverine, reed bed,

alluvial & river sand

Bidens

pilosa L. *, 284, T, 0.2, -, disturbed (overgrazed & trampled, garden),

sandy loam, loam

Flaveria

bidentis (L.) Kuntze *, §, T, 0. 3-0.4, riverine, river bed, river sand

Schkuhria

pinnata (Lam.) Cabrera *, 1209, T, 0.2, -, disturbed (old lands, roadside,

garden), sandy loam

Tagetes

minuta L. *, §. T, 0.5-0.7, -, disturbed (overgrazed & trampled), sandy

loam

Senecio

pleistocephalus S. Moore, §, L, 4.0, koppie, rock outcrop, sandy gravel

viminalis Bremek., 875, L, 1. 5-3.0, riverine, river bank, loam

Emilia

transvaalensis (Bolus) C. Jeffrey

(= Senecio transvaalensis Bolus), 1391, 1725, T, 0.4, -, disturbed

(roadside), shallow sandy loam

Kleinia

longiflora DC.

(= Senecio longiflorus (DC.) Sch.Bip), §, Ch, 0.5-0.8, open

parkland, seepline complex, clayey loam

Hirpicium

bechuanense (S. Moore) Roessler, 378, 618, H, 0.4, Combretum wood-

land. disturbed (roadside), gravel, sandy loam

Sonchus

oleraceus L. *, 283, T, 0.5, grassland, vlei, disturbed (garden), loam

Bothalia 24, 1 : 55-66 ( 1 994)

External fruit morphology of southern African Arundineae (Arundinoideae:

Poaceae)

N.P. BARKER*

Keywords: achene, Arundineae, Arundinoideae, caryopsis, embryo, external morphology, hilum, Poaceae, surface sculpturing, systematics

ABSTRACT

Fruits of a number of taxa of all indigenous southern African arundinoid genera were examined by means of SEM. Size,

shape, compression, surface sculpturing, embryo and hilum features were recorded and fruits of all genera are illustrated. Results

are compared to existing descriptions. The genera are placed in five informal groups according to similarities noted in the fruits.

U1TTREKSEL

Vrugte van ’n aantal taksons van alle inheemse Suider-Afrikaanse arundinoiede genusse is deur middel van

aftaselektronmikroskopie ondersoek. Grootte, vorm, samedrukking, oppervlakskulptuur, embrio- en hilumkenmerke is

aangeteken en vrugte van alle genusse word geillustreer. Resultate word met bestaande beskrywings vergelyk. Die genusse word

volgens ooreenkomste wat by die vrugte waargeneem is, in vyf informele groepe geplaas.

INTRODUCTION

In his book of grasses of the British Isles, which is

now in its third edition, C.E. Hubbard (1984) included a

section on the ‘seeds' of the grasses, which gave illustra-

tions of the grains of various genera, as well as a key to

the genera based mainly on characteristics of their fruits

(Hubbard 1984, 1st edn 1954). Despite this emphasis, fruit

morphology is still a neglected aspect of grass systematics.

In this study, observations on the fruits of the southern

African Arundineae are presented.

The tribe Arundineae

The tribe Arundineae belongs to the subfamily Arun-

dinoideae which Kellogg & Campbell (1987) consider to

be polyphyletic — an assemblage of basal groups and ev-

olutionary dead ends.

The classification by Clayton & Renvoize (1986),

which is followed in this work, provides a broad definition

for the tribe Arundineae which encompasses most of the

genera in the subfamily. The tribe is a fragmented, hetero-

geneous group of numerous isolated or weakly linked gen-

era. Other workers divide the subfamily into numerous

smaller tribes on the basis of phenetic similarity (Watson

1990) or breeding systems (Conert 1987).

The Arundineae are cosmopolitan. Of the approxi-

mately 40 genera in the tribe, 16 occur in southern Africa.

Two of these, Arundo L. and Phragmites Adans.. are pan-

demic, and species of a third, Cortaderia Stapf, were in-

troduced into South Africa to control soil erosion on mine

tailings dumps (Robinson 1984).

* Botany Department, University of Cape Town, Private Bag, Ronde-

bosch 7700.

MS. received: 1992-08-14.

Terminology

There is as yet no general agreement on the terminol-

ogy to be used for describing the fruits of the Poaceae.

In the Poaceae the ovary is uniloculate. After pollination

and fertilisation (ignoring examples of apomixis) the ovule

develops into a seed, and the ovary wall becomes a fruit

coat or pericarp. In the majority of grasses, the seed and

pericarp are fused, forming a grain (Clifford & Watson

1977). These authors equate this term to the term

‘caryopsis’ but point out that the grain may take the form

of one of a number of different structural types, such as

‘achenes’, ‘nuts’ or ‘berries’. In the fruit of some grasses,

the seed is free from the pericarp. Clayton & Renvoize

(1986) refer to such a fruit as an achene. However,

Sendulsky et al. (1987) consider it a modified caryopsis,

formed by the collapse of either the endocarp or the endo-

and mesocarps at a late stage of development, and rec-

ommend that use of the term ‘achene’ in grass biology be

discontinued.

In the light of the uncertainty in terminology, the grass

grain will be referred to as a fruit in this study. When the

necessary distinction between a grain with a free pericarp

and a grain with an adnate pericarp has to be made, the

term ‘achene’ will be used for the former, and ‘caryopsis’

for the latter grain type.

The fruit of the Arundineae

Clayton & Renvoize (1986) describe the fruit of this

tribe as a caryopsis, sometimes with a free or separable

pericarp, rarely an achene. They name three genera within

the tribe which possess the achene fruit type: Pyrr-

hanthera Zotov, Dregeochloa Conert and Pentameris P.

Beauv. The latter two are endemic to southern Africa and

were included in this study.

SEM studies on the fruits of arundinoid genera by

Barker (1986, 1989, 1990, 1993, and in prep.), have cov-

roups as presented in the discussion

symmetry of shape of fruit in outline; D-V, dorsiventral; Lat., lateral; + embryo and hilum lengths are fractions of total fruit length.

Bothalia 24,1 (1994)

57

ered fruits of most species of Pentameris P. Beauv. and

Pseudopentameris Coned. Davidse (1988) examined the

fruits of the genus Prionanthium Desv. These studies are

discussed below. In this paper the external morphology of

the fruits of the southern African arundinoids is explored.

MATERIAL AND METHODS

Fruit collection

Mature fruits of several taxa in each genus were col-

lected from herbarium specimens housed in the National

Herbarium in Pretoria (PRE). Taxa representing all en-

demic southern African genera were sampled, as well as

taxa of the pandemic genus Phragmites Adans. Fruits of

Arundo L. were unfortunately not available for study. The

specimens and species from which fruit were obtained are

listed below.

As the fmits were dry, no additional desiccation pro-

cedures were followed. Specimens were mounted on the

SEM stubs by means of two-sided tape. They were then

coated in gold-palladium and examined using an ISI-SX-

25 Scanning Electron Microscope. Photographs were

taken using 60 x 70 mm format black and white Ilford

FP4 100 ASA film.

Specimens examined

Centropodia

glauca (Nees) T.A. Cope: De Winter & Hardy 8053 ; Dinter s.n.

(= PRE 331 15); Merxmtiller & Giess 30681, 32064.

mossamedensis (Rendle) T.A. Cope: Boss s.n. (TM 35977).

Chaetobromus

dregeanus Nees: Merxmtiller & Giess 32059.

involucratus (Schrad. ) Nees: Boucher 2542.

Dregeocliloa pumilla (Nees) Conert: Dinter 6391; Schaeffer 12991.

Elytrophorus

globularis Hack.: Erens 342; Smith 1847.

spicatus (Willd.) A. Camus: Ellis 3718.

Karroochloa

cur\>a (Nees) Conert & Tiirpe: Lichtenberg 7718A.

purpurea (L. f.) Conert & Tiirpe: Flanagan 1669; CL Reid 1253.

schismoides (Stapf ex Conert) Conert & Tiirpe: Hardy 607; Munro

s.n.; CL Reid 1271.

tenella (Nees) Conert & Tiirpe: Esterhuysen 23506.

Merxmuellera

drakensbergensis (Schweick.) Conert: Hoener 2184.

dura (Stapf) Conert: Poggenpoel 8274.

stricta (Schrad.) Conert: Adanson 3604; Boucher 1837; Kinges 3482.

Pentaschistis

airoides (Nees) Stapf subsp. airoides: Davidse 33245.

ampla (Nees) McClean: Taylor 11054.

cirrhulosa (Nees) H.P. Linder: Davidse 33801.

curvifolia (Schrad.) Stapf: Davidse 34061 .

natalensis Stapf: Braun 279.

pusila (Nees) H.P. Linder: Esterhuysen 22763, 24195.

Phragmites

australis (Cav.) Steud.: Burtt Davy 358; Le Roux 558; Ward 4225.

mauritianus Kunth: Jacobsen 2405; Loeb 489; Miller B/1169.

Schismus

barbatus (Loefl. ex L.) Thell.: Giess 3225; Liebenberg 6651: Ver-

doorn 1046.

inermis (Stapf) C.E. Hubb.: Lovemore s.n.; Tyson s.n.

scaberrimus Nees: De Winter & Verdoom 9037; Oliver, Toelken &

Venter 635; Van Rensburg 123.

Styppeiochloa gynoglossa (Gooss.) De Winter: Killick & Vahrmeijer

3609.

Tribolium

amplexum Renvoize: Ihlenfeldt 1661.

brachystachyum (Nees) Renvoize: Esterhuysen 1699.

uniolae (L. f.) Renvoize: Kruger KR902; Liebenberg 7719.

Urochlaena pusilla Nees: Davidse 33398, 34021 .

OBSERVATIONS AND DISCUSSION

The salient features of the fruits examined are listed

in Table 1, and these features are illustrated in Figures

1-5. Although only a few of the species in the larger gen-

era were examined, the illustrations show that the fmits

of different species within a genus are relatively uniform.

The differences at generic level are more distinct, al-

though some genera display strong similarities. These

similarities in fruit morphology may be used to form

groups of genera on a purely phenetic basis, where char-

acter and structural homologies are assumed. Four such

groups were assembled on the basis of shared fruit char-

acters. A fifth group comprises three remaining genera,

the fmits of which resemble no other genera in the tribe.

As the tribe is considered to be polyphyletic, these groups

may represent various monophyletic lineages within the

Arundineae. These informal groups, and the genera that

are placed in each, are discussed below, and the salient

features of these groups and genera are presented in Table

2

TABLE 2. — The groups of genera, their fruit characteristics and the figures illustrating the fruits of each genus

58

Bothalia 24.1 (1094)

FIGURE I . — Fruit of Karroodtloa and Schismus spp. (Group I ). A, D, K. purpurea: A, hilar side of fruit showing broadly obovale shape and remains

of style branches; D, detail of surface features. B, K. schismoides , embryo side. C, K. tenella, lateral view showing degree of compression.

E, Schismus scaberrimus, glabrous, broadly obovate, showing large embryo. F, H, S. inermis: F, alveolate surface features; H, hilar side. G,

S. barbatus, lateral view, showing asymmetrical outline. Scale bars: A, 220 pm; B, 1 80 pm; C, 210 pm; D, 80 pm; E, 85 pm; F, 150 pm; G,

170 pm; H, 230 pm.

Bothalia 24,1 (1994)

59

Group 1: Karroochloa , Schismus and Dregeochloa

Genera in this group share fruits which are broadly

obovate, strongly dorsiventrally compressed, generally

glabrous or almost so, have embryos 1/3 to Vi the fruit

length, and punctiform hila.

Karroochloa Conert

Fruit of all four species of this genus were examined

and illustrated in this study (Figure 1A-D). The differ-

ences between the fruit of the species examined is slight,

but a larger sample size may reveal that these differences

intergrade.

Karroochloa was separated from Danthonia DC. on

the basis of, among other characters, differences in the

fruit (Conert & Tiirpe 1969). The latter genus has fruit

which are described as having a hilum half to three quar-

ters the length of the fruit and an embryo no longer than

a third the length of the fruit. Further differences in the

fruit morphology between Karroochloa and Danthonia in-

clude the size of the fruit, surface features, and nature of

the endosperm (Conert & Tiirpe 1969).

Clayton & Renvoize (1986) include Karroochloa ,

along with Merxmuellera Conert and several other non-

African genera, in Rytidosperma Steud. However, the de-

scription of the fruit of Rytidosperma does not match that

observed in Karroochloa in this study. Thus the placement

of Karroochloa in this genus ought to be questioned, es-

pecially in the light of the strong apparent similarities be-

tween Karroochloa and Schismus, which is not included

in the broad Rytidosperma complex.

Schismus P. Beauv.

Schismus comprises five species, four of which occur

in southern Africa. Fruit of three of the five species were

examined (Figure 1E-H). They are all very similar and

closely resemble those of Karroochloa. Results presented

in Table 1 agree essentially with descriptions provided by

Clayton & Renvoize (1986) and Conert & Tiirpe ( 1974).

Dregeochloa Conert

Dregeochloa was created by Conert (1966) to accom-

modate two Danthonia species with an unusual fruit mor-

phology: an obovate caryopsis with a punctiform hilum.

and a membranaceous, removable pericarp. As mentioned

in the introduction, Clayton & Renvoize (1986) consider

the fruit of this genus to be an achene. The fruits of D.

pumila were examined (Figure 2 A, B). The only other

species in the genus, D. calviniensis Conert, is extremely

rare.

The pericarp appears to be flaking and easily remov-

able from the fruit (Figure 2A). It is possible that the fruit

surface was damaged during manipulations, but the fra-

gility of the pericarp and the apparent ease of removal

suggests that it is free from the layers beneath. This and

other features recorded in Table 1 agree with the descrip-

tions by both Conert (1966) and Clayton & Renvoize

(1986). Conert (1966), however, describes the pericarp as

membranaceous whereas Clayton & Renvoize (1986)

refer to it as thick.

Dregeochloa is placed in this group on the basis of the

shape and compression of the fruit and the large embryo.

Conert (1971) observed that the fruit of this taxon is unlike

that of any other arundinoid. The somewhat mgose surface

and the fact that the fruit appears to be an achene, as

noted by Clayton & Renvoize (1986) and corroborated

here, suggests that the relationship of this taxon to the

Schismus-Karroochloa alliance is somewhat tentative.

Group 2: Chaetobromus and Pseudopentameris

The genera in group 2 have narrowly elliptic fruits with

reticulate sculpturing, narrow, canaliculate hila extending

almost the entire length of the fruit and embryos which

are a fifth to a quarter the length of the fruit. These two

genera have not been previously considered as closely re-

lated.

Chaetobromus Nees

Chaetobromus comprises either two (Barker in Gibbs

Russell et al. 1990), three (Clayton & Renvoize 1986) or

four species (Chippindall 1955). Spies et al. (1990) have

suggested that the genus may be a polyploid series, divis-

ible into two phenotypic groups: C. dregeanus and C. in-

volucratus. Fruits of both these groups were examined

and found to be identical (Figure 2C), and corroborate the

descriptions of Clayton & Renvoize (1986).

Pseudopentameris Conert

Barker (1986, 1989. 1990) found the fruits of all spe-

cies to be almost identical, differing only in size. Figure

2D shows the embryo and surface features of the fruit of

this genus. Fruits of this genus are unique in having stig-

mata- like apical appendages, termed pseudostigmata by

Barker (1990, and in prep.). These structures, visible on

the apices of the ovary, developing and mature fruit, are

readily deciduous, being best observed in mature ovaries

and developing fruit. These structures have not been ob-

served in Chaetobromus, but no immature fruits of this

genus have been examined. The descriptions of the fruit

of Pseudopentameris by Clayton & Renvoize ( 1 986) and

Gibbs Russell et al. (1990) agree with details given in

Table 1 . The decision to transfer Pentameris obtusifolia

(Hochst.) Schweick. to Pseudopentameris (Barker in

prep.) was made partly on the basis of strong similarities

of the fruit characters.

Group 3: Tribolium and Uroehlaena

The fruits of genera in this group are ovate, strongly

dorsiventrally compressed, with a flaking pericarp. The

embryo is a quarter to half the length of the fruit.

Tribolium Desv.

This genus comprises either 1 1 (Gibbs Russell et al.

1990) or nine species (Spies et al. 1992). Clayton &

Renvoize (1986) consider this genus to be an outlier in

the Arundineae, with at least superficial similarities to

genera in the Eragrostideae.

The fruits of this genus are described by Clayton &

Renvoize (1986) as having a reluctantly separable peri-

carp. This description agrees with that given by Gibbs

60

Bothalia 24.1 ( 1004)

FIGURE 2. — Fruit of various genera in Groups 1 , 2 & 3. A, B, Dregeochloa pumila (Group 1): A, hilar side showing flaking pericarp; B, details ol

rugose surface. C, Chaetobromus dregeanus (Group 2), hilar side, showing long, canaliculate hilum and scalariform-reticulate surface

features; D. Pseudopentamerisbrachyphylla (Group 2), embryo, showing scalariform-reticulate surface. E-G, Tribolium spp. (Group 3): E,

T. uniolae , hilar view, showing rugose surface and stylar remains; F, T. amplexum, embryo side, broadly elliptic in outline; G, T.

brachystachyum , surface, showing flaking outer layers and rugose surface beneath. H, Urochlaenapusilla (Group 3), showing large embryo

and extensive flaking of outer layers. Scale bars: A, 350 pm; B, 60 pm; C. 560 pm; D, 285 pm; E, 300 pm; F, 425 pm; G, 1 15 pm; H, 490

pm.

Bothalia 24, 1 (1994)

61

Russell et al. (1990), who describe the fruits as being

small, with a short hilum and embryo, and a fairly loosely

adherent pericarp. These descriptions match observations

made in this study, where the fruits of three species in

this genus were studied (Figure 2E-G). As shown in Fig-

ure 2G, the rugose outer layer of the fruit is separable,

matching the decription given by Clayton & Renvoize

(1986). The exposed surface below the pericaip is also

rugose.

Urochlaena Nees

The fruit of the monotypic genus Urochlaena is de-

scribed by Clayton & Renvoize (1986) as a caryopsis with

a free pericarp, an observation repeated by Gibbs Russell

et al. (1990). The embryo is described by these latter au-

thors as being large, while the hilum is short but large.

The fruits of Urochlaena (Figure 2H) are similar to those

of Tribolium.

Tribolium and Urochlaena thus form a natural group

on the basis of all the fruit characters examined. In par-

ticular, the nature of the flaking and separable pericarp

indicate a close relationship between the two genera.

These observations augment the data from other studies

suggesting strong similarities between Urochlaena and

Tribolium (Clayton & Renvoize 1986; Ellis 1988; Spies

et al. 1992).

Group 4: Centropodia, Merxmuellera , Styppeiochloa ,

Prionanthium and Pentaschistis

The fruits of the genera in this group share features

such as a reticulate surface, a hilum approximately half

the length of the fruit, an embryo about a quarter to a

third the length of the fruit and only slight compression

in either plane. However, the genera in this group are

probably, at best, only distantly related, given the differ-

ences in floral and vegetative morphology.

Centropodia Rchb.

Two of the four species in this genus were examined.

Conert (1962) describes the fruits of Centropodia as being

naked caryopses, with linear hila and embryos one third

the length of the fruits. This description approximates the

observations made in the present study (Figure 3A, B),

although Conert’s (1962) description of the hilum as being

linear differs from the present study where it is shown to

be shallowly punctiform (Figure 3B).

In view of the differences in photosynthetic pathways

and thus leaf anatomy (Ellis 1984), the relationship of

Centropodia to the other genera in this group should be

considered as tentative and distant.

Merxmuellera Conert

At present, 20 species are recognised in this genus,

two of which are known only from Madagascar. The fruits

of three southern African species of the genus were stud-

ied. Each of these species came from a different part of

the distribution range of the genus and from a different

habitat. The fruits of all three species studied were re-

markably similar (Figure 3C-E).

The fruits of Merxmuellera have been described as

brown, free, almost terete caryopses, two to three milli-

metres in length (Conert 1970). This description, though

sparse, agrees with observations made in this study. Clay-

ton & Renvoize (1986) include Merxmuellera in Rytido-

sperma, and their description of the fruits of this latter

genus is considered too broad to be applicable to Merx-

muellera alone.

Styppeiochloa De Winter

Styppeiochloa gynoglossa, the only southern African

species in the genus, was investigated. De Winter (1966)

described the fmits of Styppeiochloa as elliptic caryopses

with a linear hilum three quarters the length of the grain

and an embryo one quarter the length of the grain, a de-

scription matching observations made here. Clayton &

Renvoize (1986) do not provide a description of the fruit.

The fmit of S. gynoglossa is illustrated in Figure 3F-H.

Pentaschistis (Nees) Spach

Pentaschistis is the largest genus in the tribe, compris-

ing 72 species (Linder & Ellis 1990). The fruits of only

six species, including those of P. pusilla {- Poagrostis

pusilla (Nees) Stapf) were studied. Despite the size of the

genus, little variation was found in the external morphol-

ogy of the fmits investigated (Figure 4A-H).

The fmits were found to conform with descriptions

given by Clayton & Renvoize (1986) and Gibbs Russell

et al. (1990). However, the pericarp description given by

the latter workers (free to loosely adherent to fused) was

found to be too broad; the pericaip appeared to be fused

in all the species studied.

Prionanthium Desv.

Prionanthium comprises three species. It was revised

by Davidse (1988) who described and illustrated the fruit

of two species. The genus shares many fmit characters

with Pentaschistis, an observation supporting the hypoth-

esis that these two genera are closely related, as proposed

by Linder et al. (1990) on the basis of the presence of

unusual glands in both these genera. In addition. Davidse

(1988) considers the base chromosome number of x = 7,

which is common to both genera, to also be an indication

of their close relationship. This is in contrast to Clayton

& Renvoize (1986), who place Prionanthium close to

Tribolium and Urochlaena.

Group 5: special cases

Three genera are placed in this group, not because of

any similarities in the nature of the fmit, but simply be-

cause their fruits resemble none of the fruits of the other

genera discussed above. These 'misfits’ are Phragmites,

Pentameris and Elytrophorus.

Phragmites Adans.

Phragmites is a pandemic genus with two species oc-

curring in southern Africa, P. australis and P. mauritianus.

The fmits of these species examined were almost identi-

cal. As no detailed description of the fmit from past work-

ers was found, the observations presented here cannot be

62

Bothalia 24.1 (1994)

FIGURE 3. — Fruit of genera in Group 4 (Centropodia, Merxmuellera and Styppeiochloa spp). A, B, C. glcutca: embryo side, showing large embryo,

reticulate surface features and remains of style; B. hilar side, showing punctiform hilum. C. M. dura , hilar region, close-up of scalariform-

reticulate surface features. D, E, M. stricter. D, embryo side, showing embryo a third the length of fruit; E, broad, fairly short hilum and

remains of styles. F-H, S. gynoglossa: F, reticulate surface features; G, embryo one third length of fruit; H, lateral view. Scale bars: A. 370

pm; B, 425 pm; C, D, 1 1 0 pm; E, 695 pm; F, 65 pm; G, H, 465 pm.

Bolhalia 24.1 (1994)

62

FIGURE 4. — Fruit of Pentaschistis spp. (Group 4). A, P. pusilla, showing embryo and regularly reticulate surface features. B. P. ampla, lateral

view, with regularly reticulate surface features. C, D, H, P. natalensis: C, embryo side of elliptic fruit; D, embryo region showing reticulate

surface features; FI, hilar side. E, P. cirrhulosa , showing long, canaliculate hilum. F, G, P. airoides subsp. airoides : F, embryo showing

rugose-reticulate surface features; G, deep, broad long hilum. Scale bars: A, 350 pm; B. 415 pm; C, 455 pm; D, 180 pm; E, 325 pm; F, 85

pm; G, 230 pm; H, 445 pm.

64

Bothalia 24,1 (1994)

compared to past descriptions. The fruits of this genus are

illustrated in Figure 5A-C.

The fruits of Phragmites are unusual within the con-

text of the southern African Arundineae. Their small size

undoubtedly assists in their dispersal by wind, the fruit

being dispersed in situ in the lloret. Further examination

of the fruits of other reed-like grasses such as Arundo ,

Hakonechloa Honda and Molinia Schrank may assist in

clarifying the relationships between Phragmites and other

related reed genera with wind dispersed fruit.

Pentameris P. Beauv.

The fruits of all except one of the nine species of Pen-

tameris have been investigated and illustrated (Barker

1986, 1989, 1990, 1993). The fruit of P oreophila N.P.

Barker is illustrated in Figure 5D.

FIGURE 5. — Fruit of genera in Group 5 (Phragmites, Pentameris and Elytropharus). A, Phragmites australis, showing large embryo, regularly

reticulate surface and remains of styles. B, C, Phragmites mauritianus: B, surface features; C, hilar side showing rugose surface features. D,

Pentameris oreopltila showing colliculate surface. E-G, Elytropharus globularis: E, surface detail; F. lateral view showing unusual shape

and raised, reticulate surface; G, embryo view. Scale bars; A, 380 mm; B, 250 mm; C, 80 mm; D, 490 mm; E, 280 mm; F, 140 mm; G, 155

mm.

Bothalia 24,1 ( 1994)

65

Pentameris possesses fruits which differ markedly

from other southern African arundinoid genera, in both

external and internal structure. The fruits of this genus are

achenes with apical appendages in the form of hairs or

hair-like structures.

These two characters clearly distinguish this genus

from the closely related and probable sister genus Pen-

taschistis. Thus in a phylogenetic system based on overall

morphology, Pentameris would belong in group 4. How-

ever, on the basis of phenetic groupings employed here,

it is treated as an exceptional taxon.

Elytrophorus P. Beauv.

Elytrophorus differs from the other genera of the

Arundineae in having dimorphic spikelets and a membra-

nous ligule (as opposed to the more usual arundinoid lig-

ule comprising a row of hairs).

Both species in this genus were examined, and the

fruits of these two taxa were identical in all but size, those

of E. globularis being far larger than those of E. spicatus

(Figure 5E-G). Schweickerdt (1942) reported that the em-

bryo is almost as long as the grain, whereas Clayton &

Renvoize (1986) consider the embryo to be only half the

length of the fruit, a size more in keeping with that ob-

served here. Schweickerdt ( l.c .) alludes to the caryopsis

as ‘showing the remains of the pericarp’, implying that

the pericarp is separable. However, Clayton & Renvoize

(1986) do not consider the fruit of this genus to be an

achene. Instead, the pericarp is described, somewhat con-

fusingly, as being free.

The shape of the fruit of these species is like that of

no other southern African arundinoid. and it is not possible

to place it in any of the other groups described above,

which only serves to enhance the confusion of this taxon's

uncertain affinities: Clayton & Renvoize (1986) consider

Elytrophorus to have uncertain affinities, and Chippindall

(1955) placed it in the Eragrosteae.

CONCLUSION

This study has revealed that the diversity in the floral

and vegetative morphologies of the genera of southern

African Arundineae is parallelled by an equally diverse

set of fruit morphologies. However, this diversity is gen-

erally minimal at infrageneric levels. Despite the diversity

in fruit morphology, certain genera are united in almost

all fruit characters examined (: Tribolium and Urochlaena ;

Karroochloa and Schismus). Where this occurs, it is in-

terpreted as indicating a strong degree of relatedness. Un-

fortunately, only this very limited phylogenetic inter-

pretation is possible using this data. This is because struc-

tural homologies are unproven, only southern African taxa

are being examined, and insufficient characters are ob-

tained from a purely SEM-based study. The extension of

this study to include non-southern African taxa may, how-

ever, provide additional data for comparison.

This study is strictly descriptive. In many instances,

the generic relationships proposed on the basis of fruit

morphology are speculative. Nonetheless, this study has

provided valuable data for incorporation in a phyloge-

netic analysis which is presently being undertaken in the

form of a combined molecular (a survey of chloroplast

DNA sequence variation) and morphological survey of the

entire tribe. This study has indicated the need for a thor-

ough examination of the nature of the fruit of grasses at

the generic level.

ACKNOWLEDGEMENTS

I would like to thank Mrs A. Romanowski (NBI), Mrs

W. Hitchcock (UCT) and Mr R. Carelse (UCT) for assis-

tance with the photographic work, Mrs S. Perold (NBI)

for assistance with the SEM. The Curator of the National

Herbarium (PRE) is thanked for allowing access to, and

SEM exmination of, the specimens cited above. Drs R.P.

Ellis, O.A. Leistner and H.P. Linder are thanked for their

comments on an earlier draft of the manuscript. This work

was carried out while the author was employed by the

National Botanical Institute.

REFERENCES

BARKER. N.R 1986. The shape and ultrastructure of the caryopsis of

Pentameris and Pseudopentameris species (Arundinoideae,

Poaceae). Bothalia 16: 65-69.

BARKER. N.R 1989. The caryopsis surface of Pentameris and Pseudo-

pentameris (Arundinoideae,' Poaceae) revisited. Bothalia 19:

134-136.

BARKER, N.P. 1990. The taxonomy of Pentameris P. Beauv. and

Pseudopentameris Conert. M.Sc. thesis. University of the

Witwatersrand.

BARKER. N.P. 1993. A biosystematic study of Pentameris (Arundineae,

Poaceae). Bothalia 23: 25—47.

BARKER, N.P. in prep. A systematic study of Pseudopentameris

(Arundineae; Poaceae).

CHIPPINDALL, L.K.A. 1955. A guide to the identification of grasses in

southern Africa. In D. Meredith, The grasses and pastures of

South Africa. Central News Agency, Cape Town.

CLAYTON, W.D. & RENVOIZE, S.A. 1986. Genera graminum.

Grasses of the world. Her Majesty’s Stationery Office, London.

CLIFFORD, H.T. & WATSON. L. 1977. Identifying grasses: data, meth-

ods and illustrations. University of Queensland Press, St. Lucia.

CONERT, H.J. 1962. Uber die Gramineen-Gattung Asthenatherum

Nevski. Senckenbergiana Biologica 43: 239-266.

CONERT. H.J. 1966. Dregeochloa, eine neue Gattung der Gramineen

(Gramineae. Arundinoideae, Danthonieae). Senckenbergiana

Biologica 47: 335-343.

CONERT. H.J. 1970. Merxmuellera : eine neue Gattung der Gramineen

(Poaceae: Arundinoideae). Senckenbergiana Biologica 51: 129—

133.

CONERT, H.J. 1971. The genus Danthonia in Africa. Mitteilungen der

Botanischen Staatssanunlung, Munchen 10: 299-308.

CONERT. H.J. 1987. Current concepts in the systematics of the

Arundinoideae. In T.R. Soderstrom, K.W. Hilu, C.S. Campbell &

M.E. Barkworth, Grass systematics and evolution. Smithsonian

Institution Press, Washington D.C.

CONERT, H.J. & TURPE, A.M. 1969. Karroochloa , eine neue Gattung

der Gramineen (Poaceae, Arundinoideae, Danthonieae).

Senckenbergiana Biologica 50: 289-318.

CONERT. H.J. & TURPE, A.M. 1974. Revision der Gattung Schismus

(Poaceae: Arundinoideae: Danthonieae). Abhandlungen der

Senckenbergischen Naturforschenden Gesellschaft 532: 1-81.

DAV1DSE, G. 1988. A revision of the genus Prionanthium (Poaceae:

Arundinoideae). Bothalia 18: 143-153.

DE WINTER. B.D. 1966. Styppeiochloa De Winter, gen. nov.

(Gramineae). Bothalia 9: 134—137.

ELLIS, R.P. 1984. Leaf anatomy of the South African Danthonieae

(Poaceae). IX. Asthenatherum glaucum. Bothalia 15: 153-159.

ELLIS, R.P. 1988. Leaf anatomy of the South African Danthonieae

(Poaceae). XVI. The genus Urochlaena. Bothalia 18: 101-104.

GIBBS RUSSELL, G.E.. WATSON, L„ KOEKEMOER. M„ SMOOK.

L„ BARKER. N.P, ANDERSON, H.A. & DALLWITZ, M.J

1990. Grasses of southern Africa. Memoirs of the Botanical Sur-

vey of South Africa No. 58.

HUBBARD, C.E. 1984. Grasses. A guide to their structure, identifica-

tion, uses and distribution in the British Isles. Penguin Books,

Harmondsworth, Middlesex, England.

KELLOGG, E.A. & CAMPBELL, C.S. 1987. Phylogenetic analyses of

the Gramineae. In T.R. Soderstrom, K.W. Hilu, C.S. Campbell &

M.E. Barkworth, Grass systematics amt evolution. Smithsonian

Institution Press, Washington D.C.

LINDER, H P. & ELLIS, R.P. 1990. A revision of the genus Pentaschistis

(Arundineae: Poaceae). Contributions from the Bolus Herbarium

12.

66

Bothalia 24,1 ( 1994)

LINDER, H R, THOMPSON, J.F., ELLIS, R.P & PEROLD, S.M. 1990.

The occurrence, anatomy, and systematic implications of the

glands in Pentaschistis and Prionanthium (Poaceae, Arundinoi-

deae, Arundineae). Botanical Gazette 151: 221-233.

ROBINSON, E.R. 1984. Naturalized species of Cortaderia (Poaceae) in

southern Africa. South African Journal of Botany 3: 343-346.

SCHWEICKERDT, H.G. 1942. A taxonomic and anatomical study of

ElvtrophorusP. Beauv. Annals of the Natal Museum 10: 191-214.

SENDULSKY, T„ FILGUEIRAS, T.S. & BURMAN, A.G. 1987. Fruits,

embryos and seedlings. In T.R. Soderstrom, K.W. Hilu, C.S.

Campbell & M.E. Barkworth, Grass systenmtics and evolution.

Smithsonian Institution Press, Washington D.C.

SPIES, J.J., DU PLESSIS, H„ BARKER, N.P & VAN WYK, S.M.C.

1990. Cytogenetic studies in the genus Cltaeiobrotnus (Poaceae:

Arundineae). Genome 33: 646-658.

SPIES, J.J., DAV1DSE, G. & DU PLESSIS, H. 1992. Cytogenetic studies

in the genus Tribolium (Poaceae: Arundineae). American Journal

of Botany 79: 689-700.

WATSON, L. 1990. The grass family, Poaceae. In G.P Chapman, Repro-

ductive versatility of the grasses. Cambridge University Press.

Bothalia 24,1:67-76 (1994)

Names of the southern African species of Rhus (Anacardiaceae) and their

etymology

R.O. MOFFETT*

Keywords: Anacardiaceae, etymology, Rhus, southern Africa

ABSTRACT

Brief notes on the origin of the scientific names of all the Rhus species recognized in the recent revision of the genus (Moffett

1993) are given, as well as a complete list of common names for each species. Recommendations are also made on those species

which should be added to or removed from the National list of indigenous trees.

UITTREKSEL

Kort aantekeninge oor die oorsprong van die wetenskaplike name van al die Rhus- spesies wat in die onlangse hersiening van

die genus (Moffett 1993) erken is, asook 'n volledige lys van volksname vir elke spesie word voorsien. Daar word ook

aanbevelings gemaak oor watter spesies by die Nasionale lys van inheemse borne gevoeg of daaruit verwyder moet word

INTRODUCTION

Rhus is one of the most widespread of all genera in

the southern Africa flora and there are few areas without

at least one species (Figure 1). Species such as R. lancea

(karree), R. leptodictya (mountain karree), R. pendulina

(white karree), R. erosa (broom karree) and R. pyroides

(common wild currant) are fairly well known and their

importance as providers of shelter, food, fencing posts,

fuel and ornamentals was highlighted in 1983 when the

Department of Environment Affairs declared the karree

'Tree of the Year’ (Esterhuyse 1983).

In my recent revision of the southern African taxa of

Rhus for the Flora of southern Africa, I recognized 72

species, 18 infraspecific taxa and a further eight provi-

sional species whose position I have yet to finalize

(Moffett 1993).

Whereas this paper deals mainly with the etymology

of the specific epithets and common names of the 90 rec-

ognized taxa, I have also included the following in the

treatment of each species: the correct name of each spe-

cies, its author and date of publication and its number in

the Flora of southern Africa ( Moffett op. cit. ), the most

important synonym(s) (if any) of each species in paren-

theses and brief descriptions of the habit and distribution

of each species.

Common names preceded by a number, as follows:

386, karree, karee, are taken from the National list of in-

digenous trees (Von Breitenbach 1990). Species of which

the suggested common names are followed by an * should

be added to the national list, whereas those marked with

+ should be removed from the list as they are not arbo-

rescent.

* Department of Botany, University of the North, Drakensberg Campus,

Private Bag XI 3, Phuthaditjhaba 9866.

MS. received: 1991-09-02.

1 am against the indiscriminate invention of common

names but having recently revised the genus, felt it nec-

essary to not only create order in the scientific nomencla-

ture but also to provide a uniform vernacular nomen-

clature. In this regard I have followed Von Breitenbach

{op. cit.) who standardized the common names of the ar-

borescent species by, in most cases, highlighting a single

English and single Afrikaans name for each species, e.g.

red currant/bostaaibos for R. chirindensis, a species given

fourteen common names in Palmer & Pitman (1973). In

order not to create further confusion I have maintained

Von Breitenbach’s {op. cit.) choice of common names de-

spite some of them being inappropriate. Where I have pro-

posed a new name it was either based on an unpublished

name already in use in the field as reflected on herbarium

sheets, or on an intimate knowledge of the species con-

cerned. Many of the new names are for lesser-known non-

arborescent species and hopefully may lead to these

species becoming better known. As far as I could ascertain

I have not proposed a new common name for an existing

published one.

Information about botanists and collectors honoured in

specific epithets was taken from Gunn & Codd (1981).

General note on common names

The most popular common names used for the southern

African Rhus species are currant, "korentebos’; karree,

'karee' and 'taaibos'.

The name currant refers to the currant-like small fruits

of the genus; among the currants one finds blue currant

(R. zeyheri), dune currant (R. laevigata) and real wild cur-

rant (R. tomentosa) to name but three. The Korenterivier

near Riversdale in the southern Cape was probably named

after the latter two species, both of which occur on its

banks.

68

Bothalia 24.1 (1994)

FIGURE 1. — Distribution of Rhus species in southern Africa.

Many karree species, with karree (R. lancea) perhaps

the best known, are the source of many place and farm

names. Among the collecting localities listed by Leistner

& Morris (1976), for example, are kareeberge, kareeboom,

kareebos, kareebult, kareefontein, kareekloof, kareekolk,

kareekop, kareekuil, kareelaagte, kareepan, kareeput,

kareerivier and kareevlakte. Other familiar karrees in-

clude bitter karree (R. marlotliii ), broom karree {R. erosa),

mountain karree (R. leptodictya ) and sour karree (R.

ciliata).

There are different views concerning the origin of the

name karee and most authors have suggested that the

name has the same source as the word karoo which means

arid, dry or hard. Nienaber (1963) stated that Karoo was

a compromise between the earlier English Karroo and the

Afrikaans Karo. He reported that its earliest use was by

Schrijver who, in 1689, referred to the Xhaeruh or

Droogerivier, (VRV 12: 1215), whereas Thunberg in 1772,

wrote ‘of the carro’.

According to Nienaber (op. cit.) some of the earliest

references to the tree included the following:

1774: kari-boom, Masson (specimen in BM).

1774: Karre-hout, Thunberg (2:191).

1778: de care-boom. Van Plettenberg (RZA, II: 67).

1791: kare. Van Reenen (RZA, II: 145).

1803: carru hout, Janssens (RZA, IV: 159); car-

ruboom en bosch. Janssens (RZA IV: 176)

1803: caree bosch. Van Reenen (VRV, 18: 242);

currubosch. Van Reenen (VRV, 18: 243).

1803-1806 Karreeboom; Karee bosch; Karee-

bosch, Lichtenstein (VRV, XI: 27).

With the simultaneous use by some of the authors of

the names carro and carre or karoo and karee for the arid

plain and its distinctive tree, it is understandable that a

link should be sought between the two. Pettman (1931)

differed, however, and stated that Karree or Kiri was de-

rived from the Hottentot ‘karib,’ meaning honey beer.

The source of the name karee which I favour is that

given in A bushman dictionary of Dorothea Bleek (1956):

kare n. = haakdoom tree; karee n. = toes, claws, heels,

hoofs, spoor made by feet; karri n. = trunk of tree. Al-

though Bleek does not link karee with the tree, it is easy

to see the connection between a karree leaf and the spoor

of a fairly large bird. A further point of interest concerning

the vernacular is that in Australia, the word karri is used

for Eucalyptus diversicolor , a tree with dark red wood

like our karree (Onions 1956). Could there be a philolog-

ical link between the bushman and the aborigine?

Although I personally support those authors who spell

karree in English with only one r, e.g. Smith (1966) and

Coates Palgrave (1977), Von Breitenbach (op. cit.) main-

Bothalia 24,1 (1994)

69

tains the traditional karree and as I have already men-

tioned, in order not to create further confusion, I have

followed the national list.

The name ‘taaibos’ is applied to several species and

Smith (op. cit.) stated that this was due to the tough 'taai'

but resilient branches. Another explanation given to me

in 1981 by a Wolseley farmer was that because of their

woody underground stems, the shrubs were difficult to

remove and therefore ‘taai’. He was referring to R. an-

gustifolia and R. undid ata. two species which prove par-

ticularly difficult to get rid of when clearing natural bush

for cultivation. Although this farmer called these two spe-

cies ‘taaibosse’, they are known by other common names

in the standardized national list which emphasizes the gen-

eral unreliability of common names. Other species known

as ‘taaibosse" include ‘blou taaibos’ (R. zeyheri), ‘bos-

taaibos’ (R. chirindensis) and ‘gewone taaibos’ (R. py-

roides).

Another common name of interest is that of the crow-

berries. Three species have this name, viz. common crow-

berry (R. pentheri), dune crow-berry (R. crenata ) and

broom crow-berry (R. refmcta). Smith (op. cit.) suggested

that the name referred to the fact that the fruit was so

inferior that it was only fit for crows. Besides karee, two

other Khoi (Hottentot) names that have become associated

with the southern African Rhus species are ‘kuni’ as in

kuni-bush for R. undulata and ‘nana’ as in nana-berry for

R. dentata. Smith (op. cit.) stated that the latter name,

which is incorrectly given in his text as ‘namabessie’ but

correctly in the systematic index, was first recorded in

about 1869.

The various common names rarely reflect taxonomic

relationships between the species. Species such as karree

(R. lancea), mountain karree (R. leptodictya) and white

karree (R. pendulina) look superficially alike but upon

close examination are morphologically very different from

one another. The situation regarding the currants is similar

and the red currant (R. chirindensis), is for example, mark-

edly different from the blue currant (R. zeyheri).

Where I have proposed new common names, I have

standardized on currant (English) and ‘taaibos’ (Afri-

kaans), except where a more suitable name has suggested

itself, e.g. Burchell’s kuni-bush for R. burchellii, a species

hitherto included in kuni-bush (R. undulata).

Classification and etymology

Family: Anacardiaceae Lindl. (1830). After the type

genus Anacardium L., a name referring to the heart-like

shape of the swollen, fleshy pedicel of the fruit (cashew

nut).

Tribe: Rhoeae Marchand (1869). After the type genus

Rhus L.

Genus: Rhus L. (1753). After Rhus or povs , an ancient

Greek name which has no Latin equivalent and whose

precise meaning is obscure. Some authors have, however,

suggested a derivation from the Greek word meaning ‘to

flow’ or ‘flowing’ or a derivation from the Celtic rhudd

or rub meaning red. Others have suggested that the name

may have been derived from the Greek for reddish or

rose-pink.

Miller ( 1768), suggested that the ‘flow’ referred to the

use of parts of R. coriaria to check haemorrhages, whereas

Sonder (1860) suggested that the ‘flow’ referred to the

resinous secretions of the plants.

The derivation from the Celtic for red, in allusion to

the colour of the fruit and the autumn leaves as suggested

by G. Don (1832) is, 1 think, dubious. In this regard, Mc-

Nair (1925) pointed out that the Celts were probably too

far north to have influenced the name of this plant which

occurred in the Mediterranean region and in Asia Minor.

It is certain that the type species, R. coriaria L.. was

well known before the time of Christ. Theophrastus in

300 B.C. wrote extensively about this species which he

called sumach, an old Arabian name. Dioscorides was

probably the first to record the name Rhus when in A.D.

± 50 he wrote of Rhus coriaria ‘which is soe called, be-

cause tanners doe use it for ye thickning of their hydes’

(Barkley & Barkley 1938), quoting from John Goodyer’s,

‘1655 Englished edition’.

With regard to the pronunciation of the name Rhus,

the Oxford English Dictionary favours Rhus as in bus (On-

ions 1956). Von Breitenbach (1983), however, pointed out

that the genitive of rhous is rhoos, that the stem is rho

and that Rhus should therefore be pronounced as in bonus.

Subgenus: Thezera (DC.) K. Koch (1853). After The-

zera which first appeared as a specific epithet in R. thezera

Pers., a species subsequently included in R. pentaphylla

(Jacq.) Desf. Persoon cited its habitat as ‘Sicily and

Barbaria’, the latter being North Africa west of Egypt, but

gave no origin for the name Thezera (Persoon 1805). I

suspect it had a geographic connotation.

De Candolle (1825) used Thezera as the name for one

of his sections of the genus Rhus and the name was sub-

sequently given subgeneric status by Koch (Koch 1853).

Rhus acocksii Moffett ( 1 988), FSA 3.

After J.P.H. Acocks ( 1911-1979), South African bota-

nist and ecologist, who collected the type specimen in

Transkei in 1947.

Proposed common names: rock red currant, klipbos-

taaibos.

A scandent shrub/climber occurring on quartzite krantzes

in southern Natal and Transkei.

R. albomarginata Sond. (1860), FSA 46.

Latin for white margin, referring to the creamy white

thickened margin of the leaflets.

Proposed common names: white-edged currant, witrand-

taaibos.

Rare dwarf shrub up to 0.5 m high, from the area between

Grahamstown and Suurberg, north of Port Elizabeth.

70

Bothalia24,l (1994)

R. angustifolia L. (1753), FSA 57.

Latin for narrow leaves, referring to the lance-shaped

leaflets.

Common names : 377 . 1 , willow currant, wilgerkorentebos.

Shrub 1-3 m high, usually found on streambanks in the

southwestern Cape.

R. batophylla Codd (1956), FSA 63.

Greek for bramble leaves, referring to the bramble-like

leaflets.

Proposed common names : bramble currant, braamtaaibos.

Shrub up to 2 m high, only found in the Steelpoort area of

the eastern Transvaal.

R. baurii Schonland ( 1930). See R. pyroides var .pyroides.

R. bolusii Sond. ex Engl. (1883), FSA 32.

After H. Bolus (1834-1911), businessman, botanist

and botanical philanthropist who collected the type spec-

imen near Graaff-Reinet in 1868.

Proposed common names : Bolus’s broom karee, baster-

besemkaree.

Shrub up to 2 m high, found scattered in the eastern OFS,

Lesotho and northeastern Cape.

R. burchellii Sond. ex Engl. (1883), FSA 43.

(= R. undulata sensu Schonland)

After W.J. Burchell (1781-1863), collector par excel-

lence, who between 1811 and 1815 gathered about 50 000

plant specimens in the Cape, 1 10 of which were numbered

Rhus taxa.

Proposed common names : Burchell’s kuni-bush, Burchell

se-koeniebos.*

Shrub up to 5 m high, common on rocky hills of the central

karroid parts of southern Africa.

R. carnosula Schonland (1930), FSA 8.

Latin for slightly fleshy, referring to the somewhat

fleshy leaflets.

Common names'. 379, false nana berry, bastemanabessie.-i-

Suffrutescent shrub up to 1 .5 m high, found mainly along

the coast and adjacent interior between Oribi Gorge, Natal

and East London.

R. chirindensis Baker f . (1911), FSA 2.

(= R. legatii Schonland & R. chirindensis forma

legatii (Schonland) R. Fern. & A. Fern.)

After Chirinda, Zimbabwe, near where the type speci-

men was collected in 1906.

Common names : 380, red currant, bostaaibos.

Shrub or tree up to 23 m high, widely distributed in moister

parts from Swellendam, northeastwards to the Sout-

pansberg and Zimbabwe.

* should be added to the national list.

+ should be removed from the national list.

R. ciliata Licht. ex Schult. (1820), FSA 21.

Latin for cilia, referring to the minute hairs along the

leaflet margins.

Proposed common names: sourkaree, suurkaree.

Shrub up to 2 m high, widespread in the central drier parts

of South Africa and also in central to northern Namibia.

R. crenata Thunb. (1803), FSA 36.

Latin for having rounded teeth, referring to the crenate

apical margin of the leaflets.

Common names: 380.1, dune crow-berry, duinekraai-

bessie.

Shrub up to 4 m high, found on the coastal and adjacent

inland dunes from the Cape Peninsula to mZumbe in

southern Natal.

R. cuneifolia L. f. (1781), FSA 50.

Latin for cuneate leaves, referring to the wedge-shaped

leaflets.

Proposed common names: Kogelberg currant, Kogelberg-

taaibos.

Dwarf xerophytic shrublet up to 0.5 m high, from the

southwestern Cape.

R. dentata Thunb. (1794), FSA 7.

Latin for sharply toothed, referring to the dentate mar-

gins of the leaflets.

Common names: 381, nana berry, nanabessie.

Shrub usually up to 2 m high, rarely reaching 5 m, wide-

spread over the eastern parts of southern Africa from

Storms River, Cape to Potgietersrus.

R. discolor E. Mey. ex Sond. ( 1 860), FSA 52.

Latin for not of the same colour, referring to the leaflets

which are grey-green above, whitish cream below.

Proposed common names: grassveld currant, grasveld-

taaibos.

Dwarf suffrutescent shrub up to 1 m high, widespread in

moister grassland from Hogsback, Cape to Pietersburg.

R. dissecta Thunb. ( 1 803), FSA 60.

Latin for deeply divided, referring to the sharply

toothed and irregularly divided margin of the leaflets.

Proposed common names: long-stalked currant, langsteel-

korentebossie.

Dwarf shrub up to 1.5 m high, found in the southwestern

Cape between Cape Town and Van Rhynsdorp.

R. divaricata Eckl. & Zeyh. ( 1 836), FSA 20.

Latin for spreading, referring to the wide-angled

branching.

Proposed common names: rusty-leaved currant, roesblaar-

taaibos.*

Shrub up to 3 m high, found along mountain ranges from

Graaff-Reinet to Heidelberg, Transvaal.

Bothalia 24, 1 (1994)

71

R. dracomontana Moffett (1993), FSA 14.

Latin for dragon mountains, referring to the Drakens-

berg mountain range.

Proposed common names: Drakensberg dwarf currant,

Drakensbergdwergtaaibos.

Dwarf suffrutescent shrub up to 1.2 m high, found along

the low Drakensberg escarpment between Van Reenen,

Natal and just north of Wakkerstroom, Transvaal.

R. dregeana Sond. ( 1 860), FSA 29.

After J.F. Drege (1794—1881), professional collector,

who between 1826 and 1834 collected about 200 000

specimens in South Africa.

Proposed common names : needle-leaved broom karee,

naaldblaarbesemkaree.

Shrub up to 2 m high, found in the northeastern Cape,

southern OFS and Lesotho.

R. dura Schonland (1930). See R. tumulicola.

R. engleri Britten (1900), FSA 23.

AfterH.G.A. Engler( 1844-1930), Professor of Botany

and Director of the Botanic Garden at Berlin-Dahlem who

was the leading authority on the Anacardiaceae and pro-

duced bench-mark works on them.

Common names : 382, velvet karree, fluweelkaree.

Shrub up to 3 m high, found in the central Transvaal.

R. erosa Thunb. ( 1 8 1 8), FSA 3 1 .

Latin for irregularly toothed, referring to the jagged-

edged margin of the leaflets.

Common names: 383, broom karree, besemkaree.

Shrub up to 3 m high, plentiful on koppies in the northeast-

ern Cape, southern and eastern OFS and Lesotho.

R. fastigata Eckl. & Zeyh. (1836), FSA 19.

Latin for high, exalted. The name, however, is probably

derived from fastigiatus meaning ‘branches clustered, par-

allel and erect, giving a narrow elongated habit'. This

species has, in fact, since 1860 been incorrectly known as

R. fastigiata.

Common names: 383.1, broom currant, besemtaaibos.+

Shrub up to 2 m high, found along the coast and adjacent

interior from near Flumansdorp to near Port Shepstone.

R. gerrardii (Harv. ex Engl.) Diels (1898), FSA 17.

(= R. montana sensu Schonland)

After W.T. Gerrard (d. ± 1866), a noted collector who

with M.J. McKen was responsible for finding many new

species in Natal. This species is listed as Drakensberg

karree under the name R. montana var. gerrardii in the

National list of indigenous trees.

* should be added to the national list.

+ should be removed from the national list.

That common name should, however, be maintained for

R. montana and 1 propose that R. gerrardii be named river

currant, riviertaaibos.*

Shrub up to 2.5 m high, found along banks of perennial

streams from the northern Transkei to the eastern Trans-

vaal highveld.

R. glauca Thunb. ( 1 803), FSA 4 1 .

Latin for glaucous, referring to the colour of the leaflets

which are often bluish green due to a waxy surface.

Common names: 383.2, blue kuni-bush, bloukoeniebos.

Shrub up to 4 m high, found along the coast and the

adjacent interior from Velddrif in the southwestern Cape

to near Kentani in Transkei.

R. gracillima Engl. ( 1 883), FSA 69.

Latin for thin, slender, referring to the slender stems and

pendulous leaves with long narrow leaflets.

Proposed common names: needle-leaved dwarf currant,

naaldvormigedwergtaaibos.

Two varieties are recognized:

var. gracillima and var. glaberrima Schonland after the

smooth, hairless leaflets.

Dwarf rhizomatous shrublet up to 0.7 m high, found in the

central and eastern Transvaal.

R. grandidens Harv. ex Engl. (1883), FSA 5.

Latin for large teeth, referring to the pronounced

toothed margins of many of the leaflets.

Proposed common names: sharp-toothed currant, skerp-

tandtaaibos.

Shrub up to 2 m high, found in a number of disjunct

localities from the eastern Transkei to the northeastern

Transvaal.

R. gueinzii Sond. (1860), FSA 38.

(= R. crispa (Harv. ex Engl.) Schonland, R. simii

Schonland & R. spinescens Diels)

After W. Gueinzius (1814—1874), apothecary and col-

lector who lived in Durban.

Common names: 384, thorny karree, doringkaree.

Shrub up to 3 m high, or sometimes a small tree reaching

8 m, ranging from the eastern Cape to the northern Trans-

vaal.

R. harveyi Moffett (1993), FSA 53.

After W.H. Harvey (1811-1866), one-time Colonial

Treasurer at the Cape and later Professor of Botany at

Trinity College, Dublin.

Proposed common names: Harvey’s currant. Harvey-se-

taaibos.

Dwarf suffrutescent shrub up to 1 m high,* confined to

grassland near Louwsburg, northern Natal and Mbabane,

Swaziland.

72

Bothalia 24,1 (1994)

R. horrida Eckl. & Zeyh. ( 1 836), FSA 7 1 .

Latin for sticking out, prickly, rough, bristly, referring

to the spinous nature of the branches.

Proposed common names: red thorn currant, rooidoring-

taaibos.

Shrub up to 1.7 m high, occurring in the Kamiesberg of

Namaqualand.

R. incisa L. f. (1781), FSA 59.

Latin for cut deeply and sharply, referring to the mar-

gins of the leaflets of var. incisa.

Common names: 385, rub-rub berry, baardbessie.

Two varieties are distinguished:

var. incisa and var. effusa (Presl) R. Fern. (= R. obovata

Sond.), after the effusive or loosely spreading inflores-

cence.

Shrub up to 3 m high, ranging from the Richtersveld,

northwestern Cape southwards to Worcester and

eastwards to East London.

R. keetii Schonland ( 1930), FSA 68.

After J.D.M. Keet (1882-1976), one-time Director of

Forestry in the Union Government.

Proposed common names: Keet’s slender currant, Keet-se-

taaibos.

Shrub up to 1.7 m high, occurring in the Waterberg,

northwestern Transvaal and in the Lydenburg Dist.,

eastern Transvaal.

R. kirkii Oliv. (1868), FSA 54.

After Sir John Kirk, pioneer naturalist and member of

Dr Livingstone’s 1858 Zambesi Expedition.

Proposed common names: Kirk’s currant, Kirk-se-taaibos.

Dwarf suffrutescent shrub up to 1 m high, occurring in

central Africa and just reaching as far south as the Capri vi

Strip, Namibia.

R. krebsiana Presl ex Engl. (1883), FSA 13.

After L. Krebs (1792-1844), plant collector and farmer

in the Bedford Dist. of the eastern Cape Province.

Common names: 385.1, false sour currant, bastersuur-

taaibos.

Shrub or small tree up to 3 m high, ranging from near

Graaff-Reinet to Loteni in the Natal Drakensberg.

R. kwazuluana Moffett (1993), FSA 33.

After KwaZulu, the place of the Zulu.

Suggested common names: Kwazulu dwarf currant, Kwa-

zuludwergtaaibos.

Dwarf suffrutescent shrub up to 1 m high, confined to

north eastern Zululand between Mkuzi, Sodwana Bay and

St Lucia.

* should be added to the national list.

R. laevigata L. ( 1763), FSA 1 1 .

(= R. mucronata Thunb.)

Latin for smooth and polished, referring to the shiny

smooth leaflets of var. laevigata.

Two varieties are recognized:

var. laevigata and var. villosa (L. f.) R. Fern. (= R. incana

Mill. & R. villosa L. f.) after the hairy leaves.

Common names: 385.2, dune currant, duinetaaibos.

var. laevigata has been divided into forma laevigata and

forma cangoana Moffett, after the Cango Caves, near

which it occurs.

Proposed common name for forma cangoana: Cango cur-

rant, Kangotaaibos.

Shrub up to 2.5 m high, distributed along the coast and

coastal foreland from Lambert’s Bay in the west to East

London and with a disjunct population near the Cango

Caves, Oudtshoom, Cape Province.

R. lancea L. f. ( 1 78 1 ), FSA 30.

Latin for a light spear or lance, referring to the shape of

the leaflets.

Common names: 386, karree, karee.

Large shrub or spreading trees, usually up to 6 in high, but

sometimes reaching 12 m. Widespread in the more arid

parts of the interior of southern Africa.

R. leptodictya Diels (1907), FSA 26.

(= R. amerina Meikle & R. gueinzii sensu Schon-

land)

Greek for fine or thin net, referring to the network of

veins in the leaflets.

Common names: 387, mountain karree, bergkaree.

Shrub or tree up to 10 m high, widespread over almost the

whole of the Transvaal, also occurring in the Orange Free

State, Botswana and Namibia.

R. longispina Eckl. & Zeyh. (1836), FSA 42.

Latin for long spines, referring to the spinous branch

ends.

Common names: 388, thorny currant, doringtaaibos.

Shrub up to 4 m high, widely distributed in the Karoo

between Worcester, Stutterheim and Queenstown. Also

found in Namaqualand.

R. lucens Hutch. (1946), FSA 27.

Latin for shining, polished, glistening, referring to the

leaflets.

Proposed common name: shiny-leaved rhus, as used in

Zimbabwe.*

Shrub or small tree up to 4.5 m high, found in the north-

eastern corner of Botswana and further in Zimbabwe and

Zambia.

R. lucida L. (1753), FSA 40.

Latin for shining, clear, transparent, referring to the

shiny smooth leaflets which are, however, not clear or

transparent.

Common names: 388.1, glossy currant, blinktaaibos.

Bothalia 24,1 (1994)

73

Shrub up to 3 m high, ranging from near Clanwilliam,

Cape in an eastward arc to the Soutpansberg in the northern

Transvaal and also Zimbabwe.

Three forms are distinguished:

forma lucida; forma scoparia (Eckl. & Zeyh.) Moffett (=

R. schlechteri Diels) after the genus Scoparia L. (Scro-

phulariaceae); and forma elliptica (Sond.) Moffett after

the elliptic shape of the leaflets.

R. macowanii Schonland (1930). See R. rehmanniana var.

glabrata.

R. magalismontana Sond. (1860), FSA 22.

(= R. cinerea R. Fern. & A. Fern.)

Latin for Magalies Mountains, referring to the Mag-

aliesberg range in southern Transvaal.

Dwarf shrublet or shrub up to 2.1 m high, occurring in the

Transvaal, northern Cape, Botswana and Zimbabwe.

Three subspecies are distinguished:

subsp. magalismontana

Proposed common names'. Magaliesberg dwarf currant,

Magaliesbergdwergtaaibos.

subsp. coddii (R. Fern. & A. Fern.) Moffett (= R. coddii

R. Fern. & A. Fern. & R. schliebenii R. & A. Fern.) after

L.E. Codd, former Director of the Botanical Research

Institute, Pretoria.

Proposed common names: Soutpansberg currant, Sout-

pansbergtaaibos.

subsp. trifoliolata (Baker f.) Moffett (= R. trifoliolata

Baker f.), after the three-digitate leaves.

R. maricoana Moffett ( 1993), FSA 66.

After Marico, the area where it occurs.

Proposed common names: Marico dwarf currant, Marico-

dwergtaaibos.

Dwarf shrublet up to 1 m high, confined to a small area

near Zeerust, western Transvaal.

R. marlothii Engl. (1888), FSA 24.

After H.W.R. Marloth (1855-1931), Cape Town based

pharmacist, analytical chemist and botanist who for three

years was also Professor of Chemistry at the Victoria

College, Stellenbosch.

Common names: 389.2, bitter karree, bitterkaree.

Shrub up to 2.2 m high, widespread in central Namibia.

R. microcarpa Schonland (1930). See R. pyroides var.

integrifolia.

R. montana Diels (1907), FSA 6.

Latin for mountains, referring to its mountainous hab-

itat.

Common names: 384.1, Drakensberg karree, Drakensberg

karee.

Shrub up to 2.2 m high, occurring in the foothills of the

* should be added to the national list.

Drakensberg between Engcobo, Transkei and Wakker-

stroom, Transvaal.

R. natalensis Bemh. ex Krauss (1844), FSA 37.

After the province of Natal.

Common names: 390, Natal karree, Natalkaree.

Semi-scandent shrub or slender tree up to 5 m high,

occurring in coastal scrub between East London and the

Zululand-Mozambique border.

R. nebulosa Schonland (1930), FSA 12.

Latin for cloudy, possibly referring to the nebulous or

cloud-like inflorescence.

Proposed common names: sand currant, sandtaaibos.*

Scandent shrub up to 4 m high, occurring in coastal scrub

from near Alexandria, Cape Province to Kosi Bay, Tonga-

land.

Two forms are distinguished:

forma nebulosa and forma pubescens Moffett after the

hairy leaflets.

R. pallens Eckl. & Zeyh. (1836), FSA 44.

Latin for pale, referring to the pallid leaflets.

Suggested common names: pale kuni-bush, bleekkoe-

niebos.*

Shrub or small tree up to 7 m high, ranging from the

southwestern Cape eastwards to Natal and also occurring

in the eastern OFS and southwestern Transvaal.

R. pendulina Jacq. ( 1 805), FSA 48.

(= R. viminalis sensu Schonland)

Latin for hanging down, referring to the pendulous or

drooping branches and leaves.

Common names: 396, white karree, witkaree.

Tree up to 10 m high, occurring naturally on the banks of

the Orange River between near Luckhoff in the OFS and

Oranjemund, Namibia. Also found on the banks of the

Olifants and Berg Rivers in the western Cape Province.

R. pentheri Zahlbr. (1900), FSA 34.

After A. Penther (1865-1931), Austrian zoologist and

collector.

Common names: 391, common crow-berry, gewone kraai-

bessie.

Shrub or tree up to 5 m high, ranging from Transkei to the

Soutpansberg, northern Transvaal.

R. pondoensis Schonland (1930), FSA 65.

After Pondoland, where the type specimen was col-

lected.

Proposed common names: many-veined currant, veel-

nerftaaibos.

Slender dwarf shrub up to 1 m high, occurring disjunctly

between Transkei and the northeastern Transvaal.

74

Bothalia 24,1 (1994)

R. populifolia E. Mey. ex Sond. ( 1860), FSA 61.

Latin for poplar leaves, referring to the shape of the

leaflets.

Proposed common names: Gariep currant, Garieptaai-

bos.*

Shrub up to 2.5 m high, occurring in the northwestern Cape

and southern Namibia, on either side of the Orange River,

in the area where it is known as the Gariep.

R. problematodes Merxm. & Roessler (1973), FSA 72.

Greek for resembling a problem, referring to the prob-

lems encountered by botanists in determining the taxo-

nomic relationships of this species.

Proposed common names: dwarf spiny currant, dwerg-

doringtaaibos.

Dwarf shrublet up to 0.6 m high, confined to the area

between Aus and Witpiitz, southern Namibia.

R. pterota Presl (1844), FSA 70.

(= R. longispina sensu Schonland)

Greek for winged, referring to the somewhat flattened

petioles.

Suggested common names: spike-thorn currant, pen-

doringtaaibos.*

Shrub usually up to 2 m high, rarely up to 4 m high, ranging

along the coast and adjacent interior from near Bredasdorp

to East London with an isolated population at Langebaan,

western Cape.

R. pyroides Burch. (1822), FSA 15.

Latin for resembling the pear tree (Pyrus), referring to

the pear-like leaflets.

Shrub or small tree up to 6 m high, occurring over virtually

the whole of South Africa excepting the western Cape

Province. Also scattered in Botswana and Namibia.

Four varieties are recognized:

var. pyroides (= R. baurii Schonland)

Common names: 392, common wild currant, gewone

taaibos.

var. dinteri (Engl.) Moffett (= R. dinteri Engl.) after M.K.

Dinter (1868-1945), pioneer South West African botanist.

Proposed common names: Dinter ’s currant, Dinter-se-

taaibos.*

var. gracilis (Engl.) Burtt Davy, after the slender branches.

Proposed common names: mountain currant, bergtaai-

bos.*

var. integrifolia (Engl.) Moffett, after the entire-margined

leaflets, (= R. fraseri Schonland, R. intermedia Schonland

& R. microcarpa Schonland). Despite the name many

specimens have indented margins.

Proposed common names: Natal currant, Nataltaaibos.*

R. quartiniana A. Rich. ( 1 847), FSA 1 6.

After R. Quartin-Dillon, French medical doctor and

naturalist who collected in Ethiopia between 1839 and

1843.

* should be added to the national list.

Common name: 393, glossy velvet karree, blinkfluweel-

karee.

Shrub or small tree up to 7 m high, occurring along river

banks from the northern parts of Botswana and Namibia

through Central Africa to Ethiopia.

R. refracta Eckl. & Zeyh. (1836), FSA 35.

Latin for bent or curved back abruptly, referring to the

almost horizontal pattern of branching.

Proposed common names: rough-leaved currant, growwe-

blaartaaibos.*

Shrub or small tree up to 4 m high, occurring in the eastern

Cape and Transkei.

R. rehmanniana Engl. (1883), FSA 18.

After A. Rehmann (1840-1917), Polish botanist and

collector.

Shrub or more commonly a tree up to 5 m high, ranging

from the southwestern Cape eastwards to the northern

Transvaal.

Two varieties are recognized:

var. rehmanniana

Common names: 393.1, blunt-leaved currant, stompblaar-

taaibos.

var. glabrata (Sond.) Moffett (= R. macowanii Schonland)

after the glabrous or hairless leaves.

Common names: 389, sour currant, suurtaaibos.

R. rigida Mill. (1768), FSA 10.

(= R. eckloniana Sond.)

Latin for rigid, referring to the fairly stiff leaflets.

Shrub up to 1 m high, ranging from the eastern Cape to the

northern Transvaal.

Three varieties are recognized:

var. rigida

Proposed common names: rock currant, kliptaaibos.

var. margaretae Burtt Davy ex Moffett, after Margaret

Moss, one-time curator of the C.E. Moss Herbarium,

Witwatersrand University.

Proposed common names: Margaret’s rock currant, Mar-

garet-se-kliptaaibos.

var. dentata (Engl.) Moffett (=R. dyeri R. Fem. & A. Fern.

& R. rupicola J.M. Wood & M.S. Evans) after the toothed

leaflets.

Proposed common names: Waterberg currant, Water-

bergtaaibos.

R. rimosa Eckl. & Zeyh. (1836), FSA 49.

(= R. rigida sensu Schonland)

Latin for full of cracks, chinks or fissures, referring to

the concentric cracks which develop in the bark of dried

branches.

Proposed common names: Cederberg currant, Sederberg-

taaibos.*

Shrub up to 3 m high, confined to the Cederberg Moun-

tains, western Cape Province.

Bothalia 24,1 (1994)

75

R. rogersii Schonland (1930), FSA9.

After Rev. F.A. Rogers (1876-1944), railway missioner

and Archdeacon of Pietersburg.

Proposed common names: Rogers’s currant, Rogers-se-

taaibos.

Shrub up to 2 m high, occurring in the eastern and north-

eastern Transvaal.

R. rosmarinifolia Vahl (1794), FSA 55.

Latin for rosemary leaves, referring to the rosemary-

like leaflets.

Proposed common names: rosemary currant, roosmaryn-

taaibos.

Dwarf shrublet up to 1 m high, ranging from the Ceder-

berg, southwestern Cape Province to the Suurberg, north

of Port Elizabeth.

R. rudatisii Engl. (1921), FSA 64.

After A.G.H. Rudatis (1875-1934), horticulturist and

farmer.

Proposed common names: Rudatis’s dwarf currant. Ruda-

tis-se dwergtaaibos.

Dwarf suffrutescent shrublet up to 0.35 m high, confined

to grassveld in southern Natal.

R. scytophylla Eckl. & Zeyh. ( 1 836), FSA 5 1 .

(= R. africana sensu Schonland)

Greek for leathery leaves, referring to the texture of the

leaflets.

Proposed common names: red-flowered currant, rooi-

blomtaaibos.

Shrub up to 2 m high occurring in the southwestern Cape

Province between Niewoudtville in the north and Caledon

in the south.

Two varieties are recognized:

var. scytophylla and var. dentata Moffett, after the

toothed leaflets.

R. sekhukhuniensis Moffett (1993), FSA 39.

After Sekhukhuniland, where it is endemic.

Proposed common names: Sekhukhune currant, Sekoe-

koenietaaibos.*

Shrub up to 3 m high, occurring between Lydenburg and

Pietersburg, northeastern Transvaal.

R. stenophylla Eckl. & Zeyh. (1836), FSA 56.

Greek for narrow leaves, referring to the leaflets.

Proposed common names: narrow-leaved currant, smal-

blaartaaibos.

Suffrutescent shrub up to 1 .2 m high, only found in the

southwestern Cape.

* should be added to the national list.

R. tenuinervis Engl. ( 1 883), FSA 25.

(= R. commiphoroides Engl. & Gilg)

Latin for thin-nerved, referring to the venation of the

leaflets.

Common names: 393.2, Kalahari currant, Kalaharitaaibos.

Shrub up to 3 m high, widely distributed in Botswana and

Namibia, also reaching the northern Cape and the western

Transvaal.

R. tomentosa L. (1753), FSA 58.

Latin for a thick and even covering of short matted

hairs, referring to the pale downy underside of the leaflets.

Common names: 394, real wild currant, korentebos.

Shrub or small tree up to 5 m high, ranging from the

Cederberg, southwestern Cape eastwards along moister

parts to the northern Transvaal and Zimbabwe.

R. transvaalensis Engl. (1883), FSA 4.

(= R. eburnea Schonland)

After the province of Transvaal.

Common names: 394.1, Transvaal currant, Transvaaltaai-

bos.

Shrub up to 2 m high, rarely a small tree reaching 4 m

occurring between northern Natal and the Soutpansberg,

northern Transvaal.

R. tridactyla Burch. (1822). FSA 28.

Greek and Latin for three fingers, referring to the

leaves.

Proposed common names: Burchell's sour karee, Burchell

se-suurkaree.*

Shrub up to 4 m high, occurring in the northern Cape.

R. tumulicola S. Moore ( 1921), FSA 1 .

(= R. dura Schonland. R. culminum R. Fern. & A.

Fern. & R. synstylica R. Fern. & A. Fern.)

Latin for inhabitant of the mounds or hills, referring to

the farm The Downs’ where the type specimen was col-

lected.

Shrub or small tree up to 3.5 m high, ranging from north-

ern Natal to the Soutpansberg, northern Transvaal and also

Mozambique and Zimbabwe.

Two varieties are recognized:

var. tumulicola

Common names: 38 1 .2, hard-leaved currant, hardetaaibos.

var. meeuseana (R. Fern. & A. Fern.) Moffett, after A.D.J.

Meeuse ( 1 9 1 4-), Dutch botanist and collector.

Proposed common names: velvet currant, fluweeltaaibos.

Two forms of this variety are recognized:

forma meeuseana (= R. ernestii Schonland) and forma

pumila Moffett, after the dwarf habit and small leaves.

R. undulata Jacq. (1798), FSA 45.

(= R. celastroides Sond. & R. excisa Thunb.)

Latin for wavy, referring to the undulate margins of the

leaflets.

Common names: 395. kuni-bush. koeniebos.

76

Bothalia24,l (1994)

Shrub up to 3 m high, occurring from the southern and

southwestern Cape northwards to southern Namibia.

R. volkii Suess. ( 1953), FSA 62.

After O.H. Volk ( 1903-), German botanist and collec-

tor.

Proposed common names : Volk’s currant, Volk-se-taaibos.

Shrub up to 1 .5 m high, confined to a small mountainous

area near Maltahohe, Namibia.

R. wilmsii Diels (1898), FSA 67.

After F. Wilms (1848-1919), German apothecary, bot-

anist and collector.

Proposed common names : Wilms’s dwarf currant, Wilms-

se dwergtaaibos.

Dwarf shrublet up to 0.5 m high, confined to a small area

near Lydenburg, Transvaal.

R. zeyheri Sond. ( 1 860), FSA 47.

(= R. glaucovirens Engl.)

After C.L.P. Zeyher (1799-1858), prolific botanical

collector.

Common names : 396.1, blue currant, bloutaaibos.

Shrub up to 2 m high, rarely a tree up to 4 m high,

distributed widely in the Transvaal interior.

REFERENCES

BAKER. E.G. 1911. Anacardiaceae. In Contribution to the flora of Gaza-

land. Botanical Journal of the Linnean Society 40: 48-50.

BARKLEY, F.A. & BARKLEY, E.D. 1938. A short history of Rhus to the

time of Linnaeus. American Midland Naturalist 19: 265-333.

BLEEK, D.F. 1956. A bushman dictionary. American Oriental Society,

New Haven.

BRITTEN, J. 1900. Notes on Rhus. Journal of Botany, London 38:

315-317.

BURCHELL, W.J. 1822. Travels in the interior of southern Africa. Vol. 1.

Longman, London.

COATES PALGRAVE, K. 1977. Trees of southern Africa. Struik, Cape

Town.

CODD, L.E. 1956. Anacardiaceae. In New and interesting records of

southern African flowering plants. Bothalia 6: 539, 540.

DE CANDOLLE, A.P. 1825. Prodromus systematis naturalis regni veg-

etabilis Vol. 2. Treuttel & Wiirtz, Amsterdam.

DIELS, F.L.E. 1898. Die Epharmose der Vegetationsorgane bei Rhus L.

Gerontogeae Engl. Botanische Jahrbiicher 24: 568-647.

DIELS, F.L.E. 1907. Anacardiaceae africanae. 4. Botanische Jahrbiicher

40: 86, 87.

DON, G. 1832. A general system of gardening and botany containing a

complete enumeration and description of all plants hitherto

known. Vol. 2. London.

ECKLON, C.F. & ZEYHER, C.L.P. 1 836. Enumeratio plantarum qfricae

australis extra-tropicae ... Part 2. Hamburg.

ENGLER, H.G.A. 1883. Anacardiaceae. In A.L.P.P. & A.C.P. de Can-

dolle, Monographiae Phane rogamarum. Vol. 4.

ENGLER, H.G.A. 1888. Anacardiaceae. In Plantae Marlothianae.

Botanische Jahrbiicher 10: 37, 38.

ENGLER, H.G.A. 1921. Die Pflanzenwelt Afrikas, inbesondere seiner

tropischen Gebiete. Vol. 3,2. Engelmann, Leipzig.

ESTERIIUYSE. C.J. 1983. Karee, tree of the year. Pamphlet 304. De-

partment of Environment Affairs, Pretoria.

GUNN, M. & CODD, L.E. 1981. Botanical exploration of southern

Africa. Balkema, Cape Town.

HUTCHINSON, J. 1946. A botanist in South Africa. Gawthorn, London.

JACQUIN, N.J. VON 1798. Plantarum rariorum horti caesarei schoen-

brunnensis: descriptiones et icones. Vol. 1. Wappler, Vienna.

JACQUIN, N.J. VON 1805. Plantarum rariorum horti caesarei schoen-

brunnensis: descriptiones et icones. Vol. 4. Wappler, Vienna.

KOCH, C. 1853. Hortus dendrologicus. Schneider, Berlin.

KRAUSS, F. 1844. Pflanzen des Cap- und NatalLandes, gesammelt und

zusammengestellt von Dr. Ferdinand Krauss. Flora 27: 346-359.

LEISTNER, O.A. & MORRIS, J.W. 1976. South African place names.

Annals of the Cape Provincial Museums 12.

LINDLEY, J. 1 830. Introduction to the natural system of botany. London.

LINNAEUS, C. 1753. Species plantarum. Vol. 1. Stockholm. Facsimile

edn, 1957. Ray Society, London.

LINNAEUS, C. 1763. Species plantarum, 2nd edn, Vol. 2. Stockholm.

LINNAEUS, C. VON (filius). 1781. Supplementum plantarum. Bruns-

wick.

MARCHAND, N.L. 1 869. Revision du groupe des Anacardiacees. J.-B.

Bailliere, Paris.

MCNAIR, J.B. 1925. The taxonomy of poison ivy. Field Museum of

Natural History, Botanical Series 4: 55, 56.

MERXMULLER, H. & ROESSLER, H. 1973. Eine aussergewohnliche

neue Anacardiacee aus Siidwestafrika. Mitteilungen der Botani-

schen Staatssammlung, Miinchen 11: 62-82.

MILLER, P. 1768. The Gardener's dictionary, 8th edn. London.

MOFFETT, R.O. 1988. Rhus acocksii (Anacardiaceae), yet another new

endemic from the Mtamvuna area. South African Journal of Bot-

any 54: 172-174.

MOFFETT, R.O. 1993. Rhus. In O.A. Leistner & E. du Plessis, The Flora

of southern Africa 19,3,1: 1-129.

MOORE, S. LE M. 1921. Anacardiaceae. In Plantae Rogersianae 6.

Journal of Botany, London. 59: 227.

NIENABER, G.S. 1963. Hottentots. Van Schaik, Pretoria.

OLIVER, D. 1868. Anacardiaceae. In Flora of tropical Africa. Vol. 1.

Reeve, London.

ONIONS, C.T. 1956. (ed.). The shorter Oxford English dictionary on

historical principles, 3rd revised edn. Oxford University Press,

London.

PALMER, E.M. & PITMAN. N. 1973. Trees of southern Africa. Vol. 2.

Balkema, Cape Town.

PERSOON. C.H. 1 805. Synopsis plantarum, seu Enchiridium botanicum.

Vol. I. Cramer, Paris.

PETTMAN, C. 1931. South African place names : past and present.

Queenstown Daily Representative, Queenstown.

PRESL, K.B. 1844. Botanische Bemerkungen. Gottlieb Haase, Prague.

RICHARD, A. 1847. Tentamen florae abyssinicae. Vol. 4. Paris.

SCHONLAND, S. 1930. The South African species of Rhus L. Bothalia

3: 3-115.

SCHULTES, J.A. 1820. Pentandra trigynia. In Systema vegetabilium,

15th edn. Stuttgart.

SMITH, C.A. 1966. Common names of South African plants. Memoirs

of the Botanical Survey of South Africa No. 35.

SONDER, O.W. 1860. Terebintaceae. In Flora capensis. Hodges &

Smith, Dublin.

SUESSENGUTH, K. 1953. Neue Taxa, Kombinationen und Vorkommen

in Siidafrika. Mitteilungen der Botanischen Staatssammlung,

Miinchen 8: 343.

THLINBERG, C.P. 1794. Prodromus plantarum capensium. Part 1. Upp-

sala.

THUNBERG, C.P. 1803. Novae species plantarum capensium ex-

aminatae et descriptae. In D.G.F. Hoffman, Phytographische

Blatter. Schroder, Gottingen.

THUNBERG, C.P. 1818. Flora capensis. Vol. 2,1. Bonnier, Copenhagen.

VAHL, M. 1794. Symbolae botanicae. Part 3. Copenhagen.

VON BREITENBACH, F. 1983. Karree and wild currant: a review of the .

arborescent members of the genus Rhus L. in southern Africa.

Journal of Dendrology 3,3 & 4: 99-160.

VON BREITENBACH, F. 1990. National list of indigenous trees. Den-

drological Foundation, Pretoria.

ZAHLBRUCKNER, A. 1900. Anacardiaceae. In Plantae Pentherianae.

Anndten des Naturhistorischen Museums in Wien 15: 52-54.

Bothalia 24,1: 77-90(1994)

Flowering phenology in the arid winter rainfall region of southern Africa

M. STRUCK*

Keywords: climatic variables, flowering sequence, Namaqualand, phenology, precipitation, seasonality, Succulent Karoo, temperature, timing, vigour

of blooming

ABSTRACT

The impact of physical factors on the flowering phenology of a succulent karroid community in the winter rainfall region of

the northwestern Cape, South Africa, based upon a three year study on permanent plots, is examined. On the permanent plots,

flowering of the shrubby species extended over a period of 4 to 4Vl months each year, while blooming of the therophytes peaked

in the first half of the flowering season. Species composition and numbers of individuals in the therophytes and geophytes

offering flowers varied greatly according to the pattern and amount of seasonal precipitation. Despite these variations a

consistent flowering sequence between the years was observed. Possible relations between the flowering phenology and the

climatic variables are discussed in detail. The present data suggest that the onset of flowering is determined indirectly by the first

drop in temperature in autumn, indicating the beginning of the rainy season and presumably the start of the growing period,

and/or by the increase of temperatures in the beginning of spring. The pattern and amount of rainfall within a given season

mainly influenced the duration of anthesis and the number of flowers produced.

UITTREKSEL

Die invloed van fisiese faktore op die blomfenologie van ’n sukkulente karoogemeenskap in die winterreengebied van die

Noordwes-Kaap, Suid-Afrika, gebaseer op ’n driejaar-studie op permanente persele, is ondersoek. Die blomperiode van die

struikagtige spesies op die permanente persele was versprei oor ’n periode van 4 tot 4'/2 maande elke jaar, terwyl die

blomperiode by die terofiete gedurende die eerste helfte van die blomseisoen ’n spits bereik het. Spesiesamestelling en aantal

individue by die terofiete en geofiete wat blom, is sterk beinvloed deur die patroon en hoeveelheid van die jaarlikse neerslag.

Ondanks hierdie variasies is "n reelmatige blomopeenvolging oor die drie jaar waargeneem. Moontlike verhoudings tussen die

blomfenologie en die klimaatveranderlikes word in besonderhede bespreek. Die huidige data dui daarop dat die begin van die

blomseisoen indirek bepaal word deur 'n afname in temperatuur gedurende die herfs, wat die begin van die reenseisoen en

moontlik die begin van die groeiseisoen aandui, en/of deur die toename in temperatuur aan die begin van die lente. Die patroon

en hoeveelheid reenval binne 'n gegewe seisoen beinvloed hoofsaaklik die duur van antese en die aantal blomme wat gevorm

word.

INTRODUCTION

A short but copious blooming is a characteristic feature

of many arid and semi-arid regions. This is particularly

true of Namaqualand in the northwestern Cape, South Af-

rica, which is renowned for its flower displays during

springtime.

For most plant species the growing season is restricted

to the cool and moist autumn-winter-spring period and the

dormant season occurs during the hot and dry summer

months. The growing period usually commences in au-

tumn (March to April) with the sprouting of the perennials

and the germination of the therophytes (Van Rooyen et

«/. 1979; Le Roux et al. 1989). The flowering season lasts

from late winter to spring (August to early October). Dur-

ing this period 90% of the shrubby perennials and virtually

all the annuals are blooming (Le Roux et al. 1989). In

comparison to other regions of winter rainfall climate an-

thesis is markedly synchronized (Orshan et al. 1989).

The overall influence of the climatic factors on the phe-

nological events in arid regions is obvious. With respect

to the phenology of flowering, special attention has been

given to the timing and synchrony of blooming in relation

* Botany Department, University of Cape Town, Private Bag, Ronde-

bosch7700.

to the influence of climatic variables (e.g. Baker et al.

1982; Beatley 1974; Noy-Meir 1973; Solbrig & Yang

1977). Generally, the importance of an opportunistic re-

sponse by the plants to water availability has been

stressed. Compared to other arid areas receiving a similar

amount of average rainfall, the predictability of seasonal

precipitation is relatively high in the arid winter rain-

fall region of the Cape Province (Hoffman & Cowling

1987). Hence, the phenology of the vegetation should

be comparatively predictable, too. However, the question

of the impact of current seasonal weather conditions on

the flowering phenology has not yet been addressed ade-

quately.

Against this background, results of a three year study

of the flowering phenology of a succulent karroid com-

munity of the northwestern Cape are presented. The fol-

lowing questions shall be addressed in particular: 1, how

does the timing and vigour of flowering vary between the

years?; and 2, can the phenological pattern be linked to

the current rainfall and/or temperature conditions, thus

indicating potential environmental cues? The present sur-

vey is clearly constrained by the limited number of plots

and years studied, and the data do not provide a robust

basis for a thorough statistical evaluation. Hence, the fol-

lowing observations should be regarded as indicating

trends.

78

Bothalia 24,1 (1994)

METHODS

The field work was carried out at the Goegap Nature

Reserve during the blooming seasons of the years 1985

(late June to October), 1986 (July to late November), and

1987 (mid-August to mid-November).

The records of the actual temperatures and precipita-

tion at Goegap were provided by the authorities of Goegap

Nature Reserve. The data were supplemented by the au-

thor with the aid of a thermohygrograph.

Phenological data were collected for a total of 112

plant species on six permanent plots of 1 00 m2, each. Five

plots were laid out on various sites on a mountain slope

with undisturbed dwarf-scrub vegetation so as to cover

some of the conspicuous variation in species composition.

One plot was marked on a sandy plain with exclusively

ephemeral vegetation showing a particularly high degree

of variation in yearly species composition and plant cover.

The permanent plots are characterized in an Appendix.

For each species onset and duration of anthesis were

recorded. Furthermore, numbers of open flowers or in-

florescences with open flowers were repeatedly counted

at time intervals of three to five (rarely up to seven) days.

Inflorescences and partial inflorescences representing the

functional unit of visitor attraction or pollination (i.e. blos-

soms sensu Faegri & Van der Pijl 1979) were taken as

equivalent to single flowers.

Plant names are mostly according to Gibbs Russell et

al. (1985, 1987).

STUDY SITE

Description of the study site

The Goegap Nature Reserve is situated 12 km east of

Springbok (between S 29°34' and 29°41' and E 17°57'

and 18°02/) in the Namaqualand Rocky Hills in the north-

western Cape, South Africa.

The area is characterized by an open dwarf-scrub vege-

tation which has been classified as ‘Namaqualand Broken

Veld' by Acocks (1988) and as part of the Succulent Karoo

Biome by Rutherford & Westfall ( 1986). Succulent and nar-

row-leaved sclerophyllous species abound. The flora is

clearly dominated by Asteraceae and Mesembryanthemaceae

(Le Roux 1984).

The study area is situated in the realm of the winter

rainfall region. About 70% of the yearly precipitation is

received from April to October and about 38% during the

wettest quarter of the year (June to August) (Table 1).

Drought conditions prevail virtually throughout the year.

All climatic factors are subject to conspicuous annual fluc-

tuations.

Weather conditions during the study

The amount of precipitation as well as the timing and

number of rain events varied considerably between the

study years (Figure 1 ; Tables 1 & 2). Annual rainfall was

highest in 1985 (244.5 mm, i.e. 73% above the average

for the years 1974 to 1987), but much lower in both of

the subsequent years (110.5 mm in 1986, i.e. 22% below

the average and 113.5 mm in 1987, i.e. 20 % below the

average). However, rainfall figures for the main growth

period (April to October) and wettest quarter of a year

(June to August) provide quite a different picture (Tables

I & 2). During the main growth period, 87 mm was re-

ceived in 1985 (35.6% of the year’s total), 81.5 mm in

1986 (73.7% of the year’s total) and 113.5 mm in 1987

(100% of the year’s total). The maximum number of rainy

days was recorded in 1986, especially between June and

mid-September. The relatively high rainfall of 1987 was

due to several exceptionally good rains during April, June,

and August. In contrast, in 1985 drought conditions pre-

vailed in June and during August to September with only

few intermediate rains.

In comparison to the rainfall pattern the annual course

of air temperatures was rather uniform (Figure 1). In each

year, winter minimum values were reached during a pe-

TABLE 1. — Rainfall (mm) in Goegab Nature Reserve 1974-1987, caleulated from data from Le Roux (1984) and unpublished weather data of the

Reserve

Bothalia 24,1 (1994)

79

Weekly

temperatures

absolute maximum

mean maximum

mean

mean minimum

absolute minimum

Temperatures for 7-day

intervals not corresponding

to calender weeks

FIGURE 1. — Rainfall and temperature conditions in Goegab Nature Reserve during the study period, A. autumnal drop in temperature; yearly

flowering period; i, rainfall (further details see text).

riod of about five months, after which temperatures in- the cool season appeared to shift from year to year. In

creased again during a period of one to two months to 1985 the mean monthly temperatures dropped consider-

reach summer values (Table 3). Nevertheless, the onset of ably during March, but only during April in both of the

80

Bothalia 24,1 (1994)

TABLE 2.-Monthly rainfall and number of rainy days (in brackets) dur-

ing the main growth period (April to October) 1985 to 1987

subsequent years. The coldest quarter of a year was June

to August. Mean and absolute minima occurred during

July in 1985 (13.8 / 1.0°C) and 1986 (12.8 / -1.0°C), but

during August in 1987 (1 1.6 / 0.5°C). Mean and absolute

maxima occurred during February (26.3 / 48.0°C) in 1985,

during December (28.2 / 45.5°C) in 1986, and during Jan-

uary (26.2 / 48.0°C) in 1987.

RESULTS AND DISCUSSION

Numbers of species in flower

In total, 112 species were observed in flower on the

permanent plots during the present study. Of these, 44%

flowered in 1985, 95% in 1986, and 79% in 1987. These

variations were mainly attributable to the phenological

performance of therophytes and geophytes (Table 4).

The number of therophytic species flowering in a given

year seems to be related to the amount and distribution

of rainfall occurring during the two months before the

onset of the flowering season. In 1985 only 25.5 mm fell

during this period, whereas in both of the subsequent years

the figure was nearly twice as high (1986: 46 mm, 1987:

50.5 mm; Table 2; Figure 1).

In 1985, the flowering was also affected by unfavour-

able weather conditions during the flowering period itself,

as shown in Figure 2. In that year, a dry period set in

after the initial rains at the onset of the blooming season

(Figure 1 and Table 2) causing drought stress particularly

for the therophytes. A good rain in mid-September, how-

ever, stimulated active growth and flowering.

Very few plant species growing on the permanent plots

flowered during summer. These are Othonna furcata

(March and April), Tylecodon wallichii (end of November

to beginning of December), the therophyte Tribulus ter-

restris (April), and the hysteranthous geophyte Erio-

spermum paradoxum (March and April; all dates after Van

Rooyen et al. 1979).

Timing and sequence of flowering

In each year, the blooming season lasted 4 to 4lA

months. However, the exact timing varied markedly be-

tween the years. In 1985, most plants flowered from early

July until the end of October, whereas in both of the sub-

sequent years blooming commenced about four weeks

later, in early August, and continued until the end of No-

vember (Figure 1).

TABLE 3.-Temperature regime in Goegab during the study period 1985 to 1987. Data from unpublished weather report Goegab Nature Reserve

and own measurements by thermohygrograph

x, mean monthly temperature; x (max.), mean maxima; x (min.), mean minima; abs. max., absolute maxima; abs. min., absolute minima; d, differ-

ences between means of consecutive months.

Bothalia 24,1 (1994)

81

TABLE 4. — Number of flowering species observed on the permanent

plots listed according to their life form

Despite the temporal fluctuation in the onset of the

flowering season the sequence of flowering of the species

involved remained nearly constant (see flowering se-

quence for plot 1 in Figure 3), i.e. independent of the

current pattern of precipitation in a given year. Such con-

sistent sequences of flowering phenologies have been de-

scribed from areas of virtually all climatic zones (Heinrich

1976; Heithaus 1974; Hocking 1968; Kratochwil 1984;

Mooney et al. 1974; Pierce 1984; Reader 1975; Stiles

1977). Considerable yearly variations in flowering dates,

but unaltered flowering sequences have also been reported

from southwestern Spain (Arroyo 1990). In Goegab the

shrubby species make up the 'backbone' of the sequence,

while therophytes and geophytes close the ranks in sea-

sons when their specific requirements in terms of moisture

and temperature conditions are met. This basic pattern re-

mained unchanged, though the vigour of blooming varied

greatly between years (see below). Taxonomically related

species usually flowered sequentially, with their flowering

periods slightly overlapping (e.g. Euphorbia, Hermannia,

Ruschia spp.; for further details see Struck 1992).

In the present observations no recurring pattern be-

tween the timing and/or amount of precipitation and the

timing of flowering in single species or the timing of the

blooming season in general were detected, though the vig-

our of flowering appeared to be stimulated by single rain-

fall events in single cases (see below).

Moreover, the present data do not indicate that flow-

ering of the therophytes was stimulated by drought. This

is in contrast to a view widely accepted for desert annuals

(Fox 1990a, b; Rathcke & Lacey 1985). Yet, recent ex-

perimental and field demographic studies (Aronson et al.

1992; Fox 1989, 1990b; Van Rooyen et al. 1991) indicate

that water stress had little or no effect on the induction

of flowering.

Conversely, a correlation was found between the au-

tumnal drop in temperatures and the onset of the blooming

season: as mentioned above, mean monthly temperatures

dropped considerably during March in 1985. but only dur-

ing April in both of the subsequent years (Table 3). The

blooming season started about three months later in each

year, namely in early July in 1985 and about early August

in both of the subsequent years. It should be noted that

the amount and distribution of rainfall during autumn dif-

fered notably between the years (Figure 1). The autumnal

drop in temperature corresponds to the beginning of the

winter rainy season and presumably marks the onset of

the growing period for the plants (Van Rooyen et al.

1979).

With regard to the weekly temperatures (or mean tem-

peratures of 7-day-intervals not corresponding to calendar

weeks) during autumn (Figure 1), the first cool interval

occurred in mid-March in 1985, during the second week

of April in 1986, and about the end of April in 1987 (with

5.0, 5.9, and 3.1°C below the means of the respective

months). These temperature events shifted for about 25

days between 1985 and 1986 and for about 10 days be-

tween 1986 and 1987.

This situation is reflected in the yearly shift of flow-

ering dates of selected perennial species (shift 1985/86:

about 3 weeks, 1986/87: 4 to 7 days, see Table 5). Though

the data show a high degree of variation, the differences

appeared to be significant. In contrast, the average shift

in flowering dates for selected annuals was 14 and 7 days,

respectively. Yet, due to the generally low proportion of

annuals in the plots, particularly during drier blooming

seasons, the results remain ambiguous (Table 5). However,

the lesser yearly shift of flowering dates of annuals may

also indicate other influences, e.g. a delay of flowering

by low temperatures during the winter period (Van

Rooyen et al. 1979). Low temperatures have been shown

to lengthen the time between flower initiation and anthesis

in therophytes (Van Rooyen et al. 1991). One could expect

such an effect for 1985 and 1987 when the cool winter

period overlapped with the onset of the blooming period,

i.e. when most annuals started to flower. Given this, the

differences in flowering dates should be less between

1985/86, but more between 1986/87 compared to the per-

ennials. This is what Table 5 indicates. However, the pos-

sibility that this is merely coincidence, cannot be ruled

out.

In some species flowering dates differed conspicuously

in various places of the study area (e.g. up to a month in

Ruschia robusta). Similar observations have also been re-

ported from other arid regions (e.g. Turner & Randall

1987 and references cited therein). Whether these differ-

ences reflect small-scale variations in soil conditions or

intra-specific genetic variability, is not known.

EIGURE 2. — Perccnluge ol species flowering simultaneously (cumula-

tive for all plots, 100%= 1 12 species). Solid line= 1985: stippled

line = 1986; dotted line = 1987.

82

Bothalia 24.1 ( 1994)

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Bothalia 24,1 (1994)

83

TABLE 5. — Yearly differences between dates of initial flowering and dates of peak bloom for selected species. Species poorly represented in the

study sites, including all geophytes, were omitted

n, sample size; x, mean; s, standard deviation; V, coefficients of variation, U, significance after Mann-Whitney U test.

Some of the species under present study were reported

to flower one to two months earlier (in 1974, Van Rooyen

et al. 1979) or later (in 1981, Le Roux et al. 1989) than

observed during the present survey. Nevertheless, the se-

quence of flowering of the species involved generally

matches the present results. Unfortunately, a further com-

parison is not possible, because the authors collected phe-

nological data in a notably larger area, notwithstanding

the small-scale variations of flowering dates. Besides, rel-

evant weather data are lacking for the above-mentioned

years.

Numbers of flowers and duration of flowering

The great majority of the observed species produce

large numbers of flowers over a time span of several

weeks (‘cornucopia’. Gentry 1974). In these species the

maximum numbers of flowers (or inflorescences, if re-

garded as the functional unit of visitor attraction or pol-

lination) offered simultaneously, varied from a few

thousand to tens of thousands (up to 40 000 per 100 m2

in Galenia sarcophylla [Aizoaceae] on plot 6 in 1986).

Conversely, all observed geophytes as well as some

chamaephytes and therophytes produced but a few flowers

per day over a period of several weeks or more (‘steady

state’. Gentry 1974). Of these the perennial climber

Microloma sagittatum (Asclepiadaceae) showed the long-

est flowering period ( 120 days). A transitional phenolog-

ical pattern was shown by some species which produced

moderate numbers of flowers over an extensive period

(e.g. the succulent shrublet Crassula muscosa subsp.

muscosa flowering more than 70 days, the therophytic Os-

teospermum hyoseroides and O. amplectens flowering for

95 days).

As already mentioned, blooming of the therophytes

was generally rather poor in 1985, but was markedly better

in 1986 and 1987. The shrubby species responded rather

differently to seasonal moisture availability: generally,

non-succulent shrubs showed the highest production of

flowers in the moist years 1986 (most Asteraceae) and

1987 (Lebeckia sericea [Fabaceae]), whereas succulents

(most Mesembryanthemaceae, Euphorbia spp.) flowered

most prolifically in the comparatively dry season of 1985.

Beside the seasonal moisture availability, single rainfall

events appeared to stimulate the vigour of flowering in

certain cases. An example is the good rain in early Sep-

tember 1985, which ended a short drought period (Figure

1). After that shower, several therophytes and chamae-

phytes produced a further minor bloom (Figure 3). Given

a causal relationship between these incidents, the species

did, nevertheless, not respond to similar rain events (in

terms of amount and timing of precipitation) in the sub-

sequent years.

Furthermore, ‘small rainfall events’ ( sensu Sala &

Lauenroth 1982) seemed to stimulate flowering in thero-

phytes: several species in plot 6 (e.g. Senecio arenarius,

Leysera tenella ) showed a slight, though remarkable in-

crease in the numbers of flowers after sparse rains (below

5 mm) occurring during September and the beginning of

October 1986 and 1987. This is in contrast to the findings

of various authors (for references see Sala & Lauenroth

1982) who only regard rain events of at least 8-10 mm

(‘effective rainfall event’ sensu Noy-Meir 1973) as eco-

logically significant.

The duration of flowering of most taxa ranged from

10 to 80 days. Only the geophytes showed short flowering

periods of 20 to 30 days in all three years. Due to the

adverse climatic conditions during the growth period of

1985, most therophytes had a comparatively short flow-

ering period (and low flower numbers) in that year. On

the other hand, duration of flowering of most therophytes

was clearly longer in 1987 than in 1986 (the year with

the maximum number of flowering therophytic species,

see Table 4).

No specific pattern of skewness in the flowering

phenologies could be detected: flowering may begin

abruptly and then tail off in one year but may start slowly

and end more quickly in another year. In species showing

the ‘cornucopia’ type of flowering pattern the duration of

full bloom (defined as the time interval with at least 50%

of a species’ maximum number of flowers in anthesis)

was longest in the year with the highest numbers of flow-

ers and shortest in the year with the lowest numbers of

flowers, i.e. a lower flower production was not compen-

sated for by an extended duration of flowering.

84

Bothalia24,l (1994)

CONCLUSIONS

A high degree of phenological variation on the species

level, and to a lesser extent between populations, is an

outstanding feature of the plant community studied. The

present observations revealed that the timing of flowering

and the vigour of blooming varied independently within

the species under study. Conversely, the flowering se-

quence of given species remained largely unaltered during

the years.

Regarding the possible impact of climatic variables, it

is inferred that the timing of flowering is indirectly deter-

mined by the first considerable drop in air temperatures

in autumn and/or by the increase of temperatures in the

beginning of spring. In contrast, species composition and

number of individuals of therophytes and geophytes in

flower are greatly affected by the pattern and amount of

seasonal precipitation. The same applies to the number of

blossoms produced. Due to fluctuations in seasonal rain-

fall the vigour of a species’ blooming may be promoted

in one year and retarded in another. Yet, potential pheno-

logical responses seem also to be related to morphological

and physiological constrains: e.g. non-succulent shrubs

produced the greatest number of flowers in moister years,

whereas succulents flowered most prolifically in a dry

year.

ACKNOWLEDGEMENTS

This paper is based on a dissertation submitted in par-

tial fulfilment of the requirements of the Dr rer. nat. at

the University of Hamburg (Germany). I am indebted to

my supervisor. Prof. Dr H.-D. Ihlenfeldt for his encour-

agement and constant support. I would like to express my

thanks to the directors of the Department of Nature and

Environmental Conservation in Cape Town and Jonkers-

hoek for permission to work in the Goegap Nature Re-

serve and the Officers in Charge for logistical support

during my stays. For identifications of plant specimens I

appreciate the assistance of Dr V. Bittrich, Dr N. Jurgens,

A. le Roux, and Dr K. Steiner. Special thanks are due to

A. and G. Cloete, C. Eardley, Dr S.K. Gess, Dr F.W. Gess,

E. van Jaarsveld, A. le Roux and Dr V. Whitehead for

their generous support during my stays in South Africa.

Furthermore, I would like to thank Drs M. Gerbaulet, S.K.

Gess, F.W. Gess, and unknown referees for comments on

drafts of the manuscript. E. van Jaarsveld and E. du Plessis

provided the Afrikaans version of the abstract. The work

was funded in part by the Deutsche Forschungs- gemein-

schaft and by the Akademisches Auslandsamt of the Uni-

versity of Hamburg, for which I am very grateful.

REFERENCES

ACOCKS, J PH. 1988. Veld Types of South Africa, 3rd edn. Memoirs of

the Botanical Survey of South Africa No. 57.

ARONSON, J„ KIGEL, J„ SHMIDA, A. & KLEIN, J. 1992. Adaptive

phenology of desert and mediterranean populations of annual

plants grown with and without water stress. Oecologia 89: 17-26.

ARROYO, J. 1990. Spatial variation of flowering phenology in the

mediterranean shrublands of southern Spain. Israel Journal of

Botany 39: 249-262.

BAKER, G.A., RUNDEL, RW. & PARSONS, D.J. 1982. Comparative

phenology and growth in three chaparral shrubs. Botanical Ga-

zette 143: 94-100.

BEATLEY, J.C. 1974. Phenological events and their environmental trig-

gers in Mojave Desert ecosystems. Ecology 55: 856-863.

FAEGRI, K. & VAN DER P1JL, L. 1979. The principles of pollination

biology, 3rd edn. Pergamon Press, Oxford.

FOX, G.A. 1989. Consequences of flowering-time variation in a desert

annual: adaptation and history. Ecology 70: 1294-1306.

FOX, G.A. 1990a. Drought and the evolution of flowering time in desert

annuals. American Journal of Botany 77: 1508-1518.

FOX, G.A. 1990b. Components of flowering time variation in a desert

annual. Evolution 44: 1404-1423.

GENTRY, A.H. 1974. Co-evolutionary patterns in Central American

Bignoniaceae. Annals of the Missouri Botanical Garden 61: 728-

759.

GIBBS RUSSELL, G.E., REID, C., VAN ROOY, J. & SMOOK, L. 1985.

List of species of southern African plants, 2nd edn, part 1. Mem-

oirs of the Botanical Survey of South Africa No. 5 1 .

GIBBS RUSSELL, G.E., WELMAN, W.G., RETIEF, E„ 1MMELMAN,

K.L., GERMISHUIZEN, G„ PIENAAR, B.J., VAN WYK, M. &

NICHOLAS, A. 1987. List of species of southern African plants,

2nd edn, part 2. Memoirs of the Botanical Survey of South Africa

No 56.

HEINRICH, B. 1976. Flowering phenologies: bog, woodland, and dis-

turbed habitats. Ecology 57: 890-899.

HEITHAUS, E.R. 1974. The role of plant pollinator interaction in deter-

mining community structure. Annals of the Missouri Botanical

Garden 6 1 : 675-691 .

HOCKING, B. 1968. Insect-flower associations in the high Arctic with

special reference to nectar. Oikos 19: 359-388.

HOFFMAN, M.T. & COWLING. R.M. 1987. Plant physiognomy, phe-

nology and demography. In R.M. Cowling & PW. Roux, The

Karoo Biome: a preliminary synthesis. Part 2 — vegetation and

history. South African National Scientific Programmes Report

No. 142: 1-34.

KRATOCHWIL, A. 1984. Pflanzengesellschaften und Bliitenbesucher-

Gemeinschaften: biozonologische Untersuchungen in einem

nicht mehr bewirtschafteten Halbtrockenrasen (Mesobrometum)

im Kaiserstuhl (Siidwestdeutschland). Phvtocoenologia 11: 455-

669.

LE ROUX, A. 1984. 'n Fitososiologiese studie van die Hester Malan-

Natuurreservaat. M.Sc. thesis, University of Pretoria.

LE ROUX, A., PERRY, P. & KYRIACOU, X. 1989. South Africa. In G.

Orshan, Plant phenomorphological studies in mediterranean type

ecosystems. Kluwer Academic Publishers, Dordrecht.

MOONEY, H.A., PARSONS, D.J. & KUMMEROW, J. 1974. Plant

development in mediterranean climates. In H. Lieth, Phenology

and seasonality modeling. Ecological Studies 8: 255-268.

Springer Verlag, New York.

NOY-MEIR, I. 1973. Desert ecosystems: environment and producers.

Annual Review of Ecology and Systematics 4: 25-51.

ORSHAN, G„ FLORET, Ch„ LE FLOC'H, E„ LE ROUX, A., MONTE-

NEGRO, G. & ROMANE, F. 1989. General synthesis. In G.

Orshan. Plant phenomorphological studies in mediterranean type

ecosystems : 389-399. Kluwer Academic Publishers, Dordrecht.

PIERCE, S.M. 1 984. A synthesis of plant phenology in the Fynbos Biome.

South African National Scientific Programmes Report No. 88.

Pretoria.

RATHCKE, B. & LACEY, E.P 1985. Phenological patterns of terrestrial

plants. Annual Review of Ecology and Systematics 16: 179-214.

READER. R.J. 1975. Competitive relationships of some bog ericads for

major insect pollinators. Canadian Journal of Botany 53: 1 300—

1305.

RUTHERFORD, M.C. & WESTFALL, R.H. 1986. Biomes of southern

Africa — an objective categorization. Memoirs of the Botanical

Survey of South Africa No. 54.

SALA, O.E. & LAUENROTH, W.K. 1982. Small rainfall events: an

ecological role in semi-arid regions. Oecologia (Berl.) 53: 30 1 —

304.

SOLBRIG, O.T. & YANG, T.W. 1977. Strategies in relation to timing of

blooming and fruiting. In G.H. Orians & O.T. Solbrig, Conver-

gent evolution in warm deserts, US/IBP Synthesis Series 3: 83-90.

STILES. F.G. 1977. Ecology, flowering phenology, and humming bird

pollination of some Costa Rican Heliconia. Ecology 56: 285-302.

STRUCK, M. 1992. Pollination ecology in the arid winter rainfall region

of southern Africa: a case study. Mitteilungen aus dem Institut fiir

Allgemeine Botanik Hamburg 24: 61-90.

TURNER, F.B. & RANDALL, D.C. 1987. The phenology of desert

shrubs in southern Nevada. Journal of Arid Environments 13:

119-128.

VAN ROOYEN, M.W., THERON, G.K. & GROBBELAAR, N. 1979.

Phenology of the vegetation in the Hester Malan Nature Reserve

in the Namaqualand Broken Veld: 1. General observations. Jour-

nal of South African Botany 45: 279-293.

VAN ROOYEN, M.W., GROBBELAAR, N„ THERON, G.K. & VAN

ROOYEN, N. 1991. The ephemerals of Namaqualand: effects of

photoperiod, temperature and moisture stress on development and

flowering of three species. Journal of Arid Environments 20:

15-29.

Bothalia 24,1 (1994)

85

APPENDIX

Habitat, species composition, percentage of plant cover and flowering phenology of the permanent plots. Where species flowered twice during a

flowering season, number of days of flowering (fourth column) are given separated by '+’ . *+’ put before the number of days indicates that flowering

started before an observation period, ‘+’ put behind means that flowering continued after an observation period. Floral units counted (sixth column)

were single flowers unless stated otherwise. Names of plant communities follow Le Roux (1984).

Plot 1. Ruschia robusta-Eriocephalus ericoides Community facies 1: open dwarf-scrub with low proportion of therophytes on NE slope, 5%

inclination, at SE side of Carolusberg

Ch, chamaephytes; NP, nanophanerophytes; T, therophytes; G, geophytes; Ca, capitula; Cy, cyathia; Inf, inflorescences or flower aggregates.

Bothalia 24,1 (1994)

Plot 2. Didelta spinosa-Ruschia cymosa Community: moderately dense scrub with low proportion of therophytes on SSE slope, 20% inclination at

SE side of Carolusberg

Ch, chamaephytes; NP, nanophanerophytes; T, therophytes; G, geophytes; Ca, capitula; Cy, cyathia; Inf, inflorescences or flower aggregates.

Bothalia 24,1 (1994)

87

Plot 3. Ruschia robusta-Eriocephcilus ericoides Community: open scrub and dwarf-scrub with moderate proportion of therophytes on WSW slope,

6% inclination, at SE side of Carolusberg

Ch, chamaephytes; NP, nanophanerophytes; T. therophytes; G, geophytes; Ca, capitula; Cy, cyathia; Inf, inflorescences or flower aggregates.

Plot 4. Ruschia robusta-Eriocephalus ericoides Community: moderately dense scrub and dwarf-scrub with high proportion of therophytes on SE

slope, 8% inclination, at SE side of Carolusberg

Bothalia 24,1 ( 1994)

Ch, chamaephytes; NP, nanophanerophytes; T, therophytes; G, geophytes; Ca, capitula; Cy, cyathia; Inf, inflorescences or flower aggregates.

Plot 5. Ruschia robusta-Eriocephalus ericoides Community facies 1: open dwarf-scrub with high proportion of therophytes on SSW slope, 6%

inclination, at SE side of Carolusberg

Bothalia 24,1 (1994) 89

Ch, chamaephytes; NP, nanophanerophytes; T. therophytes; G. geophytes; Ca, capitula; Cy, cyathia; Inf. inflorescences or flower aggregates.

Plot 6. Diverse ephemeral vegetation with therophytes and some geophytes on disturbed NW slope, 1% inclination, on alluvial plain (glacis)

90

Zaluzianskya benthamiana T

Tribulus zeyheri T

Tetragonia microptera T

Ifloga paronychioides T

Oncosiphon suffruticosum T

Aizoon canariense subsp. paucandrum T

Conicosia elongata G

Oxalis grammopetala G

Gladiolus orchidiflorus G

Albuca cooperi G

Homeria schlechteri G

Ornithoglossum vulgare G

Omithogalwn secundum G

Total cover [%]

Bothalia 24,1 (1994)

* dried fructiferous plants of the previous season.

Ch, chamaephytes; NP, nanophanerophytes; T, therophytes; G, geophytes; Ca, capitula; Cy, cyathia; Inf, inflorescences or flower aggregates.

25

50

25

Bothalia 24,1: 91-99 (1994)

Miscellaneous notes

VARIOUS AUTHORS

ASTERACEAE

CHROMOSOME COUNTS FOR SEVEN SPECIES OF CINERARIA (SENECIONEAE)

Cineraria L. belongs to the ‘senecioid- genera in the

tribe Senecioneae of the family Asteraceae, which have

the basic chromosome number of x = 10 (Nordenstam

1977). Grant (1971 : 228) notes that 533 taxonomic species

in the Dicotyledonae have the gametic (haploid) chromo-

some number of 10.

Counts for only two species of Cineraria have been

published in the chromosome indices of Moore (1973)

and Goldblatt (1981): C. aspera Thunb. 2x = 20 and 4x

= 40 (Nordenstam 1969); C. grandiflora Vatke (now C.

deltoidea Sond.) 2x = 20 (Turner & Lewis 1965) and 4x

= 40 (Hedberg & Hedberg 1977).

MATERIALS AND METHODS

Mitotic counts were made from root tips of germinated

seeds. Standard methods of pretreatment in 0.2 M 8-

hydroxyquinoline for four hours at room temperature (±

19°C) (Darlington & La Cour 1976), storage in Carnoy’s

fixative (absolute alcohol: glacial acetic acid : chloroform:

formalin in the ratio 5:1: 1:0.5), and staining in lacto-pro-

pionic orcein (Dyer 1963) were used. The root tips were

hydrolysed in 5N HC1 for 35^10 minutes at room tem-

perature, followed by two rinses in distilled water.

Meiotic counts were made from young anthers follow-

ing the method described by Turner & Lewis (1965). The

buds were collected in the field and placed in a freshly

mixed solution of chloroform : absolute alcohol : glacial

acetic acid (4:3: 1), and then processed through a graded

alcohol series and stored in 70% alcohol. Staining was

achieved using acetocarmine and iron acetate (Darlington

& La Cour 1976).

As the chromosomes were very small (1.5-4.0 pm),

repeated counts of pollen mother cells and of root tips

were made to ensure accuracy. Voucher specimens, as

listed in Table 1, are housed in the C.E. Moss Herbarium

(J), University of the Witwatersrand.

RESULTS AND DISCUSSION

The results of the chromosome counts from mitotic

and meiotic preparations are summarised in Table 1 . They

corroborate the published chromosome numbers for Cine-

raria. The most common chromosome number in the spe-

cies investigated was a diploid number of 20. Three of

the species show evidence of polyploidy with 2n = 40.

C. grandibracteata possibly has both levels of ploidy

within one population. There is no evidence of aneuploidy

in any of the species investigated thus far.

The population of C. deltoidea from the Natal Mid-

lands ( Cron & Scott-Shaw 11) was found to have a chro-

mosome number of n = 20, the same as that of the popu-

lations from Kenya and Tanzania examined by Turner &

Lewis (1965), and by Hedberg & Hedberg (1977).

Polyploidy is common in the Asteraceae (Grant 1971)

and it is evident that further sampling of populations of

the different species is required to ascertain whether dip-

loidy and polyploidy exist in the same populations and/or

species. This would also be a prerequisite before differ-

ences in chromosome number could be used to distinguish

different species.

The frequency of polyploidy in higher plants has been

positively correlated with increase in latitude and altitude

(Grant 1971; Love & Love 1967). C. albicans and C.

grandibracteata are both frequently found at high alti-

tudes, the latter species often in the mist belt of mountain

tops. The fact that polyploidy does exist in C. albicans

indicates that speciation may occur more rapidly in this

species than in those species shown thus far to be diploid.

Stebbins (1950: 359) notes that polyploidy is ‘widely

TABLE 1. The chromosome numbers obtained from root tips and pollen mother cells for species of Cineraria

92

Bothalia 24,1 (1994)

recognised as one of the principal methods for the forma-

tion of new species among higher plants’, but that the

species originating from this process are usually ‘very

similar to their diploid ancestors in external morphology

and ecological preferences’. C. albicans is a highly vari-

able species (Hilliard 1977; Cron 1991) and may well be

a species complex or in the process of speciating (Cron

1991). Characters such as the size and persistence of the

petiolar auricles, the glabrescence of the involucral bracts

and the indumentum of the cypselae show great variation.

(Note; the ‘species A’ of Hilliard ( 1977) is here considered

to be part of C. albicans.) Apparent isolation of popula-

tions of C. albicans in the gorges comprising valley

bushveld in the Natal midlands and coastal areas, and in

Transkei has evidently resulted in variability becoming

fixed in certain areas (Dubinin 1940). Similarly, isolation

on mountain tops in Lesotho, Natal and in the eastern

Cape has resulted in marked variation in the features noted

above.

C. deltoidea , as described by Jeffrey ( 1986), is the most

wide-ranging of the species, occurring in the eastern high-

lands of Ethiopia, Kenya, Tanzania, and Malawi, the

Zoutpansberg in the Transvaal, and the Natal midlands

and coastal regions of Natal and the Transkei. It is a very

variable species, with especially distinctive variants oc-

curring on some mountain massifs (Jeffrey 1986) and it

is therefore not surprising that polyploid populations exist.

The presence of polyploidy in the genus Cineraria is

thus confirmed, with a diploid chromosome number of 20

in six species and a polyploid chromosome number of 40

in three species.

ACKNOWLEDGEMENTS

This work was made possible by bursaries from the

FRD and the University of the Witwatersrand while the

senior author was reading for an M. Sc.

REFERENCES

CRON, G.V. 1991. A systematic study of certain species of Cineraria L.

(Asteraceae). M.Sc. thesis, University of the Witwatersrand. Jo-

hannesburg.

DARLINGTON, C.D. &LACOUR, F.L. 1976. The handling of chromo-

somes. George Allen & Unwin, London.

DUBININ, N.R 1940. Darwinism and the genetics of populations.

Uspekhi Sovremennoi Biologii 13: 276-305.

DYER, R.A. 1963. The use of lactopropionic orcein in rapid squash

methods for chromosome preparations. Stain Technology 38: 85-

90.

GOLDBLATT. P. 1981. Index to plant chromosome numbers 1975-1978.

Monographs in Systematic Botany 5. Braun-Brumfield, Ann

Arbor, Michigan.

GRANT, V.E. 1971. Plant speciation. Columbia University Press, New

York and London.

HEDBERG, I. & HEDBERG, O. 1977. Chromosome numbers of

afroalpine and afromontane angiosperms. Botaniska Notiser 1 30:

1-24.

HILLIARD, O.M. 1977. Compositae in Natal. University of Natal Press,

Pietermaritzburg.

JEFFREY, C. 1986. The Senecioneae in East tropical Africa. Kew Bulletin

41: 873-943.

LOVE, A. & LOVE, D. 1967. Polyploidy and altitude: Mt Washington.

Biologisclies Zentralhlatt. Supplementary Volume : 307-312.

MOORE, R.J. 1973. Index to plant chromosome numbers 1967-1971.

Regnum vegetabile 90. Oosthoek's Uitgeversmaatschappij B.V.,

Utrecht, Netherlands.

NORDENSTAM, B. 1969. Chromosome studies on South African vascu-

lar plants. Botaniska Notiser 122: 398—408.

NORDENSTAM, B. 1977. Senecioneae and Liabeae — systematic re-

view. In V.H. Heywood, J.B. Harbome & B.L. Turner, The biol-

ogy and chemistry of the Compositae. Vol. 2: 799-830. Academic

Press, London and New York.

STEBBINS, G.L. 1950. Variation and evolution in plants. Columbia

University Press, New York.

TURNER, B.L. & LEWIS, W.H. 1965. Chromosome numbers in the

Compositae. IX. African species. Journal of South African Botany

31:207-217.

G.V. CRON*. B-E. VAN WYK** and P.L.D. VINCENT*

* Department of Botany, University of the Witwatersrand, Private Bag 3,

Wits 2050.

** Department of Botany, Rand Afrikaans University, P.O. Box 524,

Johannesburg 2000.

MS. received: 1993-03-01.

POACEAE

A CYTOTAXONOMIC STUDY OF SOME REPRESENTATIVES OF THE TRIBE CYNODONTEAE (CHLORIDOIDEAE)

Poaceae is divided into five major subfamilies. One of

these subfamilies, Chloridoideae Rouy, is represented by

approximately 50 genera and 232 species in southern Af-

rica (Gibbs Russell et al. 1990). One of the tribes of the

Chloridoideae, Cynodonteae Dumort., commonly occurs

in unstable communities (Clayton & Renvoize 1986).

Cynodonteae has four subtribes, of which two were stud-

ied, namely Chloridinae Presl and Zoysiinae Benth. The

following genera of the subtribe Chloridinae are included

in this study; Chloris Sw., Cynodon Rich., Eustachys

Desv., Harpochloa Kunth, Microchina R. Br., and Rendlia

Chiov. The only genus in the sublribe Zoysiinae studied,

is Tragus Haller.

The aim of this study is to establish the chromosome

numbers of the different genera and species.

MATERIAL AND METHODS

The material was collected and fixed in the field.

Voucher herbarium specimens are housed in the Geo Potts

Herbarium, Department of Botany and Genetics, Univer-

sity of Orange Free State, Bloemfontein (BLFU) or the

National Herbarium, Pretoria (PRE).

SPECIMENS EXAMINED

Chloris gayana Kunth: n = 10.

TRANSVAAL. — 2428 (Nylstroom): Soutpan Experimental Station,

(-CD), Spies 3727. 2528 (Pretoria): Sphinx Station, (-CA), Spies 2021.

Bothalia 24,1 (1994)

93

C. virgata Swartz: n = 10.

ORANGE FREE STATE.— 2827 (Senekal): 6 km from Clocolan to

Peka bridge, (-DC), Spies 4799. 2926 (Bloemfontein): on the U.O.F.S.

campus, (-AA), Spies 5151, 5161, 5164, 5165, 5174', 26 km from

Dewetsdorp to Hobhouse, (-DB), Spies 4783.

CAPE. — 3026 (Aliwal North): Aliwal North, (-DA), Spies 5245,

5249.

Cynodon dactylon (L.) Pers.: n = 18.

CAPE. — 3026 (Aliwal North): Aliwal North, (-DA), Spies 5248.

3318 (Cape Town): on the top of Botmaskloof Pass, (-BD), Spies 4424.

3420 (Bredasdorp): De Hoop Nature Reserve, (-E>C), Spies 4626.

Eustachys paspaloides (Vahl) Lanza & Mattei: n = 20.

TRANSVAAL. — 2530 (Lydenburg): 10 km from Boshoek to

Buffelsvlei, (-AC), Spies 1521.

Harpochloa falx (L. f.) Kuntze: n = 20, 25, 30.

TRANSVAAL. — 2430 (Pilgrim’s Rest): 4 km from Pilgrim's Rest to

Graskop, (-BA), Spies 5134 (n = 20); 25 km from Sabie to Lydenburg,

(-BA), Spies 5140 (n = 20). 2530 (Lydenburg): Nederhorst turnoff on

Lydenburg-Roossenekal road, (-AA), Spies 5128 (n = 30); 1 1 km from

Dulistroom to Lydenburg via Frischgewaagd, (-AC), Spies 5118 (n =

25); 16 km from Dulistroom to Lydenburg via Frischgewaagd, (-AC),

Spies 5125 (n = 30); 5 km from Belfast to Dulistroom, (-CA), Spies

5113 (n = 20).

ORANGE FREE STATE. — 2729 (Volksrust): 53 km from Harrismith

to Newcastle via Normandien Pass, (-DC), Spies 5063 (n = 20); 92 km

from Harrismith to Normandien Pass, (-DC), Spies 5065 (n = 20); 97

km from Harrismith to Normandien Pass, (-DC), Spies 5078 (n = 20).

CAPE. — 3027 (Lady Grey): 45 km from Barkly East to Rhodes,

(-DD), Spies 3986 (n = 20); 52 km from Rhodes via Lundeansnek,

(-DD), Spies 4729 (n = 25). 3028 (Matatiele): 65 km from Rhodes via

Naudesnek, (-CC), Spies 4695 (n = 20); 69 km from Rhodes, (-CC),

Spies 4701 (n = 20). 3128 (Umtata): 38 km from Maclear to Elliot,

(-AC), Spies 4712 (n = 30).

Microchloa caffra Nees: n = ± 50.

TRANSVAAL. — 2430 (Pilgrim’s Rest): 4 km from Pilgrim’s Rest to

Graskop, (-BA), Spies 5132', 25 km from Sabie to Lydenburg, (-BA),

Spies 5141. 2530 (Lydenburg): 49 km from Lydenburg to Machadodorp,

(-CB), Spies 5146.

CAPE. — 3128 (Umtata): 38 km from Maclear to Elliot, (-AC), Spies

4714.

Rendlia altera (Rendle) Chiov.: n = 20.

TRANSVAAL. — 2430 (Pilgrim’s Rest): 4 km from Pilgrim's Rest to

Graskop, (-BA), Spies 5133', 25 km from Sabie to Lydenburg. (-BA),

Spies 5142. 2530 (Lydenburg): Nederhorst turnoff on the Lydenburg-

Roossenekal road, (-AA), Spies 5129', 11 km from Dulistroom to Lyden-

burg via Frischgewaagd, (-AC), Spies 5120\ 49 km from Lydenburg to

Machadodorp, (-AD), Spies 5147.

ORANGE FREE STATE. — 2729 (Volksrust): 93 km from Harrismith

to Normandien Pass, (-DC), Spies 5072; 97 km from Harrismith to

Normandien Pass, (-DC), Spies 5077.

CAPE. — 3028 (Matatiele): 69 km from Rhodes, (-CC), Spies 4700.

3128 (Umtata): 38 km from Maclear to Elliot, (-AC), Spies 4713.

Tragus berteronianus Schult.: n = 10.

SWAZILAND. — 2631 (Mbabane): Pikiti in the Lebombo Mountains,

(-BB), Spies 2605.

CAPE. — 3026 (Aliwal North): Aliwal North, (-DA), Spies 5246.

Tragus racemosus (L.) All.: n = 10.

CAPE. — 3026 (Aliwal North): Aliwal North, (-DA), Spies 5244.

Young inflorescences were fixed in Camoy’s fixative

for 24—48 hours and the fixative was subsequently re-

placed by 70% ethanol. Anthers were squashed in 2%

aceto-carmine (Darlington & La Cour 1976) and small

aliquots of iron acetate. Slides were made permanent by

freezing them with liquid CCD (Bowen 1956), followed

by dehydration in ethanol and mounting in Euparal. Mei-

otic chromosome behaviour for each specimen, was ex-

amined during diakinesis, metaphase I, anaphase 1 and

telophase I. At least 20 cells, representative of each of

these meiotic stages, were examined per specimen.

RESULTS AND DISCUSSION

The genus Chloris (Cynodonteae Dumort.; Chloridinae

Presl) usually has a chromosome base number of ten, oc-

casionally nine (Gibbs Russell et al. 1990). Both Chloris

gayana and C. virgata have somatic chromosome num-

bers of 20 (Figure 1A-G). We accept that the basic chro-

mosome number is ten, because that is the lowest haploid

chromsosome number observed in this study, and de-

scribed for this genus.

The chromosome numbers published, range from 2n =

20 to 40 for C. gayana and from 2n = 14 to 36 for C.

virgata , with 2n = 20 being the most frequent (Darlington

& Wylie 1955; Omduff 1967-1969; Fedorov 1969; Moore

1970, 1971, 1972, 1974, 1977; Goldblatt 1981, 1983,

1985, 1988; Goldblatt & Johnston 1990, 1991). The genus

Chloris is either diploid, as observed in this study, or poly-

ploid, with the polyploid levels ranging from triploid to

tetraploid (Spies & Jonker 1987). There were almost no

meiotic abnormalities in any of the specimens.

The Cynodon dactylon (Cynodonteae; Chloridinae)

specimens studied have haploid chromosome numbers of

9 (Figure 1H), 18 and 20. The basic chromosome number

is nine, because most published chromosome numbers are

multiples of nine. These chromosome numbers ranged

from 2n = 18 to 54 (Darlington & Wylie 1955; Malik

1967; Omduff 1967-1969; Fedorov 1969; Moore 1970-

1977; Goldblatt 1981-1988; Goldblatt & Johnston 1990.

1991). Malik (1967) described three cytological races in

this species: diploid (2n= 18 + 0- 3B), tetraploid (2n =

36 + 0 - 2B) and hexaploid (2n = 54).

A few cells of Spies 4626 were diploid and in all these

cells some extent of desynapsis occurred (Figure 1H). Just

why desynapsis should occur in the diploid cells, is not

known. Since some of the cells in this specimen contained

40 chromosomes, the specimen is regarded as a tetraploid

specimen, with four additional chromosomes. The addi-

tional chromosomes were similar in size to the euchro-

mosomes. During metaphase I these univalents lay on the

metaphase plate and showed no lagging. Since the number

of additional chromosomes varied from 0-4 per cell, we

have regarded them as B-chromosomes.

The polyploid specimens in this study form only biva-

lents and no multivalents and can, therefore, be regarded

as allopolyploids. The fact that only rod bivalents were

observed, however, indicates that only one chiasma forms

per chromosome. Multivalent formation is thus impossi-

ble, since a multivalent requires more than one chiasma

per chromosome. There were no meiotic abnormalities in

94

Bolhalia 24.1 (1994)

FIGURE 1 . — l-’holomicmgraph.s ol mciolie chromosomes in CynodoiUcac. A. Clilnns gayana, Spies .1727. n = 10. diakmesis with lOn; B. C. C.

virgata, Spies 4783, n = 10, diakinesis with 1 On; D, E, C. virgata, Spies 4799, n = 10, metaphase 1 with 1 On; F, C. virgata, Spies 4783, n = 10,

metaphase 1 with ] On; G, C. gayana. Spies 3727, n = 10, anaphase I with 10-10 segregation; H. C. dactylon, Spies 4626, n = 9, metaphase I

with 5[[8i; I. C. dactylon, Spies 4424, tetrad after meiosis II; J, K, Eustachys paspaloides, Spies 1521, n = 20, metaphase I with 20n. Scale

bar; 20 pm.

the specimens studied, except for desynapsis in the few

diploid cells observed in Spies 4626.

The genus Eustachys (Cynodonteae; Chloridinae) was

represented by Eustachys paspaloides with a haploid chro-

mosome number of 20 (Figure 1 J, K). Therefore the basic

chromosome number of ten described by Gibbs Russell

et al. (1990), is substantiated. De Wet (I960), however,

reported a specimen with 2n = 36. This could possibly be

attributed to loss aneuploidy, as these grasses are usually

tetraploid. Spies 1521 is an allopolyploid, since no mul-

tivalents were observed. Only rod bivalents were observed

and the evidence for alloploidy is, consequently, not con-

clusive. There were no meiotic abnormalities.

Harpochloa falx (Cynodonteae; Chloridinae) has hap-

loid chromosome numbers of 20, 25 and 30 (Figure 2).

The basic chromosome number is ten and ploidy levels

range from tetraploid to hexaploid. Only rod bivalents

were observed, which is an indication that only one chias-

ma forms per bivalent and, therefore, no multivalents can

be formed. Therefore, it is impossible to determine the

type of polyploidy present since the number of chiasmata

play a restricting role. Abnormalities observed included

univalents (Figure 2M), laggards (Figure 2N-P) and an

anaphase I bridge (Figure 2Q). These abnormalities oc-

curred at very low frequencies.

Rendlia altera (Cynodonteae; Chloridinae) has haploid

chromosome numbers of 18 and 20 (Figure 3A-D). The

basic chromosome number can be either nine or ten. No

other chromosome numbers were determined or pub-

lished. Loss aneuploidy from a basic chromosome number

of ten can result in a somatic chromosome number of 36,

or gain aneuploidy from a basic chromosome number of

nine can result in a somatic chromosome number of 40.

There were no meiotic abnormalities. Further studies are

Bothalia 24,1 ( 1994)

95

needed to determine the basic chromosome number and

to establish the range of polyploid levels.

Tragus (Cynodonteae; Zoysiinae) has a basic chromo-

some number of nine or ten (Gibbs Russell et al. 1990).

Tragus berteronianus has a haploid chromosome number

of 10 (Figure 3E, F).

In conclusion, this study indicates that Cynodonteae

have polyploid or agamic complexes, with the somatic

chromosome numbers ranging from diploid to hexaploid.

Two different basic chromosome numbers (i.e. 9 and 10)

are present. Further studies are needed to determine the

phylogenetic relationship between the two base numbers

and to establish the polyploid levels in each taxon.

ACKNOWLEDGEMENTS

We thank the National Botanical Institute for material

of Chloris virgata and Eustachys paspaloides provided

during this study. Financial support by the Foundation for

FIGURE 2. — Photomicrographs of meiotie chromosomes in Harpochlou falx. A, Spies 4695. n = 20, diakinesis with 20ii; B, Spies 5065, n = 20,

diakinesis with 19ii2i; C,Spies4712,n = 30, diakinesis with 30n; D, Spies 5128, n = 30, diakinesis with 30ii; E, Spies 5125, n = 30, metaphase

I with 30n; F, Spies 5128, n = 30, metaphase I with 30n; G, Spies 5078, n = 20, anaphase I with a 20-20 segregation: H. Spies 5128, n = 30,

anaphase I with a 30-30 segregation; I, Spies 5125, n = 30, anaphase I with a 30-30 segregation; J, Spies 5078, n = 20, anaphase I with 40

chromosomes; K, Spies 5078, n = 20, anaphase I with a 20-20 segregation; L, Spies 4695, n = 20, metaphase I with desynapsis; M, Spies

4729, n = 25, metaphase I with a B-chromosome in each pole; N, Spies 4695, n = 20, anaphase I with two laggards; O, Spies 4712 , n = 30,

anaphase I with laggards; P, Spies 4695, n = 20, anaphase I bridge; Q, Spies 4712, n = 30, anaphase I bridge. Scale bar; 20 pm.

96

Bolhalia 24.1 (1994)

FIGURE 3. — Photomicrographs of meiolic chromosomes in Rendlia ahem and Tragus beneronianus. A-D. K. ahem: A, Spies 5077. n = 20,

diakinesis with 20ii; B. Spies 5077, n = 20, metaphase I; C, Spies 5072, n = 20, metaphase I with 20n; D, Spies 5077, n = 20, anaphase I

with a 20-20 segregation. E, F, T. berteronicinus, Spies 2605, n = 10, diakinesis with 10n. Scale bar = 20 pm.

Research and Development and the University of the Or-

ange Free State is gratefully acknowledged.

REFERENCES

BOWEN. C.C. 1956. Freezing by liquid carbon dioxide in making slides

permanent. Stain Technology 3 1 : 87-90.

CLAYTON, W.D. & RENVOIZE, S.A. 1986. Genera graminum. Kew

Bulletin Additional Series XIII: 1-389.

DARLINGTON, C.D. & LA COUR, L.F. 1976. The handling of chromo-

somes. Allen & Unwin, London.

DARLINGTON, C.D. & WYLIE, A.P. 1955. Chromosome atlas of flow-

ering plants. Allen & Unwin, London.

DE WET, J.M.J. 1960. Chromosome numbers and some morphological

attributes of various South African grasses. American Journal of

Botany 47: 44-50.

FEDOROV, A. A. 1969. Chromosome numbers of flowering plants. Acad-

emy of Science, Leningrad.

GIBBS RUSSELL, G.E., WATSON, L„ KOEKEMOER, M„ SMOOK,

L„ BARKER, N.P., ANDERSON, H.M. & DALLWITZ, M.J.

1990. Grasses of southern Africa. Memoirs of the Botanical Sur-

vey of South Africa No. 58.

GOLDBLATT, P. 1981. Index to plant chromosome numbers. Mono-

grams of Systematic Botany 5.

GOLDBLATT, P. 1983. Index to plant chromosome numbers. Mono-

grams of Systematic Botany 8.

GOLDBLATT, P. 1985. Index to plant chromosome numbers. Mono-

grams of Systematic Botany 13.

GOLDBLATT, P. 1988. Index to plant chromosome numbers. Mono-

grams of Systematic Botany 23.

GOLDBLATT. P. & JOHNSON, D.E. 1990. Index to plant chromosome

numbers. Monograms of Systematic Botany 30.

GOLDBLATT, P. & JOHNSON, D.E. 1991. Index to plant chromosome

numbers. Monograms of Systematic Botany 40.

MALIK, C.P. 1967. Polyploidy in Cynodon dactylon (L.) Pers. Chromo-

somal Information Service Tokyo 1: 6,1 .

MOORE, R.J. 1970. Index to plant chromosome numbers. Regnum Veg-

etable 68.

MOORE, R.J. 1971. Index to plant chromosome numbers. Regnum Veg-

etable 77.

MOORE, R.J. 1972. Index to plant chromosome numbers. Regnum Veg-

etable 84.

MOORE, R.J. 1974. Index to plant chromosome numbers. Regnum Veg-

etable 91.

MOORE, R.J. 1977. Index to plant chromosome numbers. Regnum Veg-

etable 96.

ORNDUFF, R. 1967. Index to plant chromosome numbers. Regnum

Vegetable 50.

ORNDUFF, R. 1968. Index to plant chromosome numbers. Regnum

Vegetable 55.

ORNDUFF, R. 1969. Index to plant chromosome numbers. Regnum

Vegetable 59.

SPIES, J.J. & JONKER, A. 1987. Chromosome studies on African plants.

4. Bothalia 17: 135, 136.

A. STRYDOM* and J.J. SPIES*t

* Department of Botany and Genetics, University of the Orange Free

State, P.O. Box 339, Bloemfontein 9300.

t Author to whom correspondence should be addressed.

MS. received: 1992-12-14.

Bothalia 24,1 (1994)

97

PTERIDOPHYTA

SYSTEMATIC STUDIES IN THE GENUS MOHRIA (ANEMIACEAE). V, KARYOLOGY

Mohria Sw. Actinostachys Wall., Anemia Sw., Lygod-

ium Sw. and Schizaea J. Smith constitute the schizaeoid

ferns. They are commonly placed in a single family, the

Schizaeaceae (Engl. & Prantl 1898-1902; Bower 1923;

Christensen 1938; Copeland 1947; Tryon & Tryon 1982;

Tryon & Lugardon 1991) because of their sporangium

morphology. The 'schizaeaceous' sporangium type, how-

ever, appears to be polyphyletic (Mickel 1974) and the

morphological differences between these genera are sig-

nificant. The subdivision of this assemblage into distinct

families, the Anemiaceae (Anemia and Mohria), Lygo-

diaceae (Lygodium) and the Schizaeaceae (Actinostachys

and Schizaea) has therefore recently gained wider accep-

tance (Nayar 1970; Bierhorst 1971: Love et al. 1977; Clif-

ford & Constantine 1980; De la Sota & Morbelli 1987).

Anemia and Mohria are dissimilar in frond morphology

but are evidently related in view of similarities in their

anatomy (Prantl 1881; Bower 1918; Roux et al. 1992),

trichomes (Mickel 1962; Roux 1992a) and spores (Nayar

1968; Roux 1992b). The similarity in their chromosome

numbers also supports this view.

The chromosome numbers of the genera also endorse

the proposed subdivision:

Anemiaceae

Anemia : n = 38 (Mickel 1962; Lovis 1977); n = 76, 114 (Mickel 1982).

Mohria : n = 38 (Mickel 1962; Lovis 1977)

Lygodiaceae

Lygodium'. 2n = 56 (Roy & Manton 1965); 2n = 58 (Manton & Sledge

1954; Roy & Manton 1965); 2n = 60 (Wagner 1963; Manton & Sledge

1954); 2n = 112 (Roy & Manton 1965); 2n = 116 (Manton & Sledge

1954); 2n = 120 (Roy & Manton 1965).

Schizaeaceae

Actinostachys: 2n = 280 (Mitui 1973); n = 325 ± 30 (Lovis in Holttum

1959); 2n = ± 1120 (Abraham et al. 1962).

Schiwea: 2n = ± 144 (Araujo in Love 1976); 2n = 154 (Lovis 1958);

2n = ± 188 (Brownlie 1965); 2n = ± 540 (Brownlie 1965).