Bothalia

A JOURNAL OF BOTANICAL RESEARCH

Vol. 28,2

October 1998

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STRELITZIA

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MEMOIRS OF THE BOTANICAL SURVEY OF SOUTH AFRICA

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ANNALS OF KIRSTENBOSCH BOTANIC GARDENS

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FLOWERING PLANTS OF AFRICA (FPA)

This serial presents colour plates of African plants with accompanying text. The plates are prepared

mainly by the artists at the National Botanical Institute. Many well known botanical artists have

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From Vol. 55, twenty plates are published at irregular intervals.

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FLORA OF SOUTHERN AFRICA (FSA)

A taxonomic treatise on the flora of the Republic of South Africa, Lesotho, Swaziland, Namibia

and Botswana. The FSA contains descriptions of families, genera, species, infraspecific taxa, keys

to genera and species, synonymy, literature and limited specimen citations, as well as taxonomic

and ecological notes.

Contributions to the FSA also appear in Bothalia.

PALAEOFLORA OF SOUTHERN AFRICA

A palaeoflora on a pattern comparable to that of the Flora of southern Africa. Much of the infor-

mation is presented in the form of tables and photographic plates depicting fossil populations. Now

available:

Molteno Formation (Triassic) Vol. I . Introduction. Dicroidium, by J.M. & H.M. Anderson.

Molteno Formation (Triassic) Vol. 2. Gymnosperms (excluding Dicroidium), by J.M. & H.M.

Anderson.

Prodromus of South African Megafloras. Devonian to Lower Cretaceous, by J.M. & H.M.

Anderson. Obtainable from: A. A. Balkema Marketing, Box 3 1 7, Claremont 7735, RSA.

BOTHALIA

A JOURNAL OF BOTANICAL RESEARCH

Volume 28,2

Scientific Editor: G. Germishuizen

Technical Editor: B.A. Momberg

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INSTITUTE

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Editorial Board

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Missouri Botanical Garden, St Louis, USA

Compton Herbarium, NBI, Cape Town, RSA

University of Utrecht, Utrecht, Netherlands

CONTENTS

Volume 28,2

New species and combinations in Bothalia 28,? (1998)

1. Five new species of Lachenalia (Hyacinthaceae) from arid areas of Namibia and South Africa. G.D.

DUNCAN 131

2. A revision of Lachenalia (Hyacinthaceae) in the Eastern Cape, South Africa. A.P. DOLD and

P.B. PHILLIPSON 141

3. FSA contributions 12: Plantaginaceae. H.F. GLEN 151

4. Studies in the liverwort genus Fossombronia (Metzgeriales) from southern Africa. 6. New spe-

cies from Lesotho, Swaziland and Mpumalanga and new records from Lesotho. S.M. PEROLD 159

5. The genus Echiutn (Boraginaceae) in southern Africa. E. RETIEF and A.E. VAN WYK 167

6. A revision of Ledebouria (Hyacinthaceae) in South Africa. 2. Two new species, L. crispa and L. par-

vifolia, and L. macowanii re-instated. S. VENTER and T.J. EDWARDS 179

7. Notes on African plants:

Acanthaceae/Orchidaceae. New records from KwaZulu-Natal, South Africa. T.J. EDWARDS

and E. HARRISON 187

Amaryllidaceae: Amaryllideae. A new species of Amaryllis from the Richtersveld, South Africa.

D.A. SNIJMAN and G. WILLIAMSON 192

Asteraceae. Key to the species of the genus Hertia in southern Africa. P.P.J. HERMAN 192

Cyperaceae. Coleochloa setifera new to the flora of KwaZulu-Natal. C. ARCHER 190

Metzgeriales-Fossombroniaceae. Fossombronia occidento-africana : is it conspecific with

F. indical S.M. PEROLD ' 183

8. A reconnaissance survey of the vegetation of the North Luangwa National Park, Zambia. P.P. SMITH 197

9. Wetland plant communities in the Potchefstroom Municipal Area, North-West, South Africa. S.S.

CILLIERS. L.L. SCHOEMAN and G.J. BREDENKAMP 213

10. Cytogenetic studies in the genus Pentaschistis (Poaceae: Arundinoideae). K.C. KLOPPER, J.J. SPIES

and B. VISSER 231

11. Obituaries:

David Spencer Hardy (1931-1998). H.F. GLEN 239

Barend Petrus Barkhuizen (1921-1995): amateur botanist, succulent plant specialist and edu-

cator. G.F. SMITH and E.M.A. STEYN 243

Mary Elizabeth Connell (1917-1997). D.J.B. KILLICK 245

12. National Botanical Institute, South Africa: administration and research staff 22 May 1998, publica-

tions 1 April 1997-31 March 1998. Compiler: B.A. MOMBERG 249

13. Guide for authors to Bothalia 261

14. The history of the Botanical Research Institute 1903-1989. DENISE FOURIE 271

New species and combinations in Bothalia 28,2 (1998)

Amaryllis paradisicola Snijman, sp. nov., 193

Fossombronia angulifolia Perold , sp. nov., 159

Fossombronia swaziensis Perold , sp. nov., 162

Lachenalia attenuata W.F. Barker ex G.D. Duncan, sp. nov., 132

Lachenalia doleritica G.D. Duncan, sp. nov., 134

Lachenalia lactosa G.D. Duncan, sp. nov., 135

Lachenalia leipoldtii G.D. Duncan, sp. nov., 137

Lachenalia nutans G.D. Duncan, sp. nov., 131

Ledebouria crispa S. Venter, sp. nov., 179

Ledebouria macowanii (Baker) S. Venter, comb, nov., 181

Ledebouria parvifolia S. Venter, sp. nov., 180

IV

Bothalia 28,2: 131-139 (1998)

Five new species of Lachenalia (Hyacinthaceae) from arid areas of

Namibia and South Africa

G.D, DUNCAN*

Keywords: Hyacinthaceae, Lachenalia , Namibia, new species. South Africa

ABSTRACT

Five new species of Lachenalia are described: L. nutans G.D. Duncan from southwestern Namibia, L. attenuata

W.F.Barker ex G.D. Duncan from the Roggeveld Plateau, Little Karoo and southern Cape; L. doleritica G.D. Duncan from

the Bokkeveld Plateau; L. lactosa G.D. Duncan from the Lowland Fynbos of the Hermanus District, and L. leipoldtii

G.D. Duncan from the Olifants River Valley and Little Karoo.

INTRODUCTION

This is the third in a series of papers on new species

of Lachenalia , and serves as a continuation of the work

of W.F. Barker (Barker 1978, 1979, 1983a & b, 1984,

1987, 1989) and the present author (Duncan 1987, 1988a

& b, 1989a & b, 1992, 1993, 1994, 1996, 1997a & b)

towards a revision of the genus.

Lachenalia nutans G.D. Duncan, sp. nov. L.

anguineae Sweet affinis propter flores similares campan-

ulatos albos tumoribus flavo-virentibus vel brunneo-

virentibus, sed floribus distincte nutantibus staminibus

declinatis folioque lanceolato vel late lanceolato pagina

inferiore glauca margine atromarronino basi subterranea

amplectenti differt.

TYPE. — Southwestern Namibia, 2715 (Bogenfels):

2.5 km NE of Schlafkuppe, on sandy gravel flats, (-BD),

21-7-1986. N.J. van Berkel 563 (NBG, holo.!).

Deciduous, winter-growing geophyte 35-110 mm

high. Bulb subglobose, 10-20 mm diam., white with thin

layer of reddish brown, membranous outer scales. Leaf

solitary, lanceolate to broadly lanceolate, 30-60 x

4-20 mm, erect or suberect, upper surface unmarked,

dark green with depressed longitudinal veins, lower sur-

face glaucous, unmarked or barred with dark green trans-

verse bands, margin dark maroon; clasping base white,

subterranean, 20-55 mm long. Inflorescence racemose,

dense, 40-55 mm long with a very short sterile tip;

peduncle erect, 25^40 mm long, sturdy, pale green,

mottled with dull purplish red spots, slightly to conspic-

uously swollen towards apex, and at base of inflores-

cence; rachis 35-45 mm long, pale green in lower half,

with dull purplish red spots, shading to white in upper

half, with sporadic, much paler spots, slightly to con-

spicuously swollen in lower half; pedicels erect to

suberect during flowering, bending downwards during

fruiting, white, 2-5 mm long; bracts white, cup-like at

base of inflorescence, becoming lanceolate above,

1.0-1. 5 mm long. Flowers oblong-campanulate, distinct-

* National Botanical Institute, Kirstenbosch, Private Bag X7, 7735

Claremont, Cape Town.

MS. received: 1997-11-14.

ly nodding or cernuous, white with yellowish green or

brownish green gibbosities; perianth tube white, 3-4 mm

long; outer tepals oblong, 4-6 x 2.5 mm, white with yel-

lowish green or brownish green gibbosities, and pale

greenish zone at base; inner tepals narrowly spathulate,

4-6 x 1.0-1. 5 mm, white with pale greenish yellow zone

near apex. Stamens well exserted beyond tip of perianth,

declinate; filaments white, 6-10 mm long; anthers yel-

low. Ovary obovoid, pale green, 2-3 mm long; style

white, 7-9 mm long, protruding well beyond stamens as

ovary enlarges. Capsule obovoid, 4-6 x 4 mm. Seed

black, glossy with granulated testa, 1.5 mm long, globose

with small ridged arillode 0.3 mm long and distinct

decurrent keel. Flowering time-: July to August. Figures

1A; 2 & 3.

Etymology: named L. nutans to describe the distinctly

nodding flowers.

Diagnostic characters

L. nutans is characterised by a dense, racemose inflor-

escence of distinctly nodding, oblong-campanulate

flowers with the inner tepals as long as, or very slightly

longer than the outer tepals. The declinate stamens are

very well exserted and the upper part of the peduncle

and the lower part of the rachis is slightly to conspicu-

ously swollen. The plant has a solitary, lanceolate or

broadly lanceolate leaf with a dark green upper surface,

a glaucous lower surface and a white, deep subterranean

clasping base.

L. nutans is related to L. anguinea Sweet which

occurs in deep red sand from the Richtersveld as far

south as the Piketberg District. L. anguinea has similar

campanulate white flowers with yellowish green or

brownish green gibbosities and well exserted white sta-

mens. It differs, however, in being a larger plant with

much smaller, patent flowers produced on very long

pedicels, and its long, arcuate, deeply canaliculate, flac-

cid leaf. The seeds of L. anguinea also differ from those

of L. nutans in having a very small, almost obsolete, ter-

minal ridged arillode, whereas the seeds of L. nutans

have a granulated testa and a distinct decurrent keel. The

two species are also separated geographically, L. nutans

being known only from southwestern Namibia.

132

Bothalia 28,2 (1998)

FIGURE 1. — A, Lachenalia nutans, Van Berkel 563; B, L. attenuata, Saunders s.n.', C, L. doleritica, Manning s.n. Scale bars: 10 mm.

Distribution and habitat

Lachenalia nutans is known from two collections

made in southwestern Namibia. In August 1929, Dinter

collected material of Lachenalia klinghardtiana Dinter at

Haalenberg which was deposited at B, K and PRE

{Dinter 6666). However, one of the two herbarium sheets

of this collection housed at B is in fact not L. klinghard-

tiana but represents the first collection of Lachenalia

nutans. The specimens of the latter species are in a fruit-

ing stage and the collecting date is given as 31st August

1929, whereas the collecting date given for the other

sheet correctly identified as L. klinghardtiana is 30th

August. Fifty-seven years later, Mrs N.J. Van Berkel col-

lected (lowering material of L. nutans further south at a

locality northeast of Schlafkuppe (Figure 4), which is

now the type material of the species. Plants grow singly

or in groups on sandy gravel flats in full sun. The typi-

cally deep-seated bulbs probably only grow and flower if

there has been sufficient seasonal rainfall.

Material examined

NAMIBIA.— 2615 (Luderitz): Haalenberg, (-DA), 31-8-1929,

Dinter 6666 (B). 2715 (Bogenfels): 2.5 km NE of Schlafkuppe, (-BD),

21-7-1986, Van Berkel 563 (NBG).

Lachenalia attenuata W.F. Barker ex G.D. Duncan,

sp. nov. L. hirtae (Thunb.) Thunb. affinis propter flores

similares pallide azureos oblongos vel oblongo-campanu-

latos segmentis interiores perianthii viridi-flavis tumor-

ibus purpureo-brunneis staminibusque inclusis, sed pedi-

cellis multo brevioribus folioque conduplicato sine setis

vel papillis et basi folii non distincte dilatata differt.

TYPE. — Western Cape, 3319 (Worcester): Keisie-

berg, Montagu Dist., (-DB), Sept. 1946, G.J. Lewis 2431

(SAM, holo!).

Deciduous, winter-growing geophyte 65-220 mm

high. Bulb globose, 8-10 mm diam., white with hard,

dark brown outer tunics. Leaf solitary, linear, attenuate,

conduplicate, clasping base of peduncle up to 50 mm

long, pale green shading to yellowish green in upper half,

with short, darker green bands on lower surface, upper

surface unmarked, spreading to suberect, 35-140 x

7-10 mm, clasping base marked with magenta bands just

below ground level, shading to brownish purple and pale

green at, and just above ground level, respectively.

Inflorescence subspicate or racemose, few to many- flow-

ered, very lax, 20-60 mm long, with very short sterile

tip; peduncle erect to suberect, very slender, 35-200 mm

long, very pale green in lower half and mottled with

brownish purple blotches, shading to yellowish green or

brownish purple in upper half, with or without brownish

purple blotches; rachis heavily mottled with brownish

purple, shading to pale blue at apex, 20-60 mm long;

pedicels white or brownish purple, spreading or suberect,

1-3 mm long; bracts ovate to obovate, 1-3 x 1-3 mm.

Flowers patent to slightly cernuous, oblong to oblong-

campanulate, pale blue and greenish yellow with pur-

plish brown markings; perianth tube pale blue, 2 mm

long; outer tepals oblong, 6-7 x 3-4 mm, pale blue at

base, shading to dull purplish brown at lips, with dull

Bothalia 28,2 (1998)

133

purplish brown gibbosities; inner tepals protruding well

beyond outer tepals, obovate, dull white at base shading

to greenish yellow at apex, with distinct brownish green

keels, 6-7 x 4-5 mm, upper two inner tepals overlap-

ping. Stamens declinate, included or very slightly exsert-

ed; filaments white, 6-7 mm long; anthers maroon prior

to anthesis, yellow at anthesis, and black after anthesis.

Ovary ovoid, pale green, 3-4 x 3 mm; style white, 7 mm

long, protruding beyond stamens as ovary enlarges.

Capsule ovoid, 6-7 x 5-6 mm. Seed globose, 1 mm long,

dull black, with a ribbed arillode 0.7 mm long.

Flowering time: August to September. Figures IB; 5.

Etymology : named L. attenuata by W.F. Barker to de-

scribe the gradually tapering leaf.

Diagnostic characters

L. attenuata is characterised by a subspicate or race-

mose, very lax inflorescence of patent or slightly cernu-

FIGURE 2. — Sheet of Lachenalia

nutans, Dinter 6666 (B),

material collected on 31-8-

1929.

ous, oblong or oblong-campanulate flowers with the

inner tepals protruding well beyond the outer. The decli-

nate stamens are included or very slightly exserted, and

the plant has a single linear, attenuate, conduplicate leaf

with an unmarked upper surface, the lower surface

marked with short green bands, and the clasping leaf

base marked with brownish purple bands above ground

level, and with magenta bands below ground level.

L. attenuata is related to L. hirta (Thunb.) Thunb. var.

hirta which has similar oblong or oblong-campanulate

pale blue flowers with greenish yellow inner tepals and

included or slightly exserted stamens, and a single linear

leaf with brownish purple and magenta bands on the

clasping leaf base. L. hirta differs in having very long

pedicels and a canaliculate leaf with distinct bristles and

papillae on the lower surface and margins. The seeds of

L. hirta differ in having an almost obsolete, terminal aril-

lode, and a finely netted testa. The distribution ranges of

the two species do not overlap, L. hirta occurring from

134

Bothalia 28,2 (1998)

FIGURE 3. — Holotype of Lachenaliu nutans, Van Berkel 563.

HKBMtnat HCSEt Al ST*t) \iW\M

FIGURE 5. — Holotype of Lachenaliu attenuata, Lewis 2431.

Namaqualand southwards to Malmesbury, and L. attenu-

ata occurring on the Roggeveld Plateau, the Little Karoo

and the southern Cape.

Distribution and habitat

L. attenuata was first collected by Dr J. Muir in

August 1933 in the Riversdale District of the southern

Cape, and represents the most easterly record of this

FIGURE 4. — Distribution of Lachenaliu nutans. A; L. attenuata, •; L.

doleritica, ★.

poorly collected, but fairly widespread species. It is also

known from three recent collections made on the

Roggeveld Plateau, and a further three records from the

Little Karoo (Figure 4). It grows in montane habitats

where it is usually encountered growing singly in south-

facing, seasonally moist, well-drained loamy clay soil

amongst rocks.

Material examined

NORTHERN CAPE. — 3220 (Sutherland): Farm Agterkop,

Gannaga Pass, (-AA), Sept. 1996, Saunders s.n. sub NBG755601

(NBG); mountain S of Voelfontein farmhouse, (-AD), Sept. 1981,

Snijman 515 (NBG); Farm Jakhals Valley, Verlatekloof Pass, (-DA),

Sept 1986, Cloete & Haselau 238 (NBG).

WESTERN CAPE. — 3319 (Worcester): Hex River Pass, W of sum-

mit, (-BD), Sept. 1974, Nordenstam & Lundgren 2059 (NBG);

Rabiesberg, NE of Nuy, (-DA), Sept. 1935, Lewis & Esterhuysen s.n.

sub BOL21949 (BOL); Keisieberg, Montagu Dist., (-DB), Sept. 1946,

Lewis 2431 (SAM); 3421 (Riversdale): Riversdale Dist., (-AB), Aug.

1933, Muir 4886 (K).

Lachenalia doleritica G.D. Duncan, sp. nov. L.

neilii W.F.Barker ex G.D. Duncan affinis propter inflo-

rescentiam similarem multifloram floribus virellis oblon-

go-campanulatis patentibus vel suberectis tumoribus

atroviridis vel brunneis, folios plerumque immaculatos

venis longiludinalibus distincte depressis, sed staminibus

inclusis foliisque ovatis arcuatis non glaucis, bulbo sine

annulo bulbillorum basi differt.

Bothalia 28,2 (1998)

135

FIGURE 6. — Holotype of Lachenalia doleritica, Thomas s.n.

TYPE. — Northern Cape, 3119 (Calvinia): Akkeren-

dam Nature Reserve, Calvinia, (-BD), 7-9-1984, M.L.

Thomas s.n. sub NBG140212 (NBG, holo.!).

Deciduous, winter-growing geophyte 120-180 mm

high. Bulb subglobose, 15-20 mm diam., white with sev-

eral layers of dark brown outer tunics. Leaves 2, ovate,

85 x 40 mm, fleshy, slightly arcuate, with leaf tips rest-

ing at ground level, bright green to yellowish green,

upper surface unmarked, with distinct, depressed longi-

tudinal veins, lower surface unmarked, margins coria-

ceous, dark maroon; clasping base white below, shading

to greenish white above, 30-40 mm long. Inflorescence

racemose, erect, many-flowered, 80-110 mm long, with

a short sterile tip; peduncle sturdy, erect or suberect,

40-100 mm long, becoming slightly swollen in upper

third, pale green and unmarked, or pale green in lower

two thirds with tiny pale brown speckles, tinged with

dull brownish mauve in upper third; rachis pale green or

tinged with dull brownish mauve, slightly swollen in

lower third; pedicels suberect, pale green or pale mau-

vish white, increasing in length towards apex of inflores-

cence, 2-4 mm long; bracts cup-shaped in lower half of

inflorescence, becoming lanceolate above, white, 3-5 x

2-4 mm. Flowers oblong-campanulate, patent or

suberect, very pale yellowish green with darker green,

very dark brown or brownish mauve markings; perianth

tube very pale yellowish green, 2-3 mm long; outer

tepals ovate, very pale yellowish green, 8-9 x 4-5 mm,

with darker green, very dark brown or brownish mauve

gibbosities, and a central yellowish green or brownish

mauve stripe; inner tepals obovate, protruding beyond

outer tepals, apices slightly recurved, 9-11 x 4-5 mm,

translucent yellowish green with a darker green or very

dark brown marking near apex, and brownish green

keels, inner lower tepals longer than two upper laterals.

Stamens included, declinate; filaments white, 9 mm

long; anthers pale maroon prior to anthesis, yellow at

anthesis. Ovary ovoid, pale green, 5x3 mm; style white,

7 mm long, protruding beyond tepals as ovary enlarges.

Capsule unknown. Seed unknown. Flowering time :

September to October. Figures 1C; 6.

Etymology, named L. doleritica to describe the heavy

doleritic clay soil in which this species occurs.

Diagnostic characters

L. doleritica is characterised by a many-flowered

racemose inflorescence of oblong-campanulate, patent or

suberect, yellowish green flowers with the inner tepals

protruding beyond the outer. The declinate stamens are

included, and the unmarked, ovate leaves are slightly

arcuate with the tips resting at ground level, and with dis-

tinct depressed longitudinal veins on the upper surface.

L. doleritica is related to L. neilii W.F.Barker ex

G.D. Duncan which has a similar many-flowered inflo-

rescence of greenish, patent or suberect, oblong-campan-

ulate flowers and usually unmarked leaves, with distinct

depressed longitudinal veins on the upper surface. L.

neilii differs in having smaller flowers with shortly

exserted stamens, lanceolate, suberect, glaucous leaves,

and the bulb produces a ring of bulbils at its base.

Distribution and habitat

The first recorded collection of L. doleritica was

made by Mrs M.L. Thomas as recently as September

1984 in the Akkerendam Nature Reserve at Calvinia, and

as far as I am aware, there are only two other records of

this species, one to the west of Calvinia towards

Nieuwoudtville, and the other to the east of Calvinia

towards Williston (Figure 4). L. doleritica appears to be

a rare species confined to heavy doleritic clay soil on the

Bokkeveld Plateau, where plants occur singly in open

situations in full sun. The apparently very restricted dis-

tribution range of this species overlaps the wider range of

the related L. neilii.

Material examined

NORTHERN CAPE. — 3119 (Calvinia): Calvinia-Nieuwoudtville

road, (R27), at side of road, (-AC), Sept. 1996, Manning s.n. (NBG,

photo.); Akkerendam Nature Reserve, Calvinia, (-BD), 7-9-1984,

Thomas s.n. sub NBG14021 2 (NBG); Calvinia-Williston road, at side

of road, (-BD), Oct. 1986, Thomas s.n. (NBG, photo.).

Lachenalia lactosa G.D. Duncan, sp. nov. L. peer-

sii Marloth ex W.F.Barker affinis propter flores albos

similares tumoribus viridi-brunneis foliosque ex mar-

roninos normaliter virides loratos, sed floribus valde

minoribus oblongo-campanulatis basibus pallide azureis

staminibusque declinatis paulo exsertis pedunculoque

leviter vel dense maculato differt.

136

Bothalia 28,2 (1998)

FIGURE 7. — A, Lachenalia lactosa, Barker 10476', B, L. lactosa, Barker 10510. Scale bars: 10 mm.

TYPE. — Western Cape, 3419 (Caledon): Honingklip

Farm, near Bot River, (-AC), 29-9-1967, W.F Barker

10510 (NBG, holo. ! ; PRE).

Deciduous, winter-growing geophyte 100-265 mm

high. Bulb globose, 8-15 mm diam., white with dark

brown outer tunics. Leaves 1 or 2, lorate, 40-200 x

5-18 mm, suberect or slightly arcuate, plain green, usu-

ally unmarked, occasionally marked with purplish brown

spots on upper surface, lower surface tinged with purple

or maroon, upper surface with distinct longitudinal

veins; clasping base white below, shading to pale

maroonish purple above, 10-20 mm long. Inflorescence

racemose, usually dense, few to many-flowered,

25-80 mm long, with short sterile apex; peduncle erect

or suberect, slender or sturdy, pale green with many

small to large purplish maroon spots or blotches,

50-130 mm long; rachis pale green with dull purplish

blue blotches; pedicels pale green, erect or suberect,

2- 4 mm long; bracts purplish maroon and cup-shaped at

base of inflorescence, becoming lanceolate and translu-

cent white above, 1-3 x 1-3 mm. Flowers small to very

small, oblong-campanulate, suberect or patent, white and

pale blue with dark magenta or greenish brown mark-

ings, fading to purplish blue; perianth tube very short,

pale blue or white, 0.5-1. 0 mm long; outer tepals ovate,

3- 4 x 5 mm, white, or pale blue at base, shading to white

above, with dark magenta or greenish brown gibbosities

and keels; inner tepals oblong obovate, 5-7 x 3 mm, pro-

truding well beyond outer segments, canaliculate, trans-

lucent white or cream with very narrow dark magenta or

greenish brown keels. Stamens shortly exserted, decli-

nate; filaments white, 4-6 mm long. Ovary ovoid, bright

green, 2-3 x 2 mm; style white, 4-5 mm long. Capsule

ovoid, membranous, 5-6 x 4-5 mm. Seed globose, shiny

black, 1 mm long, with a terminal, inflated arillode

0.5 mm long. Flowering time : September to October.

Figures 7A, B; 8.

Etymology, named L. lactosa to describe the milky white

colouring of the flowers.

Diagnostic characters

L. lactosa is characterised by a dense raceme of small

or very small, oblong-campanulate, suberect or patent,

white and pale blue flowers with shortly exserted, decli-

nate stamens. The peduncle is usually heavily marked

with purplish maroon spots or blotches, and the one or

two lorate, suberect or slightly arcuate leaves are plain

green, usually unmarked, with depressed longitudinal

veins on the upper surface, and tinged with purple or

maroon on the lower surface.

L. lactosa is related to L. peersii Marloth ex W.F.

Barker, which has a similar many-flowered inflorescence

of white flowers with greenish brown markings and one

or two lorate, unmarked leaves tinged with purple or

maroon. L. peersii differs, however, in being a larger

plant with much larger, urceolate flowers without pale

blue bases and with distinctly recurved inner tepals,

included, spreading stamens and an unspotted peduncle.

The two species have similar globose seeds with a termi-

nal, inflated arillode.

Bothalia 28,2 (1998)

137

FIGURE 8. — Holotype of Lacheruilia lactosa, Barker 105 10.

Distribution and habitat

Material of L. lactosa was first collected by Miss W.F.

Barker in October 1933 near Kleinmond, where most of

the records of this species have been made, and it has

since been collected on the Houw Hoek Pass, as well as

at Bot River and beyond Baardscheerdersbos, which is

the most easterly record (Figure 9). It is a fairly rare

species with a restricted distribution, occurring in

Lowland Fynbos on flats and gentle slopes in full sun.

Plants occur singly, and unless in flower are extremely

difficult to detect in their natural habitat, due to their rel-

atively small size compared to the surrounding fynbos

vegetation. As with the related L. peersii, which occurs

in the same area, veld fires would most probably benefit

the flowering performance of L. lactosa , resulting in

more robust specimens more easily noticeable in their

habitat. Unfortunately, the populations of this species in

the Kleinmond area are now under real threat due to

housing development.

Material examined

WESTERN CAPE — 3419 (Caledon): Houw Hoek Pass, (-AA),

Sept. 1946, Lewis 2434 (SAM); Honingklip Farm, near Bot River,

(-AC), Sept. 1966, Barker 10476 (NBG); 29-9-1967, Barker 10510

(NBG); near Kleinmond, (-AC), Oct. 1933, Barker 163 (BOL);

Kleinmond, along rocky coast, (-AC), Oct. 1946, De Vos 238, (NBG);

Kleinmond, W of Malherbe Street, between Piet Le Roux and Bob

Laubscher Streets, (-AC), Oct. 1996, Mostert 194 (NBG); Shaw’s

Pass, (-AD), Oct. 1955, Van Niekerk672 (BOL); beyond Baardscheer-

dersbos, on Elim Road, (-DA), Oct. 1966, Chater s.n. sub NBG93633

(NBG).

Lachenalia leipoldtii G.D. Duncan, sp. nov. L.

physocaulotis W.F.Barker affinis propter inflorescentiam

similarem dense subspicatam floribus oblongo-campan-

ulatis staminibus valde exsertis declinatis pedunculo

dense maculato saepe tumido, sed floribus viridi-flavis et

albis folioque lanceolato canaliculate margine undulato

coriaceo basin conspicue dilatatanti, basi folii arete am-

plectenti differt.

TYPE. — Western Cape, 3219 (Wuppertal): 4.8 km N

of Citrusdal, (-CA), 11-9-1933, T.M. Salter 3608 (BOL,

holo.!, BM; K).

Deciduous, winter- growing geophyte 100-280 mm

high. Bulb subglobose, 10-15 mm diam., white with dark

brown outer tunics. Leaf solitary, lanceolate, canalicu-

late, pale green or yellowish green, suberect or slightly

arcuate, 30-95 x 3-12 mm, upper surface unmarked or

with few purplish magenta spots or blotches, lower sur-

face marked with purplish magenta spots or blotches in

lower third, margin coriaceous and undulate; clasping

base tight, white in lower half with magenta spots or

blotches, yellowish green in upper half with purplish

magenta spots or blotches. Inflorescence subspicate,

many-flowered, erect or suberect, dense, 35-130 mm

long, with a short or long sterile apex up to 20 mm long;

peduncle slender or sturdy, erect or suberect, yellowish

green, lightly to densely marked with purplish magenta

spots or blotches, slightly to conspicuously swollen in

upper half, 45-60 mm long; rachis pale brownish mauve,

slightly swollen in lower third; pedicels white or pale

green, very short, 1-2 mm long; bracts ovate in lower

third, becoming lanceolate above, white, 2-3 x 1-2 mm.

Flowers oblong-campanulate, suberect, patent or slight-

ly cernuous, pale creamy white or pale greenish yellow

and white, sometimes fading to purplish maroon on

herbarium sheets, with greenish brown or pinkish brown

markings; perianth tube pale blue, 1-2 mm long; outer

tepals ovate, pale greenish yellow, 4-5 x 2-3 mm, with

greenish brown or pinkish brown gibbosities; inner

tepals obovate, 6-7 x 2-3 mm, translucent white with

greenish yellow keels and recurved tips. Stamens well

FIGURE 9. — Distribution of Lachenalia lactosa, •; and L. leipoldtii.

138

Bothalia 28,2 (1998)

FIGURE 10. — Lachenalia leipoldtii , drawn from Salter 3608, repro-

duced from the original watercolour by Miss W.F. Barker.

exserted, declinate; filaments white, 9-10 mm long.

Ovary ovoid, pale green, 2x3 mm; style white, protrud-

ing beyond stamens, 9-10 mm long. Capsule ovoid,

membranous, 5x4 mm. Seed globose, 1 mm long, matt

black, with terminal inflated arillode 0.8 mm long.

Flowering time: August to September. Figures 10 & 11.

Etymology, named L. leipoldtii after Dr C. Louis

Leipoldt, celebrated South African naturalist, physician

and poet, who collected this species at Piekenierskloof in

1931, and whose fondness for lachenalias from the

Clanwilliam-Citrusdal area is reflected in his famous

Afrikaans poem ‘Oktobermaand’.

Diagnostic characters

L. leipoldtii is characterised by a dense, subspicate

inflorescence of patent or slightly cernuous, oblong-cam-

panulate, pale creamy white or pale greenish yellow and

white flowers with greenish brown or pinkish brown gib-

bosities. The declinate stamens are well exserted, and the

lower surface of the solitary, lanceolate, canaliculate leaf

is marked with purplish magenta spots or blotches. The

tightly clasping leaf base is white in the lower half and

yellowish green in the upper half, with purplish magenta

spots or blotches. The leaf margin is thickened, and

slightly to distinctly undulate. The specimens recorded

from the Olifants River Valley, Piekenierskloof and

Bidouw only have a slightly swollen peduncle and

rachis, whereas those from Ceres, Karoopoort and Little

Karoo have a conspicuously swollen peduncle and

rachis.

L. leipoldtii is closely related to L. physocaulos

W.F.Barker which has a similar dense subspicate inflor-

escence, oblong-campanulate flowers, well-exserted,

declinate stamens and usually a distinctly swollen

peduncle and rachis. It differs from L. leipoldtii in hav-

ing a subterranean, very loosely clasping leaf base, a lin-

ear, conduplicate, glaucous leaf without markings or

thickened margin, and pale magenta inner tepals and fil-

aments. The two species have similar very small, globose

seeds with a short, terminal inflated arillode.

Distribution and habitat

Material of L. leipoldtii was collected for the first time

by C.L. Zeyher in 1931 on the farm Brakfontein in the

Olifants River Valley (Gunn & Codd 1981) and it has sub-

sequently been recorded from Piekenierskloof and

Bidouw Valley, and to the southeast near Ceres and

I*

FIGURE 1 1 . — Holotype of Lachenalia leipoldtii, Salter 3608.

Bothalia 28,2 (1998)

139

Karoopoort, and further north between Touwsrivier and

Montagu (Figure 9). Plants occur in colonies in sandy soil

in full sun.

Material examined

WESTERN CAPE. — 3218 (Clanwilliam): Farm Brakfontein,

Olifants River Valley, (-DB), Ecklon & Zeyher 45 (SAM); Pieke-

nierskloof, (-DB), Aug. 1931, Leipoldt s.n. sub BOL20456 (BOL).

3219 (Wuppertal): Bidouw, (-AB), Aug. 1945, Bolus s.n. sub

BOL23641 (BOL); 4.8 km N of Citrusdal, (-CA), 11-9-1933, Salter

3608 (BM, BOL, K). 3319 (Worcester): 38.6 km on Ceres-Calvinia-

Sutherland road, (-BA), Sept. 1971, Thomas s.n. (NBG); Karoo Poort,

(-BC), Sept. 1941, Barker 1110 (NBG); Hottentots Kloof, Ceres Dist.,

(-BC), Sept. 1944, Barker 3064 (NBG). 3320 (Montagu): Tweedside,

Laingsburg Dist.. (-AB), Sept. 1932, Barker 162 (BOL); Touwsrivier,

(-AC), Sept. 1974, Obermeyer s.n. sub NBG108599 (NBG). Without

precise locality: between Montagu and Touwsrivier, Sept. 1933, Lewis

& Leighton s.n. (BOL).

ACKNOWLEDGEMENTS

I am very grateful to the staff of the Compton

Herbarium, particularly Dr J.C. Manning, Dr D.A. Pater-

son Jones and Mrs S.E. Foster for their assistance at var-

ious stages of this study. I am also much indebted to Dr

O.A. Leistner for compiling the Latin translations of the

diagnoses, Mrs J. Loedolff for taking the black and white

photographs of the herbarium sheets and Mrs B.A.

Momberg for technical assistance, as well as Ms K. Behr,

Mr L. Mostert, Dr J.P. Rourke and Mr and Mrs R.

Saunders for assistance in the field.

REFERENCES

BARKER, W.F. 1978. Ten new species of Lttchenalia (Liliaceae).

Journal of South African Botany 44: 91^-18.

BARKER, W.F. 1979. Ten more new species of Lachenalia (Liliaceae).

Journal of South African Botany 45: 193-219.

BARKER, W.F. 1983a. A list of the Lachenalia species included in

Rudolf Schlechter's collections made on his collecting trips in

southern Africa, with identifications added Journal of South

African Botany 49: 45-55.

BARKER, W.F. 1983b. Six more new species of Lachenalia

(Liliaceae). Journal of South African Botany 49: 423-444.

BARKER, W.F. 1984. Three more new species of Lachenalia and one

new variety of an early species (Liliaceae). Journal of South

African Botany 50: 535-547.

BARKER, W.F. 1987. Five more new species of Lachenalia (Liliaceae -

Hyacinthoideae), four from the Cape Province and one from

southern South West Africa\Namibia. South African Journal of

Botany 53: 166-172.

BARKER, W.F. 1989. New taxa and nomenclatural changes in

Lachenalia (Liliaceae) from the Cape Province. South African

Journal of Botany 55: 630-646.

DUNCAN, G.D 1987. Lachenalia macgregoriorum. The Flowering

Plants of Africa 49: t. 1951.

DUNCAN, G.D 1988a. The Lachenalia handbook. Annals of

Kirstenbosch Botanic Gardens 17. National Botanical Institute,

Cape Town.

DUNCAN, G.D 1988b. Lachenalia arbuthnotiae. The Flowering

Plants of Africa 50: t. 1961.

DUNCAN, G.D. 1989a. Lachenalia. In B Jeppe, Spring and winter

flowering bulbs of the Cape. Oxford University Press, Cape

Town.

DUNCAN, G.D. 1989b. Lachenalia. In N. Du Plessis & G. Duncan,

Bulbous plants of southern Africa. Tafelberg, Cape Town.

DUNCAN, G.D. 1992. Lachenalia: its distribution, conservation status

and taxonomy. Acta Horticulturae 325: 843-845.

DUNCAN, G.D 1993 Lachenalia thomasiae. The Flowering Plants of

Africa 52: t 2061.

DUNCAN, G.D. 1994. The genus Lachenalia, and the discovery of a

beautiful new species from the Western Cape Province of South

Africa. Shin-Kaki 163: 32-35.

DUNCAN, G.D. 1996. Four new species and one new subspecies of

Lachenalia (Hyacinthaceae) from arid areas of South Africa.

Bothalia 26: 1-9.

DUNCAN, G.D. 1997a. Five new species of Lachenalia (Hya-

cinthaceae) from and areas of South Africa. Bothalia 27: 7-15.

DUNCAN, G.D. 1997b. Lachenalia rosea. Flowering Plants of Africa

55: t. 2126.

GUNN, M. & CODD, L.E. 1981. Botanical exploration of southern

Africa Balkema, Cape Town.

■

Bothalia 28,2: 141-149 (1998)

A revision of Lachenalia (Hyacinthaceae) in the Eastern Cape, South

Africa

A.P. DOLD* and P.B. PHILLIPSON*

Keywords: Eastern Cape, Hyacinthaceae, Lachenalia Jacq.f. ex Murray, South Africa, taxonomy

ABSTRACT

A taxonomic account of the genus Lachenalia Jacq.f. ex Murray in the Eastern Cape, South Africa, is given. Eight

species are recognised, and descriptions of these are amended and elaborated as necessary, three taxa have been reduced to

synonymy and five species erroneously recorded within the province are excluded. Three of the species are endemic to the

province. An identification key is provided.

INTRODUCTION

The genus Lachenalia is confined to southern Africa

and consists of a little over 100 species (Arnold & De

Wet 1993). The majority of these species occur in the

winter rainfall region of the Western Cape in the sclero-

phyllous shrublands or ‘fynbos’ of the Cape floristic

region (sensu White 1983). The geographic range of a

few species extends from this region into the western

parts of the Eastern Cape and one even occurs as far

northeast as the Free State. The few remaining species

are endemic to the Eastern Cape. The genus has attracted

attention as an horticultural subject, and some of the

more decorative species have become well known

through cultivation. Despite this, no revision of the genus

has been attempted since Baker’s (1897) treatment in

Flora capensis; however the work of Barker (see Arnold

& De Wet 1993 for references) and Duncan (1988, 1996,

1997), has resulted in the species from Western Cape

becoming reasonably well known taxonomically. The

same cannot be said for the Eastern Cape species. They

have mostly been unknown in cultivation, and they are

generally represented only by few, rather old herbarium

specimens.

In the course of field work in the Eastern Cape it

became clear that Baker’s treatment was far from ade-

quate. Observations of populations of certain species

indicated that they are very variable, and that many of

Baker’s diagnostic characters are unreliable. It was clear

that a knowledge of the plants in the field would be

essential in producing an effective treatment for the

genus in the province, and that a wider range of charac-

ters should be considered.

Key to the species of Lachenalia in the Eastern Cape

la Perianth more than 15 mm long, always yellow-green; flower erecto-patent (45° or less to inflorescence axis) at

and after anthesis; confined in Eastern Cape to coastal grassland 1. L. algoensis

lb Perianth less than 15 mm long, usually white to deep pink or purple, sometimes dull green with brown markings

or rarely yellow-green (see L. bowkeri): flower cemuous to patent at anthesis, sometimes erecto-patent after

anthesis; not confined to coastal grassland:

2a Inner perianth segments always recurved at tips, usually exceeding outer by at least 1.4 mm:

3a Leaf irregularly marked with brown, green or maroon blotches on upper surface, margin coriaceous; stamens

pale to dark maroon, well exserted 8. L. karooica

3b Leaf not marked on upper surface, margin not coriaceous; stamens white, included or exserted:

4a Stamens nearly twice as long as inner perianth segments, exserted by 3-13 mm, spreading; perianth cam-

panulate, inner lobes exceeding outer lobes by up to 2 mm 5. L. latimerae

4b Stamens included or exserted up to 3 mm, declinate; perianth oblong-campanulate, inner lobes exceeding

outer by 2 mm or more:

5a Stamens not or very slightly exserted; outer perianth lobes recurved, lower inner lobe broadly flared; leaf

banded dark green below, confined to an area extending from Eastern Cape to Karoo 7. L. perryae

5b Stamens exserted by ± 2 mm or more; outer perianth lobes not recurved, lower inner lobe not broadly

flared; leaf not banded below; extending from coastal areas to Karoo 2. L. bowkeri

2b Inner perianth segments straight or slightly spreading but not recurved at tips, ± equal in length to outer seg-

ments:

6a Stamens included; perianth broadly cup-shaped, as long as broad, ± 5 mm long, segment apices not spread-

ing; flower cemuous at anthesis; inflorescence lax, 3-5 mm, between adjacent pedicels .... 4. L. convallarioides

6b Stamens shortly exserted (up to 2 mm); perianth campanulate, longer than broad, ± 5.5 mm long or more,

segment apices spreading; flower patent at anthesis; inflorescence very dense with less than 2 mm

between adjacent pedicels:

7a Perianth up to 5.6 mm long, campanulate; endemic to high mountains of Eastern Cape ... 3. L. campanulata

7b Perianth at least 8 mm long, oblong-campanulate; confined in Eastern Cape to coastal areas in extreme

west 6. L. youngii

* Selmar Schonland Herbarium, Department of Botany, Rhodes University, P.O. Box 94, 6 1 40 Grahamstown, South Africa. BOTD@RHOBOTRU.AC.ZA

MS. received: 1997-12-10.

142

Bothalia 28,2 (1998)

In addition to more conventional macro-morphologi-

cal characters, Barker (1978) found seed morphology to

be particularly useful in delimiting the different taxa

in the Western Cape. Ornduff & Watters (1978) and

Johnson & Brandham (1997) have provided chromosome

counts for several species. The value of these characters

in understanding the Eastern Cape species has not previ-

ously been assessed.

MATERIALS AND METHODS

The present account is based on a study of herbarium

specimens from relevant herbaria (see acknowledge-

ments), and of living plants in the field and in cultivation.

A total of 16 populations representing five of the Eastern

Cape species were located and detailed descriptive notes

of living plants were made. Particular attention was paid

to differential characters used by previous researchers

and to characters that appeared to be variable within each

population. Representative specimens were collected for

cultivation and as herbarium specimens. Seeds of all

species were examined with a JEOL-JSM 840 scanning

electron microscope.

1. Lachenalia algoensis Schonland in Transac-

tions of the Royal Society of South Africa 1: 443 (1910);

Fourc.: 101 (1941); A. Batten & Bokelmann: 16 (1966);

E.Gledhill: 70 (1981); A.Moriarty: 34 (1982); Bond &

Goldblatt: 51 (1984); G.D. Duncan: 35 (1988). Type:

Eastern Cape, Port Elizabeth, 13 August 1903, Drege 64

(GRA, syn. ! ); Redhouse, Paterson 92 (syn. non vide).

Lachenalia algoensis was described in detail by

Schonland in 1910. No elaboration of Schonland's mor-

F1GURE 1 . — Seeds of Lachenalia. A, L. algoensis, Tail 155', B, L. bowkeri, Dold 280', C, L. campanulata, Dold 1233', D, L. convallarioides, Dold

1018, E, L latimerae, Dold 1802', F, L. youngii. Hops 33. Scale bars: 100 pm.

Bothalia 28,2 (1998)

143

FIGURE 2. — Inflorescences of Lachenalia. A, L. algoensis, plant from Linton Grange, Port Elizabeth (no voucher). B-D, L. bowkeri : B, Dold

2267\ C, Dold 2269\ D, Dold 1766. E, L. campanulata, Dold 1233\ F, L. convallarioides, Dold 1018. Scale bars: 10 mm.

phological description is necessary, except to note that

the seeds are ± 2.2 x 1.1 mm, black and shiny, with a

smooth testa and a blunt decurrent arillode, ± 1.0 mm

long (Figure 1A). It is relatively well known and is clear-

ly distinct from all other species that occur in the Eastern

Cape, being by far the largest flowered species in the area

(Figure 2A).

Distribution and habitat

In the Eastern Cape it ranges from the Knysna District

in the west to the Bushman’s River in the east, always

near the coast (Figure 3). According to Duncan (1988), it

extends to Worcester in the Western Cape.

Vouchers: Burrows 3355 (GRA); Cruder. 388 (PRE); Dold 691 (GRA);

Schonland s.n. (GRA); Tait 3761 (PEU).

2. Lachenalia bowkeri Baker in Flora capensis 6:

427 (1897); Bond & Goldblatt: 52 (1984); G.D.Duncan:

32 (1988); Urton & D.Page: 124 (1993). Type; Somerset

Division, Bowker s.n. (K, holo.!).

L. subspicata Fourc.: 79 (1934); Fourc.: 101 (1941). Type: Humans-

dorp Division, Flats Zuur Anys, on road to Kouga, Fourcade 3044

(BOL, holo.!).

Plant (50—) 1 1 0(— 2 1 0) mm tall. Bulb globose, 8-17

mm diam., inner flesh translucent, white, outer tunics

dry, coarsely papery, dark brown, often flaking, inner

tunic minutely white-spotted, sometimes forming a short,

coarse, brush-like neck of 5-10 mm, just visible at ground

level; leaf sheath membranous, translucent, extending

above neck by 5-15 mm, loosely clasping. Leaves

1— 2( — 4), (45— )60— 1 20(— 300) x (5-)8-10(— 30) mm, but

frequently eaten by herbivores and thus appearing un-

naturally short, deeply channelled, erect, slightly fleshy,

144

Bothalia 28,2 (1998)

FIGURE 3. — Distribution of Lachenalia algoensis , •; L. lairooica, A;

and L. perryae, H

smooth, pale green, sometimes with faint purple margin,

sometimes with bluish blush; base often clasping pedun-

cle up to 20-40 mm, extreme base white, occasionally

very faintly purple-spotted just above ground; apex

bluntly acute, often dry; keel indistinct. Inflorescence

erect, (40— )60— 80(— 1 40) mm tall, spicate to subspicate;

peduncle and inflorescence axis usually slender, up to

2-3 mm thick, thickest just below first flower, pale

green, very pale at base, sometimes spotted dull purple;

fertile part ( 1 0— )30— 50(— 70) x 15-25 mm, with 4-25

(often about 10) flowers; sterile tip tufted, 2-6 mm long;

bracts cupped, deltoid, with slightly winged margins,

white to greenish white, 1.1-1. 8 x 1.1-1. 8 mm, with

basal spur up to 0.7 mm long, apex acute, 0.7-0. 9 mm

long, facing downwards or curling under. Flowers some-

times (but not always) sweetly scented; pedicels held at

40° to stem, 0. 5-2.0 x 0.3-0. 4 mm, white to pale green;

perianth patent or more usually somewhat cernuous,

oblong campanulate, (6.5— )9.0(— 1 1.0) mm long, (3.0-)

3.6(-4.0) mm diam. at base, slightly swollen at base,

varying from white to pale green or very pale purple to

purple with green or purple distal keels, becoming dark-

er with age; inner lobes obovate, 6-10 x 3. 0-4. 5 mm,

spreading outwards, all 3 inner lobes of equal length,

exceeding outer by (1 .5— )2.0(— 3.0) mm; apex obtuse,

recurved; base cuneate, sometimes speckled with blue;

outer lobes ovate, 4. 6-6.0 x 2. 1-3.1 mm, not spreading

outwards; apex obtuse; base truncate with very slight

gibbosities which are most pronounced on upper lobe,

unevenly speckled or blotched with pale purple or pale

blue. Stamens declinate; filaments white, 7-9 mm long;

anthers 0.8-1. 4 x 0.5-0. 8 mm, maturing in stages, low-

est pair first and upper pair last, included within perianth

before maturity and purple, then exserted by ( 1 — )2(— 3)

mm and yellow when mature and finally becoming

black, and eventually, as fruit develops, being drawn

back into perianth. Ovary green, 1.7-3. 2 x 1. 5-2.0 mm;

style exserted, 6. 3-9.0 mm long, white. Capsule ovoid,

2. 4-4.0 x 1. 5-4.0 mm, papery, brown. Seed 1 .4-2.2 x

1.1-1. 5 mm; testa reticulate or smooth, black, shiny;

arillode bluntly pointed, 0.45-0.5 mm sometimes short-

ly decurrent. Chromosome number. 2n = 16 ( Dold 280,

Kew reference 95-30, Kew accession 1995-26, Johnson

& Brandham 1997). Figures IB; 2B-D.

Duncan (1988) described L. bowkeri as a poorly

known species. We interpret it as an extremely variable

species, which is actually well represented in herbaria

compared to most of the other Eastern Cape species. The

variation in L. bowkeri is so great that during the course

of the present study we were convinced for a long time

that it represented at least three distinct species. Variation

occurs in several characters, notably: overall size, num-

ber of leaves, flower orientation, pedicel length, perianth

colour and degree of exsertion of the stamens. Variation

in these characters can be seen among individual plants

in a population, and is certainly attributable in part to

phenotypic responses to minor variations in microhabitat.

Considerable phenotypic plasticity has also been noted in

plants brought into cultivation. However, different popu-

lations of L. bowkeri do show certain relatively constant

and distinct features and these appear to some extent to

be related to geographical distribution. It may eventually

be possible to describe infraspecific taxa within L. bowk-

eri and thus delimit these variations formally, but at pre-

sent we understand too little of the nature of the species’

variability to enable us to do so confidently. Further cyto-

logical studies of the different populations will be done at

Kew and may help to clarify this variation.

Interpretation of herbarium material is complicated by

the tendency in old flowers for the stamens to be drawn

into the perianth and for the inner perianth lobes to

become constricted at the mouth (Figure 2C). In this con-

dition the flowers closely resemble those of certain other

species and present a very different facies to that of care-

fully pressed specimens with freshly opened flowers. At

anthesis, stamen exsertion varies from 1 to 3 mm, but the

type specimen is a plant with old flowers and thus in

Baker’s (1897) original description, L. bowkeri is stated

to have included stamens.

Baker also states that the leaf is solitary. However,

occasional 2-leaved plants were found in the populations

we studied. Very rarely, exceptionally vigorous bulbs

were found that had three or even four leaves and two

inflorescences. Plants of L. bowkeri have regularly been

misidentified in herbaria (often as L. orchioides (L.)

Aiton), since they are frequently at variance with Baker’s

description with respect to the exsertion of the stamens

and the number of leaves.

Plant height, leaf dimensions and the number of flow-

ers per inflorescence vary considerably within popula-

tions and from one population to another depending on

exposure. Plants found in open karroid grassland are

smaller than those shaded by grass clumps in open

coastal grassland. Plants found in salt pans are more

robust but not necessarily taller. In several localities

robust plants with particularly thick peduncles have been

collected. The bulb tunic appears to be influenced by the

type of substrate and plants growing in hard, dry, rocky

ground will always have thick, dry flaking scales with a

short brush-like neck, whereas in loam or sandy soil the

tunics will be less scaly and most often without a neck at

all. The base of the leaf often tightly clasps the peduncle

for up to a third of its length and then opens abruptly.

This is seen predominantly in karroid areas, whereas

under favourable conditions leaves are broader and sel-

dom clasping, sometimes in pairs, one always much

smaller than the other. From observations in the field and

Bothalia 28,2 (1998)

145

from herbarium specimen labels it is also clear that L.

bowkeri is variable in coloration. The perianth of plants

from the dry Subtropical Thicket of the Fish River valley

vary from white, tinged with green, with pale mauve dis-

tal keels, to entirely purple. Those from Naude’s Hoek in

the Keiskamma River valley are pale green to dull yel-

low-green with dull purple markings, while further west

at Ghio Marsh Nature Reserve, the perianth is white with

green speckling on distal keels and blue speckling on the

base of the outer lobes, and older flowers already closed

become tinged with dull purple. Within populations the

flower colour seems to be more or less uniform,

although, as in other species in the Eastern Cape, the

individual flowers become darker in colour with age,

tending to fade to dull purple as the fruit develops.

Flower size, orientation and the extent to which the peri-

anth lobes are recurved or spread, particularly at the tips,

is somewhat variable (Figure 2B-D), but tends to be rel-

atively constant within populations. Pedicel length shows

little variation, from sessile to only 2 mm long.

Seed morphology is variable from one population to

another. Seeds from herbarium sheets and live plants

have been examined. Figure IB is representative of

plants from the Fish and Keiskamma River valleys. Seed

from collections further west are of similar size, but tend

to have a rather variable arillode, which is often more

blunt and less strongly decurrent.

Lachenalia subspicata was described in 1934 by

Fourcade, who compared his new species only with L.

orchioides and L. youngii Baker. It appears that he over-

looked the similarity between his plant and L. bowkeri ,

possibly because Baker (1897) incorrectly described the

latter as having 'quite sessile flowers’. On examination

of the type, a few very short pedicels can be seen on one

inflorescence. Following ‘Pretoria National Herbarium

practice’, Arnold & De Wet (1993) have placed L. sub-

spicata into synonymy with L. bowkeri. On examination

of the type collections and consideration of the variabili-

ty of the species we concur with this practice.

Distribution and habitat

Lachenalia bowkeri is virtually confined to the

Eastern Cape, with only two known collections from the

Western Cape. The type locality is ‘Somerset’ (presum-

ably Somerset East), but this is the only collection known

from there, the majority of collections are from the

Albany District, with others from the Alexandria and Port

Elizabeth Districts (Figure 4). All collections are from

below 850 m and by far the majority of these are found

at altitudes below 600 m. It usually occurs in open grass-

land and often on margins of salt pans usually partially

shaded under grass clumps, or, less commonly, in drier,

rocky areas of Subtropical Thicket on south-facing

slopes. Populations are small and localised and plants

occur singly or in small groups of two or three in areas

not exceeding 10 m* 2.

Vouchers: Dold 2267 , 2268. 2418 (GRA); 2269, 2419 (GRA, PRE).

3. Lachenalia campanulata Baker in Journal of

Botany 12: 6 (1874); Baker: 432 (1897); Bond & Gold-

blatt: 52 (1984). Type: Somerset East, In lapidosis sum-

mii Montis Boschberg, 4800ft, (-DA), MacOwan 1836

(K, holo.!; GRA, iso.!).

21 22 23 24 25 26 27 28 29 30 31

FIGURE 4. — Distribution of Lachenalia bowkeri , ▲; and L. latimer-

ae, •.

L. rhodantha Baker: 430 ( 1 897). Type: Graaff Reinet, Sides of grassy

mountains, Oudeberg, Graaff Reinet, 4300ft, (-DD), Bolus 719 (K.

holo.!).

Plant (50—) 1 1 0(— 230) mm high depending on amount

of shade. Bulb globose, 5-15 mm diam., white, fleshy,

outer tunics dry, membranous, pale brownish, peeling in

places, not neck-forming. Leaves 2, rarely 1, linear, erect,

( 1 5.0— )8.6(— 30.0) x 1-7 mm, not sheathing, smooth,

entire, drying at tips, base white with purple spots, blade

light green and unmarked to densely covered with minute

red spots, subterete, keel indistinct. Inflorescence subspi-

cate to racemose, 10-20 x 2— 3( — 5) mm, flowers dense,

number of flowers (4—) 1 6(— 34); peduncle erect with

slight curve, white at base to green often densely to

entirely covered with minute red spots, thickest at base,

2. 0-3. 5 mm, becoming thinner closer to inflorescence;

sterile tip 2-3 mm, not tufted; pedicels spreading, rarely

sessile, 2.5(— 3.0) mm, white to pale red; bracts 2 mm

wide, with acute spreading apex, cup-shaped, 1.5 mm

deep, slightly spurred at base, white tinged green and

minutely red-spotted; lower bracts ± 2 mm long; upper

bracts ± 1 mm long. Flowers unscented, perianth spread-

ing to horizontal, not becoming cernuous in lower half of

inflorescence, campanulate, base rounded, slightly

swollen, 5.6 mm long, 3.3 mm at broadest point, inner

segments oblong, 5.5 x 1.3 mm, outer segments oval

elliptic, 5.5 mm long, 2.6 mm wide at base, inner seg-

ments not exceeding outer, segments spreading in distal

end, ranging from pure white to red at anthesis, darken-

ing soon afterwards to purple; gibbosities indistinct,

tinged faintly greenish. Stamens shortly exserted by up to

2 mm, becoming included soon after anthesis, yellow;

filaments white. Ovary ovoid, green, 2.1 x 1.4 mm; style

2.7 mm, finally exserted, persistant. Capsule ovoid, 3.8 x

2.4 mm, papery, brown. Seed black, shiny, ovoid, ± 1.7 x

1.2 mm, testa reticulate, arillode short, tapering, assym-

metrical, decurrent, 0.2 mm long. Figures 1C; 2E.

Lachenalia campanulata can be distinguished by the

combination of its spreading to horizontal (lowers about

5.6 mm long with exserted stamens and subequal peri-

anth lobes, and its (usually) paired leaves. In the Eastern

Cape it most closely resembles L. convallarioides Baker

which has flowers of similar size and also has subequal

146

Bothalia 28,2 (1998)

perianth lobes. The two species differ in several charac-

ters. L. campanulata has a relatively dense inflorescence

and the thickest point of its peduncle is at the base; in L.

convallcirioides the inflorescence is rather lax and the

thickest point of its peduncle is immediately below the

first flower. In L. campanulata the perianth is campanu-

late and the lobes are reflexed, whereas in L. convallari-

oides it is cup-shaped and the lobes are not reflexed. In L.

campanulata seeds are ± 1.7 mm long and have a short,

tapering, assymmetrical, decurrent arillode, and a reticu-

late testa; those of L. convallarioides are ± 2 mm long,

and have a rounded, symmetrical, terminal, arillode, and

a smooth testa.

Plants of L. campanulata are variable in size depend-

ing on the degree of exposure and individuals 230 mm

tall can be found in sheltered positions in close proximi-

ty to others only 60 mm tall in exposed positions. Colour

is also very variable from pure white to pink to deep red

at anthesis, darkening to purple.

Baker’s description of L. rhodantha is based only on

the type collection. Bolus 719. It was said to be distin-

guished from L. campanulata by having a single leaf

rather than two, lower Bowers cernuous rather than hori-

zontal, stamens exserted, not included, and being bright

red rather than white tinged with red. Not having seen

these plants in their natural habitat Baker could not have

known of the variability of these supposedly diagnostic

characters, and plants closely matching the type speci-

mens of both species can be found growing together. In

all three populations we have studied in the field, most of

the plants were 2-leaved, whereas occasional 1 -leaved

and even 3-leaved plants were found in each. All of the

specimens we have examined (living and preserved),

including the type of L. rhodantha, have spreading to

horizontal flowers in the lower part of the inflorescence.

Baker’s description of the orientation of the lower flow-

ers of L. rhodantha is an error, as they are clearly hori-

zontal. The type collection of L. campanulata, MacOwan

1836, consists of plants past anthesis which have their

stamens included and style excluded, dark perianths and

ovaries obviously swollen. In the field we have recorded

that young plants consistently have spreading flowers

with a slightly swollen base, an open mouth and exserted

stamens. After anthesis the flower becomes horizontal

and darker in colour, the perianth lobes close at the

mouth and the stamens become drawn into the perianth

and the base of the flower swells with the ovary.

Distribution and habitat

Lachenalia campanulata is confined to the mountains

of the interior of the Eastern Cape between 24° and 29°

E and 30° to 33° S, and has been recorded at altitudes

from 1 200 to 2 400 m on steep, open slopes (Figure 5).

Vouchers: Cotterrell 52 (GRA); Dold 1233. 1993 (GRA, PRE);

Hilliard & Burn 14718 (NU, PRE); Van Der Wall 256 (PRE).

4. Lachenalia convallarioides Baker in Journal of

the Linnean Society 1 I; 407 (1871); Martin & Noel: 26

(1960); Dold: 29 (1994). Type: Eastern Cape, Kreli’s

Country, Caffraria, Bowker 444 (K, holo.!; TCD, iso.!).

L. convallarioides Baker var. robusta Baker: 407 (1871). Type:

Albany, Williamson s.n. (TCD, holo.!).

21 22 23 24 25 26 27 28 29 30 31

FIGURE 5. — Distribution of Lachenalia campanulata, •; L. conval-

larioides, A; and L. youngii, ■

Plant (80-)135(-250) mm high, depending on amount

of shade. Bulb globose 10-15 mm diam., white fleshy,

outer tunics dry, membranous, pale brownish, peeling in

places, not neck-forming. Leaves 1 (—2), 80-280 mm

long, as long as inflorescence or occasionally up to one

third of its length longer than inflorescence, especially in

more robust specimens; sheathing up to 20 mm, 8-20

mm wide at broadest point, linear to lorate, margin entire,

apex acute, erect with a shallow 40° curve away from

peduncle, fleshy, often with a slight twist in axis, persis-

tent apex dry and shrivelled, deeply U-shaped in cross

section; keel indistinct, smooth, glabrous, pale green,

extreme base white or densely, minutely spotted purple

red (BCC 822). Inflorescence racemose, flowers sparse

(5—) 1 9(— 60), (20— )40(— 1 10) mm long; peduncle erect,

slender, (50-)95(-140) mm, shallowly curved, thickest

point 1-2 mm, immediately below first flower, pale

green to pale pink near apex; sterile tip 2 mm, not tufted;

pedicels erect-patent, spreading, 2. 5-5.0 mm, white to

pinkish; bracts 2 mm across, deeply cupped, 1 mm long,

with acute patent apex, 0.9 mm, and basal spur. Flowers

unscented; perianth horizontally spreading to cernuous at

anthesis and then horizontal in older flowers, campanu-

late, rounded at base, 5 mm long, 4 mm diam. at base,

inner segments obovoid, 5 mm long, 3.5 mm at broadest

point, shorter than the outer, outer segments white, 2.5

mm long; anthers white at anthesis, black after anthesis.

Ovary ovoid, green, 3-locular, 3.0 x 2.5 mm; style 2.5

mm long. Capsule ovoid, 4x3 mm, tissue-like, finely

net-veined, golden yellow. Seed black, shiny, smooth,

ovoid, 2.0 x 1.2 mm; terminal arillode symmetrical,

blunt, 0.56 mm. Chromosome number : 2n = 30 ( Dold

1018, Kew reference 95-31, Kew accession 1995-27).

Figures ID; 2F.

Lachenalia convallarioides is recognised by its rather

lax inflorescence of nodding flowers with included sta-

mens and subequal perianth lobes, and its (usually) sin-

gle leaf. Other characters that serve to distinguish it from

L. campanulata, the Eastern Cape species it most closely

resembles, are given under that species.

L. convallarioides is very variable in leaf and inflor-

escence size depending on the plant’s degree of exposure

Bothalia 28,2 (1998)

147

to the sun. The most exposed plants are the most con-

spicuous in natural populations but as a result of their

exposure are also the smallest, and these match the type

collection. Plants growing in shaded places, partly hid-

den by other vegetation are rather inconspicuous, and can

be much larger. Baker, not having seen this variation

within populations, assigned varietal status to William-

son’s collection (71 Williamson s.n.) which clearly match-

es shade-growing plants in populations we have seen.

The chromosome number of 2n = 30 is unusual in the

genus Lachenalia, where the common basic number is x

= 7 or 8, although x = 5, 9, 10, 11, 12, 13 and 15 have also

been found. Johnson & Brandham (pers. comm.), have

suggested that the 2n = 30 (x = 15) could be an allote-

traploid derived from taxa with x = 7 and x = 8 following

hybridisation and chromosome doubling.

Distribution and habitat

Lachenalia convallarioides is only known from the

Albany District near Grahamstown in the Eastern Cape,

where it grows on rocky outcrops of Witteberg Quartzite,

and from the Mount Arthur Range about 180 km to the

northeast, at altitudes of around 17 to 1 800 m (Figure 5).

Populations are very localised resulting in their being

easily overlooked even in well-explored localities such

as on Mountain Drive, Grahamstown. This locality is

within walking distance from the Botany Department at

Rhodes University but, remarkably, only three collec-

tions have ever been made from this generally well-col-

lected area.

Bowker’s type collection from Kreli’s Country,

Caffraria, and Barber’s Zuurberg collection, both num-

bered 444, are almost certainly the same collection from

what was known in 1856 as the Zuurberg in the extreme

northwest of Chief Kreli’s territory in Caffraria (Hall

1856). These mountains are now known as the Mount

Arthur Range. Bowker, a soldier, was stationed in this

area and collected plant specimens for his sister, Mrs

M.E. Barber (nee Bowker), (Thorpe 1977; Gunn & Codd

1981), who sent them to Harvey at TCD. Baker saw this

material much later (Dr John Parnell, Trinity College,

Dublin, pers. comm.). The Bowker/Barber collection is

the only known collection from outside the Albany

District.

Vouchers: Barber 444 (TCD); Borman s.n. sub RUH10632 (GRA);

Dold 1018 (GRA. K); Pym s.n. (GRA); Williamson s.n. (TCD).

5. Lachenalia latimerae W.F. Barker in Journal of

South African Botany 45:196 (1979); Bond & Goldblatt:

52 (1984); G.D. Duncan: 60 (1988). Type: Ferndale Farm,

Patensie, (-DD), July 1949, Courtenay-Latimer s.n. sub

NBG72287 (NBG, holo.!).

Lachenalia latimerae was described by Ms W.F.

Barker in 1979 from the type and from cultivated mater-

ial collected near Patensie ( Bayliss 7102). Recently we

have located two new populations of this species.

Barker’s description needs no further elaboration. In

common with most of the other species in the Eastern

Cape it should be noted that overall size varies consider-

ably. Seed morphology of our collections conforms to

that of the type collection. Figures IE; 6.

FIGURE 6. — Inflorescence of Lachenalia latimerae. Dold 1802. Scale

bar: 10 mm.

Distribution and habitat

L. latimerae is found in the Kouga Mountains near

Patensie in the Eastern Cape and it has also been collect-

ed at the Cango Caves near Oudsthoorn in the Western

Cape. It has been found at altitudes from 320 to 1 000 m

(Figure 4). The two populations of L. latimerae that have

been examined in the field were both very localised, with

less than 50 plants occurring in an area of ± 8 m2 in each

case. Both were situated at the base of a very steep south-

facing rocky slope with dense pockets of bush. The

plants occur in open vegetation, often in moss-covered

patches of about 1 m2 in full shade.

Vouchers: Bayliss 7102 (NBG); Clark 1031 (GRA, PRE), DC1014

(GRA, K); Dol'd 1802 (GRA, PRE), 2417 (GRA).

6. Lachenalia youngii Baker in Flora capensis 6:

433 (1897); Fourc.: 101 (1941); Bond & Goldblatt: 54

(1984); G.D. Duncan: 42 (1988). Type: Oudtshoorn,

Montagu Pass, (-CD), 1200 ft, September 1889, Young

5545 (K, holo.! and iso.!; BOL, iso.!).

Lachenalia youngii was described by Baker (1897)

and by Duncan (1988). No further elaboration is neces-

sary.

Baker (1897) only cited a single sheet of Young 5545

under L. youngii. He identified a second sheet labelled

148

Bothalia 28,2 (1998)

Young 5545 as L. unicolor. We have examined both sheets,

and conclude that they are without any doubt duplicates

and they conform perfectly to Baker’s description of L.

youngii. Baker’s misidentification of the sheet accounts

for erroneous distribution records for L. unicolor in the

literature. It is uncertain whether seed morphology is

diagnostic or not as only a single specimen has been

examined (Figure IF).

Distribution and habitat

Lachenalia youngii has been recorded from numerous

localities in the mountains of the southeastern part of the

Western Cape, and extends into the Eastern Cape near

Humansdorp (Figure 5), growing in fynbos.

Vouchers: Fourcade 347 (BOL, GRA), 5385 (BOL); Rogers 27962

(GRA); Vlok 1207 ( K); West 231 (GRA).

7. Lachenalia perryae G.D. Duncan in Bothalia

26: 3 (1996). Type: Western Cape, Karoo National

Botanical Garden veld reserve, Worcester, Aug. 1985,

Perry s.n. (NBG, holo.; PRE).

Duncan’s (1996) description needs no elaboration.

Lachenalia perryae may be recognised as follows: it usu-

ally has a single narrowly lanceolate, banded leaf, which

does not widen abruptly at the base; patent or slightly

cernous, oblong-campanulate flowers with pale blue

outer perianth segments and whitish inner segments, and

included stamens.

Distribution and habitat

The species is essentially a Western Cape species with

the majority of collections from the Karoo National

Botanical Garden at Worcester. It has been recorded from

Worcester District south to Port Beaufort and eastwards

to Albertinia (Duncan 1996). One isolated record from

Kommadagga in the Eastern Cape is a wide disjunction

(Figure 3). The occurrence of the plant in the Eastern

Cape needs further study, especially since the species

appears to be quite closely related to L. bowkeri.

Vouchers: Bayliss 5919 (PRE).

8. Lachenalia karooica W.F.Barker ex G.D. Dun-

can in Bothalia 26: 1 (1996). Type: Free State, Faure-

smith, veld, 0.5 km on road from Fauresmith to

Koffiefontein, 6 Aug. 1976, Chaplin s.n. (NBG, holo.).

Duncan’s (1996) description needs no elaboration.

Lachenalia karooica is easily recognised in the Eastern

Cape by its maroon stamens and the coriaceous margins

of its leaves together with its northwestern distribution.

Distribution and habitat

Lachenalia karooica has been recorded from scattered

localities in the Great Karoo in the Western Cape, the

Northern Cape and the Free State, and extends into north-

ern parts of the Eastern Cape just south of the Orange

River (Figure 3). At the type locality plants grow on

south-facing aspects on dolomite outcrops.

Vouchers: Burrows 2157 (PRE); Meyer 206 (PRE).

EXCLUDED SPECIES

Lachenalia aloides (L.f.) Engl.

Baker (1897) cited a specimen from Hill Park, Port

Elizabeth {East South Central African Herbarium 49 K!)

as L. aloides. The specimen was wrongly identified by

Baker — it is L. algoensis. Duncan (1988) regards L.

aloides as restricted to the Western Cape, a view with

which we agree.

Lachenalia orchioides (L.) Aiton

Baker (1897) cited two specimens from what is now

the Eastern Cape, both from Grahamstown: MacOwan

1337 (GRA!) and Galpin 228 (PRE!). On examination of

the specimens it is immediately clear that they are both L.

bowkeri. A view shared by Barker ( determinavit labels,

no dates). We have not seen any collections of true L.

orchioides from our area, and this species appears to be

confined to the extreme southwestern part of the Western

Cape. A large number of other specimens of L. bowkeri

have been misidentified in herbaria as this species.

Lachenalia pustulata Jacq.

A Western Cape species. Baker (1897) cited a single

specimen for the summer rainfall area: Ecklon & Zeyher

38 (PRE!), Zwartkops River, Uitenhage. This specimen

is L. bowkeri Baker, a view shared by Barker {determi-

navit label, no date).

Lachenalia reflexa Thunb.

Baker (1897) cited Bolus 2635 (K!) from the Zuurberg

Range, Alexandria Div. in the Eastern Cape. This was

incorrectly determined and is L. algoensis. A view shared

by Barker ( determinavit label, 9/5/1967).

Another collection, just out of the province, Fourcade

1447 (K!), (locality Knysna), is also L. algoensis incor-

rectly determined as L. reflexa.

Lachenalia trichophylla Baker

Baker originally cited the type of L. trichophylla as

MacOwan 2197 from Somerset East, i.e. within the

Eastern Cape, and cited no other specimens. In Flora

capensis (Baker 1897), however, he cited a single speci-

men for the species: Mader sub herb. MacOwan 2167

from Clanwilliam (in the Western Cape). The reason for

this change is unknown but MacOwan in his list of dis-

tributed collections recorded MacOwan 2197 as Catha

edulis , whereas MacOwan 2167 is recorded as Gladiolus

spathaceus. No specimens of a Lachenalia bearing the

collection number 2197 have been traced in relevant

herbaria; however, specimens of Mader sub herb.

MacOwan 2167 are present in K and SAM. The descrip-

tion of L. trichophylla does not match any known Eastern

Cape material, but does agree with the Mader specimens,

and other material from the Western Cape referred to as

L. trichophylla. It is concluded that the original citation

contains two errors, firstly the collection number should

have been 2167 not 2197, and the specimen was not col-

lected in Somerset East.

Bothalia 28,2 (1998)

149

SPECIMENS EXAMINED

Acocks 16543 (3) PRE; 21174 (6) PRE; 21461 (1) PRE. Anon. s.n. (4)

GRA. Archibald 4388 (2) GRA; 4522 (1) GRA; 5983 (2) GRA.

Barber 444 (4) TCD. Barnard 574 (2) PRE. Bayliss 5919 (7) PRE;

7102 (5) NBG. Bolus 719 (3) K. Borman s.n. sub RUH10632 (4) GRA.

Botha 5881 (2) PRE. Bowker s.n. (2) K; 444 (4) K. Burrows 2157 (8)

PRE; 5355(1) GRA.

Clark DC1014 (5) GRA, K; 1031 (5) GRA, PRE. Cook 20452 (2) BOL.

Cotterrell 52 (3) GRA. Courtenay-Latimer s.n. sub NBG72287 (5)

NBG. Cruden 5a (2) GRA; 22 (2) GRA; 388 (1) PRE; 453 (2) GRA.

Dold 233 (3) GRA, PRE; 280 (2) GRA, K; 691 (1) GRA; 1018 (4)

GRA. K; 1765 (2) GRA; 1766 (2) GRA; 1802 (5) GRA. PRE; 1993 (3)

GRA, PRE; 2076 (3) GRA, K, PRE; 2267. 2268. (2) GRA; 2269 (2)

GRA, PRE; 2270 (2) GRA; 2272 (2) GRA; 2273 (5) GRA; 2417 (5)

GRA; 2418 (2) GRA; 2419 (2) GRA, PRE; 2420 (2) GRA. Drege s.n.

(1) GRA; s.n. (1) GRA : s.n. (2) GRA; 61 (1) GRA; 64 (1) GRA, PRE

Dyer 2085 (2) GRA.

Ecklon 38 (2) PRE. Esterhuysen 10702 (6) BOL.

Fourcade 347 (6) BOL, GRA; 3044 (2) BOL; 4694 (2) BOL; 5350 (6)

BOL; 5385 (6) BOL; 6213 (6) BOL. Fries 182 (1) PRE.

Galpin 228 (2) PRE; 1689 (3) PRE; 6280 (3) BOL, GRA, PRE. Giffen

782a (2) BOL, UFH. Gilfillan sub Herb Galpin s.n. (8) PRE. Gillett

1266 (6) BOL.

Hilliard & Burn 14718 (3) NU, PRE; 18667 (3) KEI, NU. PRE. Hops

33 (6) BOL: sub Bolus 12393 (6) PRE, BOL. Hutton s.n. (2) GRA

Jacot-Guillarmod 9431 (2) GRA. Jessop 1071 (2) GRA. Johnson 1032

(2) GRA.

Koopowitz s.n. (2) GRA.

Laughton 47 (6) BOL. Liebenberg 7248 (8) PRE. Linger 2099 (2) PRE.

Long s.n. (1) PEU; 436 (1) GRA, PEU, PRE.

MacOwan 1337 (2) GRA; 1836 (3) GRA, K. Martin 9252 (2) GRA.

McGaffin RUH1391 (6) GRA. Meyer 206 (8) PRE.

Paterson 10 (2) PRE; 42 (2) BOL; 91 (2) GRA. Pym s.n. (4) GRA.

Rodin 1025 ( 1 ) PRE. Rogers 3092 ( 1 ) GRA); 3093 (2) GRA; 23796 (6)

PRE; 27962 (6) GRA.

Schonland s.n. (1) GRA. Seagrief s.n. (6) GRA. Skinner 5 (2) GRA.

Story 2773 (2) PRE.

Tait 155 (PEU); 376i. 376ii (1 ) PEU.

Urton 962 (2) GRA.

Van Der Walt 256 (3) PRE. Vlok 1207 (6) K.

West 231 (6) GRA; 338 (2) GRA; 339 (1) GRA. White 351 (2) GRA.

Williamson s.n. (4) TCD.

Young sub Bolus 5545 (6) BOL. K.

ACKNOWLEDGEMENTS

We wish to thank the following people: Mr B. Burtt,

Mr D. Clarke, Ms M. Cocks, Mr W. Cowley, Dr G.

Hutchings, Mrs D. Page and Mr J. Sephton for assistance

in finding Lachenalia populations; Ms M. Johnson,

Royal Botanic Gardens, Kew, for cytology; Dr J. Parnell,

Trinity College, Dublin, for archival information; Prof.

C.E.J. Botha, Rhodes University Botany Department,

and Rhodes University Research Committee for funding;

Mr E. Kruger for photographs; the curators of BOL,

GRA, K, KEI, NBG, NU, PEU, PRE, TCD and UFH for

loans of their material.

REFERENCES

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. National Botanical Institute, Pretoria.

BAKER, J.G. 1871. Lachenalia convallarioides. Journal of the Lin-

nean Society 11: 407.

BAKER, J.G. 1874. Lachenalia campanulata. Journal of Botany 12: 6.

BAKER, J.G. 1897. Liliaceae. In W.T. Thiselton-Dyer, Flora capensis

6: 421-436. Reeve, London.

BARKER, W.F. 1978. Ten new species of Lachenalia. Journal of South

African Botany 44: 391-418.

BARKER, W.F. 1979. Ten more new species of Lachenalia. Journal of

South African Botany 45: 193-219.

BATTEN, A. & BOKELMANN, H. 1966. Wild flowers of the eastern

Cape Province. Books of Africa, Cape Town.

BOND, P & GOLDBLATT. P. 1984. Plants of the Cape flora. A

descriptive catalogue. Journal of South African Botany, Suppl.

Vol. 13: 51-54.

DUNCAN, G.D 1988. The Lachenalia handbook. Annals of Kirsten-

bosch Botanical Gardens 17. National Botanical Institute, Cape

Town.

DUNCAN, G.D. 1996. Four new species of Lachenalia (Hyacin-

thaceae) from arid areas of South Africa. Bothalia 26: 1-9.

DUNCAN, G.D. 1997. Five new species of Lachenalia (Hyacin-

thaceae) from arid areas of South Africa. Bothalia 27: 7-15.

FOURCADE, H.G. 1934. Lachenalia haarlemensis and L. subspicata.

Contributions to the flora of the Knysna and neighbouring divi-

sions. Transactions of the Royal Society of South Africa 21:

75-102.

FOURCADE, H.G. 1941. Check list of the flowering plants of the divi-

sions of George, Knysna, Humansdorp and Uniondale. Memoirs

of the Botanical Survey of South Africa No. 20.

GLEDHILL, E. 1981. Eastern Cape veld flowers. Creda Press, Cape

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GUNN. M. & CODD, L.E. 1981. Botanical exploration of southern

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HALL, H. 1856. Map of the eastern frontier from military and other

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JOHNSON, M.A.T. & BRANDHAM, P.E. 1997. New chromosome

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Bathurst. Rhodes University, Grahamstown.

MORIARTY. A. 1982. South African wild flower guide 2. Outeniqua,

Tsitsikamma and eastern Little Karoo. Botanical Society of

South Africa, Cape Town.

ORNDUFF. R. & WAITERS, P.J. 1978 Chromosome numbers in

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Bothalia 28,2: 151-157 (1998)

FSA contributions 12: Plantaginaceae

H.F. GLEN*

Annual or perennial herbs. Leaves simple, usually all

in a basal rosette or sometimes (in perennials) cauline,

spirally arranged, venation parallel; stipules 0. Inflores-

cence a dense to lax spike; peduncle usually longer than

inflorescence. Flowers bisexual, regular, small, brac-

teate. Calyx 4-lobed. membranous, imbricate. Corolla 4-

lobed. Stamens 4, inserted on corolla tube. Ovary > 2-4-

locular, superior; style 1, filiform, long, often exserted;

ovules 1-many in each locule. Fruit a circumscissile cap-

sule. Seeds mostly 2-4 per capsule (± 15-25 in P. major),

mostly ± boat-shaped with ventral hilum.

A family of three genera, only one of which is repre-

sented in southern Africa.

8116000 PLANTAGO

Plantago L., Species plantarum edn 1: 112 (1753);

T.Cooke: 388 (1910); Pilg.: 1 (1937); Levyns: 729 (1950);

Verde.: 1 (1971); R.A.Dyer: 601 (1975). Type species: P.

major L.

Description as for family.

A cosmopolitan genus of over 250 species, of which 12

and one variety occur in southern Africa. Five species are

restricted to the Flora area, another five are introduced

and two are weedy plants of such wide distribution as to

make it impossible to determine whether they are indige-

nous or naturalised here. Two species of doubtful occur-

rence in the Flora area are appended to this treatment.

Note: in the descriptions and notes below, the word

‘scape’ is used to mean the whole inflorescence including

both peduncle and spike.

la Leaves petiolate, or at least narrowed towards base:

2a Leaves broad (up to 4 times as long as wide):

3a Seeds 15-25 per capsule; leaves with scattered hairs,

usually drying greenish 1 . P. major

3b Seeds 2-4 per capsule; leaves glabrous or with isolated

hairs, usually drying brown to almost black

3. P. longissima

2b Leaves narrow (more than 4 times as long as wide):

4a Spike less than half as long as peduncle 10. P. lanceolata

4b Spike more than half as long as peduncle:

5a Leaves densely shaggy I P. virginica

5b Leaves with scattered hairs 9. P. myosuros

lb Leaves amplexicaul, not narrowing towards base:

6a Leaves distinctly cauline;

7a Bracts long, conspicuous, projecting well beyond the

flowers; leaves not succulent 12. P. aristata

7b Bracts short, inconspicuous, not projecting beyond the

flowers; leaves semi-succulent 6 . P. crassifolia

6b Leaves all in a basal rosette:

8a Leaves lorate, margins with a few small teeth:

9a Leaves glabrous 4. P. remota

9b Leaves villous 8. P. rhodosperma

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

MS. received: 1994-06-23

8b Leaves linear-pinnatifid or linear-filiform:

10a Leaves pinnatifid; spike over 20 mm long ... 5. P. coronopus

10b Leaves linear-filiform, not pinnatifid; spike less than

20 mm long:

1 1 a Plants hirsute 1 1 . P cafra

lib Plants glabrous 2. P. bigelovii

1. P. major L., Species plantarum edn 1, 112

(1753); Decne.: 694 (1852); T.Cooke: 388 (1910); Pilg.:

41 (1937); Levyns: 729 (1950); Verde.: 2 (1971). Type:

perhaps Sweden, LINN 144.1 (LINN).

P. dregeana Decne.: 695 (1852); T.Cooke: 389 (1910). Type: Cape,

near Nieuwjaarsfontein, Drege s.n. (P).

Perennial herbs, very variable. Leaves in a basal

rosette, elliptical, ovate or obovate, 70-400 x 24-125

mm, glabrous, rarely pubescent; petiole flattened,

25-200 mm long. Scapes several per rosette, 150-300

mm long; spikes 40-135 mm long. Bracts usually

glabrous, sometimes pubescent, apices acute. Calyx lobes

elliptical, ±2x1 mm, winged, acute. Corolla lobes del-

toid, ± 1.2 x 0.7 mm, acute. Capsule ellipsoid, ±3x2

mm (Figure ID). Seeds rounded-tetrahedral, 15-25 per

capsule, black to dark brown, 1.0-1. 2 x 0.6-0. 9 mm,

hilum scar at apex, small. Flowering time: from spring to

autumn.

A common weed of moist, sunny places; indigenous

or naturalised throughout most of the world. Known from

Northern Province to Western Cape and in Namibia

(Figure 2).

Vouchers: Archibald 4947 (PRE); Galpin 2945 (PRE); Mogg 4838

(PRE); Muir 1911 (PRE); Thode All 85 (K, NH, PRE).

This plant is easily confused with P. longissima (no.

3), but may be distinguished from that species by its

many-seeded capsules. Leaves of this species usually dry

olive-green and have scattered hairs, whereas those of P.

longissima usually dry various shades of brown to black,

and are glabrous or have only isolated hairs.

Plants of P. major are occasionally seen on sale in the

herb sections of nurseries in South Africa. The medicinal

uses of this species are similar to those of P. lanceolata',

Watt & Breyer-Brandwijk (1963) and Roberts (1983)

may be consulted for further details.

2. P. bigelovii A. Gray in Whipple, Exploration and

surveys for a railroad route from the Mississippi River to

the Pacific Ocean 4: 117 (1857); Pilg.: 73 (1937); Levyns:

730 (1950). Type: California, Benicia, Bigelow s.n. (GH).

Annual herbs. Leaves in a basal rosette, linear to fili-

form, 30-40 x 0.3-0. 6 mm, glabrous, amplexicaul.

Scapes few per rosette, 30-150 mm long; spikes 1. 5-5.0

mm long, few-flowered. Bracts pilose, obtuse. Calyx

lobes elliptic, ± 1.5 x 1.3 mm, winged, subobtuse.

152

Bothalia 28,2 (1998)

FIGURE 1. — A-C, Plantago lanceo-

lata L.: A, habit, x 0.5; B,

young fruit, x 5; C, dehisced

fruit and seeds, x 5. D,

Plantago major L., fruits and

seeds, x 5.

Corolla lobes deltoid to elliptic, ± 0.6 x 0.4 mm, acute.

Capsule ellipsoid, ± 3 mm long. Seeds not seen.

A western United States species, recorded once from

the Cape Peninsula by Levyns (Figure 3).

Voucher: Levyns 5092 (BOL, K).

This is the smallest species of Plantago occurring in

our area. While it would be difficult to confuse it with

any other southern African species of Plantago except P.

cafra (no. 1 1), it bears a striking superficial resemblance

to a species of Kyllinga (Cyperaceae). The most striking

difference between this species and P cafra is that in this

species all parts are glabrous, whereas in the latter the

peduncle in particular (but also the leaves) is covered

with brownish hairs.

3. P. longissima Decne. in DC., Prodromus 13:

720(1 852); T.Cooke: 390(1910); Pilg.: 81 (1937). Type:

Eastern Cape, between Umtata and Umsamwubo, Drege

4711 (P).

P. burchellii Decne.: 720 (1852). P. longissima Decne. var. burkei

Pilg.: 82 (1937). Type: KwaZulu-Natal, Mooi River, Burke s.n. (K).

P. longissima Decne. var. densiuscula Pilg.: 82 (1937). Type:

Mpumalanga, near Lydenburg, Wilms 1247 (B).

Perennial herbs. Leaves cauline, often with woolly

bases, elliptical, ovate or obovate, 190-725 x 45-150

mm, glabrous; petiole flattened, 50-400 mm long. Scape

one per rosette, 0.40-1.12 m long; spikes 0.15-0.50 m

long. Bracts glabrous, apices acute. Calyx lobes ellipti-

cal, ± 2.5 x 1 .0 mm, winged, acute. Corolla lobes del-

toid, ± 2.0 x 1 .2 mm, acute. Capsule ellipsoid, ± 3.0 x 1 .5

Bothalia 28,2 (1998)

153

FIGURE 2. — Distribution of Plantago major in southern Africa.

mm. Seeds rounded-tetrahedral, 2-4 per capsule, black to

dark brown, 1-2 x 0.6-0. 8 mm, hilum scar at apex,

small. Flowering time : from October to February.

FIGURE 3. — Distribution of Plantago bigelovii, •; and P virginica,

▲. in southern Africa.

Burser 10: 89 (UPS) (fide C. Jarvis & L.L. Dreyer pers.

comm.)

Endemic in Eastern Cape, KwaZulu-Natal, Mpuma-

langa, Gauteng and Northern Province (Figure 4).

Vouchers: Moss 8718 (BM); Phillips 570 (PRE); Rodin 3854 (K.

MO. PRE); Rudatis 1783 (PRE. STE).

This species is often confused with P. major (no. 1);

see there for distinguishing characters.

4. P. remota Lam., Tableau encyclopedique et

methodique 1; 341 (1791); Decne.: 721 (1852); T.Cooke:

391 (1910); Pilg.: 96 (1937); Levyns; 730 (1950). Type:

Cape of Good Hope, Sonnerat s.n. (P).

P. capensis Thunb.: 29 (1794); Thunb.: 148 (1823). Type: Cape of

Good Hope, Thunberg 3503 (UPS, holo.; PRE, fiche!).

Perennial herbs. Leaves in a basal rosette, linear to

lanceolate or oblanceolate, 50-210 x 2-9 mm. glabrous,

amplexicaul. Scapes few per rosette, 200-400 mm long;

spikes 30-125 mm long. Bracts glabrous, apices acute to

acuminate. Calyx lobes narrowly deltoid, ±3x1 mm,

narrowly winged, acute. Corolla lobes deltoid, ± 1.4 x 0.8

mm, acute. Capsule ellipsoid, ±2x1 mm. Seeds boat-

shaped, 2-4 per capsule, black to dark brown, 2. 2-3.0 x

1.0-1. 9 mm, hilum scar in a ventral depression almost as

long as seed. Flowering time', throughout the year.

Endemic to the Western and Eastern Cape (Figure 4).

Vouchers: Britten 6485 (PRE); Dyer 2400 (PRE); Esterhuysen 15454

(BOL, PRE); Glass 443 (NBG); Wolley Dod 2403 (BM, BOL, K, PRE).

This species is similar to P. crassifolia (no. 6) in that it

has semi-succulent, linear leaves, but differs from that

species in having all the leaves basal and glabrous except

for a tuft of copper-coloured woolly hairs at the base, and

in its generally more inland distribution.

5. P. coronopus L., Species plantarum 115 (1753);

Decne.; 732 (1852); Pilg.: 126 (1937); Levyns: 730

(1950); Chater & Cartier: 40 (1976). Lectotype: Herb.

Annual herbs. Leaves in a basal rosette, appressed-hor-

izontal, linear-pinnatifid, 40-200 x 2-5 mm, pubescent,

amplexicaul. Scapes many per rosette, 70-320 mm long;

peduncles horizontal below, erect above; spikes erect,

20-120 mm long. Bracts glabrescent, apices acute. Calyx

lobes narrowly elliptic, ±2x1 mm, acute. Corolla lobes

deltoid, ± 1.0 x 0.7 mm, subacute. Capsule ellipsoid, ±

2.5 x 2.0 mm. Seeds rounded-tetrahedral, 2-4 per cap-

sule, greenish brown, 1.0-1. 3 x 0. 6-0.7 mm, hilum scar

at apex, small. Flowering time : September to December.

A cosmopolitan weed, introduced from Europe; in our

area only known with certainty from the Western Cape,

where it is very common, if not locally dominant in small

areas, on paths leading to beaches, picnic areas and other

disturbed areas from Elands Bay on the west coast to

Pearly Beach near Bredasdorp, and possibly further

afield. Flanagan 1377 (PRE) from near Komga (Eastern

Cape) probably belongs here (Figure 5).

Vouchers: Glen 1518 (PRE); Moss 9082 (J); Oliver 3718 (PRE);

Salter 8516 (BOL, K, NBG); Walgate 617 (BOL, NBG).

FIGURE 4. — Distribution of Plantago longissima. ▲; and P. remota,

O, in southern Africa.

154

Bothalia 28,2 (1998)

FIGURE 5. — Distribution of Plantago coronopus, •; and P. myosuros,

A. in southern Africa.

This species is immediately distinguishable from all

others in southern Africa by its linear-pinnatifid leaves.

6. P. crassifolia Forssk ., Flora aegyptiaco-arabica

62 (1775); Pilg. : 160 (1937). Type: Egypt, Alexandria,

Forsskdl 261 (C).

P carnosa Lam.: 342 (1791); Decne.: 729 (1852); T.Cooke: 393

(1910); Levyns: 730 (1950). Type not cited.

P litoraria Fourc.: 96 (1934). Type: Western Cape, Eerste River,

Fourcade 1961 (BOL!).

Perennial herbs or suffrutices. Leaves cauline, linear to

narrowly lanceolate, 30-260 x 4-10 mm, pubescent,

amplexicaul. Scapes few per rosette, 90-300 mm long;

spikes 30-75 mm long. Bracts with scattered villi, apices

subacute. Calyx lobes elliptical, ± 2.5 x 1.5 mm, narrowly

winged, subacute. Corolla lobes deltoid, ± 1 .5 x 1.0 mm,

subacute. Capsule ellipsoid, ± 3.0 x 1.5 mm. Seeds boat-

shaped, 2-4 per capsule, brown, 1.8-2. 3 x 0.9-1. 2 mm,

hilum scar in a central depression about half as long as

seed. Flowering throughout the year, mostly in summer.

Two varieties can be distinguished:

la Leaves long and narrow, rarely less than 100 mm long . . .

6a. var. crassifolia

lb Leaves short and wide, rarely over 50 mm long ... 6b. var. hirsuta

6a. var. crassifolia.

Description as for species.

Occurs in the Mediterranean Basin and in southern

Africa, where it is known from the Western and Eastern

Cape (Figure 6).

Vouchers: Acocks 17610 (PRE); Boucher 3004 (PRE); Bourke 698

(NBG, PRE); Taylor 3178 (PRE, STE); Wells 2678 (GRA, K, PRE).

This species is usually a beach-dweller, in contrast

with P. remota (no. 4), which usually occurs in more

inland situations. In addition to the distinctions men-

tioned under that species, the present one may be disting-

uished from it by having entire or very sparsely, coarse-

ly dentate leaf margins. In P. remota the leaf margins are

sparsely minutely dentate.

6b. var. hirsuta (Thunb.) Beg. in Bulletino della

Societa botanica italiana 1901: 252 (1901); Pilg.: 162

(1937). Type: Cape of Good Hope, Thunberg 3518 (UPS,

holo.; PRE, fiche !).

P. hirsuta Thunb.: 29 (1794).

P thunbergii Poir.: 431 (1816). Type: Jacquin, Hort. Schoenbr. 3: t. 258

(PRE, icono.!).

Plants smaller in all parts than the typical variety.

Leaves more densely rosulate.

Occurs in the Western and Eastern Cape (Figure 6).

Vouchers: Bayliss 4182 (NBG); H. Hall s.n. in NBG97433 (NBG);

Schlechter 10551 (BOL, GRA).

7. P. virginica L., Species plantarum edn 1, 113

(1753); Michx.: 94 (1803); Decne.: 722 (1852); Pilg.:

213 (1937); Rahn: 154 (1974). Type: probably North

America, Kalm s.n. in LINN 144.8 (LINN).

Perennial herbs. Leaves in a basal rosette, obovate or

narrowly spathulate, 33-220 x 9-40 mm, densely vil-

lous; petiole flattened, 15-50 mm long. Scapes few per

rosette, 170-360 mm long; spikes 90-200 mm long.

Bracts glabrous, apices acute. Calyx lobes elliptical, ± 2

x 1 mm, subacute. Corolla lobes deltoid, ± 2.5 x 1.3 mm,

acute. Capsule ellipsoid, ± 3.5 x 2.0 mm. Seeds narrow-

ly tetrahedral, 2-4 per capsule, greenish brown, 1.4- 1.9

x 0.7-1 .0 mm, hilum scar at apex, small. Flowering time:

throughout the year, mainly in summer.

Originally introduced from North America (see Rahn

1974 for the indigenous distribution of this species), P.

virginica is now a widespread weed in southern Africa

(Figure 3).

Vouchers: Acocks 11450 (NH, PRE); Balsinhas 2905 (K, MO,

PRE); Code l 6455 (PRE); Jacob si. 2147 (NBG, PRE); Strey 3033 (K,

PRE, SRGH).

FIGURE 6. — Distribution of Plantago crassifolia var. crassifolia, A;

and P. crassifolia var. hirsuta, O, in southern Africa.

Bothalia 28,2 (1998)

155

FIGURE 7. — Distribution of Plantago rhodusperma , •; P. cafra, ▲;

and P. aristata, ★, in southern Africa.

The densely villous leaves distinguish this species

from all others in southern Africa. In shape, the leaves

resemble a miniature form of P. major (no. 1).

7.0-12.5 mm, glabrous, rarely pubescent; petiole flat-

tened, 25-100 mm long. Scapes few per rosette, 60-180

mm long; spikes 20-66 mm long. Bracts densely pilose,

apices acute. Calyx lobes narrowly elliptic, ± 2.5 x 1.0

mm, not winged, acute. Corolla lobes narrowly deltoid,

±2.5x1 .0 mm, acute. Capsule ellipsoid, ± 3.0 x 1.5 mm.

Seeds narrowly tetrahedral, 2-4 per capsule, pale yel-

lowish brown, 1.4-1. 6 x 0. 8-1.0 mm, hilum scar at apex,

small. Flowering time: January.

A weed introduced from South America (see Rahn

1974 for indigenous distribution); known in our area

from the Eastern Cape, and Mpumalanga (Figure 5).

Vouchers: Codd 9885 (PRE); Galpin 8230 (PRE); Hilliard & Burn

7557 (PRE); Jacot Guillarmod 8900 (GRA, PRE); Sim 19557 (PRE).

This species is superficially similar to P lanceolata

(below), from which it may be distinguished by its in-

florescences, in which the spikes are about as long as the

peduncles, whereas in P. lanceolata they are much short-

er than the peduncles. The hilum scar on the seeds is

raised in this species, but depressed in P. lanceolata and

P. rhodosperma (no. 8).

8. P. rhodosperma Decne. in DC., Prodromus 13:

722 (1852); Pilg.: 217 (1913); Pilg.: 12 (1928); Pilg.: 214

(1937); Rahn: 134 (1974). Type: Texas, Anon. s.n. in

herb. De Candolle (G).

Annual herbs. Leaves in a basal rosette, linear,

115-120 x 5-8 mm, densely villous, amplexicaul.

Scapes many per rosette, 50-300 mm long; spikes

10-180 mm long. Bracts villous, apices acute. Calyx

lobes narrowly elliptic, ±2x1 mm, subacute. Corolla

lobes broadly deltoid, ± 2.0 x 1.7 mm, acute. Capsule

ellipsoid, ±3x2 mm. Seeds boat-shaped, 2-4 per cap-

sule, greenish brown, 1.4— 1.9 x 0. 6-1.0 mm, hilum scar

in a ventral depression almost half as long as seed.

Flowering time : December.

This species was introduced from North America

(see Rahn 1974 for indigenous distribution); in our area

it is only known from the Eastern Cape (Figure 7).

Vouchers: Rattray 229 (GRA, K); C.A. Smith 3690 (PRE).

While the leaves and peduncles of P. myosuros

(below) have scattered hairs, those of the present species

are densely villous. P. rhodosperma differs from P.

lanceolata (no. 10) in that the spikes are as long as or

longer than the peduncles in the former species but much

shorter in the latter. It differs from P. remota (no. 4) in

having densely villous leaves and dense spikes, rather

than glabrous leaves and lax spikes. P. rhodosperma dif-

fers from P. virginica (above) in having narrower leaves

(both absolutely and in proportion to their length), vil-

lous (not glabrous) bracts and broader corolla lobes.

10. P. lanceolata L., Species plantarum edn 1:113

(1753); Decne.: 714 (1852); T.Cooke: 389 (1910); Pilg.:

313 (1937); Levyns: 730 (1950); Verde.: 6 (1971);

Troupin: 505 (1985). Type: specimen of Plantago angus-

tifolia major in Hort. Cliff, p. 36 no. 3 (BM, lecto.).

Perennial herbs, very variable. Leaves in a basal

rosette, linear to narrowly lanceolate or oblanceolate,

65-370 x 6-35 mm, glabrous or pubescent; petiole

30-125 mm long. Scapes many per rosette, 250-750 mm

long; spikes 15-70 mm long. Bracts glabrescent, apices

subobtuse. Calyx lobes elliptic, ± 2.5 x 1.3 mm, winged,

obtuse. Corolla lobes deltoid, ± 2.0 x 1.2 mm, acute.

Capsule ellipsoid, ± 3.5 x 1.5 mm. Seeds boat-shaped,

2-4 per capsule, dark brown, 24-2.8 x 1.2-1 .6 mm,

hilum scar in a ventral depression almost as long as seed.

Flowering throughout the year, but mainly in summer.

Figure 1A-C.

A very common weed of cultivation and disturbed

places; a typical habitat for this species is between

paving stones on urban pavements (sidewalks). This

species is indigenous or naturalised almost throughout

the world and is widely distributed in southern Africa

(Figure 8), even reaching Tristan da Cunha.

Vouchers: Boucher 1732 (PRE, STE); Dahlstrand 3570 (MO, PRE);

Marloth 10044 (PRE); Reid 307 (PRE); Van Jaarsveld 3127 (PRE).

Differences between this species on the one hand and

P. myosuros (above) and P. rhodosperma (no. 8) on the

other, are dealt with under those species. The medicinal

uses of this species are similar to those of P. major (no. 1).

9. P. myosuros Lam., Tableau encyclopedique et

methodique 1: 342 (1791); Decne.: 723 (1852); Pilg.:

244 (1913); Pilg.: 16 (1928); Pilg.: 226 (1937); Rahn:

115 (1974). Type: Uruguay, Commerson s.n. (P).

Annual or perennial herbs. Leaves in a basal rosette,

linear to narrowly lanceolate or oblanceolate, 90-120 x

11. P. cafra Decne. in DC., Prodromus 13: 719

(1852); T.Cooke: 389 (1910); Pilg.: 463 (1922); Pilg.:

352 (1937); Friedr.-Holzh.: 1 (1968). Type: Western

Cape, Riebeeck Kasteel, Drege s.n. (P).

R capillaris E.Mey. ex Decne.: 719 (1852); T.Cooke: 390 (1910). P.

cafra Decne. forma capillaris (E.Mey.) Pilg.: 464 (1922); Pilg.: 352

(1937). Type: Western Cape, Olifants River, Drege s.n. (K!).

156

Bothalia 28,2 (1998)

FIGURE 8. — Distribution of Plantago lanceolata in southern Africa.

Annual herbs. Leaves in a basal rosette, linear to fili-

form, acuminate, 32-144 x 0. 2-2.0 mm, pubescent,

amplexicaul. Scapes several per rosette, 12-130 mm

long; spikes 4-17 mm long. Bracts with scattered villi,

acuminate. Calyx lobes narrowly elliptic, ± 2.0 x 1.5

mm, winged, obtuse. Corolla lobes narrowly deltoid, ±

1.2 x 0.4 mm, acuminate. Capsule ellipsoid, ± 4.5 x 2.5

mm. Seeds boat-shaped, 2-4 per capsule, 2.3-4. 1 x

0.8-1.25 mm, brown or greenish brown, hilum scar in a

ventral depression almost as long as seed. Flowering

time : August-September.

Endemic to the Western and Northern Cape (Figure

7). According to Pilger (1937) and Friedrich-Holzham-

mer (1968), also known from one collection from

Warmbad District, Namibia.

Vouchers: Acocks 14836 (PRE); Goldblatt 2274 (NBG, PRE);

Riisch & Le Roux 467 (PRE); Schlechter 4908 (BM, GRA, PRE, SAM,

STE); Strey 3881 (PRE).

Differs from P. bigelovii (no. 2) in that the leaves and

particularly the peduncles of the scapes are hirsute-vil-

lous rather than glabrous, and from all other southern

African members of the genus by its small size and lin-

ear-filiform leaves.

12. P. aristata Michx., Flora boreali-americana 1:

95 (1803); Decne.: 714 (1852); Pilg.: 366 (1937). Type:

USA, Illinois, Michaux s.n. (P).

Perennial herbs. Leaves cauline, linear, acuminate,

100-142 x 2-7 mm, thinly hirsute to glabrous, amplexi-

caul. Scapes many per stem, 200-300 mm long; spikes

100-150 mm long. Bracts glabrous or with few villi,

very narrowly deltoid, conspicuous. Calyx lobes narrow-

ly elliptic, ±4x1 mm, not winged, obtuse. Corolla lobes

± 2.0 x 1 .5 mm, obtuse. Capsule ellipsoid, ±3.5x2 mm.

Seeds boat-shaped, 2-4 per capsule, 1.9-2. 6 x 1. 1-1.3

mm, brown, hilum scar in a ventral depression almost as

long as seed. Flowering time: January-February.

A weed, indigenous to the United States of America;

rather rare in our region, known only from the Free State

and KwaZulu-Natal border area, and from one specimen

from Pretoria, Gauteng (Figure 7).

Vouchers: Burtt Davy 7104 (K, PRE); Haygarth s.n. in NH15976

(NH); Medley Wood 8858 (NH, PRE); Ward 5968 (NU, PRE).

The bracts of this species are about 20 times as long

as wide, whereas in all other southern African species

they are about twice as long as wide.

DOUBTFUL SPECIES

P. laxiflora Decne. in DC., Prodromus 13: 699

(1852); Pilg.: 81 (1937). Type: Eastern Cape, ‘between

Gekan and Baxh’, Drege s.n. (P).

Annual (?) herbs. Leaves apparently in a basal rosette,

narrowly lanceolate, 50-150 x 20-45 mm, coarsely den-

tate, glabrescent; petioles flattened. Scapes few per

rosette, 200-650 mm long; spikes 120-300 mm long.

Bracts slightly villous, acute. Calyx lobes elliptic, ± 3

mm long, not winged, acute. Corolla lobes rotund or

ovate, ± 1.4 mm long, acute. Capsule ellipsoid. Seeds

boat-shaped, dark brown to black, to 3 mm long, hilum

scar in a long ventral depression.

Pilger states that this species occurs in the Eastern

Cape, but no specimen referable to this species was seen.

The description above is based on that of Pilger.

Material matching this description should not be iden-

tifiable using the key above, which should fail at couplet

8, where neither lead describes this species, though it

matches lead 6b.

P. afra L., Species plantarum edn 2: 168 (1762);

Verde.: 6 (1971 ). Type: Malta and North Africa, Morison

Hist. 3: 262, sect. 8, t. 17/4 (syn.).

P. psyllium sensu L.: 167 ( 1762); Pilg.: 422 (1937) et auett. mult., non

L.: 115 (1753).

Annual erect herbs. Leaves opposite, subopposite or

spirally arranged, narrowly lanceolate, 21-29 x 0.9-2. 0

mm, glabrous, obtuse. Scapes several at a node, 20-50

mm long; spikes 6-11 mm long, few-flowered. Bracts

glabrescent, acuminate. Calyx lobes elliptic, ± 3.0 x 1.5

mm, narrowly winged, subacute. Corolla lobes deltoid, ±

2.0 x 1.2 mm, acute. Capsule ellipsoid, ±4x2 mm.

Seeds boat-shaped, 2-4 per capsule, 2. 7-3. 2 x 1.0-1. 4

mm, greenish brown, hilum scar in a ventral depression

almost as long as seed.

Notes attached to a specimen from Kew indicate that

this species was once cultivated experimentally in

Pretoria. No material of it as a weed or naturalised in

southern Africa was seen, though there are two cultivat-

ed specimens in the main herbarium at PRE. The above

description was prepared from the Kew specimen.

A specimen, L.L. Britten 1613 (GRA), is similar to this

species but has much denser leaves and shorter pedun-

cles. These latter give the inflorescence the overall effect

of a compound capitulum rather than a compound umbel.

It is distinguished from all other southern African

species by its erect habit and usually opposite to subop-

posite leaves. It has been cultivated in order to extract an

oil (psyllium oil) from the seeds. For opinions on the

Bothalia 28,2 (1998)

157

nomenclature of this species see Pilger (1937) and

Verdcourt (1971).

REFERENCES

BEGUINOT, A. 1901. Intomo a Plantago crassifulia Forskal ed a

Plantago weldenii Rchb. nella flora italiana. Bulletino della So-

cietd botanica italiana 1901: 252-261.

CHATER, A.d & CARTIER. D. 1976. Plantaginaceae. Flora euro-

paea 4: 38^14.

COOKE. T. 1910. Plantaginaceae. Flora capensis 5,1: 387-392.

DECAISNE. J. 1852. Plantaginaceae. In A.P de Candolle, Prodromus

13: 693-737.

DYER, R.A. 1975. Genera of southern African flowering plants, Vol.

1. Government Printer, Pretoria.

FORSSKAL, P. 1775. Flora aegyptiaco-arabica. Moller, Kjobenhavn.

FOURCADE, H G. 1934. Contribution to the flora of Knysna and

neighbouring divisions. Transactions of the Royal Society of

South Africa 21 . 75-102.

FRIEDRICH-HOLZHAMMER, M. 1968. Plantaginaceae. In H. Merx-

mtiller, Prodromus einer Flora von Siidwestafrika 134: 1.

GRAY, A. 1857. Plantago bigelovii. In L.A. Whipple, Exploration and

surveys for a railroad route from the Mississippi River to the

Pacific Ocean. War Department, Washington DC.

LAMARCK, J.B.A.P.M. DE 1791. Tableau encyclopedique et me-

thodique. Pancoucke, Paris.

LEVYNS, M.R. 1950. Plantaginaceae. In R.S. Adamson & T.M. Salter,

Flora of the Cape Peninsula. Juta, Cape Town.

LINNAEUS, C. 1753. Species plantarum, edn 1. Salvius, Stockholm

LINNAEUS, C. 1762. Species plantarum. edn 2. Salvius, Stockholm.

MICHAUX, A. 1803. Flora boreali-americana. Crapelet, Paris.

PILGER, R. 1913. Uber Plantago sectio Plantaginella Decne.

Botanische Jahrbiicher 50: 61-71.

PILGER, R. 1922. Beitrage zur Kenntnis der Gattung Plantago. Fed-

des Repertorium 18: 449^175.

PILGER, R. 1928. Die Gattung Plantago in Zentral- und Siidamerika.

Botanische Jahrbiicher 62: 1-112.

PILGER, R. 1937. Plantaginaceae. In H.G.A. Engler, Das Pflanzen-

reich, Heft 102.

POIRET, J.L.M. 1816. Encyclopedic methodique Supplement. Agasse,

Paris.

RAHN, K. 1974. Plantago section Virginica : a taxonomic revision of a

group of American plantains. Dansk Botanisk Arkiv 30: 1-180.

ROBERTS, M. 1983. Margaret Roberts' book of herbs. Jonathan Ball,

Johannesburg.

THUNBERG, C.P 1794. Prodromus plantarum capensium. Edman,

Uppsala.

THUNBERG, C.P 1823. In J A. Schultes, Flora capensis. Cottae,

Stuttgart.

TROUPIN, G. 1985. Plantaginaceae. Flore du Rwanda 3: 504, 505.

VERDCOURT. B. 1971. Plantaginaceae. Flora of tropical East Africa.

Crown Agents for Oversea Governments and Administrations,

London.

WATT, J.M. & BREYER-BRANDWIJK, M.G. 1963. Medicinal and

poisonous plants of southern and eastern Africa, edn 2.

Livingstone. Edinburgh.

INDEX

Plantago L., 151

afra L., 1 56

anstata Michx., 156

bigelovii A. Gray, 1 5 1

burchellii Decne., 152

cafra Decne., 155

forma capillaris (E.Mey.) Piig., 155

capensis Thunb., 153

capillaris E.Mey. ex Decne., 155

carnosa Lam., 154

coronopus L., 153

crassifolia Forssk., 154

var. crassifolia, 154

var. hirsuta (Thunb.) Beg., 154

dregeana Decne., 151

hirsuta Thunb., 1 54

lanceolata L., 155

laxiflora Decne., 156

litoraria Fourc., 154

longissima Decne., 152

var. burkei Pilg., 152

var. densiuscula Pilg., 152

major L., 151

myosuros Lam., 155

psyllium sensu L., 156

remota Lam., 153

rhodosperma Decne., 155

thunbergii Poir., 154

virginica L., 154

Bothalia 28,2: 159-165 (1998)

Studies in the liverwort genus Fossombronia (Metzgeriales) from

southern Africa. 6. New species from Lesotho, Swaziland and

Mpumalanga and new records from Lesotho

S.M. PEROLD*

Keywords: Fossombronia , F. angulifolia, F. swaziensis, Hepaticae, Lesotho, Metzgeriales, new records, new species, southern Africa, Swaziland

ABSTRACT

Two new species of Fossombronia are described: F. angulifolia Perold from Lesotho and F. swaziensis Perold from

Swaziland and Mpumalanga. F. angulifolia is characterised by erect or semi-erect, 2— 4-lobed, angular leaves, by mostly

incompletely reticulate spores and by rather short elaters. F. swaziensis can be recognised by large, rounded or sometimes

shallowly notched leaves, which are rather stiff and bi- to multistratose basally, by quite large perigonial bracts in the male

plants and by spores which have irregular areolae containing tubercular inclusions.

1. Fossombronia angulifolia Perold , sp. nov.

Plantae repentes aggregatae. Folia subimbricata vel

subcontigua, superne plerumque bilobata, lobis angu-

laribus vadosis vel profundis. Rhizoidea purpurea.

Forsan dioicae; plantas masculas non vidi. Pseudo-

perianthium nonnihil brevius foliis contiguis, superne in

5 lobis irregularibus divisum. Sporae 27.5-45.0 pm

diametro, superficie distali lamellis 10 vel 11 irregu-

laribus ramosis et anastomosantibus, interdum areolas

formantibus; superficie proximali plerumque sine nota

triradiata, cum cristis pluribus pertenuibus irregularibus

granulatis. Elateres 52.5-90.0 pm longi, bis vel ter spi-

rales, 7.5-10.0 pm lati, interdum apices versus contracti.

TYPE. — Lesotho, 2927 (Maseru): 7.5 km west of

Roma along main tarred road and then south on dirt road

toward Korokoro, across river, on hillside above maize-

field, on soil between rocks, (-BC), S.M. Perold & M.

Koekemoer 3711 (PRE, holo.). Same locality, S.M.

Perold & M. Koekemoer 3712a (PRE, para.).

Plants smallish to medium-sized, creeping, in crowd-

ed, overlying stands, green, margins of some young

leaves tinged with red, upper part of older leaves fre-

quently dry and turning yellow, proximally becoming

colourless, occasionally several small, deep pink leaves

closely clasping basal part of stem; shoots mostly simple,

8.0-12.0 mm long, 1.2-1. 4 mm high, ± 1.9 mm wide,

rarely once furcate toward apex, sometimes with ventro-

lateral innovations. Stems prostrate, green, in cross sec-

tion apically (Figure IQ), ±310 pm (11 cell rows) high,

± 350 pm wide, basally (Figure 1R), ± 250 x 330 pm,

plano-convex. Rhizoids purple, 15.0-17.5 pm wide.

Leaves (Figure 1A-N) slightly overlapping to almost

contiguous, erect to semi-erect (Figure 2A, B), obliquely

inserted succubously, sinuous above and mostly 2-lobed,

rarely 3- or 4-lobed, with shallow or deep angular lobes,

dissimilar or similar in size and shape, lateral sides of

leaves ± parallel or not, distally smaller, gradually

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

MS. received: 1998-03-26.

enlarging proximally, (975-)1225-1450 x (500-)875-

1375 pm, margins with 6-8, generally one-celled slime

papillae at angulations as well as elsewhere, closer

together at ‘trailing’ edge. Leaf cells (Figure lO) thin-

walled, at upper margins subquadrate to rectangular

across, 20.0-25.0 x 27.5-57.5 pm, at lateral margins

short- to long-rectangular, 30.0-92.5 x 12.5-27.5 pm;

upper laminal cells (4-)5- or 6-sided or polygonal, 35.0-

47.5 x 27.5-50.0 pm; middle laminal cells 50.0- 57.5 x

30.0- 40.0 pm; basal cells 50.0-62.5 x 37.5-50.0 pm. Oil

bodies hyaline (Figure IP), slightly granular, 9-18 per

cell, round, oval or irregularly shaped, 3-6 pm diam.;

chloroplasts numerous, ± 5 pm diam.

?Dioicous; no male plants seen, only females.

Archegonia dorsally scattered along stem, naked (Figure

2C), few becoming fertilised. Pseudoperianths (Figure

2D-F) single, ± 1500 pm long (Figure 1U), ± 250 pm

shorter than adjacent leaves, divided above into 5 irregu-

lar lobes (Figure IV), up to 1685 pm wide across mouth,

with 15 or 16 spinous processes topped by a slime papil-

la; cells comparable in shape and size to those of leaves,

except for those in basal part, which are larger,

75.0- 125.0 x 27.5-35.0 pm. Capsule globose, ± 600 pm

diam., capsule wall bistratose, cells in inner layer irregu-

larly shaped, variable in size, 30.0-50.0 x 25.0- 32.5

pm, each cell wall with 1-3 nodular (Figure IS) and very

rare semi-annular thickenings. Seta delicate, ± 1.4 mm

long, in cross section (Figure IT) 200 x 150 pm, 8 cells

across. Spores light brown, hemispherical or occasional-

ly ± oval-shaped in outline, (27. 5-)32. 5-45.0 pm diam.,

including low spines projecting around periphery (Figure

3C); distal face convex, with 10 or 1 1 irregular lamellae

across it (Figure 3A, B), branching and anastomosing,

sometimes forming areolae, 5.0-12.5 x 5.0 pm, walls

slightly raised at corners, surface between lamellae fine-

ly granular (Figure 3D); proximal face (Figure 3E) most-

ly lacking distinct triradiate mark, with numerous very

fine, irregular ridges liberally sprinkled with granules,

around periphery many fine spines, shorter than 2.5 pm.

Elaters (Figure 3F) yellow-brown, (52. 5-)60. 0-70.0

(-90.0) pm long, often 3-spiral and uniformly up to 10

pm wide or tapering slightly toward tips, sometimes 2-

spiral, 7.5-10.0 pm wide, tips 5.0 pm wide.

160

Bothalia 28,2 (1998)

FIGURE I — Fossombmnia angulifolia. A-N, leaves; O, detail of leaf margin; P, median leaf cells with oil bodies (solid lines) and chloroplasts

(dotted lines); Q, c/s of stem apex; R, c/s of stem base; S, cells in capsule wall; T, c/s of seta; U, pseudoperianth from side; V, opened

pseudoperianth A-V, S.M. Perold & M Koekemoer 37 1 1 Scale bars: A-N, U, V, 500 pm; O, T, 100 pm; P, S, 50 pm; Q, R, 250 pm. Artist:

G. Condy.

Bothalia 28,2 (1998)

161

FIGURE 2. — Fossombronia angulifolia. A, arched terminal segment of thallus and leaves from side; B, shoot and leaves partly from side; C, archego-

nia seen from above; D, pseudopenanth (in culture stem continued growth beyond pseudoperianth); E, different shoot with pseudo-

perianth; F, close-up of pseudoperianth. A-F. S.M. Perold & M. Koekemoer 3711 . A, x 15.7; B, x 14.2; C, E, x 19.7; D, x 1 1; F, x 37.6.

Fossombronia angulifolia is known only from the type

locality in the Mountain Kingdom of Lesotho (Figure 4)

which receives rain in summer and occasional snowfalls in

winter. The two specimens referred to here were collected

in autumn after a season of good rains. Part of the holotype

specimen had to be kept in cultivation for almost two

months to allow the spores in several capsules to ripen. No

plants with antheridia were observed, despite thorough

searching. Mosses growing together with this new species

are Bartramia hampeana C.Miill., Bryum canariense

Brid., Pseudoleskeopsis cf. claviramea (C.Miill.) Ther.,

Gigaspermum repens (Hook.) Lindb. as well as the liver-

wort, Targionia hypophylla L. The soil is derived from

weathered sandstone and is finely to somewhat coarsely

granular. The vegetation type is Afromontane Grassland

(Low & Rebelo 1996). Some of the angular two-lobed

FIGURE 3. — Fossombronia angulifolia. Spores and elaters. A, distal face; B, side view of distal face; C, detail of areolae at margin of distal face;

D, surface between lamellae on distal face; E, proximal face; F, elaters. A-F, S.M. Perold & M. Koekemoer 3711. A, x 1094; B, x 1200;

C, x 2589; D, x 3533; E, x 1 120; F, x 753.

162

Bothalia 28,2 (1998)

FIGURE 4. — The distribution of F. angulifolia , •; and F swaziensis,

A; and some new records of F. gemmifera, 4; and F. glenii, □,

in Lesotho, Mpumalanga and Swaziland.

leaves in F. angulifolia are rather similar to those found in

F. spinifolia Steph. (Perold 1997b), but the spores in the

two species differ in their ornamentation, with those in F.

spinifolia having fewer and coarser ridges and some spin-

ous processes in between.

2. Fossombronia swaziensis Perold, sp. nov.

Plantae repentes aggregatae. Folia superne concava

vel inflexa, late patescentia, marginibus superioribus lat-

eralibusque rotundatis, basin versus bis vel multistratosa.

Rhizoidea purpurea. Dioicae. Antheridia dorsaliter bise-

riata in caule, bracteis perigonalibus concavis 1- vel 3-

lobatis obtecta. Pseudoperianthium plerumque supra

folia reflexa eminens; campanulatum; pedicellatum; ore

subpectinato, undulato. Sporae 47.5-55.0 pm diametro,

superficie distali lamellis 8 vel 9 rectis vel subcurvatis,

parallelis vel radiatis, irregulariter interconnexis areolas

imperfectas formantibus, unaquaeque tubercula 1 vel

2(3) continens; superficie proximali sine nota triradiata,

cum papillulis vel spiculis humilibus, congestis, perir-

regularibus, centro maioribus, circum peripheriam usque

ad 21 spinulis humilibus perisporio non connexis.

Elateres 80-150 pm longi, bis vel medio ter, apices ver-

sus bis, spirales, 10-15 pm lati, plerumque apices versus

contracti.

TYPE. — Swaziland, 2631 (Mbabane): Hhohho Dist.,

Forbes Reef, Umbuluzi River at Forbes Reef Store, on

clay soil of steep streambank in wattle forest, (-AA),

H.F. Glen 3130 (PRE, holo.). Same locality, H.F. Glen

3127 (PRE, para.).

Plants medium-sized to fairly large, creeping, in

crowded stands; shoots with proximal parts seemingly

from previous year, greyish brown, dead, or else growth

interrupted by dry conditions, at apex fresh, green, most-

ly simple, 8-10 mm long, ± 2 mm high, 4 mm wide (up

to 4.5 mm wide at pseudoperianth). Stems prostrate,

green, occasionally peripheral cell walls in some areas of

dorsal surface reddish, ventrally purple, in cross section

apically ± 400 pm (10 or 11 cell rows) high, ± 700 pm

wide in male plants (Figure 5N), becoming increasingly

fleshy, at ± midlength, 600 pm (16 cell rows) high, 900

pm wide, tapering toward base (Figure 50), 250 x 340

pm, plano-convex; female stems in cross section apically

± 350 x 550 pm (Figure 5P). Rhizoids purple, 17.5-27.5

pm wide. Leaves apically smaller, concave to inflexed

above, becoming widely spreading, inserted succubously,

upper and lateral margins rounded, occasionally shallow-

ly notched, often rather stiff, basally bi- to multistratose

(Figure 6C), sometimes up to ± midlength in a wide

median arch (Figure 5F), proximally increasing in size, in

male plants (Figure 5A-F) 1250-2000 x 1500-2125 pm,

somewhat larger in female plants (Figure 5G-K), shorter

than wide, 1500-2150 x 2625-3000 pm, margins at inter-

vals with 6-8 one-celled, papillae (Figure 6B), ± 22.5 x

12.5 pm, sessile or at lower ‘trailing’ edge raised on

( 1 — )2 — 4 basal cells. Leaf cells thin- walled, not apprecia-

bly different in male and female plants, at upper margins

(Figure 5L) subquadrate to rectangular across, 25.0-42.5

x 50.0-82.5 pm, at- lateral margins long-rectangular,

62.5- 97.5 x 25.0-30.0 pm; upper laminal cells 5- or 6-

sided, 40.0-57.5 x 27.5-55.0 pm; middle laminal cells

50.0- 87.5 x 30.0-62.5 pm; basal cells 60.0-97.5 x

55.0- 70.0 pm. Oil bodies disintegrated, remains execut-

ing Brownian movement on wetting; chloroplasts granu-

lar, up to 5 pm diam. (Figure 5M).

Dioicous. Antheridia dorsal on stem, often in 2 rows,

short-stalked, ± 250 pm wide, shielded by concave

perigonial bracts (Figures 5Q-S; 6D), single, 600-650 x

300-350 pm, or double, sometimes 3 lobes laterally

fused together, up to 480 x 850 pm, each one topped by

a papilla which is usually raised on a basal cell, margin-

al cells 50-75 x 45-50 pm, cells in interior 70.0-1 12.5 x

37.5- 42.5 pm. Archegonia naked, in a well-spaced dor-

sal row. Pseudoperianth (Figures 5U, V; 6E, F) ± 2 mm

proximal to apex, campanulate, raised on stalk up to

1000 pm long, mostly projecting above reflexed leaves,

1875-2250 pm long, 2750-2875 pm wide across mouth,

sometimes partly slit along side, margin slightly scal-

loped, undulating, very few or no papillae seen, toward

base with 2 irregularly thickened lateral outgrowths or

wings; cells not appreciably different in shape and size

from those in leaves. Capsules globose, ± 1000 pm

diam., wall bistratose, cells of inner layer irregularly

shaped, 30.0-37.5 x 25.0-30.0 pm, each cell wall with

1-3 dark brown nodular and some semi-annular thicken-

ings (Figure 5T). Seta ± 2.5 mm long, 200 pm diam., 9

cell rows across. Spores light brown to darker brown,

47.5- 55.0 pm diam., including lamellae projecting

around periphery, hemispherical; distal face (Figure

7A-C) convex, ornamentation irregularly lamellate and

incompletely reticulate, with 8 or 9 straight or slightly

curved parallel or radiating lamellae, their uneven crests

running across face, up to 3 pm high, 5. 0-7. 5 pm apart,

irregularly interconnected forming mostly incomplete

areolae of variable size and shape, each one containing 1

or 2 (3) tubercles (Figure 7D) with very faint ridges radi-

ating from their bases; proximal face (Figure 7E) slight-

ly concave, lacking triradiate mark, with low, crowded,

highly irregular small papillae or spicules, becoming

larger toward centre and linking up with one another,

around spore periphery up to 21 low ‘spines’, which are

the ‘ends’ of the lamellae from the distal face extending

over the sides and not connected by a perispore. Elaters

(Figure 7F) light brown, 80-150 pm long, 10-15 pm

Bothalia 28,2 (1998)

163

FIGURE 5. — Fossombronia swaziensis. A-F, male leaves; G-K, female leaves; L, detail of leaf margin; M, median leaf cells with chloroplasts

(dotted lines), oil bodies disintegrated; N, c/s of apical part of male stem; 0, c/s of basal part of male stem; P, c/s of apical part of female

stem; Q-S, bracts; U, pseudoperianth from side; V, opened pseudoperianth. A-F, L-O, Q-V, H.F. Glen 3130\ G-K, P, H.F. Glen 3127.

Scale bars; A-K, U, V, 500 pm; L, 100 pm; M, T, 50 pm; N-P, Q-S, 250 pm. Artist: G. Condy.

164

Bothalia 28,2 (1998)

FIGURE 6. — Fossombronia swaziensis. A, simple stem toward apex; B, apical leaves in close-up; C, leaf base showing cell layers; D, perigonial

bracts; E, pseudoperianth from side; F, pseudoperianth from above. A-C, E, F, H.F. Glen 3127\ D, H.F. Glen 3130. A, x 9; B, x 29; C, x

83; D, x 19; E, x 1 1 ; F, x 10.5.

wide in centre, mostly tapering to tips and ending in a

loop, 5 pm wide, sometimes not tapering, ends blunt, ±

12.5 pm wide, bispiral or centrally trispiral and then

bispiral toward ends.

Fossombronia swaziensis is known from the two type

collections growing on clayey soil on the banks of the

Umbuluzi River in a wattle forest at Forbes Reef Store

(Figure 4), Hhohho District, Swaziland, with vegetation

type North-Eastern Mountain Grassland (Low & Rebelo

1996), receiving >1000 mm rain annually. A third sporu-

lating specimen was recently collected on the banks of

Sterkspruit, De Kuilen, east of Lydenberg in Mpuma-

langa. The species can be recognised by its wide, round-

ed leaves, sometimes slightly indexed above, by quite

large perigonial bracts in male plants and by spores

which have one to several tubercular processes in the

complete or incomplete areolae. This ornamentation is

FIGURE 7. — Fossombronia swaziensis. Spores and elater A, B, distal face; C, side view of distal face; D, detail of areolae on distal face; E, prox-

imal face; F, elater. A, H.F. Glen 3127\ B-F, H.F. Glen 3130. A, x 715; B, x 795; C, x 780; D, x 1400; E, x 811; F, x 681.

Bothalia 28,2 (1998)

165

somewhat similar to that seen in F. straussiana (Perold

1997a), but there the rhizoids are hyaline, not purple as

in F. swaziensis. The latter is also a larger plant, with ±

entire, nonconvoluted leaves and discrete perigonial

bracts subtending the antheridia. F. swaziensis may be far

more common than the only two sporulating specimens

from Forbes Reef and the single one from De Kuilen

suggest, but all the other collections lack spores which

are needed to confirm the determinations. This new

species is also rather similar in general appearance to

robust forms of F. crispa which, however, has reticulate

spore ornamentation with the areolae lacking inclusions

(Perold 1997c). The multistratose leaf bases in F.

swaziensis may be due to the growing conditions, as

Scott & Pike (1988) surmised the case to be in F. crassi-

fola Spruce, where this also occurs.

SPECIMENS EXAMINED

Held at PRE. Bracketed numbers after citation of collectors’ names

and collecting numbers refer to the species described in the text in

alphabetical order, namely: F. angulifolia (1) and F. swaziensis (2).

Glen 3127 (2) (paratype), 3130 (2) (holotype).

Perold & Koekemoer 3711 (1) (holotype), 3712a (1) (paratype), 4064

(2).

NEW RECORDS OF FOSSOMBRON1A SPECIES IN LESOTHO

(HELD AT PRE)

F gemmifera Perold (Perold 1997a: 19). LESOTHO, 2927: (-BC),

Perold & Koekemoer 3733.

F. glenii Perold (Perold 1997a: 20). LESOTHO, 2828: (-CC), Perold &

Koekemoer 3737, 3739. 2927: (-BA), Perold & Koekemoer 3736.

2927: (- BC), Perold & Koekemoer 3713, 3719.

ACKNOWLEDGEMENTS

I sincerely thank Dr E.O. Campbell for refereeing this

article and for her helpful suggestions; Ms M.

Koekemoer, curator of the National Herbarium, Pretoria,

for her help with fieldwork in Lesotho and Mpumalanga

as well as Dr H.F. Glen for collecting the specimens of F.

swaziensis, for suggesting the specific epithet and for

translating the diagnoses into Latin. I also extend my

gratitude to Ms G. Condy for the drawings, Mrs A.

Romanowski for developing and printing the pho-

tographs and to Ms D. Maree for typing the manuscript.

REFERENCES

LOW, A.B. & REBELO, A. 1996. Vegetation of South Africa, Lesotho

and Swaziland. Dept. Environmental Affairs & Tourism,

Pretoria.

PEROLD, S.M. 1997a. Studies in the liverwort genus Fossombronia

(Metzgeriales) from southern Africa. 1 Three new species from

Northern Province, Gauteng and Mpumalanga. Bothalia 27:

17-27.

PEROLD, S.M. 1997b. Studies in the liverwort genus Fossombronia

(Metzgeriales) from southern Africa. 3. An amendment to F.

spinifolia. Bothalia 27: 39-42.

PEROLD, S.M. 1997c. Studies in the liverwort genus Fossombronia

(Metzgeriales) from southern Africa. 4. A re-examination of F.

crispa, F. leucoxantha and F. tumida. Bothalia 27: 105-115.

Scott, G A M. & PIKE, D C. 1988. Revisionary notes on Fos-

sombronia. The Bryologist 9: 193-201.

Bothalia 28,2: 167-177 (1998)

The genus Echium (Boraginaceae) in southern Africa

E. RETIEF * and A.E. VAN WYK**

Keywords: Boraginaceae, Echium L., indumentum, Lobostemon Lehm., phytogeography, pollen, relationships, southern Africa, taxonomy, weed

ABSTRACT

The genus Echium L. (Boraginaceae) comprises about 60 species, mainly from Macaronesia Europe, western Asia and

North Africa. Two species E. plantagineum L. and E. vulgare L. were introduced into southern Africa and have become

naturalised. The species occur mainly as roadside weeds in the region. Echium is closely related to Lobostemon Lehm. (inch

Echiostachys Levyns), endemic in the southwestern Cape region. Pollen morphology shows a remarkable similarity

between these genera, even suggesting that they could be merged. However, other characters, such as bilobed styles

(Echium) versus undivided ones (Lobostemon) and the presence of an annulus, composed of a minute collar or 5-10 minute

hairy lobules, at the bottom of the corolla tube inside (Echium), in contrast to hairs and/or scales at the base of the filaments

(Lobostemon) contradict the pollen structure, and Echium and Lobostemon are therefore regarded as two separate genera.

Significant taxonomic characters, an identification key, full descriptions, illustrations and distribution maps of E. plan-

tagineum and E. vulgare are given.

CONTENTS

Abstract 167

Introduction 167

Materials and methods 167

Historical outline 167

Morphological characters of taxonomic significance . 168

Trichomes 168

Corolla 168

Pollen 170

Style 170

Fruit 171

Phytogeography 171

Key to species 173

Echium L 173

1 . Echium plantagineum L 173

2. Echium vulgare L 174

Specimens examined 174

Acknowledgements 176

References 176

INTRODUCTION

The genus Echium L. (1753) comprises about 60

species, distributed in Macaronesia, Europe, western

Asia and North Africa (Mabberley 1997). It belongs to

the Boraginaceae, a family characterised by an indumen-

tum mainly of trichomes with swollen, multicellular

bases, inflorescences of scorpioid or helicoid cymes

which uncoil and elongate as the flowers mature, a ter-

minal or gynobasic style, and fruit consisting mostly of

four nutlets. Although cosmopolitan, the members of

Boraginaceae are particularly well represented in tem-

perate and subtropical regions, with centres of diversity

in the Mediterranean region and western North America

(Zomlefer 1994).

Two species of Echium, E. plantagineum L. and E.

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

** H.G.WJ. Schweickerdt Herbarium, Department of Botany,

University of Pretoria, 0002 Pretoria.

MS. received: 1998-03-23

vulgare L., were introduced into southern Africa and are

now naturalised in the Western and Eastern Cape, eastern

Free State, Lesotho, high mountainous areas of KwaZulu-

Natal and in Mpumalanga. Both species occur mainly as

roadside weeds in the region. This study forms part of an

ongoing revision of the Boraginaceae of southern Africa

which is currently in progress.

The purpose of this paper is to present a revision of

two species of Echium in South Africa and Lesotho.

Diagnostic characters, an identification key, full descrip-

tions, illustrations and distribution maps are provided.

The generic description of Echium and also the species

descriptions of E. plantagineum and E. vulgare are based

on behaviour in southern Africa only.

MATERIALS AND METHODS

Herbarium specimens in BM, BOL, E, GRA, K,

NBG, NH, NU, PRE, PRU and SAM were studied to

gather data on morphological characters, flowering time

and distribution. Both untreated and acetolysed pollen

grains were studied. Pollen was acetolysed according to

the standard method of Erdtman (1960). For SEM, sam-

ples were coated with gold and studied with an ISI-SX-

25 microscope. For LM, pollen grains were mounted in

glycerine jelly.

HISTORICAL OUTLINE

According to Smith (1966), E. plantagineum was

introduced to South Africa from Europe as part of stock

feed. A note on Britten 520 , a herbarium specimen in

PRE dated 1917, E. plantagineum might have become

established in the region after mixed birdseed had been

sown. Wright (1904) records a specimen of Villet (with-

out locality) which may have been collected about

1825-1827. Exactly how and when E. vulgare entered

the country is not known. The oldest specimens exam-

ined date back from 1912 to 1915 when Mary Page, a

botanical artist, collected this species in Lesotho.

168

Bothalia 28,2 (1998)

Lehmann (1818), like Linnaeus (1762, 1771), Thun-

berg (1794), Jacquin (1797) and others, published new

species under Echium from the Cape region. In 1830,

however, the genus Lobostemon (Boraginaceae) was

described by Lehmann, based on a plant grown in the

Botanic Gardens at Hamburg, specimens of it then

named by him L. echioides. Lehmann emphasised the

presence of a scale-like outgrowth at the base of the sta-

mens as diagnostic in Lobostemon (Levyns 1934). Buek

(1837) transferred 32 South African species to Lobo-

stemon, most of which had formerly been included in

Echium. He extended Lehmann’s genus to include cer-

tain species in which staminal scales were absent, but in

which hairs were present on the lower parts of the fila-

ments. De Candolle (1846) also recognised the genus

Lobostemon. He described some new species, but refer-

red all those without definite scales to the genus Echium.

In Flora capensis , Wright (1904) reverted to Buek’s ear-

lier concept of Lobostemon in that he once more returned

those taxa without scales to the genus. However, he

retained one species, Echium formosum Pers. in the

genus Echium. Today E. formosum is treated as conspe-

cific with Lobostemon regulareflorus (Ker Gawl.)

M.H.Buys (Buys & Van der Walt 1994).

Johnston (1924) was of the opinion that Lobostemon

should be a synonym of Echium. He published new com-

binations in Echium for 21 species of Lobostemon ,

described by Buek (1837), De Candolle (1846) and

Wright (1904). Some years later, Johnston (1953, 1954)

once again separated Lobostemon from Echium on the

position of the annulus and the fact that in Echium the

style is always 2-lobed, whereas in Lobostemon it is sim-

ple. His account on Echium and Lobostemon, published

in 1954, appeared after Levyns’s (1934) major revision of

Lobostemon. In the latter publication Levyns described

the new genus, Echiostachys, based on three species, all

formerly placed under Echium. Johnston (1954), howev-

er, rejected the erection of the genus Echiostachys.

According to him, this proposed segregate genus differs

from the remaining Lobostemon species only in their

herbaceous habit and the very dense cylindrical thyrse,

differences which are no more striking or important than

those existing between groups of species in Echium.

Both Phillips (1951) and Dyer (1975) distinguished

Echium and Echiostachys from Lobostemon, the first two

genera with the filaments sometimes hairy at the base,

but never with scales or protuberances at the base.

Lobostemon is retained for those taxa that have filaments

with a densely hairy scale, ridge or protuberance at the

base. Echium, on the other hand, is separated from

Echiostachys because of its irregular, bilabiate corolla.

Dyer (1975) was definitely not aware of Johnston’s

(1953) account on Echium and Lobostemon, where he

separated the genera based on the position of an annulus

inside the corolla tube.

The present authors follow Johnston (1953) and others

in regarding Echium and Lobostemon as two separate

genera. Although very similar in pollen morphology,

other characters like the structure of the style and a dis-

junct distribution pattern reveal two distinct genera. For

the purpose of the present paper, Echiostachys is regard-

ed as synonymous with Lobostemon, in agreement with

Johnston’s view above.

MORPHOLOGICAL CHARACTERS OF TAXONOMIC

SIGNIFICANCE

Trichomes

The indumentum of the stem, leaf and calyx in

Echium can be described as hispid or strigose. Three

morphologically different types of trichomes have hither-

to been reported in the genus: (1) tubercle-based tri-

chomes (setae), often with a concentric ring of epidermal

cells surrounding the base, (2) unicellular trichomes, and

(3) glandular, multicellular trichomes. These trichome

types are simple (unbranched). Glandular trichomes, how-

ever, do not occur on the various plant parts of E. plan-

tagineum and E. vulgare.

The trichome complement of E. plantagineum con-

sists of tubercle-based setae, each with a uniseriate, con-

centric ring of epidermal cells surrounding the base.

These setae vary from long and slender to short, and are

often stout and sparsely to moderately densely arranged

(Figure 1A-C). Furthermore, a sublayer of much shorter,

soft, unicellular trichomes is present on the stems, mar-

gins and veins of the leaves, as well as the calyx lobes

(Figure IB). E. vulgare is characterised by tubercle-

based seta with a ring of cells at the base, similar to that

of E. plantagineum, interspersed with tubercle-based

setae, which are short and stout, without a ring of cells at

the base (Figure ID, E). Although the base of shorter

setae are usually swollen, there is generally little or no

modification of the surrounding epidermal cells (Figure

IE). The setae with a ring of cells occur very sparsely,

whereas the shorter ones without prominent rings form a

moderately dense sublayer (Figure ID, E). The combina-

tion of trichome types is distinctly different in the two

southern African species, and a most useful character for

identifying specimens.

If the collective trichome cover of E. plantagineum

and E. vulgare is compared with those of Lobostemon

species, to which they are closely related, similar but not

exactly the same states are found. The nature, composi-

tion and relative abundance of trichomes are also of tax-

onomic importance in other genera of the Boraginaceae.

An overview of trichome characters in all southern

African taxa of the family is in preparation.

Corolla

Flowers of Echium are borne in thyrsoid inflores-

cences composed of many lateral helicoid cymes, the lat-

ter enlarging in fruit. The shape and colour of the corol-

la, together with the degree of stamen exsertion, are flo-

ral characters of prime taxonomic importance in the

European species of Echium (Gibbs 1971). E. plan-

tagineum and E. vulgare belong to two different groups.

The corolla of E. plantagineum is broadly funnel-shaped

with two stamens long-exserted (Figure 2). E. vulgare

has a broadly funnel-shaped corolla with four stamens

long-exserted. The stamen number can be used to distin-

guish between flowering material of the species con-

cerned. In a key to Echium species, Gibbs (1972) distin-

guished two main groups, namely species with the flow-

ers flesh-coloured or yellowish or bluish white, and those

Bothalia 28,2 (1998)

169

FIGURE 1. — Echium leaf surface. E. plantagineum : A, upper surface; B, lower surface; C, trichomes. E. vulgare: D, upper surface; E, lower sur-

face; F, trichomes. A-C, Hilliard & Burtt 6999\ D-F, Sidey 3977. A, x 8; B, x 7; C, x 21.5; D, x 22; E, x 15; F, x 57.

with flowers blue, reddish purple or pink-carmine turn-

ing blue-purple. E. plantagineum and E. vulgare belong

to the second group. The corolla in E. vulgare is pilose

(Figure 3A), whereas that of E. plantagineum is

glabrous, except for sparse, simple, long, unicellular tri-

chomes on veins and margins. The corolla indumentum

of E. vulgare is also characterised by simple, long, uni-

cellular trichomes on the veins and margin, but with sim-

ilar, shorter trichomes in the intercostal areas of the

corolla (Figure 3A, B).

Corollas in the family Boraginaceae are actinomor-

phic, rarely zygomorphic. In Echium and to a lesser

extent Lobostemon, however, irregular corollas occur.

Johnston (1924) was of the opinion that the development

of zygomorphy furnishes no substantial basis for the

recognition of the tribe Echieae in which Echium and

Lobostemon are placed. He pointed out that between the

conspicuously irregular corollas of certain species of

Echium and the regular corollas of typical species of

Lithospermum there is every intermediate stage of irre-

gularity. According to Johnston (1924), Echium italicum

has regular and subregular corollas, and in a number of

the echiums of the section Lobostemon, the corolla is

also regular or practically so. The corolla of both E. plan-

tagineum and E. vulgare is zygomorphic. On the other

hand, evidence from palynology supports the retention of

the tribe Echieae (see pollen).

Johnston (1924) regarded Lobostemon as a synonym

of Echium , but later changed his opinion. In a key to the

genera of the Lithospermeae, Johnston (1953) separated

the two genera on the basis of the position of what is

called an ‘annulus’. In the case of Echium, the annulus,

borne 1 mm or less above the base of the corolla tube, is

composed of a minute collar or ring of 5-10 minute hairy

lobules (Figure 3C). Lobostemon is characterised by the

presence of an annulus above the corolla base. It is com-

posed of five distinct, densely villous swellings or five

squamose appendages borne below the attachment of

each stamen. The two naturalised Echium species can

therefore not be confused with any member of Lobo-

stemon in southern Africa.

In some species of Echium and also Lobostemon , the

margin of the theca of the opened anthers is ± ciliate with

FIGURE 2. — Inflorescence of Echium plantagineum, Stirton 8218

(PRE). Scale bar: 10 mm.

170

Bothalia 28,2 (1998)

FIGURE 3. — A-C, corolla. E. vulgare: A, outer surface; B, simple, unicellular trichome; C, hairy lobule. D-F, pollen. D, E. vulgare, pollen grain.

E, F, E. plantagineum ; E, granular colpus; F, tectum with supratectal gemmae. G-I, style of E. vulgare: G, stigmas; H, papillae caps; I, tri-

chomes on style. A-C, I, Sidey 3977 ; D, G, H, Hilliard & Burtt 18686 ; E, Bayer 6005 ; F, Balsinhas 3546. A, x 35; B, x 155; C, x 68; D,

x 2199; E, x 4058; F, x 7546; G, x 91; H, x 1067; I, x 84.

very slender elongate incurving hairs (Johnston 1953).

According to Johnston (1953) similar slender hairs may

also arise from along the midline of the open, empty

theca. These characters are not uncommon, although

they are usually best developed in specimens of E. plan-

tagineum and its allies.

Pollen

Pollen grains heteropolar, with one pole ‘/3 to ‘A wider

than the other, broad end obtuse, narrow end more acute

in equatorial view, the long sides concave, tricolporate,

pear-shaped, P = (2 1 — )27— 29(— 37) pm, E (at widest point)

= (14-)22-25(-29) pm. Shape in polar view with broad

end rounded-triangular, narrow end more deeply 3-lobed;

in equatorial view ovate to ovate-triangular, with protrud-

ing ora (Figure 3D). Colpi narrow, elongated, extending

over about 4A the length of polar axis with more or less

rounded to pointed ends, somewhat sunken, always free

at ends, surface with gemmae (Figure 3E). Endo-aper-

tures (ora) situated closer to the broader pole, more or less

circular, aperture membrane covered with gemmae.

Tectum microreticulate with densely spaced lumina of

different shape and size. Supratectal gemmae confined to

margins of lumina (Figure 3F).

Pollen morphology is remarkably similar in the var-

ious species of Echium (Reille 1995). Pollen of Echium

in southern Africa is not sufficiently different from that

of Lobostemon. In equatorial view, grains are triangular

to somewhat pear-shaped in outline (Figure 3D), similar

to grains of Lobostemon. Two shape classes, namely sub-

spheroidal and prolate, were distinguished in the tribe

Echieae (Retief & Van Wyk 1997). Grains of E. plan-

tagineum as- well as E. vulgare are subprolate.

Palynological evidence indicates a very close rela-

tionship between Echium and Lobostemon , even suggest-

ing that they might be congeneric. Both genera have tri-

colporate, anisopolar grains. We follow Johnston (1953)

and treat Echium and Lobostemon as two separate,

although closely related, genera. A comparison of the

pollen of Echium , Lobostemon and Lithospermum , (type

genus of Lithospermae) strongly supports the retention

of the tribe Echieae (Retief & Van Wyk 1997).

Style

The gynobasic style in Echium is almost always

bilobed in the upper part with two capitate stigmas

(Figure 3G). These stigmas are covered with papillae,

Bothalia 28,2 (1998)

171

which end in prominent contiguous caps with crenulate

margins (Figure 3H). The surface of the papillae caps is

covered with cuticular irregularities (Figure 3H).

According to Heslop-Harrison (1981), pollen grains of

compatible size are wedged between the papillae by

bees, whilst grains resting on the surface of the cap do

not hydrate or germinate; the papillae caps thus function

as a series of juxtaposed parasols. The styles of all

Echium species are pubescent (Al-Shebaz 1991). Only

simple, unicellular trichomes are present (Figure 31).

The two Echium species in southern Africa differ

from all Lobostemon species in having styles prominent-

ly bilobed, unlike those of species of Lobostemon which

are undivided. Those species of Echium with cleft styles

show similarity to members of Ehretia and Cordia also

with divided styles. Echium, however, differs from

Cordia in which twice bifid styles occur with linear or

clavate stigmatic branches — not a bilobed style with cap-

itate stigmas. The style of Echium is similar to that of

Ehretia , both with bilobed styles and capitate stigmas.

The family Boraginaceae is sometimes split into two sep-

arate families, namely Boraginaceae s.str. and Ehreti-

aceae. Ehretiaceae includes both Ehretia and Cordia.

Echium, a member of the tribe Boragineae (Boragi-

nioideae) has style and stigma characteristics similar to

those of members of the Ehretiaceae and this supports

the view that the family Boraginaceae is a natural entity

and should not be divided into two or even more segre-

gate families. Stigmas in other genera of the Boragineae

are conic, discoid, subglobose, obtuse, entire or 2-lobed,

thus differing in morphology from those of Echium.

Fruit

The fruit in Echium consists of four nutlets which are

ovoid, ovoid-trigonal, oblong-bipyramidal or subglobose

(Edmondson 1978). A broad attachment scar and an erect

or incurved beak are also present. The outer surface of the

nutlet can be smooth or variously rugose, tuberculate,

scrobiculate or reticulate. E. plantagineum and E. vulgare

differ very little in the structure of their nutlets. Both have

ovoid-trigonal nutlets with rugose, tuberculate outer sur-

faces with the tubercles varying in size (Figure 4A-F).

Many taxonomic decisions in the Boraginaceae are

based on fruit characters. Fruit in Wellstedia (capsule),

Ehretia (drupaceous) and Cordia (drupe) differ markedly

from those of other genera in the family characterised by

nutlets. The outer surface of the nutlets varies considerably

in sculpture and vesture. The nutlets of E. plantagineum

and E. vulgare show similarities to those of Lobostemon ,

Anchusa and Heliotropium. Species of Myosotis have

smooth and shiny nutlets, whereas nutlets of species of

Cynoglossum are characterised by simple, barbed tri-

chomes. An overview of the different fruit types present in

the southern African Boraginaceae is in preparation.

PHYTOGEOGRAPHY

Echium is a genus concentrated in the Mediterranean

region. Its greatest morphological diversity is centred in

Macaronesia. According to Bramwell (1972), 22 species

are indigenous to the Canary Islands, three to the Cape

Verde Islands and two to Madeira. About 26 species are

native to northwestern Africa, the highest concentration

in Morocco, and about 15 in the Iberian Peninsula (Gibbs

1971). Several herbaceous species, like E. plantagineum

and E. vulgare, have been introduced by man to various

other parts of the world such as southern Africa and

Australia where they are now naturalised and widespread.

As pointed out above, Echium is very closely related

to the South African genus Lobostemon. Lobostemon,

FIGURE 4. — Echium nutlet A, E. plantagineum , dorsal face. B-F, E. vulgare : B, ventral face with prominent attachment scar; C, dorsal face;

D-F, outer surface with large and numerous small tubercles. A, Germishuizen 4102\ B-F, Zietsman 1480. A, x 13.5; B, C, x 16; D, x 23;

E, x 44.5; F, x 148.

172

Bothalia 28,2 (1998)

FIGURE 5. — Echium plantagineum A, habit, x 0.8; B, flower, x 2; C, longitudinal section of corolla, x 1.5; D, style, x 22; E, nutlet, x 14, F, lower

surface of leaf, x 3. A-D, Acocks 9250\ E, Germishuizen 4102, F, Hilliard & Burn 6999.

Bothalia 28,2 (1998)

173

confined to the Cape Floristic Kingdom, and several

other taxa in the southern African flora, have a southern

Africa-North Africa, Mediterranean and Eurasian dis-

junct distribution pattern. For example, there are four

endemic genera of the Fumariaceae in southern Africa,

geographically widely separated from other members of

the family in the northern temperate parts of the Old

World (Hutchinson 1921). Another example is Campto-

loma (Scrophulariaceae) with three vacariant species that

form part of the disjunct Afro-arid element in African

phytogeography (Hilliard 1994). The genus has one

species in Namibia-Angola, another in the Canary

Islands and the third in a small area around the Horn of

Africa. The interpretation of disjunct distributions is

often complex and impossible to explain (Stott 1981). It

can, for example, be speculated that Echium and Lobo-

stemon are derived from a once widespread, common

ancestor which was a component of a temperate flora on

the African continent. Climatic changes took place and

Echium and Lobostemon remained as relicts, evolving

separately in a Mediterranean type climate, but now

absent in the vast dry parts of Africa. However, there is

no satisfactory paleobotanical evidence to verify the

above assumption. Long-distance dispersal may also

have played a role, but this type of explanation is usual-

ly considered highly unlikely for clarifying wide dis-

junctions on continental areas (Poynton 1983).

Echium L., Species plantarum: 139 (1753); L.:

175 (1754); Juss.: 130 (1789); DC.: 13 (1846); Benth.:

863 (1876); Giirke: 128 (1897); C.H.Wright: 43 (1904);

E. Phillips: 632 (1951); Riedl: 213 (1967); P.E.Gibbs: 97

(1972); R. A. Dyer: 513 (1975); J.R.Edm.: 318 (1978);

Toelken: 1 155 (1986); Al-Shebaz: 1 37 (1991); Verde.: 81

(1991). Type: E. italicum L. [lecto., see Al-Shebaz: 138

(1991)].

Isoplesion Rafin.: 86 (1836-38).

Megacaryon Boiss.: 203 (1879).

Annual, biennial or perennial herbs or shrubs; scabrid,

hispid or canescent. Leaves alternate, with basal leaves

rosulate, petiolate or sessile, densely hairy. Flowers in

helicoid cymes, grouped in many-branched thyrses or

panicles, elongating considerably in fruit, bracteate;

bracts small and foliaceous. Calyx deeply 5-lobed, some-

what accrescent. Corolla blue or purplish blue; zygo-

morphic; tube narrowly to broadly funnel-shaped,

straight, outside pubescent or glabrous, inside usually

with a ring of 5-10 distinct scales or lobes or an undu-

late, entire, somewhat fleshy collar-like membrane at

base; throat open, without any appendages; limb marked-

ly oblique; lobes equal or unequal, short, obtuse.

Stamens 5, variously exserted, inserted on corolla tube at

various levels; filaments linear, glabrous or with long

hairs; anthers versatile. Ovary 4-lobed, with a single

ovule in each locule; style gynobasic, linear, bifid at

apex, pubescent, long-exserted; stigmas 2, capitate. Fruit

of 4 nutlets; nutlets convex on one face, keeled on the

other, ± triangular at base, rugose, tuberculate.

A genus of about 60 species, mainly confined to

Madeira, the Azores and Canary Islands but also found in

Europe, western Asia and North Africa. Some herba-

ceous species have been introduced in various parts of

the world where they are now naturalised. Two species,

E. plantagineum and E. vulgare, are naturalised in south-

ern Africa; recorded from the Western and Eastern Cape,

Free State, Lesotho and Mpumalanga.

Key to species

Corolla glabrous, except for sparse simple unicellular trichomes

on veins and margin; two stamens long-exserted, three

enclosed; basal leaves petiolate, broadly ovate, with

prominent lateral veins; indumentum of sparse, simple,

long, slender tubercle-based setae with a sublayer of

much shorter, fine, simple, unicellular trichomes on leaf

margins and especially on midrib of lower leaf surface

1 E. plantagineum

Corolla softly hairy, with simple, unicellular trichomes promi-

nently longer on veins; four stamens long-exserted, one

enclosed; basal leaves subsessile, linear-lanceolate or

narrowly oblong with lateral veins obscure; indumen-

tum of sparse, simple, long tubercle-based setae with a

sublayer of much shorter tubercle-based setae, simple as

well as unicellular trichomes on midrib of lower leaf

surface 2. E. vulgare

1. Echium plantagineum L., Mantissa plantarum

altera: 202 (1771); C.H.Wright: 44 (1904); P.E.Gibbs: 57

(1971); Bramwell: 64 (1972); P.E.Gibbs: 99 (1972);

Piggin: 217 (1977); J.R.Edm.: 322 (1978); Toelken: 1 156

(1986); Verde.: 81 (1991). Type: Barrelier, Plantae per

Galliam, Hispaniam et Italiam observatae, iconibus

exhibitae 145, 1. 1 026 (1714), see Verde.: 83 (1991).

E. lycupsis auct. non L.

E. violaceum auct. non L. (1767).

Annual or biennial herb with 1-many flowering

stems, up to 0.8 m high. Indumentum of soft, appressed,

long (up to 3 mm) tubercle-based setae, each trichome

with a concentric ring of cells at base, sometimes stems,

leaf margins and veins on lower surface of leaf with a

sublayer of soft, much shorter, simple, unicellular tri-

chomes. Stems erect or decumbent. Basal leaves ovate to

oblanceolate, 50-300 x 10-70 mm, with prominent lat-

eral veins on both surfaces; lamina narrowing rather

abruptly into petiole; petiole up to 55 mm long. Cauline

leaves oblong to lanceolate, uppermost ± cordate at base.

Cymes usually distinctly stalked, lengthening after flow-

ering to 250-300 mm. Calyx up to 12 mm long in fruit.

Corolla blue or deep purplish blue, broadly funnel-

shaped, 10-20 mm long, glabrous except for sparse hairs

on veins and margins of outside surface, with a ring of 10

distinct pilose lobes within tube. Stamens : lower pair

long-exserted, the others included or only slightly exsert-

ed; filaments with long hairs. Nutlets greyish brown, tri-

quetrous, rugose, tuberculate with tubercles varying in

size, 2. 0-3.0 x 2. 0-2. 5 mm. Chromosome number : n = 8

(Britton 1951). Figure 5.

Distinguishing characters'. E. plantagineum differs from

some Echium species in the colour of the corolla. E.

italicum L. and E. boissieri Steud., for instance, have

yellowish or bluish white or flesh-coloured corollas,

whereas E. plantagineum has blue or purplish blue flow-

ers. The subglabrous corolla with hairs on veins and mar-

gins only, and basal leaves usually broadly ovate to

174

Bothalia 28,2 (1998)

oblanceolate with prominent lateral veins distinguish E.

plantagineum from E. vulgare and a number of other

species, including E. albicans Lag. & Rodr., E. creticum

L. and E. tuberculatum Hoffmanns. & Link. E. vulgare

has a corolla with long, simple, unicellular trichomes on

the veins and margins and smaller ones on the corolla

itself. The basal leaves of E. vulgare are linear-lanceolate

to narrowly oblanceolate, attenuating into a short petiole.

Distribution: E. plantagineum is native to western

Europe including southwestern Britain and the

Mediterranean area. It has been introduced into other

parts of Europe, southern Russia and the Caucasus; it is

a widely distributed, troublesome weed in Australia

(Verdcourt 1991). In southern Africa this introduced

species occurs from the Western Cape, extending more

or less along the coast, to the Eastern Cape. It is also

found in the temperate mountainous areas of the Free

State, Lesotho and KwaZulu-Natal, with a few records

from southern Mpumalanga (Figure 6).

Habitat: although E. plantagineum is most noticeable

along roadsides, it also occurs in waste ground and as a

weed of cultivation. It has also been recorded from rocky

slopes in grassland.

Flowering time: September to March.

Common names: bloudissel (doring), Paterson’s curse,

salvation Jane and purple bugloss.

Illustrations: Jacq.: t 45 (1771); Butcher: t. 995 (1961); Verde.: 82

(1991).

2. Echium vulgare L., Species plantarum 139

(1753); P.E. Gibbs: 54 (1971); P.E.Gibbs: 99 (1972);

Piggin: 224 (1977); J.R.Edm.: 322 (1978); Toelken: 1156

(1986). Type: LINN 191.19.

For synonymy see Gibbs (1971 ).

Biennial, or sometimes annual herb, up to 1 m high.

Indumentum of sparse, stout, appressed to patent hairs

with a multicellular base and a dense sublayer of much

shorter hairs with base not so prominently multicellular.

FIGURE 6. — Distribution of Echium plantagineum.

Stem simple or with many decumbent, flowering stems;

usually branched if grazed or damaged. Basal leaves lin-

ear-lanceolate to narrowly oblanceolate, 50-260 x 10-30

mm, attenuating into a short petiole (subsessile), lateral

veins not prominent. Cauline leaves narrowly lanceolate,

sessile. Calyx lobes 6-8 mm long in fruit. Corolla bril-

liant blue, fading to mauve, broadly funnel-shaped,

10-15 mm long, puberulous and with sparse long, simple

trichomes on veins and margins outside. Stamens: four

long-exserted, one short-enclosed. Nutlets brownish

grey, tuberculate with large and numerous small tuber-

cles, triquetrous, with prominent dorsal and ventral

keels, beak narrow, acute. Chromosome number: n = 16

(Britton 1951). Figure 7.

Echium vulgare is a very variable species in other

parts of the world. The synonymy of Gibbs (1971) fol-

lows a rather broad species concept in his revision of the

Spanish members of the genus.

Distinguishing characters: E. vulgare can be distin-

guished from other Echium species by its prominently

exserted stamens, in contrast to, for example, E. parvi-

florum Moench and E. arenarium Guss., which have all

the stamens included in the corolla. It differs, like E.

plantagineum , from another group of Echium species in

having a blue corolla and not one which is flesh-

coloured, yellowish white or bluish white. Some Echium

species, like E. italicum L., have a narrowly funnel-

shaped corolla with the tube not markedly widening

towards the apex. E. vulgare, however, belongs to those

taxa with a broadly funnel-shaped corolla in which the

tube widens markedly towards the apex.

Distribution: E. vulgare, native to Europe, has been

introduced to various countries worldwide, often becom-

ing a weed. In southern Africa the species is found in

Mpumalanga, Free State, Lesotho and the Eastern Cape

(Figure 8).

Habitat: E. vulgare is a weed of old cultivated lands,

waste places and roadsides, but occasionally also occurs

in montane grassland.

Flowering time: November to March.

Common names: blue devil, blue weed, viper’s bugloss.

Illustrations: Lam.: t. 94 (1791); Butcher: t 994 (1961).

SPECIMENS EXAMINED

Specimens held at PRE, unless otherwise indicated.

The numbers in brackets signify the identity of the spec-

imens: (OF. plantagineum , (2) E. vulgare.

Acocks 9250 ( 1 ) K, PRE; 16282 (2) K.

Balsinhas 3546 (1). Barnes 94 (1). Bayer 6005 (1) Bayliss 73 (1)

GRA, K, PRE; 1133, 6132 (1); 24276 (1) NBG. Bester 710 , 955 (1)

NH. Bolus 4989 (1) BOL, K. Bos 259 (1) NBG. Boucher 3941 (1);

3395 (1) NBG. Boucher & Ahmed 7/5(1). Brink 239 ( 1) GRA, K, PRE,

NBG; 265 (2) GRA, K, PRE. Britten 520 (1); 2969 (1) GRA. Brown

sub PRE41060 (1). Brusse 5601 (2). Burgoyne 971 (1). Burrows 2198

(1).

Case sub PRE654738 ( 1 ). Chelchinsky 16 (2) GRA. Codd & Dyer 6233

(2) K. Coleman 524 ( 1) NH, PRE. Comins 1899 (1) GRA, PRE.

Bothalia 28,2 (1998)

175

FIGURE 1 .—Echium vulgare. A, lower part of plant, x 0.8; B, inflorescence, x 0.5; C, flower, x 2; D, longitudinal section of corolla, x 1.5; E,

style, x 20 ; F, nutlet, x 15; G, lower surface of leaf, x 3. A, B, Codd & Dyer 623 3\ C-E, Van Wyk 5268', F, Zietsman 1480', G, Sidey 3977.

176

Bothalia 28,2 (1998)

Dahlstrand 541, 3094 ( 1 ) GRA. Davidse 6721 (1). Davies s.n. ( 1 ) GRA.

De Vries 9 (1) GRA. Dieterlen 1030 (2) PRE, SAM. Dold 1987, 2026

(2) GRA. Drege 442 (1) GRA. Dreyer 14 (2). Drijfhout 4137 (1) NBG,

PRE.

Edwards, Cadman & Norris 3163 (2) BOL, NU, PRE. Engelbrecht

JE650 (2) J.

Eourcade 1886(1) BOL, K, NBG; 5S25(1) NBG. Fugler 100(1) K,

PRE

Gafney sub PRE10201 (2). Galpin 121 ( 1 ) K, NBG; 212(1) PRE; 4351

(1) GRA, PRE; 7939 (1). Garrett 60 (2) E, NU. Garside 147(1) K.

Germishuizen 4102 ( 1 ). Giffen 7/5(1); 809 ( 1 ) GRA, PRE. Greef 2(1).

Guthrie 3114 (1) NBG.

Hilger 44(1). Hilliard & Burn 6999 (1) NU, PRE; 18686 (2) E, K, NU,

PRE. H Hilary sub NH 37460 ( 1 ) NH. Hoare 31 (1) GRA. Hoener 1705

( 1 ). Hutton s.n. ( 1 ) GRA.

Jacobs?. 311 (2) PRE; 2082 (2) NBG, PRE. Jacot Guillarmod 3211 (2);

7341 ( 1 ) GRA; 9837 ( 1 ) GRA, PRE. Joffe 873 ( 1 ) NBG, PRE. Jordaan

1 (1).

Kerfoot 6216 ( 1 ). Killick & Marais 2107 ( 1 ) K, NU, PRE.

Leendertz 8184 ( 1 ). Loubser 3867 ( 1 ). Low 754 ( 1 ) NBG.

MacCabe 3149 (2). Marais 567 ( 1 ) GRA, NBG, PRE; 807 ( 1 ) GRA, K,

PRE. Maree 12 (1) Mar loth 7201 (1) PRE; 10864 (1) NBG, PRE.

Marsh 75 (2) K, NH, PRE. Mauve & Tblken 4520 (2) K, PRE. Moll

2663 (1) K, NU.

Nel 77/9(1) GRA.

Page sub BOL16500 (2). Parker 4376 (1) BOL, K, NBG. Pegler 2138

(1) BOL, PRE. Phelan 200 ( 1) NU. Pienaar & Kok 1225 (1) PRE,

PRU. Pretorius 3(1) NBG.

Relief 373 (I). Rogers 26933, sub TRV3007 (1).

Saaiman 615 (2). Scharf 1763 (1) NBG, PRE. Schlieben 7607 (1) K.

Schlieben & Ellis 12331 (1) K, PRE. Schonland 3631 (1) GRA, K.

Sidey 3848 ( 1 ); 3977 (2) K, NH, PRE. Steyn 27(1). Stirton 8151, 8218,

8389 ( 1 ). Story 2084 (2) GRA, PRE. Strever 721 (2). Strey 7882 ( 1 ) K,

NH, NU, PRE.

Taylor 5485, 5784, 5978 (1) NBG. Tenant sub PRE14149 (2). Theron

708 (2) K, PRE; 2189 (2) Thode 62716 ( 1 ) K, PRE; sub PRE19418 ( 1 ).

Turner 267 (1).

Van Breda 12 (1) Van Rensburg 1957 (1) K. Van Wyk 5268 (2) PRE,

PRU; 5279 (I) PRE, PRU; 12059 ( 1) PRU. Van Wyk & Abbott 12059

(2) . Venter 7244 (2).

Ward 303 ( 1 ) NH Welman 780 (2) GRA, PRE. West 1514 ( 1 ). Williams

296, 3155 (1) K Wolley Dod 1893 (1) K, PRE. Wurtz 464 ( 1 ) NBG.

Zietsman 1480 (2).

ACKNOWLEDGEMENTS

The assistance of the artist, Ms G. Condy, the photog-

rapher, Mrs A. Romanowski and Mrs E. du Plessis of the

National Botanical Institute in preparing the manuscript,

is gratefully acknowledged. The directors, curators and

staff of the cited herbaria are thanked for allowing the

authors to study their material and for sending specimens

on loan.

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Bothalia 28,2: 179-182 (1998)

A revision of Ledebouria (Hyacinthaceae) in South Africa. 2. Two

new species, L. crispa and L. parvifolia , and L. macowanii re-instated

S. VENTER* * and T.J. EDWARDS **t

Keywords: Hyacinthaceae, Ledebouria, new combination, new species. South Africa, taxonomy

ABSTRACT

Two endemic Ledebouria Roth species are described with notes, diagnostic characters, distributions and alliances. L.

crispa appears to be allied to L. socialis (Baker) Jessop with which it shares densely gregarious, epigeal bulbs and L. undu-

lata (Jacq.) Jessop of the Northern and Western Cape which shares crispate leaf margins. L. parvifolia is endemic to

Mpumalanga (eastern Transvaal) and was recorded from a single locality near Graskop; it differs from L. cooperi (Hook.f.)

Jessop in having hairy upper leaf surfaces. L. macowanii (Baker) S. Venter is re-instated; it is distinguished from L. cooperi

by its tightly clasping bulb scales and single cataphyll.

INTRODUCTION

Ledebouria Roth is distributed across Africa into the

southern tip of India and the northern tip of Madagascar

and includes 33 species. The genus is very well repre-

sented in South Africa with high species densities in the

central and eastern regions. This paper deals with two

new species recorded from Northern Province and

Mpumalanga respectively.

1. Ledebouria crispa S. Venter, sp. nov. L. sociali

(Baker) Jessop similis sed statura minore, marginibus

crispatis foliorum et bracteolisque flores subtendentibus

differt.

TYPE. — Northern Province, 2329 (Pietersburg): Farm

Majebeskraal, (-DC), Venter 11202 (PRE, holo.; NU,

UNIN).

Plants gregarious. Bulbs epigeal to semi-epigeal,

15-30 x 10-20 mm, ovoid to obovoid; dead bulb scales

pale brown, membranous, apices attenuate, live bulb

scales fleshy, tightly appressed, lacking threads when

torn; bulblets often abundant. Leaves 4-10, synanthous,

linear to linear-lanceolate, 20-70 x 4-6 mm, erectly

spreading, slightly succulent, glaucous, veins obscure;

margin crispate; base canaliculate; apex acute. Inflores-

cence solitary, erect, 40-50 mm long, 8-12-flowered;

scape terete, glaucous, glabrous, 40-50 mm long; bracts

and bracteoles membranous 1.2-1. 5 x 0.4 mm, linear-

lanceolate, glaucous. Pedicels spreading, 2^1 mm, green

suffused with pink. Tepals reflexed in distal half, oblong,

4. 0-5.0 x 1.0-1. 5 mm, pink, keel dull green, apex acute.

Stamens erect, 3. 0-3. 5 mm long; filaments pink, epite-

palous; anthers 0.5 mm long, mauve. Ovary depressed

ovoid, 2.5 x 3.0 mm, lobes obtusely deltoid, glabrous,

apex tapered; stipe 1 mm long. Style 3. 0-3. 5 mm, white.

t To whom correspondence should be addressed.

** Department of Botany, University of Natal, Private Bag X01, 3209

Scottsville.

* Part of this research was undertaken at the Department of Botany,

University of the North, Private Bag 1106, 0727 Sovenga, South

Africa.

MS. received: 1998-01-05.

Capsule symmetrical, globose-ovoid, base truncate. Seed

3.0 x 1.5 mm, drop-shaped, wrinkled, black. Figure 1.

L. crispa has only been recorded from hills surround-

ing Pietersburg. The substrates from which specimens

FIGURE 1. — Ledebouria crispa. A, habit, x 1.3; B, leaf lamina trans-

verse section, x 4.6; C, tepal apex, x 6.5; D, bract and bracteole,

x 10; E, flower, x 6.5; F, tepal with stamen, x 4.6; G, gynoeci-

um, lateral view, x 5. Drawn from Venter 11202 by F. Venter.

180

Bothalia 28,2 (1998)

were collected are light brown, gritty loams derived from

talcose schists of the Mothiba Formation (Pietersburg

Group) (SACS 1980). Precipitation in the area occurs

predominantly in summer and ranges between 400-600

mm (Barnard et al. 1972). Plants grow in scattered

groups of 5 to 40 individuals in typical Pietersburg

Plateau False Grassveld (Acocks 1988). The species

flowers from October to December.

L. crispa is allied to a number of species with promi-

nent bulb necks and lacking extendable xylem ‘threads’

in their torn leaves. This complex includes L. concolor

(Baker) Jessop, L. crispa , L. undulata (Jacq.) Jessop, L.

ovalifolia (Schrad.) Jessop, L. petiolata (Van der Merwe)

S. Venter, L. socialis (Baker) Jessop and L. macowanii

(Baker) S. Venter.

L. crispa is distinguished from its allies by its diminu-

tive size, synanthous leaves with crispate margins, epi-

geal bulbs and solitary inflorescences. L. undulata, which

is limited in distribution to the Northern and Western

Cape, also has wavy leaf margins and solitary inflores-

cences but is a much larger, hysteranthous plant with a

hypogeal bulb. In addition the leaf undulation in L. undu-

lata is variable and plants from shaded situations tend to

have smooth margins. This differs from L. crispa in which

the crispate margins are unaffected by microhabitat. L.

socialis approaches L. crispa in a number of characters.

Its densely gregarious habit and epigeal bulbs are simi-

lar, but the former has maculate leaves with smooth mar-

gins and pedicels which are ebracteolate. In addition, the

species are allopatric; L. socialis is limited to the Eastern

Cape (Figure 2).

Specimens examined

NORTHERN PROVINCE. — 2329 (Pietersburg): Pietersburg Game

Reserve, (-CD), Venter 1 3209a (PRE, UNIN); Lebowa, Majebeskraal

Farm, (-DC), Venter 11202 (PRE, NU, UNIN).

2. Ledebouria parvifolia S. Venter, sp. nov. L.

cooperi (Hook.f.) Jessop arete affinis sed foliis appressis

FIGURE 2. — Geographical distribution of L. crispa, •; and L. parvi-

folia, M

superficie adaxiali trichomatibus basibus latis in seriebus

longitudinalibus dispositis differt.

TYPE. — Mpumalanga, 2430 (Pilgrims Rest): Graskop,

near Lisbon Falls, Farm Lisbon 531, (-DD), Venter

12609a (PRE, holo.).

Plants gregarious. Bulb hypogeal, 10-25 x 10-25 mm,

obovoid; dead bulb scales membranous, brown, apices

attenuate, with threads when torn, live bulb scales fleshy,

tightly appressed; bulblets usually present. Leaves en-

veloped by a single cataphyll, synanthous, 2 or 3, humi-

fuse, lanceolate to oblong, 15-30 x 15-20 mm, with

threads when torn, fleshy, dull green, adaxial surface with

longitudinal rows of papillae, venation obscure; margin

ciliate; base canaliculate; apex acute. Inflorescence soli-

tary, 25-35 mm long, erect, lax, 8-12-flowered; scape

terete, purple, glabrous, 10-15 mm long; bracts deltoid,

0.5 x 0.5 mm, pink to purple; bracteoles absent. Pedicels

cernuous, 2-3 mm long, pink. Tepals recurved distally,

equal, oblong, 2. 5-3.0 x 1.0 mm, pink with a green keel,

apex obtuse. Stamens erect, 3 mm long; filaments pink

epitepalous; anthers 0.5 mm long, pale violet. Ovary

ovoid, 3-lobed, 1x2 mm, lobes narrowly transversely

oblong, shoulders rounded; stipe 0.5 mm long. Style 3 mm

long, triangular in section, glabrous. Seed 3 mm long,

drop-shaped, strongly wrinkled, brown. Figure 3.

FIGURE 7>. —Ledebouria parvifolia. A, habit, x 0.6; B, leaf lamina

transverse section, x 0.9; C, tepal apex, x 7.6; D, bract, x 12.7;

E, tepal with stamen, x 7.6; F, flower, x 6.3; G, gynoecium, lat-

eral view, x 6.3. Drawn from Venter 12609a by F. Venter.

Bothalia 28,2 (1998)

181

This species was recorded from a single locality on

dolomites of the Malmani Formation in the Chuniespoort

Group (SACS 1980). Soils derived from these substrata

are fine-grained, dark brown clays. The species is a con-

stituent of montane grassland with scattered Dombeya

rotundifolia and Faurea saligna. L. parvifolia flowers

from October to December.

L. parvifolia is closely related to L. cooperi (Hook.f.)

Jessop but differs in producing appressed leaves with

longitudinal rows of broad-based hairs on their adaxial

surfaces. This indumental character is stable under a

wide range of environmental conditions. Both L. cooperi

and L. parvifolia produce a single basal cataphyll, a char-

acter not found elsewhere in the genus.

3. Ledebouria macowanii (Baker) S. Venter , comb.

nov.

Scilla macowanii Baker in The Gardener's Chronicle 3: 748 (1875).

Type: Cape, Somerset Division, Boschberg, MacOwan 1841 (GRA!,

lecto. here designated; BOL!; PRE!; Z photo.).

Scilla nelsonii Baker: 488 (1896). Type: Cape, Vaal River, Nelson

167 (K!, holo.; PRE!).

Plants weakly gregarious. Bulb hypogeal, 15-30 x

15-30 mm, ovoid; dead bulb scales tightly clasping,

lacking threads when tom; bulb neck 5-10 x 10 mm.

Leaves synanthous, 3 or 4, spreading, linear-lanceolate,

75-100 x 6-8 mm, without threads when torn, fleshy,

dull green, abaxial surface green with occasional purple

spots in the lower half, venation obscure; margin

smooth; base canaliculate; apex obtuse to acute.

Inflorescences 1 or 2, lax, cylindrical, 25-60 mm long,

flaccid, 20-40-flowered; scape base terete, spotted

40-100 mm long; bracts deltoid, 0.5 x 0.5 mm, green to

pink; lacking bracteoles. Pedicels 6-7 mm long, pink to

purple. Tepals oblong, slightly recurved, 4.0 x 1.5 mm,

apex obtuse, green suffused with pink. Stamens spread-

ing, 3.8 mm long; filaments white with purple above;

anthers 0.5 mm long, pale violet. Ovary globose, 6-

lobed, 1.5 x 2.0 mm, lobes obtusely deltoid with a basal

ridge; stipe 0.5 mm long. Style 2.0 mm long, triangular in

transverse section, purple. Seed drop-shaped, 3-4 mm

long, surface strongly wrinkled, brown.

L. macowanii is closely allied to L. socialis but differs

in its hypogeal bulbs and prominent nectariferous ridges

on the carpel bases. Jessop (1970) reduced this taxon to

synonymy with L. cooperi, however, it is distinguished

from L. cooperi by its tightly clasping bulb scales and

single cataphyll. In the Eastern Cape these species are

sometimes sympatnc but the range of L. macowanii ex-

tends up to the Northern Province (Figure 4). The species

appears to occur in areas of reasonably high precipita-

tion, hence its limitation to the coast and elevated areas

in which orographic rainfall occurs. Plants occur in

damp, shallow lithosols derived from quartzites, conglo-

merates or sandstones. The species usually occurs in

semi-shade. The flowering period of L. macowanii

extends from October to March with a peak in November

and December.

Specimens examined

NORTHERN PROVINCE. — 2231 (Pafuri): Punda Maria, Dzun-

dwini Hill, (-CC), Codd & Dyer 4598 (KNP, PRE). 2329 (Petersburg):

Louis Trichardt, (-BB), Koker 3 (PRE). 2331 (Phalaborwa): Letaba

Camp, (-DC), Codd 4675 (KNP, PRE). 2427 (Thabazimbi): Rooiberg,

(-DD), Van der Merwe 2044 (PRE).

MPUMALANGA. — 2431 (Acomhoek): Skukuza, (-DC), Codd &

De Winter 5076 (PRE); Mabase, (-DC), Lang s.n. (PRE).

NORTH-WEST. — 2527 (Rustenburg): Rustenburg, (-CA), Van der

Merwe 1558 (PRE).

SWAZILAND. — 2631 (Mbabane): Komati Bridge , (-AA), Compton

29391 (PRE); Usuthu Dam, (-CA), Compton 27120 (PRE). 2632 (Bela

Vista); Mbuluzi Nature Reserve, (-AA), Culverwell 1389 (PRE).

FREE STATE. — 2827 (Senekal); Doornkop Farm, (-DD), Goosens

777 (PRE). 2828 (Bethlehem): Clarence, (-CB), Van Hoepen 18270

(PRE). 2924 (Fauresmith): Farm Wanda, (-CB), Smook 3045 (PRE).

2927 (Maseru): Clocolan, Hillcrest Farm, (-BA), Crosby 420 (PRE).

KWAZULU-NATAL. — 2732 (Ubombo): Phelendaba Crossing, (-BA),

Germishuizen 3533 (PRE). 2832 (Mtubatuba): Enseleni Nature

Garden, (-CC), Venter 6115 (PRU).

NORTHERN CAPE. — 2824 (Kimberley): Riverton Pont, (-DB),

Leistner 2016 (PRE). 2923 (Douglas): Douglas, Herbert Dist., (-BB),

Orpen 57 (BOL).

WESTERN CAPE. — 3421 (Riversdale): Riversdale, (-AD), Bohnen

7461 (STE); Still Bay, (-AD), Bolus s.n. (BOL), Ferguson s.n. (BOL);

Albertinia, Ystervarkpunt, (-BC), Willemse 107 (STE). 3423 (Knysna):

Knysna, (-AA), Breyer 23711 (PRE).

EASTERN CAPE. — 3127 (Lady Frere): Glen Grey, Nzebanya

Mountain, (-CC), Galpin 1914 (PRE). 3226 (Fort Beaufort): Katberg

Mountain, (-DB), Bayliss 7442 (NBG). 3227 (Stutterheim): Pirie,

(-CC), Taylor 1760 (BOL); King Williams Town, (-CD), Sim 628

(NU), Sim 1075 (BOL), De Victoria 15 (BOL). 3228 (Butterworth):

Kwenqura River, (-CC), Galpin 5814 (PRE). 3326 (Grahamstown):

Peddie, Committee’s Drift, (-BB), Van der Merwe 2132 (PRE);

Blaauw Krantz, (-BD), Daly 1026 (BOL). 3327 (Peddie): East London,

(-BB), Wood 2817 (BOL), Rattray s.n. (BOL), Pamphlett 77 (NBG).

ACKNOWLEDGEMENTS

Financial assistance from the Natal University

Research Fund and the FRD is gratefully acknowledged.

The cited herbaria are thanked for the loan of material

examined during this study. Mr M. Lambert and Dr O.A.

Leistner are thanked for the Latin diagnoses. Sandy

Burrows is thanked for drawing the original plates from

which Figures 1 & 2 were modified by F. Venter. Mr K.

Gamble is thanked for his assistance in the field.

182

Bothalia 28,2(1998)

REFERENCES

ACOCKS, J.P.H. 1988. Veld Types of South Africa, 3rd edn. Memoirs

of the Botanical Survey of South Africa No. 57.

BAKER, J.G. 1875. New garden plants. The Gardener's Chronicle 3:

748.

BAKER, J.G. 1896. Liliaceae. In W.T. Thiselton-Dyer, Flora capensis

6: 253-528. Reeve, London.

BARNARD, W.S., SMIT. P. & VAN ZYL, J.A. 1972. Suid-Afrika: die

land en sy streke. Nasou. Cape Town.

JESSOP, J.P. 1970. Studies in the bulbous Liliaceae: 1. Scilla , Scltizo-

carphus and Ledebouria. Journal of South African Botany 36:

233-266.

SACS (SOUTH AFRICAN COMMITTEE FOR STRATIGRAPHY)

1980. Stratigraphy of South Africa, Part 1. Flandbook of the

Geological Survey of South A frica.

Bothalia 28,2: 183-196 (1998)

Notes on African plants

VARIOUS AUTHORS

METZGERIALES — FOSSOMBRONIACEAE

FOSSOMBRONIA OCCIDENTO-AFRICANA'. IS IT CONSPECIFIC WITH F INDICA1

INTRODUCTION

Fossombronia occidento-africana S.W. Amell (1952)

was described as a new species from his collections from

Mount Aureol [Mont Oriel] at Freetown, Sierra Leone,

West Africa. He cited the following collection numbers:

2409 , 2413, 2418, 2431, 2448, 2449, 2468, 2476 and

2497, but did not designate a type specimen. Arnell’s

only illustration of the new species is a simple line draw-

ing of a cross section of the stem, with a single down-

ward-projecting rhizoid.

Because Jones & Harrington (1983) considered

Arnell’s description to be inaccurate in certain aspects,

they presented a more thoroughly investigated descrip-

tion. They also selected the specimen Arnell 2497 as the

lectotype for the following reasons: they considered it to

be Arnell’s most copious collection with numerous

mature sporangia; in addition, Arnell had made a micro-

scope preparation of it. They cited several other Arnell

specimens, namely 2409, 2413, 2448, 2517, as well as

2231 and 2249, but according to Arnell’s numbers given

above, the last two numbers must be incorrect and should

read 2431 and 2449 respectively. From Sierra Leone they

also cited several Harrington specimens: 504, 555, 599 &

640 and one Richards specimen, R7086, as well as Sellar

B35 p.p. From Ghana, three Hall specimens were cited,

and from Abuja in northern Nigeria, Jones (1985) cited

his own collection, Jones 885a. Jones & Harrington

(1983) found F. occidento-africana to be the most com-

mon species of F ossombronia in lowland West Africa.

Augier (1985) also reported it from Cameroun, after first

assigning his specimens to F. crozalsii (Augier 1972) and

Tixier (1989) cited Brunei 8838 from Togo. Neither

Jones nor Jones & Harrington published an illustration of

the species. Scott & Pike (1988), in their wide-ranging

study of Fossombronia species, published two spore

micrographs of F. occidento-africana (their figs. 25, 26)

and placed it in synonymy under F. indica Steph.

Evidently unaware of the lectotypification of F. occiden-

to-africana by Jones & Harrington, they selected Arnell

2249 as the lectotype. This number, in all probability,

should be 2449. The incorrectly cited numbers by both

Jones & Harrington and Scott & Pike are a curious coin-

cidence, and it suggests that the said numbers were

incorrectly copied on the herbarium packets held at (S).

As the holdings on loan from the Swedish Museum of

Natural History (S) had already been returned by me, I

could not check these numbers myself. However, Prof.

R. Stotler (in litt.) agrees with me that the Arnell 22 xx

numbers are ‘surely a mistake by Arnell, since all of his

other nine Fossombronia collections (of F. occidento-

africana) are 24 xx’.

This paper is presented because F. occidento-africana

has not been illustrated, except for ArneH’s single figure

of the stem cross section and the two spore micrographs

by Scott & Pike (1988). I also needed to investigate the

placement of F. occidento-africana in synonymy under

F. indica implemented by Scott & Pike.

Fossombronia occidento-africana S.W. Arnell in

Botaniska Notiser 3: 317 (1952); Jones & Harrington:

248 (1983). Type: Sierra Leone, Freetown Peninsula,

Mount Aureol (‘Oriel’), Amell 2497 (S, lecto.!).

Plants in crowded stands, green; shoots smallish, sim-

ple (Figure 2A), up to 7 or 8 mm long, 1.3-1. 4 mm high,

± 2.6 mm wide, occasionally once furcate, apical seg-

ments moderately divergent, up to 3 mm long. Stems

prostrate, tapering proximally, chlorophyllose, some-

times with a young ventrolateral branch (Figure 2B),

planoconvex in cross section, toward apex (Figure IK) ±

350 pm (11 cell rows) high, ± 420 pm wide, basally

(Figure 1L) ± 250 x 300 pm. Rhizoids purple, 12.5-22.5

pm wide. Leaves (Figure 1A-H) overlapping, suc-

cubously inserted, apically small, soon becoming larger,

oblong to broadly obovate or rounded, mostly shorter

than wide, but often somewhat narrower below, apex

truncate or rounded, sometimes shallowly notched,

slightly undulate, (875— )950— 1 275 x 1050-1400 pm,

margins with 4-7 well-spaced papillae, 30 x 20 pm, usu-

ally sessile, sometimes at base of leaf raised on 1-3

elongated cells, the latter arranged end to end. Leaf cells

thin-walled, at upper margins (Figure II) rectangular

across, 25.0-42.5 x 30.0-52.5 pm, at lateral margins

long-rectangular, 50.0-57.5 x 25.0-30.0 pm; upper 1am-

inal cells 5- or 6-sided, 37.5-50.0 x 22.5-30.0; middle

laminal cells 37.5-62.5 x 35.0-37.5 pm; basal cells

57.5-75.0 x 27.5-37.5 pm. Oil bodies disintegrated in

the material seen; chloroplasts (Figure 1J) numerous,

mostly rounded, ± 2.5 pm diam.

Monoicous. Antheridia dorsal on stem, at apex appar-

ently naked, intact, short-stalked, globose (Figure 2C), ±

160 pm diam., further along branch disintegrated, but

perigonial bracts (Figure 2A, D) remaining, irregularly

shaped (Figure 1M-P), base up to 350 x 140 pm, with

one or occasionally two, tapering, apical appendage(s),

250-500 pm long, a papilla at the tip(s). Archegonia in a

row dorsally along stem (Figure 2A). Pseudoperianth

(Figure IQ, R) near stem apex (Figure 2E), funnel-

shaped, up to 1175 pm long, width across flaring mouth

(Figure 2F) ± 1500 pm wide, with ± 8 triangular lobes,

each topped by a papilla, constricted toward base, ± 500

pm wide, sometimes deeply cleft on one side, free mar-

gins overlapping slightly; cells comparable in size and

184

Bothalia 28,2 (1998)

FIGURE I — Fossombronia occidento-africana. A-H, leaves; I, margin of leaf with 2 papillae; J, middle laminal cells with chloroplasts, oil bod-

ies disintegrated; K, c/s of stem apex; L, c/s of stem base; M-P, perigonial bracts; Q, pseudoperianth from side; R, opened pseudoperianth;

S, c/s of seta; T, cells in capsule wall with thickenings. A-T, S.W. Arnell 2448. Scale bars: A-H, Q, R, 500 pm; I, S, 100 pm; J, T, 50 pm;

K, L, M-P, 250 pm. Artist; M. Steyn.

Bothalia 28,2 (1998)

185

FIGURE 2. — Fossombronia occidento-africana. A, simple branch with row of archegonia and single perigonial bract (indicated by arrow); B,

branch with young ventrolateral shoot; C, apex of branch with antheridia (indicated by arrows) and archegonia; D, apex of branch with

archegonium and papillae around its base; perigonial bract in foreground (indicated by arrow); E, pseudoperianth near apex of branch; F,

pseudoperianth from above and part of rather shrivelled seta (indicated by arrow). A-F, S.W Arnell 2448. A, x 12; B, x 32; C, x 50; D, x

57; E, x 19; F, x 46.

shape to those of leaves. Capsules globose, ± 675 pm

diam., wall bistratose, inner cell layer with irregularly

rectangular cells, 27.5^4-2.5 x 22.5-25.0 pm, on both

longitudinal and tangential walls, with 2^1 deep brown,

heavy subnodular thickenings, some of which are pro-

longed to form complete or incomplete semi-annular

bands (Figure IT). Seta up to 3 mm long, ± 100 pm

diam. (Figure IS). Spores light brown, ± hemispherical,

40-50 pm diam.; distal face (Figure 3A-D) convex, with

8 or 9 tall and thin, very irregular, rather wavy lamellae

running ± longitudinally or diagonally across face,

spaced ± 5 pm apart and interconnected by transverse

ridges, forming irregular, complete or mostly incomplete

areolae, walls smooth above or rather ragged, sometimes

raised into processes at the angles; proximal face (Figure

3E) lacking triradiate mark, ornamented with irregular,

FIGURE 3. — Fossombronia occidento-africana. Spores and elaters. A, B, distal face; C, side view of distal face; D, close-up of distal face mar-

gin; E, proximal face; F, elaters. A-F, S. W. Arnell 2497. A, x 832; B, x 940; C, x 987; D, x 1757; E, x 825; F, x 555.

186

Bothalia 28,2 (1998)

faint and low ridges or granules that are interconnected,

around spore periphery numerous spines that are not

readily individually distinguishable nor joined by a dis-

tinct perispore. Elaters (Figure 3F) light brown, 102-167

pm long, 10.0-12.5 pm wide in middle, and trispiral,

tapering to tips, 5. 0-7. 5 pm wide, frequently ending in a

bispiral loop, occasionally branched.

DISCUSSION

Arnell (1952) was able to study fresh plants of F. occi-

dento-africana and described them as dark green; in old

herbarium specimens the colour may have faded to a

lighter green.

Jones & Harrington (1983) could not confirm the

presence on the leaf cuticle of ‘1-2 long striae per cell’

and ‘sometimes small trigones’ (at the cell corners pre-

sumably) that Arnell reported; neither could I. Arnell

(1952) described F. occidento-africana as dioicous, the

male plants ‘with antheridia on the dorsal side near the

apex, covered by rather long narrow bracts with deeply

bidentate apex. Antheridia about 100 pm diam., pale yel-

low, globose’. Jones & Harrington (1983) on the other

hand, described the species as ‘monoecious, but proba-

bly protandrous’, and do not mention antheridia or

perigonial bracts at all. As I have now demonstrated, the

species is monoicous and perigonial bracts do in fact

occur. Jones & Harrington (1983) reported ‘massive

radial thickened bands’ on the cells of the inner spo-

rangium wall, with which I agree. Arnell (1952) found

the spore ‘pattern’ of F. occidento-africana to be the

same as that of F. stephani (sic) from Reunion (lamellis

interrupts vermicularibus), but the spores of the former

are smaller in size, according to him. Scott & Pike (1988:

figs 29, 30) illustrated the spores of F. stephanii and

remarked that they ‘are very similar indeed to those of F.

texana (= F. mexicanaf the disjunction from Central

America to Reunion Island (off East Africa) seems phy-

togeographically improbable, so that we hesitate to

equate the two without further investigation’. In the same

study, Scott & Pike (1988: figs 33, 34) remark that F. tex-

ana has sometimes been confused with the F. foveolata

complex (Scott & Pike 1987). In following this train of

thought the spores of F. occidento-africana should be

similar to those of F. foveolata which I don’t think to be

the case, except that they are both reticulate. Jones &

Harrington (1983) admit that F. occidento-africana ‘may

indeed prove to be F. crozalsii Corbiere, which is, how-

ever, imperfectly known’. Furthermore, they state, ‘we

prefer to keep the two species (i.e. F. crozalsii and F.

occidento-africana ) separate until the range of variation

of F. crozalsii is adequately known’. British collections

of F. crozalsii have been referred to F. pusilla var. mar-

itima by Paton (1973).

Scott & Pike (1988: figs 25, 26) reduced F. occiden-

to-africana to synonymy under F. indica, describing the

spores of the former as ‘very distinct, with a very delicate

reticulum of low lamellae over the distal face and a cob-

web-like network over the proximal face. In this as well

as vegetative features it seems identical to F. indica

Stephani, an earlier name, to which we here reduce it as

a synonym’. I hesitate to accept this reduction for the fol-

lowing reasons: the type (and only known) specimen of

F. indica (leg. Pfleiderer s.n.) is a small and delicate

plant, its dimensions agreeing with those given by

Srivastava & Udar (1975), i.e. ‘stem 3. 5-5.0 mm long’.

This is rather shorter than the 8 or 10 mm long thallus

branches reported by previous authors for F. occidento-

africana and considerably shorter than Stephani ’s (1917)

length of 13 mm for F. indica. The very spinous spore

drawing in Stephani’s (1985) leones no. 003006 is total-

ly unlike the low, incomplete and irregularly areolate

spore markings of the type specimen of F. indica (Figure

4A-C). A note on the leones reads ‘fehlt Original’ but the

meaning of this is not clear. Srivastava & Udar wrote

that, ‘ Fossombronia indica St. (M.S.) Lucknow

University Hepatic Herbarium (a part of the materials of

the original collection presented to Prof. S.K. Pande by

Rev. P. Pfleiderer) India Orientalis, Mangalore. Legit.

Pfleiderer (part of type?) Det.: Stephani 1917. F. indica

was instituted by Stephani (1917) on the basis of the

materials collected by Pfleiderer from Mangalore, South

India. It has neither been collected nor investigated

again, after Stephani’s (1917) original publication’.

The spores of F. indica as measured by Stephani are

36 pm, although he had written on the leones, beneath

the spore drawing, 45 pm. Srivastava & Udar report

them to be ‘38.4-48.0 pm in diameter’, but I found them

to be 30.0-37.5 pm and the poorly developed lamellae

rather low, irregular and interrupted. The dimensions of

the ± oblong leaves are 750-875 x 625-875 pm, also

rather smaller than those of F. occidento-africana.

Until more material of F. indica is collected and stud-

ied, I think that it is advisable to reject the synonymy of

FIGURE 4. — Fossombronia initial Spores. A, distal face; B, side view of distal face; C, proximal face. A-C, Pfleiderer s.n. A, x 1078; B, x 11 89;

C, x 1147.

Bothalia 28,2 (1998)

187

F. occidento-africana under F. indica and to reinstate the

much better known F. occidento-africana S.W.Arnell.

SPECIMENS EXAMINED

Amell 2409 , 2431 S, 2448 PRE, 2497 (lectotype) S.

Jones 854 PRE.

Pfleiderer s.n. C(16585).

ACKNOWLEDGEMENTS

I wish to sincerely thank the curators of G and S as

well as Prof. R. Stotler, ABSH, for the loan of specimens.

Prof. Stotler is also thanked for checking the collection

numbers on four Arnell specimens. My thanks to Ms D.

Maree for typing the manuscript, Mrs M. Steyn for the

drawings and Mrs A. Romanowski for developing and

printing the photographs.

REFERENCES

ARNELL, S.W. 1952. Hepaticae collected in South and West Africa

(1951). New and little known species. Botaniska Notiser 105:

307-329.

AUGIER, J. 1972 Groupements de bryophytes terricoles sur le campus

universitaire de Yaounde. Annates de la Faculte des Sciience

Universitaire fed. Cameroun 9: 73-85.

AUGIER, J. 1985. Cles de determination pour les Hepatiques et

Anthocerotes au Cameroun forestier. Contribution a la Flore du

Cameroun (Cryptogamie) — essai destine aux etudiants et aux

chercheurs en Rue et regions voisines W — et centre-africaines.

JONES, E.W. 1985. Bryophytes of forest and savanna in northern

Nigeria. Cryptogamie, Bryologie & Lichenologie 6: 259-277 .

JONES, E.W. & HARRINGTON, A.J 1983. The hepatics of Sierra

Leone and Ghana. Bulletin of the British Museum of Natural

History (Botany) 11: 215-289.

PATON, J.A. 1973. Taxonomic studies in the genus Fossombronia

Raddi. Journal of Bryology 7: 243-252.

SCOTT, G.A.M. & PIKE, D.C. 1987. The Fossombronia foveolata

complex. Lindbergia 13: 79-84.

SCOTT, G.A.M. & PIKE, D.C. 1988. Revisionary notes on Fossom-

bronia. The Bryologist 91: 193-201.

SRIVASTAVA, S.C. & UDAR, R. 1975. The genus Fossombronia

Raddi in India. Nova Hedwigia 26: 790-845.

STEPHANI, F. 1917. Species hepaticarum. Bulletin de V Herbier

Boissier 6: 73.

STEPHANI, F. 1985. leones hepaticarum. Microfiche. Inter

Documentation Company bv, Leiden.

TIXIER, P. 1989. Bryophyta exotica — 8. Recoltes de J-F. Brunei au

Togo (1983-1985). Candollea 44: 493-511.

S.M. PER OLD*

* National Botanical Institute, Private Bag XI 01, Pretoria 0001.

MS. received: 1998-04-15.

NEW RECORDS FROM KWAZULU-NATAL, SOUTH AFRICA

INTRODUCTION

Eastern coastal links between the flora of southern

Africa and tropical East Africa are well established but

authors have concentrated on arborescent taxa (White

1965, 1971; Moll & White 1978). The southern exten-

sion of tropical species on the eastern coast of Africa is

largely the result of the ameliorating effects of the

Mozambique Current. Moll & White (1978) briefly out-

lined the vegetation of the Indian Ocean Coastal belt and

drew attention to the strong links which exist between

Tongaland-Pondoland and Zanzibar-Inhambane woody

floras. Recently Microcoelia obovata Summerh. (Orchi-

daceae) and Pseuderanthemum hildebrandtii Lindau

(Acanthaceae) were collected in northern KwaZulu-Natal.

These records further reflect the vegetation links between

the northern areas of KwaZulu-Natal and tropical East

Africa. The new records also highlight the importance of

amateur botanists, such as Mr E. Harrison, in the further

discovery of our flora.

ACANTHACEAE

1. Pseuderanthemum hildebrandtii Lindau in

Botanischer Jahrbiicher 20: 39 (1895a); Lindau: 330

(1895b); C.B.Clarke: 1 72 ( 1 899); Milne-Redh.: 260 (1936).

Type: Tanzania, Zanzibar, at Kidoti, Hildebrandt 981 (syn.,

designated by Milne-Redhead l.c. as ‘type’).

The genus Pseuderanthemum Radik, includes about

120 species and has a pantropical distribution (Dyer

1975). The genus was created to include species of the

tribe Justiceae which had formerly been placed in

Eranthemum (Ruellieae), Eranthemum sensu stricto does

not occur in Africa (Milne-Redhead 1936). Pseuderan-

themum is most closely related to Ruspolia, the latter

being distinguished by its monothecous anthers (Figure

5). In general the African species of Pseuderanthemum

occur in closed forests and produce long, narrow corollas

which are usually white or insipidly coloured. Formerly

only one species was recorded from South Africa, name-

ly P. subviscosum (C.B.Clarke) Stapf which occurs as a

herb of subtropical forest and is characterised by its long,

narrowly tubular white flowers. P. hildebrandtii (Figure

5) diverges from this typical pattern in producing brick-

red flowers very similar to those of Ruspolia hypocrater-

iformis (Vahl) Milne-Redh. from which it is distin-

guished by its bithecous anthers, slender habit and small-

er leaves. The floral similarity to R. hypocrateriformis

transcends colour to the extent that Clarke (1899) omit-

ted a detailed description of floral dimensions in his

description of Eranthemum hildebrandtii C.B.Clarke (-

Pseuderanthemum hildebrandtii) substituting ‘otherwise

as E. hypocrateriforme Roem. & Schultes’. This uncanny

floral convergence correlates with a similarity in habitat for

the two species and probably indicates shared pollinators.

P. hildebrandtii occurs in woodland and along forest mar-

gins (not on forest floors like P. subviscosum)-, this differ-

ence in habitat is likely to be associated with different pol-

linators and different floral pigments.

The sculpturing of the seeds of P. hildebrandtii is also

atypical for Pseuderanthemum in that the inner and outer

seed surfaces have different patterns (Figure 6), a trend

common in Ruspolia.

Although the distributions of other African species of

Pseuderanthemum are patchy, no major disjunctions

occur. By contrast, the collection of P. hildebrandtii in

South Africa, produces a disjunction which mirrors that of

188

Bothalia 28,2 (1998)

FIGURE 5. — A, C, D, Pseuderanthemum hildebrandtii, Harrison 1041: A, flowering branch, x 0.8; C, anthers, x 47; D, calyces, x 8. B, Ruspolia

hypocrateriformis, S Venter 12915, monothecous anther, x 31

Ruspolia hypocrateriformis . The main concentration of P.

hildebrandtii occurs in eastern Tanzania and eastern and

central Kenya between 0° and 10° south of the equator.

The main populations of Ruspolia hypocrateriformis tra-

verse similar but wider latitudes on the West Coast from

Senegal in the north, to northern Angola in the south

(Figure 7). Both species then display large disjunctions to

their southern outliers. The southern populations of P.

Bothalia 28,2 (1998)

189

FIGURE 6. — Seed of Pseuderanthe-

mum hildebrandtii, Harrison

1041 . A, outer surface; B, inner

surface. Scale bars: 2 mm

hildebrandtii occur in the Lebombo Mountains of

KwaZulu-Natal, some 2 400 km to the south. Likewise

the southern variety of R. hypocrateriformis is recorded

from Northern Province (Arnold & De Wet 1993) dis-

playing a disjunction of some 2 300 km. The Acanthaceae

are characterised by explosive capsules with elastic funi-

cles (jaculators) which are effective short-distance dis-

tributors of seed and it is difficult to conceive that the

species are capable of long-distance dispersal. From their

similar flowers, dispersal capabilities and habitat, it

seems likely that similar historical events may have led to

the vicarious distributions these two allies display.

KWAZULU-NATAL. — 2731 (Louwsburg): between Ingwavuma and

Jozini Dam, (-AA), Harrison 1041 (NU, PRE).

ORCHIDACEAE

2. Microcoelia obovata Summerh. in Botanical

museum leaflets 11: 253 (1945); Piers: 276 (1968);

FIGURE 7. — Distribution of Ruspolia hypocrateriformis, •; Pseude-

ranthemum hildebrandtii, ■

Jonsson: 83 (1981). Type: Tanzania, Lushoto, Lwengera

Valley, Moreau 445a (K, holo.).

Microcoelia Lindl. includes about 27 species and is

limited in distribution to Madagascar and tropical and

southern Africa (Cribb 1989). Currently this leafless

genus of epiphytes is represented by a single species, M.

exilis Lindl. in South Africa. This species is widespread in

Uganda, Kenya, Tanzania, Zaire, Zambia, Malawi,

Mozambique, Zimbabwe, South Africa and Madagascar.

Unlike the distributional patterns of African Pseuderan-

themum species, disjuncts are common in Microcoelia.

Species such as M. macrorrhynchia (Schltr.) Summerh.,

M. konduensis (De Wild.) Summerh. and M. microglossa

Summerh. are scattered patchily through tropical Africa,

whereas M. koehleri (Schltr.) Summerh. displays wide

tropical disjunctions. In fact one species, M. physophora

(Rchb.f.) Summerh., traverses the ocean barrier between

mainland Africa and Madagascar (Jonsson 1981). These

patterns are probably the product of seed vagility.

190

Bothalia 28,2 (1998)

M. obovata was recorded from Kenya, Tanzania and

northern Mozambique in a recent revision (Jonsson

1981). The South African collection was made at Lake

Sibaya some 1 440 km to the south of the southernmost

point recorded in Jonsson (l.c.) (Figure 8). In South Africa

the species appears to be restricted to sparse populations

in fairly dense coastal forest at Lake Sibaya, Veld Type

no. 1, Coastal Thornveld & Forest (Acocks 1988).

M. obovata is distinguished by its semicircularly in-

curved spur, (2.0-)3.5-5.6 mm long and conspicuously

elongate labellum, (4.9-)7.6(-9.0) x 3. 1-6.0 mm, which

exceeds the remaining perianth parts by a third to half

their length.

KWAZULU-NATAL.— 2732 (Ubombo): Lake Sibayi, (-BC),

Harrison & Rolfe 1044 (NU).

ACKNOWLEDGEMENTS

The Natal University Research Fund is thanked for

their financial assistance and the National Herbarium

(PRE) is thanked for the loan of specimens. The keeper

of the Herbarium at the Royal Botanic Gardens, Kew (K)

is thanked for access to their collections of P. hilde-

brandtii.

REFERENCES

ACOCKS, J PH. 1988. Veld Types of South Africa, 3rd edn. Memoirs

of the Botanical Survey of South Africa No. 57.

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.

CLARKE, C.B. 1899. Acanthaceae. In W.T. Thiselton-Dyer, Flora of

tropical Africa 25: 172.

CRIBB, P. 1989. Orchidaceae. In R.H. Polhill, Flora of tropical East

Africa: 511-521.

DYER, R.A. 1975. The genera of South African flowering plants, Vol.

2. Government Printer, Pretoria.

JONSSON, L. 1981. A revision of the genus Microcoelia (Orchi-

daceae). Symbolae botanicae upsalienses 23,4: 1-150.

LINDAU, G. 1895a. Acanthaceae africana II. Botanischer Jahrbiicher

20: 39.

LINDAU, G. 1895b. Acanthaceae. Die natiirlichen Pflanzenfamilien 4,

3B: 330.

MILNE-REDHEAD, E. 1936. Eranthemum of the ‘Flora of tropical

Africa’. Kew Bulletin: 255-274.

MOLL, E. & WHITE, F. 1978. The Indian Ocean Coastal Belt. In

M.J.A. Werger, Biogeography and ecology of southern Africa 1 :

561-598. Junk, The Hague.

PIERS, F. 1968. Orchids of East Africa, 2nd edn. Germany.

SUMMERHAYES, V.S. 1945. African orchids XV. Botanical museum

leaflets 1 1 : 249-260.

WHITE, F. 1965. The savanna woodlands of the Zambesian and

Sudanian Domains. An ecological and phytogeographical com-

parison. Webbia 19: 651-681.

WHITE, F 1971. The taxonomic and ecological basis of chorology.

Mitteilungen aus der Botanischen Staatssammlung, Miinchen

10: 91-112.

T.J. EDWARDS* and E. HARRISON**

* Botany Department, University of Natal, Private Bag X01, 3209

Scottsville.

** P.O.Box 104, 3935 Mtubatuba.

MS. received: 1998-01-05.

CYPERACEAE

COLEOCHLOA SETIFERA NEW TO THE FLORA OF KWAZULU-NATAL

Recently, Niels Jacobsen conducted a botanical sur-

vey on and near the site of a proposed new dam in the

Vryheid District of KwaZulu-Natal. The area is botani-

cally almost unexplored, as it has a very rugged terrain.

In February 1997 Coleochloa setifera (Ridl.) Gilly was

collected, representing yet another addition to Ross’s

Flora of Natal and Gordon-Gray’s Cyperaceae in Natal

(1995). For workers who are unfamiliar with the plant,

a brief description (based partly on Nelmes 1954) fol-

lows.

Coleochloa setifera (Ridl.) Gilly in Brittonia 5:14

(1943); Nelmes: 378 (1954); Compton: 73 (1976);

Haines & Lye: 363 (1983); Forbes: 39 (1987); Kativu: 36

(1994); Lebrun & Stork: 173 (1995). Syntypes:

Madagascar, Deans Cowan s.n.\ Hilsenberg & Bojer s.n.

Fintelmannia setifera Ridl: 337 (1883). Eriospora setifera (Ridl.)

C.B. Clarke: 676 (1894); Chermezon: 270 (1937).

Eriospora rehmanniana C.B.Clarke: 297 (1898); Schdnland: 65

(1922). Coleochloa rehmanniana (C.B. Clarke) Gilly: 14 (1943). Type:

Northern Province, Houtbosch, Rehmann 5624 (K, holo. [PRE, frag-

ment]).

Mat-forming perennial up to 400 mm tall, leaves and

culms sparsely to densely hairy. Shoots densely tufted,

intravaginal. Leaves 2-ranked, striate; sheaths open,

lower glossy reddish brown, upper reddish brown at

extreme base, otherwise green; ligule comprising a line

of hairs; blades convolute, ± 1 mm diam., upper surface

glabrous, lower sparsely to densely villous, ultimately

blades deciduous. Culm striate, rounded, ±1.5 mm diam.

Inflorescence a slender, 1-3-noded, suberect panicle.

Spikelets in rounded pedicelled clusters 4-5 mm long;

glumes chartaceous, pale to reddish brown, distichous, 4

or 5 per spikelet, upper 2 or 3 fertile, sometimes male

only, or female below and male above, uppermost some-

times empty. Style becoming pyramidal, hollow; stigmas

3. Nutlet whitish, 3-4 mm long, with pericarp not fused

to endosperm (thus resembling perigynium in Carex),

subtended by a cupule of whitish hairs less than, to

equalling length of nutlet. Flowering and fruiting mostly

mid- to late summer (October to March). Figure 9.

KWAZULU-NATAL. — 2731 (Louwsburg): Vryheid Dist., Farm

Dipka 590, ± 23 km north of Coronation on Manzana River, 720 m,

(-CA), Jacobsen 5484 (PRE).

This new record for KwaZulu-Natal is not altogether

unexpected, as C. setifera is quite widespread in hard

rocky outcrops (composed of e.g. quartzite, granite or

norite) in Northern Province, North-West, Gauteng,

Mpumalanga and Swaziland (Figure 10). It also occurs

Bothalia 28,2 (1998)

191

FIGURE 9. — Coleuchloa setifera. Habit, x 0.3. Photograph by A.

Romanowski.

in eastern tropical Africa (Mozambique, Zimbabwe,

Zambia, Malawi. Kenya, Tanzania) and on Madagascar.

In KwaZulu-Natal the species was observed growing in

crevices in exposed quartzite on a steep east-facing slope

above the Manzana River on Dipka Farm. It was also

observed in the vegetative state on the neighbouring No.

1 of Skurwerand 277 Farm, Paulpietersburg District.

The genus is confined to Africa and Madagascar, rep-

resented by s^ven species and one variety distributed in

FIGURE 10. — Distribution of Coleochloa Gilly in southern Africa (speci-

mens in PRE). C. setifera, •; C. pallidior , ■

the eastern half of tropical Africa. The other southern

Africa species, C. pallidior , occurs on the Soutpansberg

in Northern Province (Figure 10), and nearby in southern

Zimbabwe.

C. setifera has some economic value, for example in

Mpumalanga the wiry leaves and culms are used to make

rope (cf. Van der Schijff729, PRE) and in Swaziland they

are woven into mats (cf. Compton 26119\ 30861, both

PRE).

There are additional reasons why the species merits

further study:

1 . Taxonomic confusion

The unique loosely attached pericarp was initially

interpreted as a perigynium; therefore Coleochloa and

related genera were once classified in the Tribe Cariceae

Kunth ex Dumort. (Clarke 1897/1898). More recently

they have been classified in other tribes including

Sclerieae Kunth ex Fenzl (Clarke 1908; Raynal 1963),

and Lagenocarpeae Gilly (Gilly 1943) but have finally

been placed in their own small tribe, Trilepideae Goetgh.

(Goetghebeur 1985), which comprises Coleochloa Gilly,

the South American Trilepis Nees and the West African

Afrotrilepis (Gilly) J. Raynal and Microdracoides Hua.

At species level, there is some doubt as to the distinc-

tion between C. setifera and its geographically nearest

neighbour, C. pallidior Nelmes. Nelmes (1954) disting-

uished them by the latter species having, inter alia,

loosely tufted extravaginal shoots, glabrous leaves and

culms, leaves 3-4 mm wide, and nutlet 3. 0-3. 5 mm long,

subtended by a cupule of hairs less than half the length

of the nutlet. Browning & Gordon-Gray (1992) and

Kativu (1994), however, suggest that the differences are

not nearly as clear-cut as indicated by Nelmes and that

there is a need for extensive fieldwork. At present in the

National Herbarium the two species are retained as sepa-

rate entities.

2. Ecophysiology

Members of the tribe colonise habitats which experi-

ence periods of extreme desiccation. The other three

genera produce aerial stem-like structures composed of

the remains of leaf bases intergrown with thick adventi-

tious roots, similar to the ‘stems’ in Xerophyta Juss.

(Velloziaceae). This stem-like development is not very

marked in Coleochloa itself, however. C. setifera, stud-

ied by the author in several field localities and in the

herbarium, was seen to form dense mats of leaf bases and

roots, filling the cracks in rocks. It has been shown

(Porembski & Barthlott 1995), that these roots possess a

velamen which is able to absorb water rapidly, thus the

plant is able to capitalise on brief showers of rain.

REFERENCES

BROWNING, J. & GORDON-GRAY, K.D. 1992. The genus Coleochloa

Gilly. Notulae cyperologicae 8. Cyperaceae Newsletter 1 1: 7-9.

CHERMEZON, H. 1937. 29e famille — Cyperacees. In H. Humbert,

Flore de Madagascar. 270. Tananarive.

CLARKE, C.B 1894. Eriospora setifera. In T.A. Durand & H Schinz,

Conspectus florae Africae 5: 676. Brussels.

192

Bothalia 28,2 (1998)

CLARKE, C.B 1897/1898. Cyperaceae. In W.T. Thiselton-Dyer, Flora

capensis 7: 149-310. Reeve, London.

CLARKE, C.B 1908. New genera and species of Cyperaceae. Kew Bul-

letin, add. sen 8.

COMPTON, R H 1976. The Flora of Swaziland. Journal of South

African Botany. Suppl. Vol. 1 1

FORBES, P.L. 1987. Cyperaceae. In T.K. Lowrey & S. Wright, The

Flora of the Witwatersrand Vol. I The Monocotyledonae:

29-64. Witwatersrand University Press, Johannesburg.

GILLY, C.L. 1943. An Afro-South American Cyperaceous complex. Brit-

tonia 5: 1-20.

GOETGHEBEUR, P. 1985. Studies in Cyperaceae 6. Nomenclature of

the suprageneric taxa in the Cyperaceae. Taxon 34: 617-632.

GORDON-GRAY, K.D. 1972. Cyperaceae. In J.H. Ross, Flora of Natal.

Memoirs of the Botanical Survey of South Africa No. 39: 99-

103.

GORDON-GRAY, K.D. 1995. Cyperaceae in Natal. Strelitzia 2.

HAINES, R.W & LYE, K.A. 1983. The sedges and rushes of East

Africa'. 363. East African Natural History Society, Nairobi.

KATIVU, S. 1994. Notes on Coleochloa Gilly (Cyperaceae) in the Flora

z.ambesiaca area. Kirkia 15: 33-37.

LEBRUN, J.-P. & STORK, A.L. 1995. Enumeration des pluntes a

fleurs d'Afrique tropicale Vol. Ill — Monoctyledones: Limno-

charitaceae a Poaceae. Conservatoire et Jardin botanique de la

Ville de Geneve, Geneve.

NELMES, E. 1954. Notes on Cyperaceae: XXXI. The African genus

Coleochloa. Kew Bulletin 1953: 373-381.

POREMBSKI, S. & BARTHLOTT, W. 1995. On the occurrence of a

velamen radicum in Cyperaceae and Velloziaceae. Nordic Jour-

nal of Botany 15: 625-629.

RAYNAL, J. 1963. Notes cyperologiques. I . Afrotrilepis , nouveau genre

africain. Adansonia , ser. 2, 3: 250-265.

RIDLEY, H.N. 1883. Descriptions and notes on new or rare

Monocotyledones plants from Madagascar, with one from

Angola. Journal of the Linnean Society Botany 20: 337.

SCHONLAND, S. 1922. Introduction to South African Cyperaceae.

Memoirs of the Botanical Survey of South Africa No. 3.

Government Printer, Pretoria.

C. ARCHER*

* National Botanical Institute, Private Bag X 1 0 1 , 0001 Pretoria.

MS. received: 1997-04-23.

ASTERACEAE

KEY TO THE SPECIES OF THE GENUS HERTIA IN SOUTHERN AFRICA

New combinations of the species of Hertia Less, in

southern Africa were made in different books or journals:

H. alata (Thunb.) Kuntze (Kuntze 1891), H. ciliata

(Harv.) Kuntze (Kuntze 1891; Merxmiiller 1967), H.

cluytiifolia (DC.) Kuntze (Kuntze 1891), H. kraussii

(Sch.Bip.) Fourc. (Fourcade 1941), H. pallens (DC.)

Kuntze (Kuntze 1891; Merxmiiller 1967). A key to only

two of the species ( H . ciliata and H. pallens) is given in

Merxmiiller (1967). A practical key is presented here to

distinguish between the southern African species for use

in the herbarium:

1 a Heads radiate H. pallens

lb Heads disciform:

2a Heads 5-20 per corymb; corymbs numerous H. alata

2b Heads solitary, terminal on branches or in leaf axils:

3a Pedicels 2-8 x longer than leaves H. kraussii

3b Pedicels shorter, ± half to one and a half times longer

than leaves:

4a Leaves obovate-elliptic, obtuse, entire, drying pale . . .

H. cluytiifolia

4b Leaves cuneate-oblong, 3-5-toothed, drying dark . . .//. ciliata

REFERENCES

FOURCADE, H.G. 1941. Checklist of the flowering plants of the divi-

sions of George, Knysna, Humansdorp and Uniondale. Memoirs

of the Botanical Survey of South Africa No. 20: 27.

KUNTZE, O. 1891. Dicotyledones. 88. Compositae. Revisio generum

plantarum 1 : 344.

MERXMULLER, H. 1967. Asteraceae. Prodromus einer Flora von

Siidwestafrika 139: 99.

P.P.J. HERMAN*

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

MS. received: 1998-04-09.

AMARYLLIDACAE: AMARYLLIDEAE

A NEW SPECIES OF AMARYLLIS FROM THE RICHTERSVELD, SOUTH AFRICA

INTRODUCTION

With a taxonomic history of just over 350 years, the

monotypic genus Amaryllis L. is considered to be one of

South Africa’s most important and widely grown orna-

mental taxa (Ferrari 1633; Dyer 1955). Amaryllis bel-

ladonna L. is currently cultivated in many countries

throughout the world. In the wild, however, the species is

well documented as endemic to the Cape Floristic

Region, where it is restricted to the Western Cape (Bond

& Goldblatt 1984). The recent discovery of a large pop-

ulation of Amaryllis plants in the Richtersveld, a particu-

larly arid region of the Succulent Karoo Biome in the

remote northwestern Cape, consequently engendered a

great deal of surprise and focused attention on the need

for a taxonomic reassessment of the genus.

Leafing bulbs of the Richtersveld population were

first collected at Paradyskloof by Prof. N. du Plessis and

Prof. G. Delpierre in 1978, by Mr E.J. van Jaarsveld in

1993, and by the first author in 1996. The cultivated col-

lections maintained in the Kirstenbosch National Botanic

Garden nursery, however, have thus far failed to flower.

Flowers were first recorded in the wild by Mr J.

Domroch, a Richtersveld Park ranger, who presented the

second author with a single pressed flower in April 1996.

This collection was added to one year later when the sec-

ond author, while completing a floristic survey of the

Richtersveld National Park, found the population at

Paradyskloof flowering en masse. From the beautiful

pink dowers the plants were tentatively identified as an

Amaryllis , and this was later confirmed by the appear-

ance of the fruits. A systematic study of the population

Bothalia 28,2 ( 1998)

193

thereafter revealed several morphological features that

separate the geographically disjunt Richters veld plants

from A. belladonna. The plants are thus newly described

as A. paradisicola, an epithet which commemorates the

species’ place of origin.

The discovery of this new species marks the first

record of a wild Amaryllis outside the Cape Floristic

Region and adds an extra, exceptionally ornamental rep-

resentative to the 363 taxa already known as unique to

the Gariep Centre of endemism (Nordenstam 1969;

Hilton-Taylor 1996).

Amaryllis paradisicola Snijman , sp. nov., A. bella-

donnae L. affinis sed foliis latioribus, 70-130 x 350-500

mm; pilis brevibus patentibus in superficiebus ambabus

foliorum saltern ubi juvenibus; floribus roseis concol-

oris; staminibus interioribus manifeste longioribus quam

exterioribus; stigmate profunde trifido ramis gracilibus

patentibus maturitate usque ad 1.5-2. 5 mm longis.

Figurae 11 & 12.

TYPE. — Northern Cape, 2817 (Vioolsdrif): Richters-

veld National Park, Paradyskloof, on rocky slopes,

(-AC), 11-4-1997, Snijman 1576 (NBG, holo.; K, KMG,

PRE).

Deciduous bulbous herbs, up to 800 mm tall when

flowering and fruiting. Bulbs solitary or sometimes clus-

tered, large, ovoidal, ± 100 x 140 mm. without a narrow

neck, sometimes more than half exposed above ground;

tunics numerous, cream-coloured, papery, producing

extensible threads when tom. Leaves 6-9, hysteranthous,

distichous, suberect when emerging, spreading and

apparently rosulate when mature; blade ligulate when

young to Ungulate when mature, up to 70-130 x

350-500 mm, somewhat channelled, more or less thin-

textured, bright green, covered with short patent hairs on

both surfaces at least while developing, producing copi-

ous brown sap when cut and numerous extensible threads

when torn; abaxial surface with a prominent midrib; mar-

gins sometimes slightly thickened, reddened and undu-

late, at least when young. Inflorescence upright; scape

solid, stout, succulent, 20-40 x 480-800 mm, somewhat

compressed, slightly ancipitous, glaucous-green, reddish

brown proximally, smooth, withering after seed disper-

sal; spathe valves 2, papery and reflexed at anthesis,

20-25 x 50-55 mm; bracteoles numerous, flattened, up

to 2 x 50 mm. Flowers 10-21, trumpet-shaped, with fun-

nel 30-40 mm long, 15-20 mm wide at throat, 70-80

mm wide at rim, in an evenly spreading, umbel-like clus-

ter, uniformly pink, without contrasting colours in veins

or throat, turning darker pink with age, scented; pedicels

evenly spreading, firm, 3-5 x 30-70 mm at anthesis,

elongating to 250 mm in fruit, mostly subterete but

slightly triangular in cross section below developing

fruit. Tepals free, broadly lanceolate, 12-20 x 65-80

mm, slightly recurved apically. Stamens declinate, short-

er than tepals, biseriate; filaments stout, pale pink, basal-

ly fused and adnate to perigone for ± 7 mm; outer whorl

FIGURE 1 1 . — Amaryllis paradisico-

la: A, flowering plants in

habitat; B, C, scape and

flower head; D, flower head

showing lateral view of

exserted trifid stigma.

194

Bothalia 28,2 (1998)

FIGURE 12. — Amaryllis paradisico-

la: A, mature fruiting head; B,

bulb in habitat during fruiting

showing scape, young leaves

and basal bulblet; C, plants in

habitat showing mature hys-

teranthous leaves.

± 35 mm long; inner whorl ± 50 mm long, basally adnate

to perigone for up to ± 5 mm beyond tube; anthers ver-

satile, dorsifixed, curved, 7-9 mm long, yellow; pollen

yellow. Ovary green, trilocular, 5 mm across; ovules 4-6

per locule, axile; style declinate, ± 65 mm long, longer

than stamens, shorter than tepals, pale pink; stigma trifid

with spreading, slender branches up to 1.5-2. 5 mm long.

Capsule loculicidal, subglobose, 20-30 mm across,

leathery. Seeds fleshy, more or less ovoid, 12-17 mm

across, glossy, whitish to pink; embryo green.

Flowering, fruiting and vegetative phenology

The bulbs bloom in early April and the flowering peri-

od of the population lasts approximately ten days. When

A. paradisicola was collected in 1997, several hundred

bulbs had flowered after rain fell in March that year but

only two bulbs flowered at the same site during a pro-

longed dry autumn the following year. Flowering is thus

profuse after good autumn rain but poor when precipita-

tion in autumn is low. Despite the profusion of flowers in

1997, only a small number set seed and many infructes-

cences had been damaged, possibly by baboons. The

seeds take approximately one month to be released and

germinate after another three to four weeks. Although the

young leaves sometimes emerge at flowering time, the

foliage is mostly absent until the start of the winter rain-

fall season. The leaves grow rapidly in May and persist

until they dry off with the onset of the summer drought,

usually at the end of September.

Diagnostic features

Amaryllis paradisicola differs from A. belladonna in

having broad, tongue-shaped leaves, 70-130 x 350-500

mm, covered with short, patent hairs on both surfaces, at

least during their development. Although the hairs

become increasingly sparsely spaced as the leaf blades

enlarge, they remain evident under a hand lens in most

plants and are only rarely absent at maturity. Studies in

the field and of the herbarium material at NBG and BOL

indicated that populations of A. belladonna in the south-

ern Cape, the Agulhas Plain and the Cape Peninsula have

widely spreading, narrow leaves, 13-26 x 270-680 mm,

whereas the western populations from Saldanha and the

Cederberg have somewhat upright, slightly broader

leaves, 36-73 x 300-675 mm, that arise from an elon-

gated basal sheath, 110-300 mm long. The leaves of A.

belladonna are therefore perceptibly longer and narrow-

er than those of A. paradisicola while the leaf surfaces

are consistently glabrous, even when young.

Leaf surface pubescence in the form of soft hairs is

rare in the family. Softly pubescent leaves is a derived

character state in Amaryllideae and is so far known only

in Hessea Herb, and Strumaria Jacq. (subtribe Amaryl-

Bothalia 28,2 (1998)

195

lidinae) but must now be considered for Amaryllis , the

basal genus in subtribe Amaryllidinae (Snijman & Lin-

der 1996). Similar leaf surface pubescence also occurs

sporadically in Haemanthus L. and Gethyllis L. (tribe

Haemantheae sensu Dahlgren et al. 1985) and Pancra-

tium L. (tribe Pancratieae).

Amaryllis paradisicola is furthermore distinguished

by several floral characters, of which flower colour and

stigma shape are most diagnostic. The 10-21 -flowered

A. paradisicola is slightly more floriferous than A. bel-

ladonna, which has 6-12 or occasionally 21 flowers, per

inflorescence; and the tepals, although darkening with

age, are uniformly pink, without the cream to lemon-yel-

low perigone throat of A. belladonna. The inner stamens

relative to the outer stamens vary from 10 mm longer to

almost equally long in A. belladonna , whereas the inner

stamens of the new species exceed the outer stamens by

15 mm or more. Lastly, the mature stigma of A. paradis-

icola is distinctly trifid with slender, spreading branches

that reach up to 1.5-2. 5 mm long when mature, unlike

the broader (up to 2 mm across), minutely trifid stigma

of A. belladonna. Although not as strongly scented, the

flowers of both species smell somewhat like those of

Narcissus.

Distribution, habitat and phytogeography

Amaryllis paradisicola is endemic to the Richtersveld

National Park, an arid, mountainous region in the North-

ern Cape. Here the species is known from a single large

population in Paradyskloof, a narrow gorge overlooking

a perennial pool along a tributary of the Abiekwa River

between Hottentotsparadys and Paradysberg. The bulbs

grow on south and east-facing, quartzitic cliffs, on nar-

row rock ledges and partially vegetated screes that are

shaded for much of the day. If the locality is not patrol-

led, the population may be endangered by unscrupulous

bulb collectors and by the large numbers of goats that

gather to drink in Paradyskloof.

The discovery of A. paradisicola extends the distrib-

ution of the genus Amaryllis approximately 450 km

beyond the Olifants River Valley mountains in the

Western Cape, where the most northerly populations of

A. belladonna are thus far known (Figure 13).

Other endemics of the Gariep Centre that occur at

Paradyskloof include three geophytes: Cyrtanthus herrei

(Leighton) R.A.Dyer, Bulbine pendens G.Will. & Baij-

nath, and an as yet unnamed Spiloxene species; also the

Paradise Toad, Bufo robinsonii Branch & Braack (1995).

The highly disjunct distribution pattern in Amaryllis

is repeated in several other geophytic genera.

Cyrtanthus herrei is confined to the Gariep Centre

whereas C. carneus (Reid & Dyer 1984) is found only in

the southwestern Cape; furthermore Moraea garipensis

and M. namaquamontana are narrowly restricted in the

Gariep Centre and M. ramosissima occurs only in the

Cape (Goldblatt 1986). Since the species in all three

genera are putative sister taxa, they probably represent

vicariants of previously widespread ancestral taxa.

Methods are not yet available to determine the age of

these species. Speciation may date from the advent of

FIGURE 13. — Known distributions of Amaryllis paradisicola, ★; and

A. belladonna, •, based on collections at BOL, PRE and NBG.

summer aridity and winter rainfall conditions in the

Pliocene (Tankard & Rogers 1978) or from the great cli-

matic fluctuations of the Pleistocene (Deacon 1983;

Tyson 1986). In particular, preliminary palynological

data from spring deposits at Eksteenfontein in the

Richtersveld showed a marked change from predomi-

nantly Chenopodiaceae-Amaranthaceae and different

Asteraceae pollen in the Late Pleistocene (before 10 700

BP) to assemblages totally dominated by pollen of the

succulent Aizoaceae-Mesembryanthemaceae type in the

early Holocene, 10 700-8 450 years BP (Scott et al.

1995). A tentative interpretation of this data is that it

reflects change from cooler to warmer conditions (Scott

et al. 1997). If this is correct, the mountains from the

Richtersveld to the Cape may have provided suitable

habitats for a widespread ancestral taxon in Amaryllis at

least during the Late Pleistocene. However, when the

available moisture reached its present level in the early

Holocene, populations in the Cape Region were proba-

bly ecologically isolated from those in the Gariep Centre

and thus diverged. We hypothesise that A. belladonna

radiated widely in the southwestern Cape, where it

became adapted to frequent disturbance by fire, where-

as A. paradisicola became narrowly restricted to rela-

tively cool, moist montane habitats in the otherwise arid

Richtersveld, where it evolved xerophilous, pubescent

leaves.

NORTHERN CAPE. — 2817 (Vioolsdrif): Richtersveld National

Park, Paradyskloof, (-AC), Snijman 1550 (NBG), Snijman 1576 (K,

KMG, NBG, PRE), Snijman 1578 (K, KMG, NBG, PRE), Williamson

5904 (NBG).

ACKNOWLEDGEMENTS

The National Parks Board kindly granted permission

to collect specimens of A. paradisicola in the Rich-

tersveld National Park. Mrs F. Williamson assisted in the

field, Mr A. de Villiers provided information on seed ger-

mination, and Dr J.C. Paterson-Jones kindly supplied the

original colour slides for Figures 11 & 12.

196

Bothalia 28,2 (1998)

REFERENCES

BOND, P & GOLDBLATT, P. 1984. Plants of the Cape Flora. Journal

of South African Botany Suppl. Vol. 13: 1-455.

BRANCH, W R. & BRAACK, H.H. 1995. A new toad from paradise.

Madoqua 19: 15-23.

DAHLGREN, R.M.T., CLIFFORD, H.T. & YEO, PF. 1985. The fami-

lies of the monocotyledons. Springer- Verlag, Berlin.

DEACON, H.J. 1983. Another look at the Pleistocene climates of

South Africa. South African Journal of Science 79: 325-328.

DYER, R.A. 1955. Amaryllis belladonna. The Flowering Plants of Afri-

ca 30: t. 1200.

FERRARI, G. 1633. Deflorum cultura libri IV: t. 121. Rome.

GOLDBLATT, P 1986. The moraeas of southern Africa. Annals ofKir-

stenbosch Botanic Garden 14: 1-224.

HILTON-TAYLOR, C. 1996. Patterns and characteristics of the flora of

the Succulent Karoo Biome, southern Africa. In L.J.G. van der

Maesen, X.M. van der Burgt & J.M. van Medenbach de Rooy,

The biodiversity of African plants. Proceedings XIVth AETFAT

Congress. August 1994. Wageningen, Netherlands: 58-72. Kluwer

Academic Publishers, Dordrecht.

NORDENSTAM, B. 1969. Phytogeography of the genus Euryops (Com-

positae). A contribution to the phytogeography of southern

Africa. Opera Botanica 23: 1-77.

REID, C. & DYER, R.A. 1984. A review of the southern African

species of Cyrtanthus. American Plant Life Society, La Jolla,

California.

SCOTT, L„ ANDERSON, H.M. & ANDERSON, J.M 1997. Vegeta-

tion history. In R.M. Cowling, D.M. Richardson & S.M. Pierce,

Vegetation of southern Africa: 62-84. Cambridge University

Press, Cambridge.

SCOTT, L„ STEENKAMP, M. & BEAUMONT, PB. 1995. Palaeo-

environments in South Africa at the Pleistocene-Holocene tran-

sition. Quaternary Science Reviews 14: 937-947.

SNIJMAN, D A. & LINDER, H P 1996. Phylogenetic relationships,

seeds characters, and dispersal system evolution in Ama-

ryllideae (Amaryllidaceae). Annals of the Missouri Botanical

Garden 83: 362-386.

TANKARD, A.J. & ROGERS, J. 1978. Late Cenozoic palaeo-environ-

ments on the west coast of southern Africa. Journal of Bio-

geography 5: 319-337.

TYSON, P.D. 1986. Climatic change and variability in southern

Africa. Oxford University Press, Cape Town

D A. SNIJMAN* AND G. WILLIAMSON**

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

7735 Claremont, Cape Town.

** Research Associate, Bolus Herbarium, University of Cape Town,

7700 Rondebosch, Cape Town.

MS. received: 1998-05-27.

Bothalia 28,2: 197-211 (1998)

A reconnaissance survey of the vegetation of the North Luangwa

National Park, Zambia

P.P. SMITH*

Keywords: Luangwa Valley, vegetation description, vegetation mapping. Zambia

ABSTRACT

A comprehensive survey of the vegetation of the North Luangwa National Park (NLNP) was earned out over a period

of two years. The main aims of the survey were to describe the major vegetation communities in the park and to produce a

vegetation map of the NLNP

Initial differentiation of vegetation units was established by the appearance of the vegetation on aerial photographs.

Further information was derived from 353 ground plots in which > 20 000 woody plants were identified and measured.

Thirteen broad vegetation types were recognised in the NLNP. Details of their physiognomy, species composition, distrib-

ution, topography and edaphic associations are given.

INTRODUCTION

Until the present study, no detailed vegetation survey

of Zambia’s North Luangwa National Park (NLNP) had

been carried out. Naylor et al. (1973) and Phiri (1989)

have described the vegetation of the Luangwa valley, but

in both of these surveys, the area covered is too large and

the classification too broad to be useful to park man-

agers. The most detailed study of the region available is

that of Astle et al. (1969), who surveyed the South

Luangwa National Park, the North Luangwa National

Park east of the Muchinga Escarpment, and surrounding

areas. Habitat classification in this study was based on

physiognomic units recognised in a series of landsystems

representing all of the topographic units present in the

survey area.

The close relationship between vegetation, climate,

landsystems, edaphic factors is well established (Cole

1982; Bell 1984), and the rationale for producing a veg-

etation-based habitat classification of the NLNP was

founded on the premise that vegetation is a readily iden-

tifiable and measurable facet of habitat (Timberlake et al.

1993). In the long term, the vegetation boundaries desig-

nated in this study will form part of a geographical infor-

mation system for the NLNP, together with soil, geolog-

ical, hydrological, topographical and other information

which, combined, will help define habitats for individual

species. Justification for studying the vegetation compo-

nent of habitat as a first step is that a vegetation type is

readily discemable to managers and park personnel, peo-

ple who are not necessarily plant ecologists. Descriptions

based on vegetation physiognomy (e.g. woodland, wood-

ed grassland, thicket) and four or five characteristic

species should be recognisable to all.

As well as providing a valuable basis for habitat

description and monitoring, floristic data are a useful

measure of biological diversity. Conservation resources

* The Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey,

TW9 3AB, UK

MS. received: 1997-04-29.

are scarce in Zambia, as in most developing countries,

and as a signatory to the Convention on Biological

Diversity, Zambia is committed to conserving as much of

her biological resource as possible. Clearly, a rational

approach to conservation needs to be devised in which

limited financial and technical resources can be used to

maximum benefit. An essential first step is to carry out a

biological resource assessment (Article 7a of the

Convention) with the aim of identifying areas of high

biodiversity for priority protection. Zambia has set aside

63 585 km2 (8.5%) of its total land mass as national park

and a further 10% is designated as forest reserve. All of

these areas require protection, but only a fraction have

been subjected to biological inventory. Without such

basic inventory, monitoring (Article 7b), identification of

adverse processes (7c) and maintenance or management

(7d) of the biodiversity resource is impossible. Due to its

great topographical diversity, North Luangwa National

Park is potentially one of the most biologically diverse of

Zambia’s protected areas. The results of the present study

will help to confirm this and will form baseline data

against which future trends in plant diversity can be mea-

sured.

This paper describes the major vegetation types pre-

sent in the NLNP, and presents a vegetation map of the

park. A comprehensive checklist of the plants collected

during this study is published elsewhere (Smith 1998).

STUDY AREA

The North Luangwa National Park (NLNP) is the

most northerly national park in Zambia’s Luangwa

Valley (Figure 1). It covers an area of 4 636 km2

(between ll°25' S to 12°20' S and 31°45' E to 32°40' E)

and is situated entirely on the west bank of the Luangwa

River, which forms its eastern boundary. In the west, the

park boundary incorporates part of the Muchinga

Escarpment which constitutes approximately 24% (1113

km2) of the park’s area. The northern boundary of the

park is formed by the Lufila River, while in the south, the

NLNP is bordered by the Munyamadzi corridor.

198

Bothalia 28,2 (1998)

RIVER

HABITAT

A1/F2 Valley riverine complex

B1 Mixed alluvial thicket

B2 Combretum thicket

C1/C2 Escarpment miombo woodlands

JjJ C2 Hill miombo woodlands

| C2/D1 Woodland mosaic

D1 Combretum-Terminalia woodland

] D2/E/C2 Wooded grassland mosaic

I ] D2/E Woodland/wooded grassland mosaic

E Mopane woodland and scrub woodland

FI Secondary grassland on alluvium

F2 Valley riverine grassland

FIGURE 1. — The vegetation of the North Luangwa National Park, Zambia. Scale approx. 1 : 900 000.

Karoo System (Permian) sedimentary rocks form the

dominant strata of the valley floor while the escarpments

to the east and west of the Luangwa valley are made up

of igneous (e.g. granite) and/or metamorphic (e.g. gneiss

and quartzite) rocks (Utting 1976). In the NLNP, the ele-

vation of the Luangwa River and its adjacent floodplain

is ± 600 m. Topographical relief in this region is only a

few metres. Further west, but still on the valley floor, the

terrain becomes more dissected (relief 5-50 m) and the

elevation rises to between 700 and 800 m. In the far

western areas of the park the Muchinga Mountains rise

600-700 m above the valley floor, with the highest peaks

within the National Park reaching between 1 200 and

1 300 m. In this area the terrain is deeply dissected with

a relief of 500-600 m.

The valley receives a moderate rainfall of between

700 mm and 900 mm per annum, the wet season extend-

Bothalia 28,2 (1998)

199

ing from November through to April. At higher altitudes,

on the Muchinga Escarpment and plateau, rainfall is cor-

respondingly higher [mean for the plateau (Mpika),

1 065 mm].

No comprehensive study of the soils of the Luangwa

valley has yet been carried out. However, in their land

classification study of the South Luangwa National Park

(SLNP), Astle et al. (1969) described and mapped the

heterogeneous soil types of the park according to their

association with the vegetation. Soil types range from the

deep, red sandy loams of the upper escarpment to the

alluvial sands and clays of the Luangwa floodplain.

The NLNP forms part of the Zambezian Regional

Centre of Endemism (White 1983) and. due to its great

topographical diversity, it probably possesses a greater

range of habitats than any other park in Zambia. In the

higher (800-1 200 m) deeply dissected terrain of the

Muchinga Mountains, miombo woodland predominates,

while in the lower (600-700 m) nearly flat valley

regions, mopane woodland is the dominant vegetation

type. Fire (Naylor et al. 1973) and elephant damage

(Caughley 1976) have been the most influential factors

affecting vegetation in the NLNP in recent years.

METHODS

Site selection and data collection

The approach employed for this survey was essential-

ly inductive, with the primary aim being vegetation

description rather than analysis (Kent & Coker 1992). As

a first step, the vegetation of the park was stratified

according to its macroscopic appearance, and subse-

quently systematic ground survey was used to provide

descriptive detail.

Initial differentiation of vegetation units was carried

out using aerial photographs and aerial survey. In this

procedure, panchromatic aerial photographs of the

NLNP (1983, scale 1: 34 000) were used to delineate nat-

ural vegetation boundaries. Interpretation of the air-

photo mosaic was carried out in order to define areas

homogeneous in tone and texture, corresponding to

homogeneous vegetation types. Thus, physiognomic

units (woodland, bushland, grassland etc.) were differen-

tiated and marked on the aerial photographs. Air-photo

information was validated and augmented by aerial sur-

vey. Homogeneous areas of vegetation (and to a certain

extent, their composition) were identified by an observer

and positions were recorded using a Garmin inboard

Geographical Positioning System (GPS). A total of 116

points were recorded in a systematic transect survey and

a further 105 points were recorded on other aeroplane

and helicopter flights over the park.

In the second part of the survey, floristic and physiog-

nomic vegetation data were collected in ground studies.

Physiognomic classification followed White (1983)

(Table 1). A total of 353 belt transects were located with-

in the vegetation types defined in the air-photo mosaic.

Placement of transects was not random as an effort was

made to cover as many vegetation units (homogeneous

areas defined on the aerial photographs) as possible.

Other considerations were accessibility and avoidance of

ecotones and atypical landscape features e.g. termite

mounds, tracks, roads (Walker 1976). Transect positions

were determined using a portable GPS (Magellan 5000).

Standard transects measured 50 x 5 m (= 250 m2), rec-

tangular plots being chosen for ease of sampling and to

maximise species diversity (Brown 1954; Condit et al.

1996). Transects incorporated a minimum of 50 trees/

shrubs or at least 15 specimens of a dominant species.

Where these criteria were not met, transects were broad-

ened or lengthened accordingly (Taylor & Walker 1978).

Within each transect, woody plants (> 1 m in height) were

identified, counted and measured (diameter at breast

height and estimated height). Herbs and grasses were

collected and/or noted at all transect sites.

Samples of all plant species recorded were collected,

pressed and dried. Identifications were carried out in

herbaria in Lusaka, Kitwe, Harare and Kew. Nomenclature

follows Flora zambesiaca, Flora of tropical East Africa and

Lebrun & Stork (1991-1997). Voucher specimens are

lodged at the Kew, Missouri and Mount Makulu herbaria. A

full checklist of the plants collected in the NLNP is pub-

lished elsewhere (Smith 1998).

In addition to the vegetation survey, a limited soil sur-

vey of the NLNP was carried out and the resulting soil

types were related to the plant communities defined

above. In order to include the main vegetation types, soil

pits were dug beside a vehicle track which transects the

park, from Mano Game Scout Camp ( 11°37' S, 32°02' E)

on the Muchinga Escarpment in the west, to the Luangwa

River ( 1 1 °5 1 ' S, 32°26' E) in the east. Additional pits

were dug between 1 1 °37' S, 32°30' E and 1 1 °33' S,

32°25' E along a cutline parallel to the Lufila River. Soil

pits (2 m deep) were dug at ± 2 km intervals within 5-10

m of the track. Soil profiles were measured and pho-

tographed and a soil sample (200 g) from each horizon

was placed in a plastic bag and retained for further analy-

sis. The displaced soil was returned to the pit and the sur-

face levelled in an attempt to minimise disturbance to the

site. Retained soil samples were measured for colour,

texture and pH. Colour was determined using Munsell

Soil Colour Charts (1949). Soil texture (stoniness, shape

of peds, structure, consistency and roots) was described

according to Courtney & Trudgill (1988); sand, silt and

TABLE 1 . — Physiognomic classification of vegetation from White

(1983)

200

Bothalia 28,2 (1998)

clay content of the soil was determined using the

hydrometer method (Pramer & Schmidt 1964). Finally,

the pH of the soil was recorded using a Kel soil tester (for

acidity and soil moisture).

Data analysis and interpretation

Ground study provided detailed information on vege-

tation structure and composition. Thus, for each vegeta-

tion unit delineated from the aerial photographs, it was

possible to: 1, list plant species; 2, calculate tree/shrub

density, expressed as n ha'1 (Brown 1954); 3, calculate

woody biomass, expressed as t ha'1 using the following

formulae developed in similar habitats in Sengwa,

Zimbabwe by Guy (1981):

Tree biomass (Kg) = 0.0549 x (diameter at breast height)25101

Shrub biomass (Kg) = 1.2102 x (canopy volume)0-9118

4, determine dominant species by calculation of impor-

tance values (Curtis & McIntosh 1951) for each species

according to the formula:

Importance value = relative frequency + relative density

+ relative dominance

Ground study data permitted further division of vege-

tation units according to the floristic criteria above and

the environmental information collected from each plot

(altitude, soil type, topography).

Vegetation types were described in terms of their typ-

ical vegetation structure, characteristic woody species,

associated grass and herb species, soils, geology, topog-

raphy, distribution and area covered.

Map production

A preliminary working vegetation map of the NLNP was

derived from the 1983 panchromatic aerial photographs.

Transparent overlays marked with the vegetation bound-

aries delineated on the aerial photographs were reduced

from a scale of 1 : 34 000 to a scale of 1 : 250 000 on a pho-

tocopier (Xerox U.K. Ltd). This reduction was then super-

imposed onto the 1 : 250 000 UTM Ordnance Survey maps

of the NLNP (no. SD-36-2 and SC-36- 14) and redrawn.

This rough map was used as a working image upon which

ground survey could be based.

A second more accurate, more detailed and up to date

vegetation map (scale 1:100 000) was produced from a

partial scene (90 x 90 km) LANDSAT TM satellite

image (Band 4, 7th July 1995), georeferenced and

geocoded to UTM map projection (Satellite Applications

Centre, Pretoria). The image was supplied by SAC as

digital data (EOSAT Fast Format) on CD-ROM, which

was processed into a photographic image (Hunting

Technical Services, UK) upon which vegetation bound-

aries could be marked by hand. The vegetation types

delineated on this image were assigned according to the

aerial and ground survey results presented below and the

end result is a map incorporating 1 1 broad vegetation

types and in which vegetation mosaics are indicated

(Figure 1). This map was digitised using PC ARC/INFO

(Kent Cassells, DICE, UK) and incorporated into the

beginnings of a Geographical Information System (GIS)

for the NLNP. More detailed vegetation data, as well as

information about geology, topography, water relations,

plant-animal interactions, etc. can all be incorporated

into this database in the future in order to build up a more

complete picture of the park ecosystem.

RESULTS AND DISCUSSION

During this study, the vegetation of the NLNP was

stratified into six floristic classes, divided into 13 distinct

vegetation types. Vegetation type distributions are shown

in Figure 1, with the exception of types A2 and F3, the

areas of which are too small to be shown at this scale.

The vegetation descriptions below are largely based

upon quantitative tree/shrub data from sample sites. The

herbaceous component is described from qualitative data

and collection material. Further division of vegetation

types into subtypes is based on substrate. A detailed

description of all vegetation types and subtypes is given

below. Key characteristics of woody vegetation types are

given in Table 2.

A. RIPARIAN FORESTS. WOODLANDS AND THICKET

This vegetation class comprises moist forests to

woodlands and thicket fringing perennial and seasonal

watercourses. At lower altitudes (600-700 m) valley

riverine fringe vegetation takes the form of woodland

(usually with a well-developed shrub layer), and thicket.

It should be noted that even on the valley’s perennial

rivers (Luangwa, Lufila and Mwaleshi), fringe woodland

and thicket is discontinuous, being interspersed with

other vegetation types such as riverine grassland (F2)

and/or non-riverine vegetation types. This is even more

noticeable on seasonal watercourses where fringe river-

ine trees are usually mingled with trees and shrubs char-

acteristic of adjacent vegetation types. At higher altitudes

(800-1 300 m), on the Muchinga Escarpment, riverine

fringe vegetation takes the form of dense evergreen for-

est which, on the larger watercourses, may extend to

adjacent swampy areas. This vegetation type tends to be

well developed and distinctive (although narrow) even

on the smaller, seasonal watercourses.

Valley riverine woodland (Al), as described in the pre-

sent study, is readily recognisable in the Luangwa valley

surveys of Astle et al. (1969), Naylor et at. (1973) and Phiri

(1989) (Table 3). Escarpment riverine forest (A2) however,

is not described in detail in any of the above studies but is

covered thoroughly by Fanshawe (1971). Vegetation type

A2 contains elements of Fanshawe’s ‘Riparian forest’ (p.

34) and ‘Swamp forest’ (p. 32).

Al . Valley riverine woodland and thicket

The Luangwa River lies at an altitude of ± 600 m and

topographical relief is only a few metres. The large trib-

utaries of the Luangwa in the NLNP are the Lufila,

Mwaleshi and Mulandashi Rivers. These rivers emerge

from the Muchinga Escarpment in the west at an altitude

of between 700 and 800 m. Topographical relief along

Bothalia 28,2 (1998)

201

TABLE 2. — Summary of woody vegetation data in the NLNP from aerial and ground survey results

* Data not available, due to the discontinuous nature of A1 Valley riverine woodland and thicket.

these watercourses varies from as much as 5-50 m near

the escarpment to only a few metres at the Luangwa con-

fluences.

Valley riverine woodland and thicket is associated

with the rich, recently deposited alluvial soils which he

adjacent to the rivers of the valley floor. This alluvial belt

may only be a few metres wide or, along the Luangwa

and its major tributaries, may extend many hundreds of

metres from the river. Riverine valley soils show consid-

erable diversity and cannot be assigned to any one soil

class. In general, these are deep, stoneless, clearly strati-

fied soils which vary in texture from sands to clays (see

vegetation type F2). Soil pH is usually around neutral but

ranges from pH 5.6 to pH 6.9. Soil colour varies from

light yellow brown to dark grey.

The fringe woodland of the valley’s perennial and sea-

sonal rivers is generally two-storeyed in structure, with

canopy trees reaching 20 m or higher and a well-devel-

oped shrub layer, which may extend to form areas of

bushland or thicket. Characteristic tall trees include

Kigelia africana, Diospyros mespiliformis, Trichilia

emetica, Lonchocarpus capassa, Colophospermum

mopane, Combretum imberbe, Faidherbia albida,

Sclerocarya birrea and Tamarindus indica (invariably

associated with termitaria). Other less frequent but typi-

cal large trees include Breonadia salicina and Khaya

nyasica. Commonly occurring small trees and shrubs are

Piliostigma thonningii , Ziziphus abyssinica, Oncoba spi-

nosa, Feretia aeruginescens, Flueggea virosa, Anti-

desma venosum, Phyllanthus reticulatus, Acacia sieberi-

ana and A. polyacantha subsp. campy lac antha. Amongst

the thicket-forming shrubs, the genus Combretum is well

represented: C. obovatum (on clay), C. fragrans and C.

imberbe (as a shrub) are all common. C. fragrans may

form areas of bushland in which it is the single dominant

species. Other frequent thicket-forming species are

Diospyros senensis (on sand), Keetia zanzibarica and

Friesodielsia obovata. Climbers in this vegetation type

include Jasminum fluminense, Abrus precatorius and

Dregea macrantha. The grass layer associated with well-

developed fringe woodland or thicket is sparse and con-

fined to shade-loving species such as Panicum maximum,

Phyllorachis sagittata and Setaria homonyma. Herbs

typical of the stratified soils of the larger rivers of the

valley floor include Senna obtusifolia, Indigofera tincto-

ria , Sida alba , Vernonia glabra and spp., Duosperma

spp., Corchorus spp. and Ocimum spp.

On the major rivers such as the Luangwa, Lufila,

Mwaleshi and Mulandashi, vegetation type A1 occurs in

a substrate-dependent mosaic with the riverine herba-

ceous habitats described in vegetation type F2.

A2. Escarpment riverine forest

This forest is found fringing the rivers and streams of

the Muchinga Mountains in the west of the NLNR

Elevation ranges from 800-1 300 m and the terrain is

deeply dissected with relief measured in hundreds of

metres. The igneous/metamorphic geology of the Mu-

202

Bothalia 28,2 (1998)

TABLE 3. — Comparison of vegetation classifications of the Luangwa valley in past and present surveys

* not applicable.

chinga Escarpment means that many of its rivers pass

through rocky terrain (granite, quartzite) and waterfalls

are common.

Soils associated with this vegetation type are very

variable in texture and tend to be alluvial on the valley

bottoms and colluvial on the valley sides.

On the Muchinga Escarpment, rivers and streams are

bounded by dense, usually 3-storeyed evergreen forest

(known as Mushitu in CiBemba) which may extend to

adjacent swampy areas. The canopy is closed and the

trees from which it is formed may be 20 m or more in

height. Characteristic tall trees in this vegetation type

include Syzygium cordatum, Breonadia salicina, Clei-

stanthus polystachyus, S. guineense subsp. afromon-

tanum, Uapcica lissopyrena, U. sansibarica and Apo-

dytes dimidiata. Other less frequent, but characteristic

large trees include Parkia fdicoideci, Sapium ellipticum,

Tabernaemontana pachysiphon and Xylopia rubescens.

Monopetalanthus trapnellii is locally common on the

Mwaleshi River and can form extensive stands of forest

in which it is the single dominant species. Characteristic

understorcy species are Antidesma vogelianum, Engle-

rophytum magalismontanum, Craterispermum schwein-

furthii, Faurea saligna, Garcinia smeathmannii and

Berscimci abyssinica. Shrubs include Diospyros natalen-

sis, Rothmannia whitfieldii, Cremaspora triflora, Rhus

longipes, Tricalysia coriacea, Lasianthus kilimandschar-

icus and Erythroxylum emarginatum. Climbers typical of

this habitat are Carissa edulis, Sabicea laurentii, Smilax

anceps, Phyllanthus muelleranus, Clematis welwitschii,

Artabotrys spp. and Strychnos spp. Oxytenanthera bam-

boo is locally common on escarpment streams and rivers.

Where A2 fringing forest is a narrow belt of riparian

trees, grasses and herbs associated with dambo margins

occur (see F3). Common grass species include Loudetia

simplex, Trachypogon spicatus , Cleistachne sorghoides.

Bothalia 28,2 (1998)

203

Hyparrhenici spp., Pennisetum spp. and Setaria spp.

Characteristic subshrubs and herbs are Acalypha

chirindica, Urena lob at a. Polygala exelliana, Phaidopsis

imbricata, Droogmansia pteropus, Helichrysum spp.,

Tinnea spp. and Dissotis spp. Where riparian forest

extends into seasonally flooded swamp forest, or

Monopetalanthus forest, the forest floor is carpeted with

moss, and ferns are common ( Asplenium , Adiantum,

Pellaea). Epiphytes may also occur. Herbs in this habitat

include Geophila obvallata, Schwartzkopffia lastii ,

Leptactina benguellensis and Biophytum spp. Sclerochi-

ton vogellii is locally common in Monopetalanthus for-

est.

B. BUSHLANDS AND THICKET

Both vegetation types described below are characterised

by thicket species which form closed stands. However, in

some places more open stands occur and therefore vegeta-

tion types B 1 and B2 are best regarded as mosaics of bush-

land and thicket. Both Mixed alluvial thicket (Bl) and

Combretum thicket (B2) are confined to the alluvial areas

adjacent to the valley's larger rivers (Luangwa, Mwaleshi).

Together, vegetation types Bl and B2 cover an area of

approximately 180 km2, and seem to occupy a niche on soils

of an intermediate nature between the low, poorly drained

clays of vegetation type FI and the higher, sandy soils of

vegetation type Dl. Mixed alluvial thicket is the character-

istic thicket vegetation type north of the Mwaleshi River,

while Combretum thicket largely occurs to the south of the

Mwaleshi. Both types are dominated by different species of

thicket-forming shrubs, but share many species in common.

The absence of C. elaeagnoides and C. celastroides in large

areas of Mixed alluvial thicket, north of the Mwaleshi River,

which led to the distinction being made in the present sur-

vey, is apparently a local phenomenon because these two

species do occur further north, on the Lufila River. The rea-

son for their absence in the middle of the park is unknown,

and requires further study.

Astle et al. (1969) make no distinction between

Mixed alluvial thicket (Bl) and Combretum thicket (B2),

referring to this vegetation type as ‘Thicket on freely

draining alluvium'. Their species list for this vegetation

type contains the dominant elements of both B 1 and B2.

B 1 . Mixed alluvial thicket

This vegetation type is found on the alluvial soils

associated with the Luangwa River in the east of the park

and is the characteristic thicket vegetation type north of

the Mwaleshi River. Elevation in this area is between 600

m and 625 m and the terrain undulates slightly with a

relief of only a few metres. This vegetation type is fre-

quently found in mosaic with Chloris-Dactyloctenium-

Echinochloa secondary grassland (FI) but occupies

higher ground and better drained, sandy soils so that

while the surrounding grassland becomes waterlogged

during the wet season, the areas of thicket remain com-

paratively dry.

Soils associated with this vegetation type are freely

draining and are typically pale brown to orange, stone-

less, sandy clay loams (60-75% sand) of slightly acid to

neutral pH (pH 6. 5-7.0).

Vegetation type B 1 comprises closed and open stands

of bushes 2-7 m high with occasional tall trees. Colo-

phospermum mopane is present to a greater or lesser

extent. It is found in clumps on patches of calcareous or

sodic clay soil but also as individual, usually tall trees

dotted throughout the habitat. Other occasional tall trees

include Xeroderris stuhlmannii, Pseudolachnostylis

maprouneifolia, Stereospermum kunthianum and Adan-

sonia digitata. Common smaller trees are Boscia angusti-

folia, Excoecaria bussei (also a shrub) Schrebera tri-

choclada and Diospyros quiloensis (occasionally a large

tree). Vegetation type Bl is dominated by the shrubs

Croton gratissimus, Friesodielsia obovata, Mundulea

sericea, Combretum obovatum, Vangueria infausta,

Markhamia obtusifolia and M. zanzibarica. Baphia mas-

saiensis is locally common. The grass and herb layer is

not well developed in Mixed alluvial thicket but contains

elements of vegetation types Dl, El and FI. Grasses

include Digit aria acuminatissima, Sporobolus cordo-

fanus , Urochloa mossambicensis, Dactyloctenium spp.

and Chloris spp. In the wetter sump areas, Brachiaria

deflexa, Sporobolus pyramidalis and Setaria spp. are

characteristic. Common herbs are Hibiscus cannabinus ,

H. calyphyllus, Cleome monophylla, C. hirta, Senna

absus, Blepharis tenniramea, Clerodendrum spp. and

Duosperma spp.

B2. Combretum thicket

This is the characteristic thicket type south of the

Mwaleshi River. Elevation is ± 600-620 m and the ter-

rain is flat with a relief of only a few metres. As with

Mixed alluvial thicket (Bl), Combretum thicket is found

on alluvial, sandy soils close to the Luangwa and Mwa-

leshi Rivers.

Vegetation type B2 is comprised of thicket-forming

shrubs, 2-7 m in height, which grow in a mosaic of

closed and open stands. The grass layer is not well devel-

oped. Tall trees are rare but species such as Manilkara

mochisia and Diospyros quiloensis may occur infre-

quently. Small trees found in Combretum thicket include

Schrebera trichoclada and Combretum collinum subsp.

gazense. The dominant shrubs are Combretum elaeag-

noides, C. celastroides, C. obovatum, Holarrhena pubes-

cens, Vangueria infausta and Markhamia spp. Grasses

and herbs associated with this habitat are the same as

those in vegetation type B 1.

C. MIOMBO WOODLANDS

The miombo woodlands of North Luangwa National

Park cover an area of approximately 1 300 km2. This is

the dominant vegetation type of the Muchinga Escarp-

ment and its attendant foothills, Chinshenda, Soma and

the Mvumvwe range. The vegetation described under the

umbrella of miombo woodlands is physiognomically

diverse, ranging from closed woodland to open wood-

land to scrub woodland. For the purposes of this classifi-

cation, miombo woodland in the NLNP has been divided

into two types, Cl Upper escarpment and C2 Lower

204

Bothalia 28,2 (1998)

escarpment/hill miombo woodland, a division largely

based on floristic composition. Further division into sub-

types is dependent on substrate.

Miombo woodland is clearly recognised in all three

of the previous valley surveys (Table 3). Further de-

scriptions are available from Trapnell (1953) and from

Fanshawe ( 1971 ).

Cl. Brachystegia-Julbernardia-Isoberlima upper

escarpment and plateau miombo woodland

At elevations over 1 000 m, this is the most important

and extensive vegetation type of the Muchinga Escarp-

ment. It can be separated into three distinct vegetation

subtypes dictated by substrate. Subtype 1 occurs over

most of the upper escarpment, on deep laterite soils.

Subtype 2 is rare in the NLNP and is found in isolated

patches on shallower plateau soils. Subtype 3 is associat-

ed with the granite outcrops dotted throughout the

escarpment terrain.

Vegetation type Cl (subtype 1) corresponds to

Trapnell’s E and El types (Trapnell et al. 1950), which

occur on the deep soils of the escarpment, as evidenced

by the dominance of Brachystegia utilis and B. spici-

formis, species which cannot tolerate shallow soils

(Fanshawe 1971). The tall trees and comparatively

sparse grass layer seen in this vegetation type are proba-

bly due to the improved drainage and better soils associ-

ated with the dissected terrain (Cole 1963). In this vege-

tation type, laterite was more commonly found as nod-

ules in the B horizon rather than the impermeable layer

characteristic of the pediplain plateau soils (Cl subtype

2). Subtype 2 corresponds to Trapnell’s P4 type (Trapnell

et al. 1950), and Subtype 3 is described by White (1983)

as ‘Zambezian rupicolous bushland and thicket’.

Subtype 1. Upper escarpment miombo woodland', is the

dominant vegetation type of the upper Muchinga Escarp-

ment in the west of the NLNP, and is found at elevations

ranging from 1 000 to > 1 300 m over deeply dissected ter-

rain where relief is measured in hundreds of metres.

Subtype 1 is associated with deep, red, stoneless

sandy loams or sandy clays. These soils arc slightly acid

(pH 6. 6-7.0) and usually contain laterite nodules and

mica aggregates in the B horizon.

This woodland takes the form of a clearly two-

storeyed woodland witli an open to lightly closed canopy

of semi-evergreen trees 15-20 m high. Characteristic

canopy trees include Julbernardia paniculata, Brachy-

stegia spiciformis, B. utilis, Isoberlinia ahgolensis, Mar-

quesia macroura, Parinari curatellifolia, B. manga and

Pericopsis angolensis. Common lower storey trees are

Brachystegia stipulata, Uapaca kirkiana and spp.,

Craterosiphon quarrel, Phyllocosmos lemaireanus,

Memecylon flavovirens, Dalbergia nitidula, Combretum

zeyheri, B. longifolia, Anisophyllea pomifera, Diplo-

rhynchus condylocarpon and Pseuclolachnostylis ma-

prouneifolia. Frequent shrubs are Keetia gueinzii, K.

venosa, Rothmannia engleriana, Landolphia parvifolia

and Protea spp. Climbers are infrequent in this vegetation

type, Strophanthus welwitschii being a notable exception.

In upper escarpment miombo woodland, the grass layer is

sparse and generally restricted to scattered clumps of pre-

dominantly tall grasses. Common species are

Andropogon chinensis, A. schirensis, Anthephora elonga-

ta, Heteropholis sulcata, Loudetia simplex, Sporobolus

sanguineus and Trichopteryx fruticulosa. The herb layer

is sparse but varied. Pteridium aquilinum subsp. centrali-

africanum is locally common, and sedges such as

Ascolepis elata, Carex echinochloe subsp. nyasensis,

Bulbostylis spp., Cyperus spp. and Scleria spp. are found

in poorly drained areas. Frequent subshrubs are

Psychotria spithamea, P. eminiana, Indigofera emargin-

elloides, Tapiphyllum cinerascens , Desmodium barbatum

and Kotschya spp. Typical herbs include Costus

spectcibilis and Siphonochilus rhodesicus at the beginning

of the rains, followed by Thunbergia kirkiana , Endo-

stemon dissitifolius, Hybanthus enneaspermus , Otio-

phora scabra, Nidorella spartioides, Rhynchotropis pog-

gei, Aspilia spp., Becium spp., Spermacoce spp., Teph-

rosia spp. and Triumfetta spp.

Subtype 2. Plateau miombo woodland: occurs in isolat-

ed patches in the far west of the NLNP, and is associat-

ed with Hat terrain. Plateau soils tend to be shallow, poor

in nutrients and humus, slightly acid, and are typically

leached, with a laterite or gley horizon near the surface.

In contrast to upper escarpment miombo woodland,

plateau miombo woodland is of single storey structure

and is characterised by stunted Brachyste gia-Julber-

nardia, interspersed with Uapaca, Protect , Faureci and

Monotes species. In addition, the shrub and grass layers

are comparatively well developed, with Hyparrhenia

and Andropogon spp. predominant. This subtype occurs

on the comparatively shallow, infertile soils of the

plateau peneplain and over large areas of Zambia

appears to be secondary miombo woodland which has

been subjected to repeated fires and cultivation (Fan-

shawe 1971).

Subtype 3. Rupicolous miombo woodland : the rocky out-

crops and granite kopjes of the Muchinga Escarpment

support a distinctive vegetation type and although many

of the taxa listed above may occur, additional species

such as Brachystegia microphylla, Pterocarpus rotundi-

folius, Sclirebera trichoclada, Kirkia acuminata,

Landolphia parvifolia and Tarenna neurophylla are typi-

cal. Carphalea pubescens is a characteristic subshrub.

C2. Julbernardia-Brachystegia lower escarpment and

bill miombo woodland and scrub woodland

This woodland covers much of the lower Muchinga

Escarpment and its attendant foothills, Chinshenda,

Soma and the Mvumvwe range. It is also found on the

upper valley Boor where il may intergrade with vegeta-

tion types D I and D2.

Vegetation type C2 can be separated into two subtypes

according to substrate and vegetation physiognomy.

Subtype 1 is miombo scrub woodland, associated with

Bothalia 28,2 (1998)

205

the thin, eroded, stony soils of the hill slopes. Subtype 2

is open woodland, found on the deeper soils of the inter-

fluves and flatter sites.

Vegetation type C2, subtype 1 (scrub woodland) is

recognised by Astle et al. (1969) who refer to it as

'Miombo scrub on shallow soils’. This type is also

described by White (1983), in a direct reference to the

Luangwa valley (p. 99). Fanshawe (1971), Naylor et al.

(1973) and Phiri (1989) all refer to this distinctive form of

miombo as ‘Scrub miombo woodland'. Vegetation type

C2, subtype 2 (woodland) is designated ‘Miombo wood-

land on deep soil' by Astle et al. (1969). Phiri ( 1989) cites

Brachystegia boehmii, B. bitssei, B. manga and Julber-

nardia globiflora as common taxa in this habitat.

Subtype 1. Brachystegia stipulata-Julbernardia globi-

flora miombo scrub woodland : occurs at elevations rang-

ing from 700-1 000 m and is the most extensive vegeta-

tion type of the lower Muchinga Escarpment, where it

occurs in mosaic with subtype 2. It also intergrades with

vegetation types D 1 and D2 on the upper valley floor.

Subtype 1 occurs on the shallow, slightly acid (pH

6. 6-6. 9) light grey to yellowish brown, generally stony

sandy clay loam soils (lithosols and shallow fersiallitic

soils) associated with the hill slopes of the lower escarp-

ment and foothills.

The vegetation takes the form of 3-5 m tall scrub

woodland, in which there are very few tall trees,

Brachystegia bussei being an infrequent exception. In this

habitat the vegetation is dominated by Julbemardia glob-

iflora, Brachystegia allenii and B. stipulata in stunted

form. B. manga is locally dominant on Chinshenda hill.

Other common small trees and shrubs are B. boehmii,

Diplorhynchus condylocarpon, Combretum zeyheri,

Pseudolachnostylis maprouneifolia and Monotes afri-

canus. The grass layer is sparse, but similar in composition

to that found in vegetation types Cl and D2. Typical grass-

es are Trichopteryx fruticulosa, Tristachya bequaertii,

Diheteropogon amplectens, Sporobolus myrianthus,

Andropogon spp. and Melinis spp. Common subshrubs

include Ochna leptoclada , O. richardsiae and Cypho-

stemma spp. Characteristic herbs are Chlorophytum rubri-

bracteatum , Ectadiopsis producta , Endostemon dissiti-

folius, Indigofera sutherlandoides, Justicia striata ,

Zygotritonia nyassana. Becium spp. and Commelina spp.

Subtype 2. Julbemardia-Brachystegia open miombo wood-

land: is found in the deeply dissected terrain of the lower

Muchinga Escarpment and its foothills, where it occurs in

mosaic with miombo scrub woodland (Subtype 1). It also

intergrades with the Combretaceae woodlands and wooded

grasslands, D1 and D2, on the upper valley floor.

Subtype 2 is found on the deep, yellowish brown

slightly acid, sandy soils associated with the hill tops and

ridges of the lower escarpment and foothills.

The vegetation takes the form of open woodland with

scattered tall trees (15-20 m) and a well-developed grass

layer, dominated by tall species. Shrub patches are occa-

sional. Characteristic tall trees include Burkea africana,

Julbemardia globiflora, Pericopsis angolensis and

Brachystegia allenii. Common small trees include

Brachystegia stipulata, Combretum zeyheri, C. psid-

ioides, Terminalia sericea, and Oldfieldia dactylophylla.

The shrub layer is reduced but includes Ozoroa insignis,

Diplorhynchus condylocarpon, Vernonia glaberrima and

Lannea discolor (also a small tree). Common grasses are

Andropogon chinensis, A. gayanus, A. schirensis , Hyparr-

henia anemopaegma, PI. filipendula, Hyperthelia disso-

luta and Tristachya superba. Like the grasses, the herb

component is related to that of vegetation types D 1 and

D2 (below); typical species include Acalypha allenii, A.

villicaulis, Barleria fulvostellata, Dicoma anomala, and

Agathisanthemum spp.

D. COMBRETACEAE WOODLAND AND WOODED GRASSLAND

This vegetation class is dominated by (distinct) spe-

cies of the genera Combretum and Terminalia. Both veg-

etation types described below have well-developed grass

layers.

Vegetation type Dl, Combretum-Terminalia wood-

land, closely resembles the ‘ Terminalia sericea tree savan-

na’ of Wild & Barbosa (1967). White (1983) refers to it as

'North Zambezian undifferentiated woodland’, comprising

miombo associates but not Brachystegia/ Julbemardia

species. Fanshawe (1971) groups this vegetation type

together with the miombo woodlands of the valley floor,

but refers to it as ‘ Erythrophleum woodland'. The classifi-

cation of the present survey is largely based on floristic

composition and our ground survey data strongly indicates

that vegetation type Dl does not fall into the miombo

class. Brachystegia/Julbernardia species do occur in the

ecotones where vegetation type C2 intergrades with

Combretum-Tenninalia woodland, but elsewhere they are

never dominant and over large areas of this woodland, par-

ticularly in the east, they do not occur at all. Astle et al.

(1969) and Phiri (1989) refer to this vegetation type as

‘ Terminalia sericea-Erythrophleum africanum woodland

savanna’ and ‘ Erythrophleum woodland’ respectively

(Table 3). This nomenclature can be related to our results

in that E. africanum is an important component of this

vegetation type. However, in the NLNP at least,

Combretum/ Terminalia species are more frequent, more

abundant and account for more woody biomass than E.

africanum. Like us, Naylor et al. (1973) refer to this veg-

etation type as ‘ Combretum-Terminalia woodland’.

Astle et al. (1969) classify vegetation type D2,

Combretum-Terminalia-Diospyros wooded grassland,

under ‘Miombo scrub on shallow soils’, although they do

make a topographic distinction between this habitat (3;1

and 5;1) and vegetation type C2 subtype 1 (4;2 and 8;1). In

Astle’s updated landsystem/vegetation map (Astle 1989) a

further land facet is added (landsystem 7; facet 6) in which

this vegetation type is described as: ‘semi-deciduous scrub-

land with local variation in species composition. A mopane

and Terminalia stenostachya association on the flatter sites,

Brachystegia stipulata, Combretum apiculatum, Julber-

nardia globiflora association in areas of greater relief.’

This is a landsystem description incorporating the floris-

tic mosaic described below, of which vegetation type D2

is the non-miombo component.

206

Bothalia 28,2 (1998)

Dl. Combretum-Terminalia woodland

Vegetation type Dl is characterised by fire-tolerant,

sandy soil species (e.g. Terminalia sericea) and bears a

close resemblance to other high grass-woodland vegeta-

tion types such as the Burkea-Erythrophleum woodlands

of western Zambia and the Chipya woodlands of the

Bangweulu region (Trapnell et al. 1950). The common

herbaceous indicators of lake basin Chipya , Aframoinum

alboviolaceum, Smilax anceps and Pteridium aquilinum

are not a feature of this habitat, but like Chipya woodland

(Lawton 1978), it is probable that vegetation type Dl is

maintained by the fierce dry season fires which sweep

through the NLNP every year. Combretum-Terminalia

woodland is found in close association with thicket

(B1/B2), both vegetation types occupying a belt of deep,

sandy soil running parallel to the Luangwa River. The

relationship between thicket and woodland is not clear,

but it is possible that the two vegetation types are deter-

mined by edaphic factors. In certain (western) areas of

the park, this woodland intergrades with hill miombo

woodland (C2). It covers large areas of the valley floor

(180 knr in addition to 313 km2 in mosaic with vegeta-

tion type C2) over elevations ranging from 650 m to 700

m. It is generally associated with flat terrain where relief

is only up to 10 m.

Soils associated with this vegetation type are deep,

light brown to orange and are mildly acidic to neutral

(pH 6. 4-7.0). These soils are characterised by a very

high sand content (70-90%) and are probably colluvial

(or old alluvial) and derived from Karoo sandstone.

Combretum-Terminalia woodland takes the form of

open 1- or 2-storeyed deciduous woodland. Canopy spe-

cies may be up to 20 m tall. The grass layer is tall and

well developed. The tall trees found in this habitat are

dominated by Terminalia sericea (up to and above 15 m

high), Pseudolachnostylis maprouneifolia, Pericopsis

angolensis , Burkea africana, Erythrophleum africanum

and Amblygonocarpus andongensis. Common small

trees and shrubs are Combretum molle, C. collinum

subsp. gazense, C. zeyheri, Bridelia cathartica, Cros-

sopteryx febrifuga and Baphia massaiensis. On the deep

soils of this type, the grass layer is well developed with

both tall and short grasses. Common tall grasses are

Andropogon schirensis, A. gayanus, A. chinensis, Hy-

parrhenia anemopaegma, H. filipendula, Pogonarthria

squarrosa and Tristachya superba. Shorter species are

Aristida scabrivalvis, Digitaria gay ana, D. acuminatissi-

ma, Heteropogon contortus and Sporobolus festivus.

Characteristic subshrubs include Senna petersiana, Cissus

cornifolia and Euphorbia matabelensis. The herb compo-

nent is dominated by ground creepers of the family Vita-

ceae and includes Cyphostemma gigantophyllum, C. her-

mannioides, C. viscosum and Cissus nigropilosa. Other

typical herbs arc Polycarpaea corymbosa, Hypoestes for-

skaolii, Tricliceras spp. and Phyllanthus spp.

D2. Combretum-Terminalia-Diospyros wooded grass-

land

In the NLNP, vegetation type D2 occurs in mosaic

with mopane (E1/E2) and hill miombo (C2) throughout

its range (see map, Figure I). The Combretum apicula-

tum/Terminalia stenostachya/Diospyros kirkii associa-

tion (vegetation type D2) is the most extensive vegeta-

tion type in this mosaic, as it occurs on the shallow, stony

fersiallitic soils which cover the gentler slopes and flatter

regions of the upper valley floor. Hill miombo (C2 sub-

type 2) occurs in isolated patches on the deeper soils of

the ridge tops, while scrub miombo (C2 subtype 1)

occurs on the thin rocky soils of the steeper slopes.

Mopane woodland and scrub woodland (El and E2)

occur on sodic or calcareous patches of soil dotted

throughout the upper valley floor. This grassland, in

mosaic with vegetation types El, E2 and C2, covers an

area of approximately 1200 km2 in the NLNP and occurs

on the upper valley floor and in the foothills of the

Muchinga Mountains. Throughout its range this vegeta-

tion type is associated with gently sloping terrain with

relief measured in tens of metres.

The soils associated with D2 wooded grassland are

stoneless to very stony soils, over siltstone and grits,

which may or may not be covered with a surface mantle

of quartzoze stones. They are highly variable in texture

but tend to be shallow, grey to reddish brown and mod-

erately acid (pH 5. 4-6. 6).

Combretum-Terminalia-Diospyros wooded grassland

is defined by a well-developed grass/herb layer scattered

with small trees and shrubs covering 10% to 40% of the

surface. It is heterogeneous in composition and form,

grading into woodland in some areas and pure grassland

in others. Dominant small trees are Diospyros kirkii,

Terminalia stenostachya, T. stuhlmannii, Combretum

apiculatum, C. frag ran s, C. zeyheri, Crossopteryx febri-

fuga and Pseudolachnostylis maprouneifolia. Shrub

species include Bauhinia petersiana, Acacia hockii, A.

gerrardii and Ximenia caffra subsp. caffra. On the shal-

low, stony soils of this habitat, the grass layer is well

developed and dominated by medium to tall, coarse

grasses. Common species are Andropogon gayanus, A.

schirensis, A. fastigiatus, Aristida scabrivalvis, Dihete-

ropogon amplectens, Heteropogon contortus, Hyparr-

henia anemopaegma, Loudetia flavida, Monocymbium

ceresiiforme, Sorghum versicolor and Zonotriche amoe-

na. Bothriochloa bladhii , Sporobolus pyramidalis,

Heteropogon contortus , Echinochloa colona and Setaria

spp. are locally common in poorly drained areas.

Subshrubs typical of this habitat are Xerophyta suave-

olens, Pachycarpus lineolatus , Cissus cornifolia and

Cyphostemma spp. Characteristic herbs include Ipomoea

welwitschii, Pterodiscus elliottii, Chlorophytum syl-

vaticum , Lapeirousia eryantha, Raphionacme longituba,

Neorautanenici mitis, Vernonia melleri , Tephrosia spp.

and Gladiolus spp. Common ground herbs are Crotalaria

cephalotes. Euphorbia oatesii , Gardenia subacaulis and

Torennia spicata. Sumps and wet areas are characterised

by Cycnium tubulosum and sedges including Ascolepis

protea var. splendida, Kyllinga alba, Scleria bulbifera,

Cyperus margaritaceus and C. macrostachys.

Trapnell believes that this vegetation mosaic is the

result of sheet erosion. He suggests that the colluvial

soils of the escarpment were more extensive in the past,

covering much of the upper valley floor in the NLNR

Evidence for this is to be found outside the park, north of

Bothalia 28.2 (1998)

207

the Lufila river, where miombo woodland covers a large

area of the upper valley floor (Trapnell et al. 1950;

Edmonds 1976) and is found on soils of a colluvial

nature containing laterite nodules. It is postulated that, in

the NLNP and further south, much of this escarpment

soil has been washed away, leaving a dissected, undulat-

ing terrain, the slopes of which are characterised by a

thin, stony soil cover. This niche is occupied by vegeta-

tion type D2 in the areas of gentler relief and miombo

scrub woodland (C2 subtype 1) on the steeper slopes.

Hill miombo woodland (C2, subtype 2) occupies the

ridge tops, where remnants of the deep escarpment soil

cover remain. The patches of sodic or calcareous soil,

associated with the mopane vegetation (E1/E2) of the

upper valley floor, are almost certainly derived from

ancient, and in some cases recent, termite activity

(Trapnell et al. 1976).

E. COLOPHOSPERMUM MOPANE WOODLAND AND SCRUB

WOODLAND

Colophospermum mopane is the single dominant tree

species in this vegetation class. In the NLNP, mopane

may grow as a tall tree of up to 15 m, or it may take the

form of a multistemmed, stunted shrub <3 m tall. The

tall form is typical of Colophospermum mopane wood-

land (El) and the shrub form is characteristic of Colo-

phospemium mopane scrub woodland (E2). These two

vegetation types may occur in discrete areas or they may

grow together in mosaic. The difference between the two

types is largely the physiognomy of C. mopane, which

can be related to browsing damage and substrate as

described below. Together, the two mopane habitats

cover an area of approximately 600 km2 in the NLNP.

Mopane woodland and scrub woodland are recog-

nised by all of the authors referred to above (Trapnell

1953; Wild & Barbosa 1967; Asti q et al. 1969; Fanshawe

1971; Naylor et al. 1973; Edmonds 1976; Phiri 1989).

El. Colophospermum mopane woodland

This woodland occurs on the alluvial soils associated

with the Luangwa River and its tributaries in the east of

the park. The terrain is flat, but due to the sparse herba-

ceous layer and the impermeable nature of the soil,

mopane woodland is usually dissected with drainage

channels and erosion gullies.

Mopane woodland soils typically consist of a shallow

(sandy loam) A horizon over an impermeable B horizon

of brown or grey, cracking, slightly acid (pH 6. 0-6. 5)

clay. These soils are poorly drained and as a result, are

waterlogged during the rainy season. Continual sheet

erosion has the effect of removing the topsoil, and the

roots of mopane trees in this habitat are frequently

exposed or undermined.

In this vegetation type, Colophospermum mopane grows

as the single dominant species in an open two-storeyed

woodland comprising a canopy layer of mature trees (10-15

m tall) and an understorey of trees in various stages of

development. Trees and shrubs associated with C. mopane

are comparatively few, mainly species found in the thicket

habitats B1 and B2. Other associated species are Afzelia

quanzensis , Balanites aegyptiaca, and Ximenia americana.

The herbaceous component is dependent on substrate. On

soils with a sandy A horizon, the herbaceous layer is sparse

and largely composed of grasses, particularly those associ-

ated with vegetation type FI (e.g. Urochloa mossambicen-

sis, Chloris spp., Dacty’loctenium spp.). Other characteristic

species are Alloteropsis cimicina, Aristida rhiniochloa, A.

scabrivalvis, Eragrostis viscosa, Microchloa indica, Sporo-

bolus cordofanus and S. panicoides. In the sumps and wet-

ter areas typical grasses are Brachiaria deflexa ,

Echinochloa colona and Setaria pumila. On alluvial black

and brown clay soils, dicotyledons make up a larger pro-

portion of the herbaceous component with the family

Acanthaceae particularly well represented (see vegetation

type F2 subtype 2). Common species are Duospenna cre-

natum, D. quadrangulare, Hygrophila auriculata,

Blepharis tenniramea , Barleria prionitis and Monechma

debile. Other common herbs are Kalanchoe lanceolata.

Senna absus, Cyphostemma spp. and Ocimum spp.

The tall ‘cathedral mopane’ woodland associated with

deep alluvial soils east of the Luangwa, outside the

NLNR is not common in the park. Instead, this two-

storeyed form of woodland has arisen due to browsing

pressure, which prevents recruitment into taller size

classes (Caughley 1976). The most influential browsers

in mopane woodland are elephants, which tend to browse

destructively, pollarding the trees as they feed (Anderson

& Walker 1974; Caughley 1976; Lewis 1991; Styles

1993). However, in the past ten years the elephant popu-

lation of the Luangwa valley has been severely reduced

due to ivory poaching (Leader- Williams et al. 1990) and

if elephants are responsible for the maintenance of this

scrub mopane, it is to be expected that without their

browsing pressure, scrub mopane trees will grow into tall

trees and set seed. In the mopane plots laid down in this

study, individual tree heights, extent of damage and seed

status were recorded. Measurements in 1993, and 1994

suggest that, in many cases, height recruitment is occur-

ring. Further visits to these plots will be necessary to

confirm this trend.

E2. Colophospermum mopane scrub woodland

This woodland occurs in discrete patches throughout

all the habitats of the valley floor forming distinct islands

of vegetation, which are clearly visible as white patches

on aerial photographs. They appear to arise due to local

soil conditions.

The soils are typically compacted pinkish grey to light

grey sandy silt loams over an impermeable calcareous or

sodic clay loam B horizon. The xerophytic conditions

created by the impermeability of the soil, and the relative

alkalinity of the B horizon produce a hostile environment

for herbaceous plant species. The resulting paucity of

herbaceous ground cover exacerbates the erosion prob-

lems in this habitat and, as with vegetation type El , scrub

mopane soils are dissected by numerous drainage chan-

nels and erosion ditches.

As in El, Colophospermum mopane is dominant in this

vegetation type, but here its growth is stunted, and it rarely

208

Bothalia 28,2 (1998)

exceeds 3 m in height. Characteristic associated shrubs are

Commiphora spp., Maerua angolensis (as a shrub) and

Lannea humilis. Climbers include Maerua juncea,

Hippocratea indica and Cissus spp. The herbaceous layer

is sparse. Typical grasses are Aristida spp., Sporobolus

cordofanus , S. panicoides and Eragrostis viscosa. Charac-

teristic subshrubs and herbs include Zanthoxylum chaly-

beum , Cadaba kirkii, Jasminum stenolobum (as a ground

herb or suffrutex), Decorsea schlechteri, Scadoxus multi-

florus, Plectranthus tettensis and Cyphostemma spp.

Ipomoea kituiensis is locally common.

There are two major factors which cause the shrub

growth form of Colophospermum mopane in the NLNP:

browser damage (see above) and/or soil conditions.

Where scrub mopane trees are intimately associated with

tall mopane trees, as in vegetation type El on the alluvial

soils in the east of the park, stunted growth appears to be

largely due to browser-pollarding. Elsewhere, both near

the river and throughout the rest of the park, compara-

tively large, discrete areas of scrub woodland occur. This

is a distinct vegetation type in which edaphic factors are

more important than browsing pressure in influencing

vegetation physiognomy (Dye & Walker 1980).

F. GRASSLANDS

The grasslands of the NLNP, defined as areas of

herbaceous vegetation with less than 10% woody vege-

tation cover, are all associated with water. Vegetation

Type FI is seasonally waterlogged, while types F2 and

F3 are associated with the rivers and dambos of the val-

ley and escarpment respectively.

Vegetation type FI is referred to by Phiri (1989) as

‘Floodplain grassland’ with Echinochloa colona cited as

the dominant species. Astle et al. (1969) describe this

vegetation type (landsystem 1; facet 7; photo 4) as ‘Short

Echinochloa grassland with Combretum obovatum

shrubs on dark cracking clays’.

Vegetation type F2 is readily recognisable in the pre-

vious valley surveys. The sandbar grassland (landsystem

1; facet lb) and pointbar deposit grassland (1; 5a) of

Astle et al. (1969) are synonymous with vegetation types

F2 subtype 1 and subtype 3, respectively, of the present

survey. Similarly, the dambos, mudflats and pointbar

grass habitats (2b, 2c and 2g, respectively) of Naylor et

al. (1973) are directly referable to vegetation type F2.

Vegetation type F3 is not dealt with by any of the val-

ley surveys, as it is an upper escarpment and plateau

habitat. However, Vesey-Fitzgerald (1963) gives a

detailed account of this vegetation type which he calls

‘Headwater valley grasslands (dambos)’. Loudetia sim-

plex and Hyparrhenia spp. are named as characteristic

species. White (1983) and Fanshawe (1971) also

describe this vegetation type, referring to it as ‘Dambo

grassland’ and ‘Bush-group grassland’ respectively.

FI. Chloris-Dactyloctenium-Echinochloa secondary

grassland

This type takes the form of short, annual grassland

punctuated with occasional clumps of Combretum obo-

vatum thicket. It is found on degraded mopane woodland

and, as a result, is often scattered with the skeletons of

dead mopane trees. Vegetation type FI covers a large

area of the NLNP (approximately 335 km2) and is found

on recently deposited alluvial soils adjacent to the

Luangwa River. The terrain is flat and low lying, and is

seasonally waterlogged during the rainy season.

Soils associated with this habitat tend to be shallow,

poorly drained, light grey, compacted neutral sandy clays

or sandy loams.

The important grasses in this habitat are short to medi-

um-sized, nutritional species such as Chloris virgata,

Dactyloctenium aegyptium, D. giganteum, Urochloa

mossambicensis, Digitaria acuminatissima and Era-

grostis gangetica. Echinochloa colona, Brachiaria

deflexa and Sporobolus pyramidalis favour the wetter

sites in this habitat. Typical herbs are Ammocharis tin-

neana, Crinum minimum, Leonotis nepetifolia, Senna

absus, Indigofera gairdnerae (ground herb), and in the

wetter areas, Heliotropium spp., Cycnium tubulosum and

sedges such as Ascolepis protea var. splendida and

Cyperus macrostachyos.

This grassland (FI) is clearly part of a dynamic suc-

cession. The extensive areas of FI grassland present in

the NLNP today were not recorded by Astle in 1965. At

that time, these areas were largely covered by mopane

woodland. It is probable that this succession has arisen

due to extensive browsing damage in mopane wood-

lands, concomitant with the well documented increase in

elephant numbers recorded in the valley during the

1970’s (Caughley 1976). Additional factors such as

waterlogging and fire may have been involved in main-

taining the grassland state. Recent observations made by

the author suggest that mopane seedlings are starting to

re-invade the FI grasslands in certain areas of the park.

This reversion to mopane woodland would be consistent

with the hypothesis that these grasslands are created and

maintained by elephants because, as stated above, ele-

phant numbers in the Luangwa valley have declined

drastically in the past ten years (Leader- Williams et al.

1990). Furthermore, recent anti-poaching efforts have

greatly reduced the incidence of man-made fires in the

park. The mechanics of this succession need to be under-

stood because this grassland is a particularly productive

habitat, which aerial survey has shown is utilised by

large numbers of grazers (NLCP census 1994). If these

grasslands revert back to mopane woodlands, an impor-

tant grazing habitat will be lost from the NLNP. Further

research is needed to investigate the grassland-mopane

succession, and the factors which influence it.

F2. Valley riverine grasslands

These grasslands are associated with the larger rivers

of the valley floor. The Mwaleshi, Mulandashi and

Luangwa Rivers all have extensive floodplains within

their meander belts as well as numerous attendant drain-

age channels, oxbow lagoons and dambos.

Various soils are associated with this vegetation type,

but all are based on the recently deposited alluvium of

the large rivers. All tend to be deep and stratified, with

Bothalia 28,2 (1998)

209

soil textures ranging from the well-drained sandy soils of

the sandbars to the cracking black clays of the flood-

plains. Each substrate supports a distinctive grass and

herb component.

Subtype 1. Cynodon-Eragrostis grassland on sandy

soils : on the sand bars and sandy deposits of the inside

curves of the valley's rivers and streams, common tall

grasses are Andropogon gayanus, Cymbopogon excava-

tus, Digitaria milanjiana, Hyparrhenia filipendula,

Hyperthelia dissoluta, Pennisetum purpureum, Phrag-

mites mauritianus, Setaria sphacelata and Themeda

triandra. Common medium-sized grasses include Dac-

tyloctenium giganteum, Eragrostis cylindriflora, E. cilia-

nensis, Heteropogon conforms, Perotis patens and

Sporobolus pyramidalis. Smaller grasses include the

stoloniferous perennial Cynodon dactylon, which fre-

quently carpets sand bars and abandoned river channels

or ‘wafwas’ on compacted sands. Other short grass

species found on sandy alluvial soils are Chloris spp..

Dactyloctenium spp., Eleusine indica, Eragrostis cil-

iaris, Perotis leptopus, Sporobolus festivus and Urochloa

mossambicensis. Herbs associated with the sandy river-

ine soils of the valley floor include Waltheria indica ,

Trichodesma zeylanicum, Tridax procumbens, Sesamum

spp. and Striga spp.

Subtype 2. Setaria-Hyparrhenia grasslands and wooded

grassland on clay: the brown and black clay loams asso-

ciated with the floodplains of the valley’s larger rivers

support distinctive tall grassland. An important species

on brown clay loam soils is Hyparrhenia rufa , whereas

on black (black cotton) clays, Setaria incrassata (Ka-

sense) grows in pure stands. Interspersed with stands of

Hyparrhenia and Setaria , riverine clay soils support

areas of herbaceous vegetation dominated by the family

Acanthaceae. Hygrophila auriculata, Duosperma quad-

rangulare and D. crenatum are common in these areas,

together with species such as Senna obtusifolia , Indi-

gofera tinctoria, Sida alba, Ocimum spp. and Corchorus

spp. Sesbania greenwayii is locally common.

Setaria-Hyparrhenia riverine grassland frequently

grades into wooded grassland, with scattered trees from

vegetation type A1 (e.g. Kigelia africana, Acacia spp.,

Combretum spp.) or Colophospermum mopane forming

the woody component.

Subtype 3. Aquatic associations: water grass associa-

tions are found on the seasonally waterlogged clays of

the Luangwa River’s oxbow lagoons and dambos. In

these areas, which remain under water for most of the

rainy season, water-loving grasses such as Oryza barthii,

Echinochloa colona, Sporobolus pyramidalis and Setaria

spp. dominate. Common sedges in this habitat include

Cyperus esculentus, C. articulatus, C. distans and

Kyllinga alba. Characteristic water-associated herbs are

Polygonum setulosum, Lindemia oliveriana, Sphenoclea

zeylanica and Heliotropium spp. When water remains in

the lagoons and dambos, the aquatic water weed Pistia

stratiotes is characteristic. Around the peripheries of

lagoons and dambos as they dry out, typical herbaceous

species are Portulaca oleracea, Ludwigia stolonifera ,

Hibiscus articulatus, Alternanthera sessilis, Mimosa

pigra and Sphaeranthus spp.

F3. Loudetia simplex-Hyparrhenia dambo grassland

On the Muchinga Escarpment and hills of the NLNP

(Chinshenda, Mvumvwe and Soma) the rivers and

streams do not have a well-developed meander belt and,

as a result, the herbaceous riverside vegetation is less

well defined than in the valley. However, the numerous

dambos and drainage channels associated with these

watercourses do have a characteristic grass and herb

component.

Dambo soils are poorly drained and compacted. They

are typically leached illuvial soils, black, dark grey or

dark brown in colour, and acid (pH 5-6).

Early in the rainy season, the dambos of the upper

escarpment are characterised by the grasses Loudetia

simplex, Setaria sphacelata and S. pumila. Later on, in

April/May, Hyparrhenia species are ascendant: H. dip-

landra, H. nyassae, H. rufa, H. schimperi, H. collina, H.

variabilis and H. welwitschii may all be found at this

time. Other late season dominants are Andropogon chi-

nensis, Diheteropogon filifolius, Monocymbium ceresi-

iforme and Pennisetum unisetum. Typical herbs associat-

ed with the dambos of the upper escarpment include

Gnidia chrysantha, Moraea bella, Satyrium carsonii,

Cynorkis hanningtonii, Eulophia cucullata. Euphorbia

cyparissioides, Gladiolus spp. and Thunbergia spp.

Common sedges are Cyperus margaritaceus, C. cyper-

oides and Scleria spp.

The dominant grasses and herbs found on the dambos

and streams of the lower escarpment and hills are similar to

those found in the upper escarpment. However, a number

of water-associated grasses and herbs from the valley may

also occur. Grasses include Brachiaria brizantha, Digitaria

milanjiana, Echinochloa colona, E. pyramidalis, Setaria

pumila, S. sphacelata, Sporobolus pyramidalis, Themeda

triandra and Urochloa mossambicensis. Subshrubs and

herbs include Urena lobata. Senna occidentalis, Aeschy-

nomene mimosifolia, Ageratum conyzoides, Acmella

caulirhiza, Tragia lasiophylla, Alternanthera sessilis,

Ludwigia stolonifera and Polygonum setulosum.

Vegetation mosaics

The representation of vegetation as a series of discrete

types, based on communities or associations is, to some

extent, artificial in that observed vegetation associations

are almost invariably part of a continuum or occur in

mosaic with other communities (Craig 1983). This is a

problem of scale. The larger the scale of the map to be

produced, the smaller the area of association that can be

represented. For practical purposes however, some vege-

tation communities will always be too small to map sep-

arately. This applies in areas of mosaic or in cases where

vegetation communities occupy a microhabitat. The veg-

etation associated with termite mounds, for example, is

distinctive (Fanshawe 1968) and may be important to the

ecology of an area, yet as a type, it is often too dispersed

210

Bothalia 28,2 (1998)

to be mapped separately. In the present study, clear

mosaics have been indicated on the map (Figure 1).

Elsewhere, dominant associations, not necessarily pure

associations, are depicted (see below).

CONCLUSIONS

Vegetation monitoring on any scale requires baseline

data from which deviations, or trends, can be measured.

Changes in woody cover over large areas, for example, can

be discerned from aerial photographs and satellite images.

In the case of the NLNP, mopane woodland for example,

can be mapped from historic aerial photographs (1952,

1965 and 1983) and compared with the 1995 LANDS AT-

derived digital data used in the present study (Figure 1 ).

Although aerial perspectives are useful records of

large-scale trends, they provide little information about

cause of change. Of much more use in this respect is

ground study information about vegetation structure and

species composition. In the present study, 353 relocat-

able plots were laid down and the precise data recorded

in each of these plots provides the basis for monitoring

vegetation changes in the future.

Finally, it is important to stress that this study presents

a coarse stratification of the vegetation of the NLNP, use-

ful for measuring changes in the park’s vegetation at this

scale. Although these vegetation units are fairly homoge-

neous, they still, in some cases, represent more than one

plant community. A more detailed study of plant associa-

tions in the park is needed if we are to begin to compre-

hend the processes involved in maintaining plant commu-

nities. At the species level, it is the local combination of

abiotic and biotic factors which make up a niche that is

important. If we arc to understand the autecology of

species, or even the ecology of ecotypes, we need to

understand the combinations of habitat components which

make up the niche. The great potential of geographical

information systems is that the detailed local picture can

gradually be built up. Spatial information about geology,

soil, topography, water relations, fire regimes, animal-

plant interactions and numerous other factors can be col-

lected over time and added to the picture. It is only

through this process that we can understand enough about

ecosystems to effectively manage them.

ACKNOWLEDGEMENTS

The author would like to thank the North Luangwa

Conservation Project for its support throughout this

study. NLCP is in turn supported by the Owens

Foundation for Wildlife Conservation and Frankfurt

Zoological Society. The Project is grateful to National

Parks and Wildlife Services, Zambia for their co-opera-

tion and assistance throughout this survey. In the

Luangwa Valley, my thanks go to David Chile and

Godfrey Chikalipe for their invaluable help in the field.

Thank you too to my wife Deborah for carrying out the

soil survey associated with this study. In Lusaka, I am

indebted to Patrick Phiri, who allowed me access to his

herbarium at UNZA. Thanks also to Mike Bingham for

his help with identifying specimens. In the UK Gerald

Pope, Krukoff Curator of African Botany at the Royal

Botanic Gardens, Kew, arranged for me to use the

herbarium and library at that institution. Many thanks to

him. Finally, I am grateful to Colin Trapnell and Ray

Lawton, who took the time to read and comment on the

first draft of this paper.

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Bothalia 28,2: 213-229 ( 1998)

Wetland plant communities in the Potchefstroom Municipal Area,

North-West, South Africa

S.S. CILLIERS*, L.L. SCHOEMAN* and G.J. BREDENKAMP**

Keywords: Braun-Blanquet, DECORANA, disturbed areas, MEGATAB, TURBOVEG, TWINSPAN, urban open spaces

ABSTRACT

Wetlands in natural areas in South Africa have been described before, but no literature exists concerning the phyto-

sociology of urban wetlands. The objective of this study was to conduct a complete vegetation analysis of the wetlands in

the Potchefstroom Municipal Area. Using a numerical classification technique (TWINSPAN) as a first approximation, the

classification was refined by using Braun-Blanquet procedures. The result is a phytosociological table from which a

number of unique plant communities are recognised. These urban wetlands are characterised by a high species diversity,

which is unusual for wetlands. Reasons for the high species diversity could be the different types of disturbances occurring

in this area. Results of this study can be used to construct more sensible management practises for these wetlands.

CONTENTS

Introduction 213

Study area 215

Materials and methods 215

Results and discussion 219

Classification 219

Description of the communities 219

Ordination 227

Conclusion 227

Acknowledgements 228

References 228

INTRODUCTION

According to the Ramsar Convention, which is also

known as The Convention on Wetlands of International

Importance, wetlands can be described as areas of marsh,

fen, peatland or water, whether natural or artificial, per-

manent or temporary (Cowan 1995). The water of wet-

lands can be static or flowing, fresh, brackish or salty,

including areas of marine water the depth of which at

low tide does not exceed six metres. Wetlands may

include adjacent riparian and coastal areas (Cowan

1995). Wetlands also refer to a mosaic of ecosystems that

typically form transition zones between uplands and

aquatic environments, although some do have discrete

boundaries thus not providing gradual transitions

(Catallo 1993).

Wetlands have many beneficial functions which are of

chemical, biological, socio-economical and physical or

hydrological nature (Williams 1990) and they play an

important role as water reservoirs, stream flow regula-

tors, flood attenuators, water purifiers and controllers of

soil erosion (Walmsley 1988). Additionally, wetlands

form specialised habitats for a great variety of plant and

animal species.

* Department of Plant and Soil Sciences, Potchefstroom University for

C H E., 2531 Potchefstroom, South Africa.

** Department of Botany, University of Pretoria, 0002 Pretoria.

MS. received: 1998-02-18.

Many natural wetlands have been destroyed in the

course of agricultural, industrial and urban development

(Archibald & Batchelor 1992), and are regarded as one

of South Africa’s most endangered ecosystem types

(Walmsley 1988). The increasing use of fertilisers and

insecticides in catchment areas of wetlands and the fur-

ther threat of pollution by industries and the almost indis-

criminate use of some areas for grazing, contribute great-

ly to the degradation of wetlands (Fuls et al. 1992). As

early as 1951, Louw made the observation that the mod-

ification of natural vegetation and the destructive conse-

quences of man’s interference in the Potchefstroom area,

is nowhere as evident as along the numerous vleis, wet-

lands and spruits.

Conservation and management of wetlands need to be

addressed to ensure the natural functioning of these areas

as well as the maintenance of species diversity (Eckhardt

et al. 1993a). The need for the conservation of urban

wetlands resulted in the development of programmes

such as the Metropolitan Open Space Systems (MOSS)

programme of which the Durban MOSS (D’MOSS) is a

prime example. This programme proposed a new holistic

approach to city planning, one whereby indigenous

plants form an integral part of the urban landscape

(Roberts 1993). Because rivers, streams, pans, marshes,

estuaries, and lagoons are critically important to both

man and wildlife (Cooper & Duthie 1992), wetlands

within urban areas should form the essential core areas of

any MOSS network. The MOSS approach is presently

also being implemented in Durban, Pietermaritzburg,

East London, Port Elizabeth, Bloemfontein, the East

Rand, Port Alfred and Empangeni.

One of the first priorities, before a system such as

MOSS can be implemented, is an extensive phytosocio-

logical survey of the plant communities within the

municipal borders (Roberts 1993). Before conservation

strategies can be successfully implemented, detailed

knowledge and information of individual biotopes, their

ecological characteristics, location, and distribution

within the municipal area should be obtained, as well as

knowledge of the faunal and floral composition of these

214

Bothalia 28,2 (1998)

Pofchefstroom University for C.H.E.

CARTOGRAPHIC SERVICES

Telephone: (0148) 299-1589

Fac simile : (0 1 48) 299-2503

E-Mail : GGPCC JVR^PUKNFT.PUK.AC.ZA

DATE : November 1997

Map : co<i<JiW\plb\sc-vs4and

LEGEND : g

| Wetland areas

I Residential and business areas

! Parks and recreation areas

j Informal green areas

University and College

Industrial area

Open Areas around City

Railway lines

FIGURE 1 . — Location of the wetlands and adjacent areas with different land use in the Potchefstroom Municipal Area, North-West, South Africa.

Bothalia 28,2 (1998)

215

biotopes (Starfinger & Sukopp 1994). The presence or

absence of some species in these biotopes may often

serve as a bio-indication of the disturbance or pollution

in these biotopes (Starfinger & Sukopp 1994).

Although phytosociological studies of wetlands in nat-

ural areas in the Grassland Bionte have been done by,

among others, Kooij et al. (1991), Fuls et al. (1992),

Bloem et al. (1993), Eckhardt et al. (1993a). Myburgh et

al. (1995) and Smit et al. (1995), no literature concerning

the phytosociology of urban wetlands exists. Bezuiden-

hout (1993) did not study wetlands in his comprehensive

syntaxonomical and synecological study of the western

Transvaal Grasslands, which include the natural areas

around Potchefstroom. The only study on the wetlands of

the Potchefstroom area was that of Louw (1951).

The objective of this study was, therefore, to conduct

a complete vegetation analysis of wetlands within the

Potchefstroom Municipal Area. Through this study it

should be possible to ascertain the extent of disturbance

due to urbanisation. This study forms part of an extensive

study of urban open spaces in a number of cities in the

North-West, South Africa, with the ultimate aim of a

phytosociological and syntaxonomical synthesis of all

urban open spaces.

STUDY AREA

Potchefstroom is situated in the Dry Sandy Highveld

Grassland (Bredenkamp & Van Rooyen 1996) of the

Grassland Biome (Rutherford & Westfall 1994).

The present study covered the wetlands within the

Potchefstroom Municipal Area (between 27° 04' and 27°

07' longitude and 26° 40' and 26° 44’ latitude) in the

North-West province (Figure 1). Potchefstroom is locat-

ed around the Mooi River, near the confluence of the

Mooi River and Loopspruit. Geologically this area is

underlain by Pretoria Group strata, namely shale and

quartzite with major diabase intrusives. The diabase sills

and the Pretoria Group shale are major sources of clay,

which may be supplemented by Karoo age clays weath-

ered from palaeokarst in Malmani Dolomite along the

upper reaches of the Mooi River (Nel et al. 1939).

The rainfall in Potchefstroom is erratic, but the mean

annual rainfall exceeds 600 mm. Summer temperatures

are high, the mean monthly maximum temperatures

exceed 32° C during October to January, whereas the

mean monthly minimum temperatures are below -1° C

during the months of June to August (Weather Bureau

1988).

The entire Potchefstroom area is drained by the Vaal

River and its tributaries, of which the Mooi River and

Loopspruit are the most important. The latter has its ori-

gin near Losberg. The Mooi River is a perennial stream,

fed by a number of strong springs in the dolomite forma-

tion, within and beyond the northern limits of the area.

The study area includes three large marshland areas

which will be referred to as the Arboretum, Van der Hoff

Park and Prozesky marshlands (Figure 1) and a number

of very small marshland areas which will not specifical-

ly be referred to. The Arboretum is a conservation area

on the city margin in the northern part of Potchefstroom,

northwest of the Potchefstroom Dam and is bordered by

natural grasslands. This area is relatively undisturbed in

comparison with the other two marshlands, although it is

invaded by large reedswamps. The Van der Hoff Park

marshland is situated between two residential areas close

to the city centre, but is heavily disturbed due to grazing

especially by horses. It is a very flat bottomland area

without any deep depressions or drainage canals. The

marshlands referred to as the Prozesky marshlands are

situated in the O.P.M. Prozesky Bird Sanctuary. This

conservation area, which stretches from relatively close

to the city centre to the southern municipal border, is

characterised by deep depressions and drainage canals

with higher-lying areas in between, which are heavily

grazed by cattle. This wetland is interrupted by a road

before it links to a number of dams of the municipal

sewage plant. The remaining parts of the banks of the

Mooi River houses recreational facilities, sports grounds,

including a trim park and a golf course, as well as a num-

ber of residential gardens (Figure 1). The Wasgoedspruit

which drains the western parts of the city, including

industrial areas, as well as natural areas, into the Mooi

River, is also part of the system. The Wasgoedspruit is

disturbed and part of it is layed out with concrete.

MATERIALS AND METHODS

Releves were compiled in 102 sample plots during the

period January to March 1996. Plot sizes were fixed at 16

nr for herbaceous communities and 100 nr for woody

communities (Bredenkamp & Theron 1978). For each

species present in the sample plots, the cover-abundance

values according to Braun-Blanquet scales were used

(Mueller-Dombois & Ellenberg 1974)

Habitat parameters recorded, included topography,

aspect, slope, soil type and various soil properties,

including physical and chemical analyses. The soil prop-

erties in the A and B horizons included percentage grav-

el, sand, silt and clay; exchangeable K+, Na2+, Mg+ and

Ca+; soil conductivity, soil pH (H2O), soil depth and the

in situ soil compaction (together with gravimetric water

content) in accordance to the Soil Classification Work

Group (1991). Different aspects of direct or indirect

anthropogenic influences (human impact) such as mow-

ing, weeding, trampling, grazing, the use of chemicals

and erosion, were also recorded as thoroughly as possi-

ble. It was impossible, though, to quantify the intensities

of these anthropogenic influences, but terms such as

heavy trampling, heavy grazing and frequent mowing are

used in the text to express extreme levels of anthro-

pogenic influences.

The TWINSPAN classification algorithm (Hill

1979a) and the BBPC suite (Bezuidenhout et al. 1996)

were used for analysing the floristic data, as first approx-

imation, and subsequently Braun-Blanquet procedures

were used to refine these results. A phytosociological

table (Table 1) was obtained through these procedures.

This approach proved to produce ecologically reliable

results in many phytosociological studies in natural areas

216

Bothalia 28,2 (1998)

Bothalia 28,2 (1998)

217

218

Bothalia 28,2 (1998)

Bothalia 28,2 (1998)

219

(Behr & Bredenkamp 1988; Bredenkamp et al. 1989).

An additional software package (TURBOVEG) was

used for capture, processing, and presentation of phy-

tosociological data (Hennekens 1996a) as well as a visu-

al editor (MEGATAB) for phytosociological tables

(Elennekens 1996b). Introduced species are clearly

marked in Table 1. The occurrence of each species in a

specific stratum is indicated with a symbol, namely, tl

for a high tree layer (> 10 m), t2 for an intermediate tree

layer (5-10 m), t3 for a low tree layer (< 5 m), si for a

high shrub layer (> 2 m), s2 for a low shrub layer (< 2 m)

and hi for a herbaceous layer, in Table 1. Differentiation

between trees and shrubs was based on definitions pro-

posed by Edwards (1983). Two different numbering sys-

tems were used in Table 1, namely a releve number

which starts at 1 and indicates the releve numbers which

were used in this particular study and a TURBOVEG

number which indicates the unique releve number in the

South African phytosociological data base.

Species which were only encountered once or twice

during the study or which are not diagnostic for a specif-

ic community or group of communities are not included

in Table 1. These species are, however, included in Table

2 and are also mentioned in the results. An ordination

algorithm, DECORANA (Hill 1979b) was applied to the

floristic data to determine floristic relationships between

communities and to detect possible habitat and/or distur-

bance gradients associated with vegetation gradients.

Taxa names conform to those of Arnold & De Wet

(1993), but are updated to November 1996 according to

the PRECIS floristic data base of South Africa, managed

by the National Botanical Institute in Pretoria. Soil

nomenclature follows the classification of the Soil

Classification Working Group (1991). No attempt was

made to formally fix syntaxa names because this is nor-

mally avoided in detailed local studies. Formal syntax-

onomy will follow after the analyses of all the different

land use types in the urban area of Potchefstroom.

Formal syntaxa which are referred to in this study were

already described by other authors. The names of these

formal syntaxa are used as they were published without

any attempt to validate invalid names or to correct any

typographical errors.

RESULTS AND DISCUSSION

Classification

The classification resulted in the recognition of one

major community, 14 communities, nine subcommuni-

ties and three variants. The specific position of some of

the communities in the three marshland areas are shown

in Figure 2.

From the phytosociological table (Table 1) the fol-

lowing communities could be recognised:

1. Salix babylonica Woodland Community

1.1. Rhus pyroides Subcommunity

1.2. Populus x canescens Subcommunity

1.3 Populus wislizenii Subcommunity

1.4 Salix babylonica-Pennisetum clandestinum Sub-

community

2 Amaranthus hybridus-Pennisetum clandestinum

Ruderal Community

3 Berkheya radula-Themeda triandra Grassland

Community

4 Hyparrhenia hirta Grassland Community

5 Cyperus longus Major Wetland Community

5.1 Cichorium intybus-Xanthium strumarium Invasive

Community

5.1.1 Paspalum dilatatum Subcommunity

5 . 1 . 1 . 1 Cynodon dacty’lon Variant

5. 1.1. 2 Ambrosia psilostachya Variant

5. 1.1. 3 Senecio inornatus Variant

5.1.2 Cyperus marginatus Subcommunity

5.1.3 Sesbania bispinosa Subcommunity

5.2 Berula erecta Community

5.2.1 Rumex conglomerate Subcommunity

5.2.2 Falckia oblonga Subcommunity

5.3 Leersia hexandra Community

6 Cyperus fastigiatus-Paspalum distichum Wetland

Community

7 Schoenoplectus corymbosus Wetland Community

8 Eleocharis palustris Wetland Community

9 Carex acutiformis Wetland Community

10 Typha capensis Reedswamp Community

1 1 Phragmites australis Reedswamp Community

12 Azolla filiculoides Floating Community

Description of the plant communities

1 . Salix babylonica Woodland Community

This woodland community occurs on the banks of the

entire Mooi River, with the exception of parts of the Van

der Hoff Park marshland (Figure 2). The soils are deep

(> 120 cm) but shallower and with lower clay content

(< 50%) than that of the marshland communities. The

dominant species in this community is Salix babylonica

(species group D, Table 1), an introduced tree which is

naturalised along watercourses in southern Africa

(Henderson 1991). These trees are planted at dams and

along riverbanks, but its extensive occurrence along the

Mooi River is most likely due to self (vegetative) propa-

gation and dispersal by floodwaters.

Although these trees are aesthetically pleasing and

beneficial to man and the environment under certain cir-

cumstances, they also have disadvantages such as being

a potential threat to the conservation of indigenous ripar-

ian and wetland vegetation (Henderson 1991). The pro-

lific lateral root system of Salix babylonica is very effec-

tive in arresting soil erosion of the river banks, but it may

also lead to the damming up of small streams and possi-

bly even flooding during high rainfall seasons.

The diagnostic species of this community are those of

species group D (Table 1), and include introduced trees

such as Salix babylonica, Celtis sinensis, More alba and

Ligustrum lucidum, as well as the declared invader Melia

azedarach and the indigenous tree, Celtis africana.

Other diagnostic species are the shrubs Solanum nigrum

and Pyracantha coccinea, the climbers Araujia seri-

cifera and Merremia tridentata and the highly invasive

rhizomatous fern-like plant, Equisetum ramosissimum

(species group D, Table 1). Other species in this commu-

220

Bothalia 28,2 (1998)

TABLE 2. — List of species which occurred only a few times and have

relatively low cover-abundance values, as well as species not

restricted to specific communities in wetlands of Potchef-

stroom Municipal Area, North-West, South Africa

introduced species.

Bothalia 28,2 (1998)

221

* introduced species

nity are the turf grass, Pennisetum clandestinum (species

group G, Table 1), the indigenous but invasive shrub.

Asparagus laricinus (species group J, Table 1) and intro-

duced annuals such as Tagetes minuta and Bidens bipin-

nata (species group R, Table 1).

Although Eckhardt et al. (1993a) mentioned the pres-

ence of huge specimens of Salix babylonica along the

Klip River and in the Seekoeivlei area, northeastern Free

State, no Salix babylonica Community was described.

Four subcommunities can be distinguished in the

Salix babylonica Woodland Community, based on spe-

cies composition:

1.1. Rhus pyroides Subcommunity

This subcommunity is situated mainly on the city mar-

gin in relatively dry areas on the river banks, but occurs on

lower-lying areas which are seasonally waterlogged, as

well. It is commonly associated with vertic soils, for exam-

ple the Arcadia soil form. It is further characterised by the

presence of a dense shrub stratum (canopy cover of 70%

and up to 3 m tall) underneath the Salix babylonica trees.

The diagnostic shrubs are the indigenous species Rhus

pyroides and Maytenus heterophylla and the introduced

species Pyracantha angustifolia (species group A, Table 1).

Another shrub with relatively high cover in this subcom-

munity is Asparagus laricinus (species group J, Table 1)

which is an important component of bush encroachment in

disturbed areas in the Grassland Biome (Friedel 1987;

Cilliers & Bredenkamp in press). The tree stratum of 70%

canopy cover and height of 8 m is equally well developed

and consists of the trees mentioned in species group D

(Table 1). One other tree, Robinia pseudoacacia and a

number of herbaceous species which occurred only once in

this subcommunity are mentioned in Table 2. The herba-

ceous stratum is poorly developed with a canopy cover of

5% and a height of 90 cm. An average number of 17

species was recorded per sample plot of which 73% were

introduced and only 23% were therophytes.

1.2. Populus x canescens Subcommunity

On wetter areas on the river banks and on small

islands in the Mooi River, the prolific invader Populus x

canescens (species group B, Table 1) forms dense stands

FIGURE 2. — Schematic illustration

of some of the communities

in three marshland areas in

wetlands of Potchefstroom

Municipal Area, North-West,

South Africa. A, Arboretum;

B, Van der Hoff Park; C,

O PM. Prozesky Bird Sanc-

tuary. All other symbols ex-

plained in text.

222

Bothalia 28,2 (1998)

due to root suckering. Although Populus x canescens can

also arrest soil erosion, it has the same disadvantages as

Salix babylonica, namely a threat to indigenous wetland

communities and it may also cause flooding. The shrub

and herbaceous strata are poorly developed, in compari-

son with the tree stratum (canopy cover of 80% and 8 m

tall). Not many other species grow in this subcommuni-

ty, apart from climbers such as Araujia sericifera and

Merremia tridentata (species group D, Table 1), and the

shrub Asparagus laricinus (species group J, Table 1). An

average number of nine species was recorded per sample

plot of which 81% were introduced and only 12% were

therophytes.

1.3. Populus wislizenii Subcommunity

This subcommunity is characterised by the presence

of the diagnostic tree Populus wislizenii (species group

C, Table 1), a very large and conspicuous shade tree

which is planted in adjacent parks. These trees propagate

by means of cuttings and do not show any great tenden-

cy to sucker and therefore, it is not sure how they are

spreading into the Salix babylonica Community from the

parks. This subcommunity, established on the higher

river banks, which are built up by man or which is the

result of deep streambed incission, rarely occurs at water

level. It is situated mainly on the man-made or anthro-

pogenous soil type, Witbank, but still with a clay per-

centage of close to 40%. The very well-developed tree

stratum with a canopy cover of nearly 100% and a max-

imum height of 12 m, makes this subcommunity very

conspicuous. The shrub and herbaceous strata are poorly

developed. An average of 14 species per sample plot was

recorded for this community of which 76% were intro-

duced and only 7% were therophytes.

1 .4. Salix babylonica-Pennisetum clandestinum Sub-

community

The Salix babylonica-Pennisetum clandestinum Sub-

community occurs mainly along the river banks in parks,

sports grounds and other recreational areas, but was

occasionally encountered in some of the marshy areas as

well. The main soil type on which this subcommunity

occurs is the vertic Arcadia soil form, but it is also asso-

ciated with the anthropogenous soil form, Witbank. The

soils are relatively deep, with a high clay content and is

sometimes heavily compacted (up to 4 kg/m2). Although

no diagnostic species occur in this subcommunity it is

characterised by the absence of the species of species

groups A, B and C (Table 1) and a relatively high cover

of the turf grass, Pennisetum clandestinum (species

group G, Table 1), which invades from adjacent lawns.

The dominant tree is Salix babylonica in the well-devel-

oped tree stratum (canopy cover of 65% and up to 8 m

high). The total lack of young trees establishing in this

subcommunity is worth mentioning. The shrub stratum

which consists mainly of Asparagus laricinus close to

the stems of the trees, has a canopy cover of 20% and an

average height of 1.5 m. Apart from Pennisetum clan-

destinum, the well-developed herbaceous stratum (70%

canopy cover) consists mainly of introduced forbs such

as Taraxacum officinale (Table 2) and the annuals, Ipo-

moea purpurea, Bidens pilosa (species group F, Table 1 ),

Tagetes minuta and Bidens bipinnata (species group R,

Table 1). Parts of this subcommunity are mown every

4-6 weeks together with adjacent lawns, according to

Mr P. Labuschagne, Department of Parks and Recrea-

tion, Potchefstroom Municipality. An average number of

15 species per sample plot was recorded of which 64%

were introduced and 13% were therophytes.

2. Amaranthus hybridus-Pennisetum clandestinum Ru-

deral Community

This community is situated on alluvial deposits in the

beds of parts of the Wasgoedspruit as well as on its gen-

tly sloping inner banks. It is also developing on debris

which is the result of an attempt to cover the beds of

some areas with concrete. The Wasgoedspruit drains

some of the natural areas on the west side of Potchef-

stroom, as well as industrial areas where a number of

chemical plants are situated. Sporadic chemical pollution

of this drainage canal was, therefore, encountered in the

past. The pollution is probably one of the reasons why no

true wetland or reedswamp communities developed in

this area. The Wasgoedspruit is mostly dry, but is heavi-

ly disturbed during floods, making this a true ruderal

community. The Amaranthus hybridus-Pennisetum

clandestinum Ruderal Community is associated with the

Witbank (anthropogenic) soil form with a clay content of

< 40%, and which are at places rather heavily compact-

ed (4.5 kg/m2).

No tree or shrub stratum exists, but the herbaceous

stratum is relatively well developed with a canopy cover

of about 50% and an average height of 1 m. The turf

grass Pennisetum clandestinum (species group G, Table

1) which invades from adjacent parks, is the dominant

species. The diagnostic species include introduced forbs

and grasses, such as the annuals Amaranthus hybridus

and Datura stramonium, which is also a declared weed

(Wells et al. 1986) and the perennials Modiola carolini-

ana and Sorghum halepense, as well as the indigenous

forbs, Solatium panduriforme and Commelina beng-

halensis (species group E, Table 1). Other species which

were only recorded once in this study are indigenous

grasses such as Chloris virgata, Urochloa mosambicen-

sis, Brachiaria eruciformis, Echinocloa colona and

Aristida congesta (Table 2). Other species in this com-

munity are the highly invasive pioneer grass Cynodon

dactylon and the conspicuous tall-growing annuals,

Tagetes minuta and Bidens bipinnata (species group R,

Table 1). An average number of 14 species was recorded

per sample plot, of which 46% were introduced and 35%

were therophytes.

3. Berkheya radula-Themeda triandra Grassland Com-

munity

The Berkheya radula-Themeda triandra Grassland

Community forms a transitional area between adjacent

higher-lying grasslands and the lower-lying wetland

communities in the study area (Figure 2). Although situ-

ated in bottomlands, it is still much higher than the

marshlands. This community occurs on clay loam to

sandy clay loam soil types, as well as on duplex soils

with a high clay content (> 50%), representing the

Bothalia 28,2 (1998)

223

Valsrivier and Rensburg soil forms, and occasionally on

rocky soils of the Glenrosa soil form.

The diagnostic species of this community are those of

species group H (Table 1). It includes the two dominant

species, the indigenous grass Themeda triandra, which is

widespread in the Grassland Biome, and the indigenous

forb Berkheya rodula which is characteristic of soils with

a high clay content. Other diagnostic species are all

indigenous grasses such as Eragrostis chloromelas, Di-

gitaria eriantha, Panicum stapfianum and Cymbopogon

plurinodis. Species which are also abundant in this com-

munity are the indigenous grass, Setaria sphacelata

(species group U, Table 1) and the introduced forb,

Physalis viscosa (species group K, Table 1 ). Forbs which

are typically found in the adjacent grasslands, such as

Crabbea angustifolia , Rhynchosia totta, Corchorus asple-

nifolius and Monsonia angustifolia (Table 2) were also

encountered in this community. This community is fur-

ther characterised by the encroachment of the shrub.

Asparagus laricinus (species group J, Table 1) forming a

canopy cover of up to 12% in some areas. No tree stra-

tum exists in this community. An average number of 13

species per sample plot was recorded of which 25% were

introduced and 11% were therophytes.

This community resembles, to a certain extent, the

Themeda triandra Variant of the Eragrostidetum planae,

an association described for the seasonally wet bottom-

lands of the Be land type in which Potchefstroom is sit-

uated (Bezuidenhout & Bredenkamp 1991). A similar

grassland community which represents a transitional

zone between relatively dry and wet grasslands, the

Themeda triandra-Eragrostis plana dry/wet grassland

was described by Eckhardt et al. (1993b) for the north-

eastern Free State.

4. Hyparrhenia hirta Grassland Community

The Hyparrhenia hirta Grassland Community is also

situated on the seasonally flooded bottomlands adjacent

to marshlands, but usually higher than them (Figure 2). It

is situated next to the Berkheya radula-Themeda triandra

Community in some areas but on the lower-lying wetter

parts on soils of the Valsrivier and Rensburg soil forms.

The dominant and also one of the diagnostic species

is the indigenous grass Hyparrhenia hirta (species group

I, Table 1). Other species in this community are those of

species groups J, K, Q, R and U (Table 1). The herba-

ceous stratum is well developed with a canopy cover of

70% and a height of 1.5 m. No tree stratum exists in this

community, but the shrub stratum with Asparagus larici-

nus and Rhus pyroides is relatively well developed

(canopy cover of up to 30% and 1.8 m tall). The abun-

dance of the invasive grass, Cynodon dactylon and the

introduced annuals, Tagetes minuta and Bidens bipinna-

ta (species group R, Table 1) is an indication of the grad-

ual degradation of this community. An average number

of 12 species was recorded per sample plot of which 47%

were introduced and 23% were therophytes.

In a syntaxonomical and synecological study of the

western Transvaal Grasslands, Bezuidenhout (1993) de-

scribed a number of wetland communities where Hy-

parrhenia hirta is dominant, and which form part of the

Eragrostido planae-Hyparrhenietea hirtae (Bezui-

denhout et al. 1994a). The Hyparrhenia hirta Grassland

Community, currently described, shows habitat and flor-

istic resemblances to the Eragrostidetum planae (Bezui-

denhout & Bredenkamp 1991) and the Hyparrhenio hir-

tae-Eragrostidetum planae (Bezuidenhout et al. 1994b),

but with much lower species diversity.

5. Cyperus longus Major Wetland Community

This major wetland community is the largest of all the

communities, fringing the reedswamps and is mainly sit-

uated in two of the three marshlands in the study area,

namely Van der Hoff Park and Prozesky (Figures 1 and

2). It occurs in a very wide spectrum of habitats, namely

from dry to seasonally wet to waterlogged areas, as well

as in different stages of habitat degradation. All these

variations are covered by the different communities, sub-

communities and variants.

With the exception of isolated and scattered occur-

rence of trees and shrubs, no real tree and shrub strata is

present in this community. The herbaceous stratum is

well developed. The diagnostic species of this major

wetland community are those of species group X (Table

1). The most conspicuous of these species are the tall-

growing perennial sedge, Cyperus longus , which shows

a great degree of plasticity as it is encountered in aquat-

ic, helophytic and mesophytic habitats. Because of the

effects of grazing and trampling, accompanied by the

development of habitats better suited for other species,

Cyperus longus is disappearing from some areas where it

was dominant some years ago. The diagnostic species

include true wetland species which are also diminishing

in certain areas, such as Ranunculus multifidus, Falckia

oblonga , Stenotaphrum secundatum and Crinum bulbi-

spermum, and invasive species such as Paspalum dilata-

tum and Senecio inornatus (species group X, Table 1).

A similar broad major wetland community, the

Echinochloa holubii-Cyperus longus Wetland was

described by Kooij et al. (1991) for watercourses, river-

banks, valley flats, flood plains and stream channels in

the Kroonstad area, Free State. Although the Cyperus

longus Major Wetland Community, currently described,

is situated on the same soil types it differs from the com-

munity described by Kooij et al. (1991) with respect to

species composition. The difference in species composi-

tion is probably due to the type and intensity of distur-

bances in urban areas.

The Cyperus longus Major Wetland Community is

divided into the following communities based on habitat

and type and intensity of the disturbances:

5.1. Cichorium intybus-Xanthium strumarium Invasive

Community

This community is highly disturbed and is situated in

the heavily grazed areas in the Van der Hoff Park and

Prozesky marshland areas (Figure 2). It is characterised

by the extensive invasion of the dominant species, the

grass Cynodon dactylon (species group R, Table 1),

224

Bothalia 28,2 (1998)

which could suggest an enriched nitrogen status of the

soil (Kooij et al., 1991), probably due to biotic effects,

e.g. from animal excretions. This community is further

characterised by the decrease of the sedge, Cyperus

longus (species group X, Table I ), which probably was

the dominant species in this area. The diagnostic species

of this community are the introduced forbs, Cichorium

intybus and the declared weed Xanthium strumcirium

(Wells et al. 1986) as well as the indigenous, perennial

grasses Helictotrichon turgidulum and Eragrostis plana

(species group P, Table 1). Other species which occur in

this community are those of species groups Q and U

(Table 1).

Three subcommunities can be distinguished in this

community based on water content of the soil and degree

of disturbance.

5.1.1. Paspalum dilatatum Subcommunity

This subcommunity is situated on the higher-lying,

somewhat drier areas occupied by the Cichorium inty-

bus-Xanthium strumarium Invasive Community (Figure

2). No diagnostic species occur but it is characterised by

the abundance of the conspicuous introduced grass,

Paspalum dilatatum (species group X, Table 1).

Three variants, which are forming interconnected

mosaics or patches with each other, could be distin-

guished in the Paspalum dilatatum Subcommunity:

5. 1.1.1. Cynodon dactylon Variant

The Cynodon dactylon Variant occurs on the flatter

areas in footpaths (in the O.P.M. Prozesky Bird

Sanctuary) and near entrance gates which are heavily

trampled by man and grazing cattle and horses. The foot-

paths are sometimes mown as well. This variant is asso-

ciated with the Witbank (anthropogenic) and Rensburg

soil forms with deeply cracking, vertic clays, and is also

heavily compacted (> 4.5 kg/m2). With the exception of

a number of herbaceous species which were recorded

once or twice (Table 2), no diagnostic species were

encountered in this variant. It is characterised, however,

by the absence of species of species groups L and M

(Table 1), the total dominance of Cynodon dactylon and

a low species diversity in comparison with the other vari-

ants. Other species which occur are typically ruderal

species which can probably withstand the effect of tram-

pling and grazing such as Physalis viscosa (species

group K, Table 1), Plantago lanceolata and the prostrate

indigenous forb, Conyza podocephala (species group U,

Table 1). An average of 12 species was recorded per

sample plot of which 44% were introduced and 20%

were therophytes.

5.1. 1.2. Ambrosia psilostachya Variant

The Ambrosia psilostachya Variant of the Paspalum

dilatatum Subcommunity is mainly associated with the

drier, overgrazed areas in the wetlands of Prozesky, on

the same soil types as the Cynodon dactylon Variant. The

diagnostic species are the forbs Ambrosia psilostachya

and Oxalis pes-caprae and the indigenous grasses

Eragrostis micrantha and Andropogon schirensis (spe-

cies group L, Table 1 ). Although no tree or shrub stra-

tums exists in this variant, the indigenous shrub

Asclepias decipiens and the introduced tree Gleditsia tri-

acanthos (Table 2) have a scattered distribution. Due to

the high levels of disturbance an unusually high average

number for wetland areas of 22 species per sample plot

was recorded, of which 42% were introduced and only

12% were therophytes.

5. 1.1.3. Senecio inornatus Variant

The Senecio inornatus Variant is encountered in the

slightly lower and wetter parts of the higher-lying areas in

the wetlands of the O.P.M. Prozesky Bird Sanctuary.

Although the diagnostic species is the indigenous shrub

Gomphocarpus fruticosus (species group M, Table 1), the

variant is better characterised by its dominant species,

namely the introduced, prostrate grass Cynodon dactylon

and the indigenous tall-growing forb, Senecio inornatus.

Although Senecio inornatus is also present in other wet-

land communities, it develops here into relatively dense

patches of conspicuous plants due to their height (2 m and

higher) and their corymbs of brightly yellow coloured

capitula. The sedge, Cyperus longus, which was the dom-

inant species in this area about two to three years ago, is

totally absent in this variant. An average number of 15

species was recorded per sample plot of which 52% were

introduced and 16% were therophytes.

5.1.2. Cyperus marginatus Subcommunity

This subcommunity is associated with lower-lying,

seasonally waterlogged areas (Figure 2) on the Rensburg

soil form with soil compaction of up to 5 kg/m2 or on the

Bloemdal soil form with soil compaction of less than 2

kg/m2. Although the annual, introduced forb Physalis

cmgulata (species group N, Table 1) is the only diagnostic

species, this subcommunity is better characterised by the

presence of the tall-growing sedge, Cyperus marginatus

(species group W, Table 1). The invasive grass, Cynodon

dactylon and the other tall-growing sedge, Cyperus longus

are still the dominant species. Louw (1951) described

Cyperus marginatus as a robust species which plays a

major role in succession in the Vaal River, but which is rel-

atively unimportant elsewhere. Although this species was

found in the Potchefstroom urban wetlands, the subcom-

munity in which it exists seems to diminish. This is prob-

ably due to the effects of trampling and grazing, because it

is only situated in relatively small patches in lower-lying,

wetter areas. The abundance of introduced annuals such as

Physalis angulata (species group N, Table 1), Tagetes

minuta and Bidens bipinnata and the declared weed

Xanthium strumarium (species group R, Table 1) (Wells et

al. 1986) is a further indication of the degraded state of the

Cyperus marginatus Subcommunity. An average number

of 1 5 species was recorded per sample plot of which 5 1 %

were introduced and 17% were therophytes.

5. 1 .3. Sesbania bispinosa Subcommunity

This subcommunity represents a more degraded form

of the Cyperus marginatus Subcommunity. The diagnos-

tic species are the introduced, annual to biannual inva-

Bothalia 28.2 (1998)

225

sive shrub, Sesbania bispinosa and the introduced forb,

Flaveria bidentis (species group O, Table 1). An average

number of 11 species was recorded per sample plot of

which 53% were introduced and 30% were therophytes.

5.2. Benda erecta Community

The Berula erecta Community is found in seasonally

wet bottomland areas which were at the time of the study

partly under standing water of up to 10 cm (Figure 2).

The soils are quite deep, undrained vertic soils with a

high clay content (> 60%). The community is associated

with areas close to or adjacent to the river, in the absence

of a tree stratum.

The diagnostic species of this community are the low-

growing, indigenous forbs, Berula erecta and Mentha

aquatica (species group T, Table 1). Other species are the

tall-growing sedge, Cyperus longus, the low-growing,

mat-forming forb, Falckia oblonga and the indigenous

vlei grass, Stenotaphrum secundatum (species group X,

Table 1 ). The Berula erecta Community is further char-

acterised by the absence of the species of species groups

P, Q and R (Table 1), although Cynodon dactylon invades

into the margins of this community. The vegetation

seems to be relatively undisturbed, but trampling by the

grazing animals keeps the wet soil in a permanently pud-

dled condition, which favours the establishment of

Phragmites australis and Typha capensis from the adja-

cent reedswamps (Communities 10 and 11). Although

not recorded during the study, blooms of the very inva-

sive water fern, Azolla filiculoides can sometimes be

found in these puddles, as well. The indigenous geo-

phyte, Crinum bulbispermum (species group X, Table 1),

although not abundant, has a scattered distribution in this

community.

Two subcommunities could be distinguished in this

community.

5.2.1. Rumex conglomeratus Subcommunity

On slightly lower-lying areas of the Berula erecta

Community, where water seems to stagnate when the

rainfall is high (Figure 2), small clumps of the perennial

forb, Rumex conglomeratus (species group S, Table 1)

establish. An average number of only eight species per

sample plot was recorded of which none were introduced

and only 8% were therophytes.

5.2.2. Falckia oblonga Subcommunity

Although no diagnostic species were recorded for this

subcommunity, it is characterised by the formation of

small but dense mats of the small, rhizomatous forb,

Falckia oblonga (species group X, Table 1) in areas

where water drainage is faster (Figure 2), and where

Rumex conglomeratus is absent. Although Falckia oblon-

ga was also found on bare areas on the higher slopes of

dry vleis, as was suggested by Louw (1951), they are

always dominated by the invasive grass, Cynodon dacty-

lon, in these areas. It is possible that Falckia oblonga

finds refuge in the wetter areas, where Cynodon dactylon

has not established so extensively. An average of 10

species per sample plot was recorded of which 34% were

introduced and only 4% were therophytes.

5.3. Leersia hexandra Wetland Community

This community was encountered in shallow depres-

sions or pools which are seasonally waterlogged in the

O.RM. Prozesky Bird Sanctuary and occasionally in

water on the river’s edge in the Arboretum, but never in

deep water (Figure 2). Although no diagnostic species

were recorded, this community is characterised by the

high cover of rhizomatous grass species such as Leersia

hexandra (species group Z, Table 1) and Stenotaphrum

secundatum (species group X, Table 1). Other species

include the tall-growing sedge, Cyperus longus and the

forb Ranunculus multifidus (species group X, Table 1),

as well as the low-growing sedge, Carex schlechteri

(species group V, Table 1). An average number of 10

species was recorded per sample plot of which 33% were

introduced and only 13% were therophytes.

A similar community, the Leersia hexandra-Schoeno-

plectus paludicola Wetland was described by Fuls et al.

(1992) for the northern Free State.

6. Cyperus fastigiatus-Paspalum distichum Wetland

Community

This community sometimes occurs on the river’s

edge, but is mainly associated with shallow to deep

drainage depressions or pools which are seasonally

waterlogged (Figure 2). In pools where water is stagnant

for long periods, this community is mainly situated at the

edges. The soils on which this community can be found

are vertic soils of the Rensburg soil form, but occasion-

ally also on the Champagne soil form.

The diagnostic species are the tall-growing sedge,

Cyperus fastigiatus (species group Y, Table 1). The dom-

inant species is the rhizomatous grass Paspalum dis-

tichum (species group BB, Table 1) which form very

dense mats, and tends to grow in deeper water as well.

Louw (1951) made the observation in the Vaal River that

when this grass migrates to deeper water, its robustness

increases and it behaves like a floating aquatic species.

This community shows an affinity with the Leersia hexa-

ndra Community of the Cyperus longus Major Wetland

Community, because of the presence of the grass, Leer-

sia hexandra and the forb Rumex lanceolatus (species

group Z, Table 1). The Cyperus fastigiatus-Paspalum

distichum Wetland Community differs, however, from

the Leersia hexandra Community with respect to the

habitat and the fact that the tall-growing sedge Cyperus

longus, is replaced by another tall-growing sedge,

Cyperus fastigiatus.

Other species occurring in this community which is

worth mentioning are the water fern Marsilea capensis

and the tall-growing forb Persicaria lapathifolia (Table

2: under the heading, no specific communities). An aver-

age number of only six species was recorded per sample

plot of which 26% were introduced and none were thero-

phytes.

226

Bothalia 28,2 (1998)

A similar community with an equally low species

diversity, the Paspalum distichum Community associat-

ed with marshes, was described in the Grootvlei area in

the eastern Transvaal (Myburgh et al. 1995).

7. Schoenoplectus corymbosus Community

This community is situated in deeper water in the

Mooi River as well as in deep depressions or pools with

stagnant water on the Champagne soil form (Figure 2).

The diagnostic species is the tall-growing, robust sedge

Schoenoplectus corymbosus (species group AA, Table

1). Occasionally the rhizomatous grass, Paspalum dis-

tichum (species group BB, Table 1) invades from the

river banks into this community. No other species were

recorded in this community.

8. Eleocharis palustris Wetland Community

This community is relatively undisturbed and occurs

only on the city margin, in the Arboretum. It is situated

on vertic soils which are seasonally waterlogged in bot-

tomland areas fringing the reedswamps (Figure 2).

The diagnostic species of this community are the

sedge, Eleocharis palustris and the forbs, Chironia pal-

ustris, Samolus valerandi and Lobelia thermalis (species

group CC, Table I ). The forb Chironia palustris, when

flowering in summer, transforms this specific communi-

ty into a conspicuous and aesthetically pleasing pink

mass, which was encountered nowhere else in the study

area. Although the Eleocharis palustris Community is

situated in the same type of habitat in which the Berulci

erecta Community occurs in other wetlands, there is no

real affinity between these two communities regarding

species composition. Eleocharis palustris, does, however,

have an isolated distribution in the Berula erecta Com-

munity. An average number of 14 species was recorded

per sample plot of which 17% were introduced and 5%

were therophytes.

9. Carex acutiformis Wetland Community

This particularly species-poor community occurs in

two of the three marshland areas (Figure 2), namely the

Arboretum and Van der Hoff Park, as well as in other

areas along the Mooi River, where it is associated with

the Typha capensis Reedswamp Community. It either

forms a narrow fringe along the reedswamps in water-

logged areas on the Champagne soil form, or it estab-

lishes in dense clumps on seasonally waterlogged, high-

er-lying areas on the Rensburg soil form. This communi-

ty seems to invade the Berula erecta Community and the

Eleocharis palustris Community.

The diagnostic and dominant, and most of the time the

only species occurring in this community is the sedge,

Carex acutiformis (species group DD, Table 1 ). All other

species which were recorded are species from adjacent

wetland communities.

10. Typha capensis Reedswamp Community

This community which occupies large areas on sea-

sonally or permanently waterlogged soils (Figure 2) was

regarded by Louw (1951) as one of the important con-

stituents of the reedswamps, together with Phragmites

australis. The Typha capensis Community, when occur-

ring together with the Phragmites australis Community,

is always situated on its outside in shallower water, act-

ing as a pioneer in reedswamp succession in some cases.

The Typha capensis Community seems to expand rapid-

ly into other communities such as the Berula erecta

Community. It occurs on the Champagne soil form but

also on vertic soils such as the Rensburg and Arcadia soil

forms. It was observed in wetlands all over the world that

small-seeded anemochorous plants such as Typha

species, colonise mudflats during periods of drawdown,

as well as areas that have been recently disturbed

(Ellison & Bedford 1995).

The diagnostic species of this community is the indige-

nous, rhizomatous bulrush Typha capensis and to a lesser

extent, the forb Hydrocotyle verticillata (species group

EE, Table 1). Other species which only occurred once in

this study includes the indigenous species Kniphofia ensi-

folia (Table 2: under the heading, no specific communi-

ties), which is situated at the edge of the Typha capensis

Reedswamp Community. An average number of only six

species was recorded per sample plot of which 27% were

introduced and 17% were therophytes.

1 1 . Phragmites australis Reedswamp Community

The Phragmites australis Reedswamp Community is

mainly situated in permanently waterlogged areas, some-

times in water as deep as 2 m, in the river bed and in all

three marshland areas (Figure 2). It colonises, together

with the Typha capensis Community, also seasonally

waterlogged and even higher-lying bare areas along the

river. These bare areas are formed due to the destabilisa-

tion of substrates in the inner city after heavy rains, when

water collects in larger volumes and passes through more

quickly than usual.

The dominant and only diagnostic species of this

community is the tall-growing reed Phragmites australis

(species group FF, Table 1). Other species occurring on

the edge of this community are ruderal species or species

of adjacent wetland communities. This community can

easily be distinguished from the other community in the

reedswamp, the Typha capensis Commnunity, as it grows

much higher (> 3 m) and is much denser in certain areas.

The very dense growth of Phragmites australis may

block waterways which lead to water stagnation and pro-

vide breeding places for mosquitoes and bilharzia snails

(Marks et al. 1994; Bromilow 1995). An average number

of five species was recorded per sample plot of which

55% were introduced and 44% were therophytes.

A similar community, the Phragmites australis Vlei

was described by Bloem et al. (1993) in the North-east-

ern Sandy Highvcld, Transvaal..

12. Azolla filiculoides Floating Community

The Azolla filiculoides Floating Community invades

stagnant water in depressions, pools and drainage canals

in some of the wetland areas in Potchefstroom (Figure

2). The only species in this community is the perennial,

mat-forming, free-floating, aquatic fern Azolla filicu-

Bothalia 28,2 (1998)

227

FIGURE 3. — Relative positions of

plant communities along first

two axes of ordination of wet-

land plant communities in Pot-

chefstroom Municipal Area,

North-West, South Africa.

hides (species group GG, Table 1). This species is a

declared invader and is regarded as one of the most dan-

gerous invaders of water bodies in temperate regions in

southern Africa (Henderson 1995). This species was not

mentioned by Louw (1951) in his study of the vegetation

of the Potchefstroom area, but none of the floating or

submerged plants observed by him were recorded in the

study area.

Ordination

Figure 3 represents the distribution of 98 of the 102

releves along the first and second axes of a DECORANA

ordination. Although the releves of the Carex acutiformis

Wetland Community and the Azolla filiculoides Floating

Community are excluded from Figure 3, they show a

very clear discontinuity with each other and with all the

other communities. This discontinuity was verified by

the ordination of the total data set which is not presented

in this paper.

With the exception of the subcommunities of the Salix

babylonica Community (Community 1) and the subcom-

munities and variants of the Cichorium intybus-Xan-

thium strumarium Invasive Community (Community

5.1), the different plant units are restricted to specific

spatial areas as shown in the scatter diagram (Figure 3).

The diagram also illustrates a gradient along ordination

axis 1 which could be related to depth of the soil water

table (position in the whole wetland system, and ability

to establish in deep water). Depth of the water table

could have a major influence on the distribution pattern

of vegetation (Eckhardt et at. 1993b). The communities

found under extreme conditions, namely away from

water or in water, occur on the periphery of the diagram

(Figure 3). The grassland communities (Communities 3

and 4) which occur on the higher-lying areas, fringing

the marshlands and the Salix babylonica Community

(Community 1) which occurs on the river banks are situ-

ated at the left of the scatter diagram. Communities

which typically establish in shallow to deep water

(Commmunities 6, 7, 10 and 1 1) are situated to the right

of the diagram. The Cyperus longus Major Wetland

Community (Community 5) is situated in an intermedi-

ate position on the scatter diagram (Figure 3) which indi-

cates that the communities, subcommunities and variants

of this major community may establish on higher-lying

drier areas as well as in lower-lying areas which are sea-

sonally waterlogged. Another community in an interme-

diate position is the Amaranthus hybridus-Pennisetum

clandestinum Ruderal Community (Community 2)

which is situated very close to the other degraded com-

munity, the Cichorium intybus—Xanthium strumarium

Invasive Community (Community 5.1) (Figure 3). The

Eleocharis palustris Wetland Community (Community

8) which occurs in the same type of habitat than the

Berula erecta Community (Community 5.2) but in less

disturbed marshlands, is also situated in an intermediate

position, close to the Berula erecta Community.

In ordinations which were separately performed on

the Salix babylonica Woodland Community (Commu-

nity 1 ) and the Cichorium intybus-Xanthium strumarium

Invasive Community (Community 5.1), none of the sub-

communities and variants were clearly distinguished

from each other.

CONCLUSION

The establishment of many of the wetland communi-

ties is the result of direct or indirect anthropogenic influ-

ences. The large areas covered by communities such as

the Salix babylonica Community show the great long-

term impact of man on the structure and species compo-

sition of wetland plant communities. The yearly expan-

sion of the Cynodon dactylon Invasive Community indi-

cates the degraded condition of the urban wetlands in

Potchefstroom, mainly because of the indiscriminate use

of these areas for cattle and horse grazing.

Some of the communities described in urban wetlands

in this study are to a certain extent, similar to those wet-

land communities described by Kooij et al. (1991); Fuls

et al. (1992); Bloem et al. (1993); Eckhardt et al.

228

Bothalia 28,2 (1998)

(1993a); Myburgh et al. (1995) and Smit et al. (1995), in

natural wetland areas. The range and intensity of distur-

bances in certain areas of the Potchcfstroom urban wet-

land, as well as the scale on which the current study was

done, probably permitted the recognition of more com-

munities which may also form small mosaics with each

other. In none of the mentioned studies on natural wet-

lands, was the abundance of the invasive grass, Cynodon

dactylon so profound, as in the current study.

Variations in those plant communities which could be

related to specific environmental conditions are mainly

ascribed to river bank wetness, period of wetness, depth

of soil water table, soil type and drainage tempo of soil.

With the exception of the anthropogenous soil form,

Witbank, all the other soils are deep, poorly drained soils

with high clay contents. No correlation between the dif-

ferent plant communities and data from either physical or

chemical soil analyses, were found.

The present study should be used as the basis for

future management and conservation of urban wetlands

in Potchefstroom, and it should also lead to the develop-

ment of a Metropolitan Open Space System (MOSS) net-

work for Potchefstroom. The development of such a net-

work, together with the involvement of the public in the

conservation of the wetlands may prove to be the salva-

tion of the Potchefstroom wetlands. One important

aspect that should immediately be addressed in the man-

agement of these wetlands is the termination of all mow-

ing and grazing practises, and the careful monitoring of

any successional changes in the vegetation, to be able to

assess the exact influence these grazing and mowing

practises have on the vegetation.

The classification and description of the wetland

communities in the Potchefstroom Municipal Area also

serve as a basis for further studies in urban wetlands.

Future research must concentrate more specifically on

the conservation status and ecological importance of cer-

tain species and plant communities, the vegetation

dynamics of wetland ecosystems under specific human

influences and habitat changes, and the effect of certain

toxic agents (ecotoxicology) on wetlands. All this infor-

mation can eventually be used in the development of a

model, such as the one of Ellison & Bedford (1995),

which can predict the consequences of anthropogenic

disturbances in wetlands.

ACKNOWLEDGEMENTS

The first author wishes to thank the Potchefstroom

University for funding this project; Prof. A.J.H. Pieterse,

Head of the Department of Plant and Soil Sciences,

Potchefstroom University for valuable comments on the

text; Dr H. Bezuidenhout, Scientific Services, National

Parks Board for the provision of the BBPC programme

which was used for the analysis of the floristic data; Mr

E van Wyk, Department of Plant and Soil Sciences,

Potchefstroom University for technical assistance; and

the National Herbarium, Pretoria and Mr B. Ubhink,

Botanical Garden, Potchefstroom University, for plant

identification.

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Bothalia 28,2: 231-238 (1998)

Cytogenetic studies in the genus Pentaschistis (Poaceae: Arundinoideae)

K.C. KLOPPER*, JJ. SPIES*f and B. VISSER*

Keywords: chromosomes, meiosis, Pentaschistis, polyploidy

ABSTRACT

Cytogenetic studies of 45 specimens, representing 16 taxa of the genus Pentaschistis (Nees) Spach confirmed two basic

chromosome numbers (x = 7, 13) for the genus. Chromosome numbers for five species and one subspecies are described

for the first time, i.e. P. airoides subsp . jugorum (n = 2x = 14), P. colorata (n = 2x = 14), P. densifolia (n = x = 7), P. lima

(n = 6x = ± 42), P. rigidissima (n = x = 7, n = 3x = 21) and P. viscidula (n = 3x = 21). Polyploidy occurs frequently and

new ploidy levels are described in four of the species, namely P. airoides (Nees) Stapf subsp, airoides (n = 3x = 21), P. cir-

rhulosa (Thunb.) McClean (n = x = 7), P. eriostoma (Nees) Stapf (n = 3x = 39+0-4B) and P. rupestris (n = 4x = 28). The

majority of species form young polyploid complexes. There seems to be no correlation between cytogenetic data and mor-

phological groupings within Pentaschistis.

INTRODUCTION

The genus Pentaschistis (Nees) Spach consists of 68

species (Linder & Ellis 1990) of which 57 are indigenous

and 40 endemic to South Africa (Gibbs Russell et al.

1990). Cytogenetic studies reveal that Pentaschistis has

basic chromosome numbers of 7 and 13 (De Wet 1954;

Hedberg 1957; De Wet 1960; Hedberg & Hedberg 1977;

Davidse et al. 1986; Spies & Du Plessis 1988; Du Plessis

& Spies 1988, 1992; Spies et al. 1994).

The aim of this study was to determine whether the

cytogenetic studies support the morphological groupings

of Linder & Ellis (1990) and Ellis & Linder (1990); to

determine the degree of polyploidy in the genus and to

use ploidy levels to determine the age of the polyploid

complexes in the genus.

MATERIALS AND METHODS

The material used during this study was collected in

the field. Voucher herbarium specimens are housed in the

Geo Potts Herbarium, Bloemfontein (BLFU) and/or in

the National Herbarium. Pretoria (PRE).

Young inflorescences were collected and fixed in

Carnoy’s fixative (Carnoy 1886). The fixative was

replaced by 70% (v/v) ethanol 24-48 hours after fixa-

tion. Anthers were squashed in 2% (m/v) aceto-carmine

(Darlington & La Cour 1976). Slides were permanently

mounted by freezing them with liquid COi (Bowen

1956), followed by dehydration in ethanol and mounting

in Euparal. At least twenty cells per specimen were stud-

ied for each meiotic stage, except where otherwise indi-

cated.

With the annotation of chromosome numbers we fol-

lowed the example set by the series 'Index to plant chro-

mosome numbers’ (Goldblatt 1981), where chromosome

* Department of Botany and Genetics, University of the Orange Free

State, P.O Box 339, 9300 Bloemfontein,

t To whom correspondence should be addressed.

MS. received: 1997-09-15.

numbers derived from meiotic studies are presented as

the gametic (n) number and chromosomes from mitotic

studies as the somatic (2n) chromosome number. The

average number of chiasmata per bivalent was consid-

ered to be the number of chiasmata per chromosome and

were calculated by dividing the mathematical average of

the chiasmata per cell by the haploid chromosome num-

ber of the plant. Chromosome configurations were cal-

culated as the average number of each configuration per

cell. The number of B-chromosomes is presented as the

minimum and maximum number of B-chromosomes

observed per cell. The numerical values of the chromo-

some abnormalities were obtained by calculating the

average number of a certain abnormality per cell.

Genomic relationships were calculated according to the

models of Kimber & Alonso (1981).

RESULTS AND DISCUSSION

Forty five specimens, representing the different

groups in Pentaschistis, except Group 5, have been stud-

ied. The results are presented alphabetically in table form

for each group (Table 1).

Number of specimens per ploidy level

All published ploidy levels (Hedberg 1957; De Wet

1960; Tateoka 1965a & b; Davidse et al. 1986; Du

Plessis & Spies 1988; Spies & Du Plessis 1988; Du

Plessis & Spies 1992; Spies et al. 1994) were used to

determine the number of specimens per ploidy level

among the different species of the genus Pentaschistis.

The frequency of specimens per ploidy level was also

plotted for the two different basic chromosome numbers

in Pentaschistis (Figure 4).

Meiotic analyses of the different Pentaschistis species

revealed haploid chromosome numbers of seven, multiples

of seven (Figures 1, 2), thirteen and multiples thereof

(Figure 3). The low number of species with a basic chromo-

some number of thirteen can be attributed to the fact that

only specimens from South Africa have been studied. All

species with a basic chromosome number of thirteen, except

P. eriostoma, grow in central Africa (Du Plessis & Spies

232

Bothalia 28,2 (1998)

TABLE 1. — List of Pentaschistis specimens studied with their respective gametic chromosome numbers, ploidy levels, figures where they are

shown, localities and voucher specimens

Gametic

Bothalia 28,2(1998)

233

TABLE 1. — List of Pentaschistis specimens studied with their respective gametic chromosome numbers, ploidy levels, figures where they are

shown, localities and voucher specimens (cont.)

Gametic

Group Taxon Chromosome Ploidy Figure

no.

Locality and vouchers

1992). The basic chromosome number of seven predomi-

nates in the southern African species, whereas x = 1 3 pre-

vails in the remainder of Africa (Du Plessis & Spies 1992).

Polyploidy is common in the genus Pentaschistis.

Nineteen of the 45 specimens studied were polyploid.

Higher ploidy levels were detected in some of the cells

of these specimens. This could be caused by cell fusion

(Spies & Van Wyk 1995) which is not uncommon in the

genus Pentaschistis (Figure 5).

Pentaschistis airoides subsp. jugorum and P. colorata

were tetraploid. The genome homology of these two

species was determined and the observed chromosome

configurations corresponded best with the expected values

for the 2:2 model (Table 2) of Kimber & Alonso ( 1 98 1 ). It

indicates that two sets of genomes are present and each set

consists of two genomes. The relative similarity of the

genomes within a set is 0.5 and the relative affinity

between the sets is expressed by an x-value, which may

vary between 0.5 (differences between sets are similar to

differences within a set) and one (sets are totally different)

(Kimber & Alonso 1981). The x-values for the specimens

studied were one or tending towards one (Table 2), thus

indicating little to no homology between the two sets of

genomes (for example AABB). Based on the specimens

used during this study, these species are alloploid.

However, because of the occurrence of an occasional

quadrivalent, P. colorata may be classified as a segmental

allotetraploid. The occurrence of quadrivalents indicates

that the genomes of this species correspond to some

extent.

FIGURE 1. — Meiotic chromosomes in Pentaschistis. A, B, P. airoides subsp. airoides: A, Spies 4388, diakinesis, n = 21, 21nR; B, Spies 4388.

diakinesis, n = 14, 10hr, 4„k. C, P. capillaris. Spies 3810 , diakinesis, n = 7, 7nR; D, P. cirrhulosa. Spies 4627, diakinesis, n = 7, 5hr, 2hk;

E, P. densifolia. Spies 3878a., diakinesis, n = 7, 6uR, lux; F, P. pallida. Spies 5368, diakinesis, n = 7+3B, 7nR, lm. G, H, P. tomentella: G,

Spies 4306, diakinesis, n = 7, 5mr, 2hK: H, Spies 3782, diakinesis, n = 14, 8hr, 6nx. Scale bar: 8.5 mm.

234

Bothalia 28,2 (1998)

FIGURE 2. — Meiotic chromsomes in Pentaschistis. A, P. aristidoides, Spies 5320, metaphase I, n = 7, 6r R, lnK; B, C, P. triseta, Spies 4413,

metaphase I, n = 7, 6n r, Irk D, E, P. rigidissima: D, Spies 5431, diakinesis, n = 7+0-2B, 6hr, Ihk; E, Spies 4388, metaphase I, n = 21,

1 4n R, 5hk, 1 iv r F, P curvifolia, Spies 4443, diakinesis, n = 7, 7hr G, H, Pentaschistis spp: G, Spies 5336, diakinesis, n = 7+0-2B, 7ur,

1 ii k; H, Spies 5768, anaphase I, n = 7+2B. Scale bar: 8.5 mm.

The genome homology of the tetraploid specimens of

P. pallida was determined. The observed and expected

values corresponded best to the 2:2 model for two of the

specimens, namely Spies 3828 and 5292, with x-values

of one or tending towards one (Table 2). This indicates

that little or no homology exists between the two sets of

genomes, thus classifying these specimens as allote-

traploids (for example AABB). However, the occurrence

of quadrivalents indicates that the genomes of this

species do correspond to some extent. Therefore, Spies

3828 is classified as a segmental allotetraploid (AAA’A1)

with a x-value of 0.7 (Table 2).

FIGURE 3. — Meiotic chromsomes in P.

eriostoma. A, Spies 5470, metaphase

I, n = 13+0-2B; B, Spies 4390,

anaphase I, n = 13+0-3B. C, D, Spies

5337\ C, metaphase I, n = 26; D,

anaphase I, n = 26. E, Spies 5319,

metaphase I, n = 39+0-4B. Scale bar:

8.5 mm.

Bothalia 28,2 (1998)

235

2x 4 x 6x 7x 8x 1 2x 1 3x

Ploidy levels

FIGURE 4. — Comparison of the

number of specimens per

ploidy level for the two exist-

ing basic chromosome num-

bers in Pentaschistis species.

The observed and expected values for the other two

P. pallida specimens ( Spies 3859 and 5381) correspond-

ed best to the 2:1:1 model, with x-values of 0.8 and 1

respectively (Table 2). The 2:1:1 model indicates that

three sets of genomes are present, one set consists of

two genomes and the other two of one genome each.

The x-values for the specimens studied were one or

tending towards one (Table 2), indicating little or no

homology among the three sets of genomes (for exam-

ple AABC or AABB'). However, because of the occur-

.rence of quadrivalents in these two specimens ( Spies

3859 and 5381 ) they are classified as segmental allote-

traploids (AAA'A"). Therefore, the specimens of P. pal-

lida are either alloploids or segmental alloploids tending

towards alloploidy (for example AABB, AABC, AABB'

or AAA'A”). The two different genomic constitutions for

this species may indicate that there is a high degree of

variation within the species.

In P. eriostoma (Figure 3A-E) three different ploidy

levels were detected, namely diploid, tetraploid and

hexaploid, that correspond with the published literature

(Spies & Du Plessis 1988; Du Plessis & Spies 1992).

Heptaploidy was also observed by Du Plessis & Spies

(1992). In the diploid (n = x = 13) specimens of P. erios-

toma 13 bivalents were observed. The genome homolo-

gy of the tetraploid specimens of P. eriostoma was deter-

mined. The observed and expected values corresponded

best to the 2:1:1 model with an x-value of 0.9 (Table 2),

indicating that little homology exists among the three

sets of genomes. This species is thus an allotetraploid

species (for example AABC). However, a low frequency

of quadrivalents occurred in the tetraploid, indicating

that the genomes of this species do correspond to some

extent. Pentaschistis eriostoma is, therefore, classified as

a segmental allotetraploid (for example AAA'A").

All the existing chromosome data of Pentaschistis

was used to determine the number of specimens per

species per ploidy level. The number of specimens per

ploidy level of the two different basic chromosome num-

bers was also determined (Figure 4). This was done to

determine the degree of maturity (it is the ratio between

diploid and polyploid specimens, as well as the level of

polyploidy) in the polyploid complexes in the different

species and in representatives of the two basic chromo-

some numbers.

The Pentaschistis species vary from young to old

polyploid complexes. Most species studied (P airoides

subsp. airoides, P. aristidoides , P. aurea subsp. aurea,

P. borussica, P. capillaris, P. cirrhulosa , P. curvifolia,

P. densifolia, P. eriostoma, P. natalensis , P. pallida,

P. papillosa, P. patula, P. rigidissima, P. tomentella.

TABLE 2. — Genomic relationships in tetraploid Pentaschistis specimens according to the models of Kimber & Alonso (1981). Values indi-

cate sum of squares between observed and expected values for each model; whereas values in parentheses indicate x-values for each model

Accepted model

236

Bothalia 28,2 (1998)

FIGURE 5. — Initial stages of cell fusion in Pentaschistis. A, P. airoides subsp. airoides , Spies 4388, n = 21; B, P. cirrhulosa, Spies 4627, n = 7;

C, P. lima, Ellis 5422, n = ± 42. D, E, P pallida: D, Spies 3860, n = 14; E, Spies 5381 , n = 14. F, P. eriostoma, Spies 3850, n = 13+0-2B.

Scale bar: 8.5 mm.

FIGURE 6. — Mciotic chromosomes in Pentaschistis with univalents. A, B, P. pallida: A, Spies 4406, diakinesis with 3 univalents, n = 7+0-1 B;

B, Spies 5381 , diakinesis with 5 univalents, n = 7. C, P. cirrhulosa. Spies 4627, metaphase I with 2 univalents, n = 7. D, E, P. eriostoma:

D, Spies 5319, metaphase I with 3 univalents, n = 39+0-4B; E, Spies 5339, metaphase I with 1 univalent, n = 26+0-2B. F, Pentaschistis

spp, Spies 5768, metaphase I with 1 univalent, n = 7+0-2B. Scale bar: 8.5 mm.

Bothalia 28,2 (1998)

237

FIGURE 7. — Meiotic chromosomes in Pentaschistis specimens with abnormalities. A, G, P. eriostoma , Spies 5319: A, anaphase I cell with five lag-

ging chromosomes; G, telophase I cells with micronuclei, n = 39+0-4B. B, P. tomentella. Spies 3773, anaphase I bridge, n = 7. C, D, P

rigidissima: C, Spies 5431, anaphase 1 bridge, n = 7+0-2B; D, Spies 5458, anaphase II bridge, n = 21 . E, F, H, telophase I cells with micronu-

clei: E, P. pallida. Spies 5393, n = 7; F, P triseta. Spies 4413, n = 7; H, Pentaschistis spp.. Spies 5768, n -- 7+0-2B. Scale bar: 8.5 mm.

P. triseta, P. tortuosa, ) were young polyploid complex-

es. In a young polyploid complex diploidy prevails, but

higher levels of ploidy do occur (Grant 1981). Poly-

ploidy prevails in the mature polyploid complexes, but

diploidy still occurs. No evidence of mature polyploidy

was found.

In old polyploid complexes only polyploid levels are

observed (Grant 1981). A few species (P. airoides subsp.

jugorum ; P. argentea ; P. aristifolia', P. barbata subsp.

barbata\ P. colorata\ P. lima', P. malouinensis', P. mannii ;

P. minor, P. aff. patula', P. rupestris & P. viscidula ), were

classified as old polyploid complexes. It is important to

note that in almost all the old polyploid species, only one

or a few specimens were examined. More specimens

should be studied to verify these hypotheses. The classi-

fication of young polyploid complexes is better support-

ed, since more specimens were examined. Representa-

tives of both the basic chromosome numbers (x = 7; x = 13)

were classified as young polyploid complexes, with sim-

ilar frequency ratios (Figure 4). The ploidy levels

obtained in this study, combined with existing data, sug-

gest that the genus Pentaschistis is a young polyploid

hybrid complex (Figure 4). Hybridisation could have

resulted in the formation of a basic chromosome number

of x = 13 for P. eriostoma and P. borussica. However,

morphological and anatomical data (Linder & Ellis

1990; Linder et al. 1990; Ellis & Linder 1990) supports

the inclusion of P. borussica in the genus Pentaschistis.

It is thus more probable that polyploidisation and subse-

quent aneuploidy of a Pentaschistis species, or hybridis-

ation between two different Pentaschistis species and

subsequent aneuploidy, resulted in P. borussica. This is

not true for P. eriostoma. Morphological and anatomical

data placed P. eriostoma in group 6. According to Linder

& Ellis (1990) and Ellis & Linder (1990) this is the group

of species with uncertain affinity. It is more probable that

P. eriostoma was formed by hybridisation of two unre-

lated species. Genomic in situ hybridisation (GISH)

analysis, using various putative parental genomes as

probes, should be used to determine the origin of the x =

1 3 basic chromosome number.

Univalents were observed in nearly all of the studied

specimens (Figure 6A-F). Lagging chromosomes were

observed during anaphase I in P. aristidoides, P. curvifo-

lia, P. eriostoma, P. lima, P. pallida, P. rigidissima, P.

rupestris, P. tomentella, P. viscidula and one of the two

unnamed Pentaschistis species (Figure 7A). A small

number of micronuclei (Figure 7E-H) was observed

throughout the whole genus Pentaschistis. Anaphase I or

II bridges (Figure 7B-D) were observed in five of the

specimens studied. This is most probably the result of

paracentric inversion. The frequency of these abnormal-

ities was so low that it should not affect the fertility of the

specimens.

In P. pallida ( Spies 4406), a diploid (n = 7+0-1 B), five

bivalents and one quadrivalent were observed. In this

diploid, seven bivalents are expected. A quadrivalent was

also observed in a diploid specimen of P. eriostoma ( Spies

5370). This phenomenon can be the result of a balanced

translocation. If complete pairing between the two non-

homologous chromosomes occurred, a quadrivalent would

result. Three types of division are possible during

anaphase I. If adjacent type I or II division occurred, there

would be no fertile pollen. During adjacent type I division,

the two centromeres belonging to non-homologous chro-

mosome pairs move to the one pole and the other two to

the other pole. During adjacent type II division, the two

centromeres from a homologous chromosome pair move

to each pole. If alternate type division occurred, all the

pollen would be fertile. During alternate type division, the

chiasmata of all the chromosome pairs lie on the

metaphase plate and the centromeres of the non-homolo-

gous pairs move to the same pole (Schulz-Shaeffer 1980).

238

Bothalia 28,2 (1998)

The fertility of P. pallida ( Spies 4406) and P erios-

toma ( Spies 5370 ) pollen is thus dependent on the type of

division that occurs. All the material in our laboratory is

fixed after collection, so no pollen germination fertility

test could be done. More specimens of these two species

must be collected to study whether this phenomenon

occurs frequently or if it was a unique event.

In conclusion, although cytogenetic differences were

observed between the different species, the cytogenetic

data did not support or reject the groupings suggested by

Linder & Ellis (1990) and Ellis & Linder (1990). The

basic chromosome numbers of x = 7 and x = 13 suggest

that P eriostoma and P. borussica must be further

removed from the other Pentaschistis species. Since P

eriostoma shows little cytogenetic, morphologic or

anatomic similarity to the other species, we do suggest

that P. eriostoma is not closely related to the other

species of the genus Pentaschistis. The data also indicate

that most Pentaschistis species form young polyploid

complexes. Species classified as old polyploid complex-

es were usually inadequately studied.

ACKNOWLEDGEMENTS

Material received from Dr R.P. Ellis and the National

Botanical Institute is greatfully acknowledged. Ms

S.M.C. van Wyk is thanked for developing the photos

used in this paper. Financial support by the Foundation

for Research and Development and the University of the

Free State are also hereby acknowledged.

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CARNOY, J B. 1886. La Cytodierese de l’oeuf. Cellule 3: 1-92.

DARLINGTON, C D. & LA COUR, L.F. 1976. The handling of chro-

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OBITUARIES

DAVID SPENCER HARDY (193 1-1998)

Ah! like gold fall the leaves in the wind, long years numberless as the

wings of trees! The long years have passed like swift draughts of the

sweet mead in lofty halls of the West ... Farewell!

J.R.R. Tolkien: The Lord of the Rings

An era has ended for those of us who have an interest

in succulents or, indeed, in all living plants of southern

Africa of any kind. To us who knew him so well, Dave

Hardy (Figure 1) seemed ageless and, sometimes in more

ways than one, timeless; yet mortality has now also

claimed him. He passed away in Pretoria on the night of

31 May 1998 of complications after surgery.

David Spencer Hardy was bom in Pretoria on 24

September 1931. His education included only four sub-

jects at Matriculation level (i.e. not a full matriculation

certificate), which makes his subsequent achievements all

the more remarkable, and also shows that academic qual-

ifications are not always a reliable indicator of underlying

intelligence. He joined the Department of Agriculture on

21 February 1951 (Gunn & Codd 1981), when he was

appointed Technician at the Veterinary Research Institute,

Onderstepoort. In 1958 he was transferred to the Division

of Botany, which subsequently became the Botanical

Research Institute and in 1989 the National Botanical

Institute. Here he was responsible for cultivating living

plant material brought in from the field by scientists, and

in due course, by himself and his extraordinary circle of

friends, acquaintances and contacts. He displayed a par-

ticular aptitude for growing succulents and greenhouse

plants. Not only did he grow them, but he studied them in

minute and painstaking detail, observing their similarities

and differences both in appearance and cultural require-

ments (Figure 2). His dedication paid dividends: plants

that others declared impossible to cultivate away from

their natural habitat, grew, flourished and became com-

monplace for him.

Dave maintained two particularly ‘special’ areas in

the nursery at the Pretoria National Botanical Garden,

two purportedly identical greenhouses situated next to

each other. In one he assembled what was at its best the

finest collection of rare and endangered Madagascan

plants in the world. In the other he started, a few years

before he retired, to re-create his vision of the Namib

Desert: a Namib not to be found in the real world, where

one would have to travel many hundreds of kilometres to

see the plants that Dave managed to persuade to grow

right next to one another (Figure 3). It is fitting that what

he referred to as the Madagascar house now bears a brass

plaque reading:

The Hardy Collection of rare and endangered plants. Named for

David Spencer Hardy, plant collector and succulent grower

extraordinary.

During his career he collected extensively in the

northwestern Cape (Northern Cape Province: Namaqua-

land, Upington, Prieska, Richtersveld etc.). He also

undertook many expeditions to Namibia and Northern

Province, South Africa, often at his own expense, as well

as to Madagascar, Angola, Comoro Islands and Mauri-

tius. He established a small garden of Madagascan plants

at Nwanedi National Park in the former Venda homeland,

where he could cultivate some of the more cold-sensitive

species out of doors. Dave enjoyed the outdoors, and his

collecting trips were, in addition to their scientific con-

tent, a welcome escape from what he saw as office

drudgery. And so he blossomed; as he relaxed with

increasing distance from Pretoria, so the stories of past

adventures came out, and new ones happened. For exam-

ple, on an expedition of which I was privileged to be a

part, he broke the ice with a tale of how he and Brand van

Breda, then of Worcester Veld Reserve, went to the

Richtersveld in a Volkswagen Beetle long before anyone

thought of made roads in that area. They stopped to ask

directions of a passing goatherd one day, and then asked

if the old lady thought they would reach their destination

in that vehicle. The old goatherd squatted down, peered

under the car and replied ‘Nee menere, in hierdie motor

sal djulle ry soos ’n bokooi wat swaar in die melk is’,

roughly translated as ‘No, gentlemen, in this car you will

go like a nanny-goat heavily in milk’.

FIGURE 1. — David Spencer Hardy (1931-1998). Studio portrait made

by A. Romanowski in 1984

240

Bothalia 28,2 (1998)

FIGURE 2. — Dave Hardy with one of

his favourite subjects of study,

an aloe. Photo: Fanie Venter

Field work was however not all humour. There were

plants to be studied, considered and collected, and when

one returned home there were papers to be prepared and

reports to be compiled. Often, the collections included

fragile flowering material for illustration and eventual

inclusion in The Flowering Plants of Africa. Many plates

published between about 1958 and Dave’s retirement in

1991 record that the original material was a Hardy col-

lection; a smaller number were written up by Dave him-

self, often with one of the Institute scientists as co-author.

Volume 48 of The Flowering Plants of Africa is dedicat-

ed to Dave with the words ‘His acute perception of the

individual cultural needs of plants and especially mem-

bers of the extremely rich succulent flora of southern

Africa and Madagascar is the underlying secret of his suc-

cess. He has given special attention to subjects suitable

for inclusion in these pages and has, on occasions, also

supplied the text’. Dave’s written output was prodigious,

especially when one considers that most of it was written

in his ‘spare’ time, apart from his horticultural duties.

Unfortunately, he was averse to recording his own activi-

ties, with the result that the attached list of 174 publica-

tions cannot be said with certainty to be complete. These

publications are supported not only by the living material

in the Pretoria National Botanical Garden and elsewhere

(a true gardener, Dave was always generous with cut-

tings, seeds and other propagules), but by over 7 000

herbarium specimens; the first set is housed in PRE, but

there are many duplicates in herbaria around the world.

His publications include two books on succulents

(Bomman & Hardy 1972; Hardy & Fabian 1992). Most

of the remainder are semi-popular, which reflects his

passion for popularising and publicising his beloved suc-

FIGURE 3. — Dave Hardy in the glass-

house for desert plants at the

National Botanical Garden,

Pretoria. In the foreground is a

member of the grape family,

Cyphostemma curorii from

Namibia.

Bothalia 28,2 (1998)

241

culents. This did not stop at the written word. Dave was

a popular and sought-after speaker at meetings of soci-

eties concerned with plants, and 18 of his talks were

included in the Department of Agriculture’s contribution

to the radio spectrum, Calling all Farmers. On several

occasions he presented survival courses to the South

African Defence Force.

After his retirement Dave concentrated on Mada-

gascar, a country he loved. In May-June 1993 he did

restoration work at Jardin Botanique et Zoologique de

Tananarive, and the next year he returned to survey

aquatic weeds in the Antananarivo area (Dorr 1997). He

also led several tour groups through the botanical glories

of the island, and was involved in a project to restore

populations of Aloe snzannae , Madagascar’s most spec-

tacular aloe.

He was a member of the South African Association of

Botanists, and was awarded their Certificate of Merit in

January 1991; this is the Association’s award in recogni-

tion of services to botany by those without a degree in

the subject. In the same year he was awarded the Fel-

lowship of the Cactus and Succulent Society of America