Bothalia

’N TYDSKRIF VIR PLANTKUNDIGE NAVORSING

A JOURNAL OF BOTANICAL RESEARCH

Vol. 21,1

May/Mei 1991

Digitized by the Internet Archive

in 2016

https://archive.org/details/bothaliavolume2121unse

BOTHALIA

’N TYDSKRIF VIR PLANTKUNDIGE NAVORSING

A JOURNAL OF BOTANICAL RESEARCH

Volume 21,1

Editor/Redakteur: O.A. Leistner

Assisted by B.A. Momberg

Editorial Board/Redaksieraad

D.F. Cutler

B.J. Huntley

P.H. Raven

J.R Rourke

M.J. Werger

Royal Botanic Gardens, Kew, UK

National Botanical Institute, Cape Town, RSA

Missouri Botanical Garden, St Louis, USA

Compton Herbarium, NBI, Cape Town, RSA

University of Utrecht, Utrecht, Netherlands

Editorial Committee

Redaksiekomitee

O.A. Leistner

B.A. Momberg

M.C. Rutherford

ISSN 0006 8241

Issued by the National Botanical Institute, Private Bag X101, Pretoria 0001, South Africa

Uitgegee deur die Nasionale Botaniese Instituut, Privaatsak X101, Pretoria 0001, Suid-Afrika

1991

CONTENTS— INHOUD

Volume 21,1

1. Erythrineae (Fabaceae) in southern Africa. E.F. FRANKLIN HENNESSY 1

2. A new species of Merxmuellera (Arundineae, Poaceae) from South Africa. N.P. BARKER and

R.P. ELLIS 27

3. Synopsis of the genera Nesaea and Ammannia (Lythraceae) in southern Africa. K.L. IMMELMAN 35

4. Studies in the genus Riccia (Marchantiales) from southern Africa. 22. R. rubricollis, now validated,

typified and described. S.M. PEROLD 51

5. Notes on African plants:

Amaryllidaceae. The correct author citation for Clivia miniata. P. VORSTER 66

Cupressaceae. Tetraclinis articulata, a hitherto unrecorded naturalised alien conifer in South Africa.

J.P. ROURKE 62

Fabaceae. A new species of Amphithalea (Liparieae). A.L. SCHUTTE and B-E. VAN WYK 59

Poaceae. Aira praecox, a new record from southern Africa. H.P. LINDER 55

Pteridophyta. An unique locality for Oleandra distenta, the first recorded for the Orange Free

State. P.J. DU PREEZ and PC. ZIETSMAN 65

Rosaceae. A new species of Cliffortia from the south-western Cape. E.G.H. OLIVER and

A.C. FELLINGHAM 60

Selaginaceae. Microdon bracteatus — the correct name for M. lucidus. I.H. HARTLEY 57

Verbenaceae. Stachytarpheta species in southern Africa. P.P.J. HERMAN 57

Zygophyllaceae. A new species of Zygophyllum from the Cape Province. E. RETIEF 55

6. Mixoploidy and cytotypes: a study of possible vegetative species differentiation in stapeliads (Ascle-

piadaceae). F. ALBERS and U. MEVE 67

7. Invasive alien woody plants of the Orange Free State. L. HENDERSON 73

8. Multivariate analysis of coastal grasslands at Mkambati Game Reserve, north-eastern Pondoland, Transkei.

C.M. SHACKLETON, J.E. GRANGER, B. MCKENZIE and M.T. MENTIS 91

9. Obituary: Hedley Brian Rycroft (1918—1990). E.G.H. OLIVER 109

10. Book reviews 115

11. Guide for authors to Bothalia 117

Bothalia 21,1: 1-25 (1991)

Erythrineae (Fabaceae) in southern Africa

E.F. FRANKLIN HENNESSY*

Keywords: Erythrina , Erythrineae, Fabaceae, Mucuna , pollination, revision, southern Africa, taxonomy

ABSTRACT

The two genera represented in the flora of southern Africa, Erythrina L. and Mucuna Adans. are revised. Keys to the

indigenous species and the commonly cultivated exotic species are provided.

U1TTREKSEL

Die twee inheemse genusse, Erythrina L. en Mucuna Adans. word hersien. Sleutels vir die inheemse spesies en vir uitheemse

spesies wat gewoonlik gekweek word, word voorsien.

CONTENTS

Erythrineae 1

Key to indigenous genera 2

Erythrina 2

Pollination and floral morphology 3

Economic uses 3

Key to the subgenera of Erythrina (world-wide) ... 3

Synonymy of all subgenera and sections represented

in southern Africa 3

Key to the indigenous subgenera and sections 4

Key to indigenous species 4

1. E. abyssinica 4

2. E. latissima 5

3. E. decora 7

4. E. mendesii 7

5. E. lysistemon 7

6. E. caffra 9

7. E. humeana 9

8. E. zeyheri 10

9. E. acanthocarpa 11

Natural hybrids 11

10. E. X coddii 11

11. E. x johnsoniae 12

12. E. X dyeri 12

13. E. x hennessyae 12

Exotic species of Erythrina common in cultivation

in southern Africa 12

Synoptic key to exotic species 12

Section Duchassaignia 13

14. E. fusca 13

Section Cristae-galli 13

15. E. crista-galli 13

16. E. falcata 14

Section Micropteryx 14

17. E. poeppigiana 14

Section Macrocymbium 14

18. E. senegalensis 14

19. E. livingstoniana 14

Section Stenotropis 15

20. E. speciosa 15

* Department of Botany, University of Durban-Westville, Private Bag

X54001, Durban 4000.

MS. received: 1990-04-01.

Section Erythrina 15

21. E. pallida 15

22. E. aff. E. x sykesii 15

Specimens examined ( Erythrina ) 16

Mucuna 17

Pollination and economic importance 18

Subgenus Mucuna 18

Subgenus Stizolobium 18

Key to indigenous species 18

1. M. gigantea 18

2. M. pruriens 19

var. pruriens 19

var. utilis 19

3. M. coriacea 21

Specimens examined ( Mucuna ) 21

Acknowledgements 23

References 23

ERYTHRINEAE

Tribe Erythrineae Hutch., The genera of flowering

plants 1: 431 (1964); Dyer: 230 (1975). Tribe Phaseoleae

DC., subtribe Erythrininae Benth.: 113 (1837); Benth.: 452

(1865); Lackey: 312 (1981). Type genus: Erythrina L.

Trees, shrubs, suffrutices, woody lianes or herbaceous

climbers. Leaves stipulate, stipellate, trifoliolate; foliar and

foliolar pulvini present. Inflorescences terminal or axillary,

paniculate, pseudoracemose, subumbellate or consisting

of fascicles of flowers scattered along a leafy axis. Flowers

non-resupinate or resupinate. Calyx gamosepalous; tube

various; limb of 5 lobes or teeth or obsolete, variously

hirsute or glabrous. Corolla papilionaceous; petals

glabrous, velutinous or hairy. Stamens (9) + 1 or (10);

anthers monomorphic or dimorphic; intrastaminal

nectariferous disc present. Ovary sessile or stipitate, few-

or many-ovulate; style terete, straight or incurved, glabrous

or proximally pubescent; stigma terminal, capitate, small.

Fruit dehiscent along both sutures or one suture; pericarp

ligneous, subligneous, coriaceous or chartaceous, glabres-

cent or hirsute. Seeds exendospermic, smooth, brown, red

or particoloured; hilum short or girdling; rim aril incon-

spicuous or conspicuous.

2

Bothalia 21,1 (1991)

An artificial assemblage of nine genera, the relationships

of which are obscure, grouped together for convenience

rather than by conviction. Representatives of two genera,

Erythrina and Mucuna are indigenous in the FSA region.

KEY TO INDIGENOUS GENERA

Carina shorter than vexillum; anthers monomorphic, sub-

basifixed; fruit glabrescent, often moniliform; stipels

glandular or, rarely, coriaceous; trees, shrubs or suf-

frutices Erythrina

Carina exceeding vexillum; anthers dimorphic, alternately

dorsifixed or versatile and sub-basifixed, the shorter

often bearded; fruit with indumentum of irritant hairs;

stipels non-glandular; woody lianes or climbing herbs

Mucuna (p 17)

ERYTHRINA

Erythrina L., Species plantarum edn 1: 706 (1753);

L.: 316 (1754a); Lam.: 392 (1786); DC.: 411 (1825);

Spreng.: 243 (1826); Guill., Per. & A. Rich.: 224 (1832);

E. Mey.: 15 (1836); A. Rich.: 213 (1847); Harv.: 236

(1862); Benth.: 531 (1865); Bak.: 181 (1871); Bak.: 188

(1876); Taub.: 363 (1894); Harms: 659 (1915); Phillips: 336

(1926b); Hutch.: 406 (1927); Bak. f.: 370 (1929); Bum

Davy: 415 (1932); Louis: 295 (1934-5); Collett: 219 (1941);

Phillips: 422 (1951); Hepper: 562 (1954); Majot-Rochez

& Duvign.: 120 (1954); Codd: 507 (1955); Torre: 247

(1966); Schreiber: 38 (1970); Verde.: 541 (1971); Hennessy:

1 (1972); Krukoff & Barneby: 332 (1974); Dyer: 269 (1975);

Compton: 277 (1976); Troupin: 306 (1982). Type species:

Erythrina corallodendrum L.

Corallodendron Mill.: 372 (1754); Kuntze: 172 (1891).

Type species: Erythrina corallodendrum L.

Tetradapa Osbeck: 93 (1754). Type species: Tetradapa

javanorum Osbeck = Erythrina variegata L.

Mouricou Rheede ex Adans.: 326 (1763). Type species:

Erythrina corallodendrum L. var. orientalis L. = E. indica

Lam. = E. variegata L.

Xyphanthus Rafin.: 103 (1812). Type species: Xyphan-

thus hederifolius Rafin. = Erythrina hederaefolia Spreng.

= E. herbacea L.

Chirocalyx Meisn.: 97 (1843). Type species: Chirocalyx

mollissimus Meisn. = Erythrina latissima E. Mey.

Micropteryx Walp. : 739 (1850). Type species: Micropte-

ryx poeppigiana Walp. = Erythrina micropteryx Poepp.

= E. poeppigiana (Walp.) O.F.Cook.

Duchassaingia Walp.: 741 (1850). Type species:

Duchassaingia glauca Walp. = Erythrina glauca Willd.

= E. fusca Lour.

Macrocymbium Walp. 149 (1853). Type species: Ma-

crocymbium vogelii (Hook, f.) Walp. = Erythrina vogelii

Hook. f.

Stenotropis Hassk.: 183 (1855). Type species: Steno-

tropis berteroi Hassk., nom. illeg. = Erythrina poianthes

Brot. = E. speciosa Andrews.

Hypaphorus Hassk.: 197 (1858). Type species: Hypa-

phorus subumbrans Hassk. = Erythrina subumbrans

(Hassk.) Merrill.

Trees, or shrubs, or suffrutices with enlarged subterra-

nean rootstocks; armed with conical or falcate prickles;

variously hirsute. Leaves synanthous or hysteranthous,

pinnately trifoliolate, deciduous, stipulate; stipules per-

sistent or caducous; leaflets stipellate; stipellae glandular

or coriaceous. Inflorescences subterminal or axillary

pseudoracemes, contemporary with the leaves or preco-

cious. Flowers subverticillate in groups of 3, bracteate;

bracts and bracteoles hirsute, caducous. Calyx gamo-

sepalous, variously hirsute; tube narrowly or broadly

campanulate, truncate or shallowly bilabiate or spatha-

ceous; limb well developed with 5 long or short lobes,

obsolescent or obsolete. Corolla papilionaceous; vexillum

conduplicate-falcate, shortly clawed, minutely velutinous

or subglabrous; alae falcate, shortly clawed, sometimes

auriculate near base; carina petals free or connate, shortly

clawed, rarely auriculate. Stamens 10, monadelphous, or

diadelphous with the vexillary stamen free or partly

coherent; anthers uniform, sub-basifixed, bithecate with

longitudinal dehiscence. Ovary stipitate, linear, multi-

ovulate, densely tomentose; style terminal, straight or

incurved, hirsute proximally, glabrous distally; stigma

small, capitate, terminal. Fruit usually falcate, moniliform,

ligneous or subligneous, smooth or aculeate, dehiscing

along adaxial suture. Seeds red or brown, smooth, elliptic;

hilum short; rim aril inconspicuous. 2n = 42.

A genus of more than 100 species distributed through-

out the tropics and subtropics. Nine indigenous species

occur in the FSA region of which E. decora and E. acan-

thocarpa are endemic while a third, E. caffra, may also

be endemic. Four natural hybrid taxa occur in the area,

one of which is ± fully fertile ( E . x dyeri ) and the other

three ± sterile.

The generic name Erythrina is derived from the Greek

erythros, meaning red, in allusion to the colour of the

vexillum and of the seed of most species.

Within the genus 5 subgenera have been recognised

(Krukoff & Barneby 1974) with 26 (25) sections. I am

unable to uphold Krukoff s (l.c.) separation of sections

Erythrina and Corallodendra which, in my opinion,

intergrade. Because of this separation of these two sec-

tions, Krukoff (l.c.) found it necessary to set aside the

original generitype, E. corallodendrum L. (which he

proposed as the type of section Corallodendra Krukoff)

and substitute E. herbacea L. as lectogeneritype. If section

Corallodendra Krukoff is sunk in section Erythrina, as

I propose, the generitype remains E. corallodendrum L.

All five subgenera are represented on the African con-

tinent, but only two have indigenous representatives

in southern Africa. Because several exotic species of

Erythrina are in cultivation in southern Africa, with a

strong likelihood of more being successfully introduced,

a synoptic key (p. 3) to all the subgenera is provided,

together with synoptic keys to the indigenous (p. 4) and

the most commonly cultivated exotic species (p. 12).

Bothalia 21,1 (1991)

3

POLLINATION AND FLORAL MORPHOLOGY

Pollination in Erythrina, a predominantly red-flowered

genus, is effected principally by birds. Southern African

and all other Old World species are pollinated by passerine

(perching) birds, whereas among New World species,

some are passerine bird-pollinated and others are

hummingbird-pollinated.

Passerine bird-pollinated species are characterised by

having the inflorescence axis usually held horizontally in

arborescent species, by inward-facing, ± gaping flowers,

by a copious supply of hexose-rich nectar (Baker & Baker

1983) and by potten grains somewhat sticky, varied in size

and ornamentation between species, almost invariably with

either small lumina or with sexinous granules present

(Hemsley & Ferguson 1985).

Hummingbird-pollinated species differ in having the

inflorescence axis vertical, the horizontally held, outward-

facing flowers ‘tubular’, smaller quantities of nectar with

a higher sucrose :hexose ratio (Baker & Baker l.c.) and

dry, powdery, medium-sized pollen grains with a simple,

regular, reticulate ornamentation and no sexinous granules

in the lumina (Hemsley & Ferguson l.c.).

Inflorescence, flower and pollen morphology of the

arborescent species, E. lysistemon, E. caffra (subgenus

Erythrina), E. latissima and E. decora (subgenus Chiroca-

lyx ) is typical of passerine bird-pollinated species with the

peduncle providing a perch and the essential whorls of the

flower directed towards the proximal end of the peduncle,

i.e. inward-facing towards the perch. The flowers of E.

lysistemon gape less widely than those of the other three

species. The inflorescence attitude of E. abyssinica

(subgenus Chirocalyx) is unusual in that the axis is vertical.

Inflorescence attitude of the four non-arborescent

species, E. humeana, E. zeyheri, E. acanthocarpa (sub-

genus Erythrina) and E. mendesii (subgenus Chirocalyx)

differs from that of the arborescent species in that the

flower-bearing part of the rhachis is erect. The inflores-

cence axis of E. humeana, E. zeyheri and E. mendesii

elongates markedly as flowering progresses. The proximal

part of the axis in taller specimens of E. humeana (a shrub)

is often horizontal with the distal part remaining vertical.

Erect, progressively elongating inflorescences in low-

growing species probably serve to keep the flowers visible

and accessible to pollinators above the level of the

surrounding vegetation.

Although birds do not appear to discriminate between

species (Jacot Guillarmod et al. 1979), any one plant being

visited by a number of local bird species, it may be

significant that sunbirds (which have finer feathers at

microscopic level than other passerine birds, particularly

on the feathers of the patches of iridescent plumage on

the throat and head of the males), are the main pollinating

agents of E. humeana and E. zeyheri. (The pollinators of

E. acanthocarpa and E. mendesii are not known.) New

World hummingbirds alone have similar plumage.

Possession of long, often curved beaks is another feature

shared by the unrelated African sunbirds and American

hummingbirds. Hemsley & Ferguson (l.c.) noted that

pollen of E. humeana and some pollen of E. zeyheri is

morphologically similar to ‘hummingbird-type’ pollen.

The erect inflorescences and defiexed, ‘tubular’ flowers

of section Humeanae they consider to be adaptations for

pollination by sunbirds, which, like hummingbirds, carry

pollen on their beak and on the specialized head and throat

feathers, whereas other passerine birds carry pollen on

the coarser chest feathers.

ECONOMIC USES

Apart from the use of Erythrina spp. as ornamental

plants and as shade plants, little commercial value has,

until recently, been attached to the genus. Various parts

of the plants have, however, long been used as ingredients

in tribal medicine and magic. Phytochemical analyses have

shown that as well as several poisonous alkaloids the seeds

contain a protein which simplifies separation and purifi-

cation of tissue plasminogen activator by acting as a

preferential inhibitor. Tissue plasminogen activator is a

possible solvent for blood clots (thromboses) in man.

Extraction of the inhibitor from seeds of Erythrina section

Cajfrae is being carried out commercially by a pharmaceu-

tical company in South Africa.

KEY TO THE SUBGENERA OF ERYTHRINA WORLD-WIDE

la Calyx dehiscing apically at anthesis, lacking true limb:

2a Calyx tube shallowly campanulate, broader than long;

carina petals connate; carina falcate, at least half as long

as vexillum; filaments connate for ± three-quarters

their length, shortly or scarcely exserted from carina;

seeds brown or blackish, sometimes mottled . I. Micropteryx

2b Calyx tube campanulate or tubular, longer than broad;

carina petals free or connate; carina more-or-less

straight, less than half as long as vexillum; filaments

connate for + two-thirds their length, long-exserted

from carina; seeds usually red:

3a Calyx symmetrically long-bilabiate at anthesis; carina

petals free; fruit inflated, longitudinally winged along

sutures; leaflets palmately lobed II. Tripterolobus

3b Calyx denticulate or asymmetrically shortly bilabiate at

anthesis; carina petals free or connate; fruit wingless;

leaflets not palmately lobed III. Erythrina

lb Calyx dehiscing longitudinally to the base at anthesis,

becoming spathaceous; limb with 5 long or short lobes;

4a Calyx tube dehiscing abaxially; carina petals free or con-

nate IV. Chirocalyx

4b Calyx tube dehiscing adaxially; carina petals free

V. Erythraster

SYNONYMY OF ALL SUBGENERA AND SECTIONS

REPRESENTED IN SOUTHERN AFRICA

I. Subgenus Micropteryx (Walp.) Bak. in Hook, f.,

The Flora of British India 2,1: 189 (1876); Krukoff &

Barneby: 339 (1974). Type species: Micropteryx poeppi-

giana Walp. = Erythrina poeppigiana (Walp.) O.F. Cook.

(3 sections; none indigenous in southern Africa.)

II. Subgenus Tripterolobus Barneby & Krukoff in

Lloydia 37: 410 (1974). Type species: Erythrina greenwayi

Verde.

(1 section; monotypic, not indigenous in southern Africa.)

4

Bothalia 21,1 (1991)

HI. Subgenus Erythrina Harv. in Flora capensis 2,1:

236 (1862); Louis: 299 (1935); Verde.: 542 (1971); Krukoff

& Bameby: 348 (1974). Type species: Erythrina coral-

lodendrum L.

(16 sections, 3 in southern Africa.)

Section Caffrae Bameby & Krukoff in Lloydia 37: 404

(1974). Type species: Erythrina caffra Thunb.

Section Humeanae Bameby & Krukoff in Lloydia 37:

407 (1974). Type species: Erythrina humeana Spreng.

Section Acanthocarpae Bameby & Kmkoff xn Lloydia

37: 408 (1974). Type species: Erythrina acanthocarpa E.

Mey.

IV. Subgenus Chirocalyx (Meisn.) Harv. in Flora

capensis 2,1: 236 (1862); Louis: 299 (1935); Verde.: 542

(1971); Krukoff & Bameby: 411 (1974). Chirocalyx Meisn.:

97 (1843). Type species: Chirocalyx mollissimus Meisn.

= Erythrina latissima E. Mey.

(5 sections, 1 in southern Africa.)

Section Chirocalyx Verde, in Flora of tropical East

Africa, Leguminosae (4) — Papilionoideae (2): 542 (1971);

Krukoff & Bameby: 418 (1974). Erythrina subgenus

Chirocalyx section Merocraspedon Louis : 302 (1935).

Type species: Erythrina latissima E. Mey.

V. Subgenus Erythraster Bameby & Krukoff in Lloydia

37: 429 (1974). Type species: Erythrina variegata L.

(1 section, not indigenous in southern Africa.)

KEY TO INDIGENOUS SUBGENERA AND SECTIONS

la Calyx dehiscing abaxially, becoming spathaceous at

anthesis; limb with 5 long or short lobes; trees or

suffrutices

IV. Subgenus Chirocalyx Section Chirocalyx (spp. 1—4)

lb Calyx dehiscing apically at anthesis, finally campanulate

and shortly bilabiate or shortly denticulate, without true

limb; trees or shrubs or suffrutices . III. Subgenus Erythrina

2a Calyx shortly bilabiate at maturity; vexillum acute; trees

Section Caffrae (spp. 5—6)

2b Calyx shortly denticulate at maturity; vexillum obtuse;

shrubs or suffrutices:

3a Calyx narrowly campanulate, distinctly 5-toothed, the

teeth arising behind the rim; vexillum scarlet, obtuse;

carina petals free or connate, shorter than alae,

included; intrastaminal nectariferous disc with ten

obtuse lobes; fruit unarmed; shrub or suffrutex

Section Humeanae (spp. 7—8)

3b Calyx broadly campanulate, obscurely 5-toothed and often

shallowly and obscurely bilabiate; vexillum red proxi-

mally, grading through orange to yellow distally with

green tip, obtuse; carina petals free, exceeding alae,

with long-acuminate, shortly exserted apices; intra-

staminal nectariferous disc with or without ten dis-

tinct lobes; fruit aculeate; much-branched shrub with

tuberous rootstock Section Acanthocarpae (sp. 9)

KEY TO INDIGENOUS SPECIES

la Calyx dehiscing abaxially ± to the base at anthesis,

spathaceous; limb with 5 long or short lobes; trees or

suffrutices:

2a Trees up to 15 m tall with stout trunks; bark thick, corky,

\vith deep longitudinal and transverse fissures; leaves

large; flowers red:

3a Calyx lobes exceeding 10 mm, filiform:

4a Inflorescence held + vertically; vexillum ± 30—40 X

20-24 mm; stamens included; terminal leaflet 25—140

x 30—160 mm 1. E. abyssinica

4b Inflorescence held ± horizontally; vexillum more than

45 X ± 30 mm; stamens exserted; terminal leaflet

100—250 X 120—300 mm 2. E. latissima

3b Calyx lobes ± 2 mm long, thickened; inflorescence held

± horizontally; vexillum 40-47 X + 20 mm; terminal

leaflet 42—110 x 45—140 mm 3. E. decora

2b Suffrutex with aerial stems up to 0,45 m tall; leaflets

rhomboid or parabolic; terminal leaflet 60-100 x 35-80

mm; inflorescence held + vertically; calyx lobes 1-3

mm long, thickened 4. E. mendesii

lb Calyx dehiscing apically at anthesis, finally campanulate

and shortly bilabiate or shortly denticulate, without

true limb; trees, shrubs or suffrutices:

5a Calyx shortly bilabiate at maturity; trees:

6a Vexillum twice as long as broad, slightly arcuate enclosing

the inner floral parts, scarlet, flesh pink or white; alae

long-clawed, less than 15 X + 5 mm; carina segment

± 10 X + 7 mm 5. E. lysistemon

6b Vexillum one and a half times longer than broad, strongly

arcuate exposing the inner floral parts, vermilion-red,

orange or cream-coloured; alae short-clawed, + 25

X ± 12 mm; carina segment ± 22 X + 15 mm ..

6. E. caffra

5b Calyx shortly denticulate at maturity; shrubs or suffrutices:

7a Calyx narrowly campanulate, distinctly 5-toothed; vexil-

lum scarlet, or very rarely, flesh pink; alae exceeding

carina segments; intrastaminal nectariferous disc

with ten rounded or truncate lobes; fruit unarmed,

seeds red:

8a Shrubs up to 3 m tall; stipules less than 10 mm long;

leaflets very variable, ovate subacute, ovate long-

acuminate or hastate, unarmed or sparingly aculeate

on the veins abaxially; calyx teeth spreading; seed

6—7 mm long 7. E. humeana

8b Suffrutex with annual aerial stems to 0,5 m long;

stipules more than 10 mm long; leaflets ovate, ovate-

rhomboid, broadly ovate or subrotund with obtuse

or subacute, or rarely, shortly acuminate apex,

aculeate on veins on both surfaces, or rarely, on

abaxial surface only; calyx teeth clasping vexillum;

seed 10—17 mm long 8. E. zeyheri

7b Calyx broadly campanulate, obscurely 5-toothed; vexil-

lum tricoloured, vermilion red proximally, grading

through orange to yellow and green distally; alae

exceeded by long-acuminate free carina segments;

fruit aculeate, seeds brown 9. E. acanthocarpa

1. E. abyssinica Lam. ex DC., Prodromus systematis

naturalis 2: 413 (1825); A. Rich.: 214 (1847); Louis: 306

(1935); Gillett: 426 (1962); Torre: 248 (1966); Verde.: 555

(1971); Hennessy: 33 (1972); Krukoff & Barneby: 427

(1974); Codd: t. 1738 (1977); Coates Palgrave : 329 (1977);

Troupin: 306 (1982); Van Rensburg: 20 (1982). Chirocalyx

abyssinicus (DC.) Hochst.: 600 (1846). Corallodendron

abyssinicum (Lam.) Kuntze: 172 (1891). Type: a plant

cultivated at Petit Trianon, Paris, from seed collected by

Bruce in N Ethiopia and sent to De Candolle by Thonin

(G, holo.— K, photo.!).

Chirocalyx tomentosus Hochst.: 600 (1846). Type: N Ethiopia, near

Adua, Schoata, Schimper 531 (B, holo. t: K, iso.!).

E. tomentosa R. Br. ex A. Rich.: 213 (1847); Bak.: 184 (1871), pro

parte; Bak f.: 373 (1929); Majot-Rochez & Duvign.: 120 (1954); White:

152 (1962). Types: Ethiopia, Salt s.n. (BM, syn.l); Schimper 531 (B,

syn. t; K, isosyn.!); Choa, Morotte, d’Hericourt s.n. (P, syn.); Taccaze

Valley, Djeladjeranne, Quartin Dillon s.n. (P, syn.); Shoho and Djeleu-

kote, Petit s.n. (P, syn.); Bruce, Voy. Abyss., t. 19 in part (syn.!).

Bothalia 21,1 (1991)

5

E. suberifera Welw. ex Bak.: 183 (1871). Corallodendron suberifera

(Welw.) Kuntze: 173 (1891). Erythrina abyssinica Lam. ex DC. subsp.

suberifera (Welw. ex Bak.) Verde. : 284 (1970), pro parte. Types: Angola,

Welwitsch 2229 (BM, syn.); Welwitsch 2230 (BM, syn.; K, isosyn.!).

E. huillensis Welw. ex Bak.: 183 (1871); Bak. f.: 372 (1929). Type:

Angola, Huilla, Welwitsch 2231 (BM, holo.).

E. comosa Hua: 52 (1898). Types: Schweinfurth Ser. 11, 60, 1799, 1868,

1882 (K, isosyn.!).

E. mossambicensis Sim: 43 (1909); Bak. f.: 370 (1929). Types: Mozam-

bique, Sim 5833 (?, f); Sim: t. 54 (1909), (lecto.! designated by Hennessy,

1972).

E. bequaertii De Wild.: B 15 (1920); Bak. f.: 376 (1929). Types: Zaire,

Lake Albert, Irumu, Bequaert 2729 (BR, syn.-K, photo.!); Ruwenzori,

Butagu, Bequaert 3948 (BR, syn.).

E. kassneri Bak. f. : 375 (1929). Type: Kenya, Kitui District, Galunka,

Kassner 794 (BM, holo.!).

E. tomentosa R. Br. ex A. Rich. var. longicauda Bak.f.: 374 (1929).

Type: Kenya, Charangani Hills, Kitale, Paget-Wilkes 0103 (BM, holo.!).

E. warneckei Bak.f: 375 (1929); Verde.: 560 (1971). Type: Tanzania,

Amani, Warnecke 330 (BM, holo.; K, iso.!).

E. platyphylla Bak. f.: 376 (1929). Types: Kenya, Machakos District,

Kilima Kiu, Decie s.n. (BM, syn.); Tanzania, Bukoba District, Karangwe,

Speke & Grant 426 (K, syn.!).

E. eggelingii Bak. f. : 238 (1938). Type: Uganda, Acholi District, Keyo,

Lamogi, Eggeling 1645 (BM, holo.- K, photo.!).

leones: Sim: t. 54 (1909); Hennessy: t. 12 (1972); Codd: t. 1738 (1977);

Troupin: t. 105 (1982); Van Rensburg: 20 (1982).

Tree with stout trunk; juvenile parts lanate-tomentose.

Leaves hysteranthous, coriaceous; leaflets broadly elliptic,

or broadly ovate-rhomboid with obtuse or emarginate

apex; stipels triangular, coriaceous. Inflorescences erect,

precocious; flowers deflexed at anthesis, not gaping. Calyx

spathaceous, scarlet; tube 10—22 mm long, split abaxially

at anthesis; lobes 10—40 mm long. Corolla scarlet;

vexillum enfolding inner floral parts at anthesis, minutely

velutinous; alae exceeding carina; carina segments free,

suborbicular, 5—6x4 mm. Stamens diadelphous with

vexillary stamen partly coherent. Fruit ligneous, falcate,

moniliform, smooth. Seeds scarlet, 9-12 x 6—7 mm;

hilum oval, prominent, blackish, 5,5— 7,0 x 3,0— 4,0 mm.

A single gathering, P.A. Smith 2928 (K, PRE) from a

rocky outcrop 63 km distant from Francistown in

Botswana, has been identified by me as E. abyssinica on

the basis of its leaf and stem morphology alone as the

material lacks flowers and fruit. The natural occurrence

of E. abyssinica in northern Botswana is not unexpected

as this species is widely distributed in west, central and

east Africa, including Angola, Zambia, Malawi and

Mozambique (Figure 1). Cultivated in Transvaal and Natal.

Late winter/spring-flowering.

This is a morphologically variable species which is

closely allied to E. latissima E. Mey. from which it is

distinguished by its usually smaller leaves; the erect

attitude of its inflorescences; the brilliant scarlet, very

conspicuous calyces; the smaller, deflexed vexilla which

are neither spread nor reflexed at maturity and which

become mahogany brown with age, contrasting strongly

with the scarlet calyx.

Voucher: P.A. Smith 2928 (K, PRE).

2. E. latissima E. Mey., Commentarium de plantis

africae australioris 1; 151 (1836); Collett: 222 (1941); Codd:

510 (1955); Batten & Bokelmann: 77 (1966); Hennessy:

12 (1972); Palmer & Pitman: 959 (1972); Van Wyk: 223

(1972); Krukoff & Barneby: 427 (1974); Compton: 227

(1976); Hennessy: t. 1710 (1976); Coates Palgrave: 331

(1977); Jacot Guill. et al 523 (1979); Van Rensburg: 8

(1982). Corallodendron latissimum (E. Mey.) Kuntze: 173

(1891). Types: South Africa, ‘Passim inter Basche et

Omtata, frequentius inter Omaziana et Omsamwubo, alt.

1000-2000 ped.’, Drege s.n. (B, holo. f); South Africa,

T3 miles W of Port St Johns’, ix. 1956, Codd 9744 (K,

neo.!, here designated; PRE!).

Chirocalyx mollissimus Meisn. : 98 (1843). Type: South Africa, Natal,

Tafelberge, Port Natal, viii. 1839, Krauss 263 (K, holo.!; PRE, iso.!).

E. sandersoni Harv.: 39 (1859). Type: South Africa, Natal, Port Natal

about Sterk Spruit, Aug.-Sept. 1858, Sanderson 44 (K, holo.!).

E. gibbsiae Bak.f.: 374 (1929). Type: Zimbabwe, Matopo Hills, Gibbs

73 (BM, holo.!).

E. tomentosa sensu Bak : 184 (1871), pro parte; sensu Wood: t. 384,

385 (1906); sensu Sim: 201 (1907), non A. Rich.

E. abyssinica sensu Marloth: 81 (1925), non DC.

leones: Harv.: t. 91, 92 (1859); Wood: t. 384, 385 (1906); Marloth:

t. 29 (1925); Batten & Bokelmann: t. 65,2 (1966); Palmer & Pitman:

958, 960 (1972); Van Wyk: 244 (1972); Hennessy: t. 12 (1972); Hennessy;

t. 1710 (1976); Van Rensburg: 8, 9 (1982).

Tree with stout bole; juvenile parts lanate-tomentose.

Leaves hysteranthous, coriaceous with minute, triangular,

coriaceous stipels; leaflets very large, oblate, broadly

elliptic or obliquely ovate with obtuse, sometimes apiculate

apices. Inflorescences ± horizontal, precocious, stout,

compact; flowers ± horizontal, gaping at anthesis. Calyx

spathaceous, dull scarlet; tube 30 x 8—10 mm; lobes

10— 16 mm long. Corolla red; vexillum scarlet becoming

crimson later, spread and partly reflexed at anthesis,

minutely velutinous; alae exceeding carina; carina petals

free, suborbicular, 18—20 x 14—15 mm. Stamens 10,

diadelphous with vexillary filament partly coherent. Fruit

ligneous, falcate, moniliform, smooth. Seeds crimson,

11- 15 x 10-1 mm; hilum oval, prominent, blackish, 7—8

x 5-6 mm.

The holotype, a Drege gathering from Transkei, has not

been located and was presumably destroyed during World

War II, nor has any isotype or syntype material been found.

FIGURE 1. —Distribution of Erythrina abyssinica, #; E. latissima, ■;

E. decora. O; E. mendesii, □.

Bothalia 21,1 (1991)

FIGURE 2. — Erythrina lalissima: 1-3, Hennessy 384 ; 4-12 Ward s.n. (flowers in spirits). 1, inflorescence; 2, infructescence; 3, seeds; 4, bract

5, bracteole; 6, calyx; 7, vexillum; 8, ala; 9, carina petal, all x 0,8; 10, 11, 12, anther, three views, X 4,7.

Bothalia 21,1 (1991)

7

Accordingly, I have chosen a Codd gathering made near

Port St John in Transkei in 1956 as neotype of Erythrina

latissima.

Trees of this species occur in open savanna, often on

rocky outcrops where they may have been afforded some

degree of protection from fire damage. Juvenile plants

seem rare in the wild which suggests that some form of

protection, especially from fire, should be afforded this

species. Erythrina latissima is recorded from Zimbabwe,

Mozambique, Swaziland, Transvaal, Natal and the eastern

Cape (Figure 1). Late winter/spring-flowering.

This species is closely allied to E. abyssinica Lam. ex

DC. from which it is distinguished by its usually larger

leaves; the ± horizontal attitude of its inflorescences; less

conspicuous calyces; larger, spreading and somewhat

reflexed vexilla which are the most conspicuous part of

the inflorescence, turning crimson with age (Figure 2).

Vouchers: Acocks 13163 (K); Codd 4418 (K, PRE); Compton 27947

(NBG, PRE); Pegler 235 (BOL, GRA, K, SAM); Scheepers 1171 (K,

PRE).

3. E. decora Harms in Engl., Botanische Jahrbiicher

fur Systematik, Pflanzengeschichte und Pflanzen-

geographie 49: 441 (1913); Schreiber: 38 (1970); Hennessy:

27 (1972); Palmer & Pitman: 962 (1972); Krukoff &

Bameby: 421 (1974); Coates Palgrave: 330 (1977); Van

Rensburg: 19 (1982). Types: South West Africa/Namibia,

Dinter 796 (B, syn. t; SAM, isosyn.!); Dinter 796a (B,

syn. BM, isosyn.; M, lecto. , designated by Krukoff &

Bameby (1974)); Dinter 1354, 1505, 1548 (B, syn. t);

Giirich 42 (B, syn. t).

leones: Hennessy: t.9. (1972); Krukoff & Bameby: 421 (1974).

Tree with very stout trunk; juvenile parts lanate-

tomentose. Leaves hysteranthous, coriaceous; leaflets

oblate, suborbicular, obovate or rhomboid-ovate with

emarginate, obtuse or shortly acuminate apices; stipels

small, conical, glandular. Inflorescences ± horizontal,

precocious; flowers ± horizontal, not gaping. Calyx

spathaceous, dull red; tube 15—20 mm long; lobes

thickened, ± 2 mm long. Corolla red; vexillum scarlet,

± enfolding inner floral parts at anthesis, minutely

velutinous; alae exceeding carina; carina petals free,

suborbicular, 6—7 x 5— 6 mm. Stamens diadelphous with

vexillary stamen partly coherent. Fruit subligneous,

falcate, moniliform, smooth. Seeds scarlet, 10—13 x 6—8

mm; hilum oval, prominent, blackish, 4—5 x 2—3 mm.

This species is known only from Namibia where it

occurs on outcrops of rock (often granite) mainly (solely?)

of the Damara sequence (Figure 1). The only other species

of Erythrina indigenous in Namibia, E. mendesii Torre,

is suffrutescent, not arborescent and occurs in

unconsolidated Tertiary to Quaternary sediments.

Although the flowers of these two species are similar, it

is not possible to confuse the identity of the plants. Spring/

summer/autumn-flowering.

Vouchers: De Winter & Leistner 5601 (WIND); Keet 1614 (WIND);

Rusch 7957 (BOL, K); Vahrmeijer & Du Preez 2555 (PRE); Walter 614

(WIND).

4. E. mendesii Torre in Boletim da Sociedade Broteri-

ana, ser. 2, 39: 212 (1965); Torre: 248 (1966); Schreiber:

38 (1970); Hennessy : 31 (1972); Krukoff & Barneby: 420

(1974). Type: Angola, ‘Chibia, entre Chibia e o rio Caculo-

var,’ xii. 1955, Mendes 1079 (LISC, holo.; BM, COI, K!,

LUA, SRGH, iso.).

E. baumii sensu Codd: t. 1412 (1963); sensu Van Rensburg: 19 (1982),

non Harms.

leones: Codd: t. 1412 (1963); Torre: t. 23 (1966); Hennessy: t. 10, 11

(1972); Van Rensburg: 19 (1982).

Suffrutex with branched subterranean horizontal stems

arising from a tuberous rootstock; aerial stems erect,

unbranched; juvenile aerial parts tomentose. Leaves

synanthous, chartaceous; stipels glandular, truncate;

leaflets equally or unequally rhomboid with obtuse apices.

Inflorescence erect; peduncle and rhachis progressively

elongating; flowers ± horizontal, not gaping. Calyx

spathaceous, red; tube 12—18 mm long; lobes thickened,

1—3 mm long. Corolla red; vexillum scarlet, ± enfolding

inner floral parts at anthesis, minutely velutinous; alae

exceeding carina, cochleariform with a triangular lobe

towards the base on adaxial side, 5-7 x 4—5 mm; carina

petals free, of similar shape to alae, 4-6 x 4-5 mm.

Stamens diadelphous, the vexillary filament partly

coherent. Fruit subligneous, falcate, moniliform, smooth,

50—90 x 15 mm. Seeds not seen.

This suffrutescent species is known from Zambia,

southern Angola, Capri vi Strip and northern Botswana

(Figure 1) where its habitat is unconsolidated Kalahari

sands usually in river valleys and seasonally dry water

courses. It is distinguished from E. baumii Harms which

occurs in savanna habitats in Angola, Zaire and Zambia

by leaflet shape (rhomboid in E. mendesii ; elliptic-ovate

in E. baumii) and ala length (±6 mm in E. mendesii ;

± 15 mm in E. baumii). When specimens of the two

species are seen together, the difference in leaflet

morphology is immediately apparent. There has been

some conftision as to the identity of the specimens

collected in Namibia and northern Botswana, but

examination of material of both species has shown that all

gatherings to date from this area are E. mendesii. Summer-

flowering.

Vouchers: De Winter 3745 (K, PRE, WIND); Henriques 238 (K); PA.

Smith 1872 (K, PRE); West 3276 (K).

5. E. lysistemon Hutch, in Kew Bulletin 1933: 422

(1933); Collett: 223 (1941); Codd: 507 (1955); Letty: 169

(1962); Batten & Bokelmann: 77 (1966); Verde.: 547 (1971);

Hennessy: 17 (1972); Palmer & Pitman: 957 (1972); Van

Wyk: 225 (1972); Krukoff & Barneby: 406 (1974);

Compton : 278 (1976); Coates Palgrave: 332 (1977); Jacot

Guill. etal.\ 523 (1979); Van Rensburg: 11 (1982). Type:

South Africa, Transvaal, Belfast, Rietvlei, Crocodile

River, vi. 1932, J.C. Smuts 66 (K, holo.!; BM, iso.).

E. caffra Thunb. var. mossambicensis Bak. f. : 238 (1938). Type:

Mozambique, Nyassa, Maniamba, Torre 523 (COI, holo.; BM, iso.).

E. caffra sensu Harv. : 236 (1862); sensu Wood: t. 542 (1912); sensu

Phillips: t. 59 (1922); sensu Marloth: 81 (1925); sensu Bak. f.: 369 (1929);

sensu Bak.f. : 238 (1938); sensu Burtt Davy: 415 (1932); sensu Collett:

223 (1941), pro parte; sensu Codd: 70 (1951), non Thunb.

leones: Wood: t. 542 (1912); Phillips: t. 59 (1922); Codd: t. 3 (1955);

Codd: t. 66 (1951); Letty: t. 84.3 (1962); Batten & Bokelmann: t. 65.1

8

Bothalia 21,1 (1991)

FIGURE 3. — Erythrina humeana , 3. 1-3.6, Hennessy 441 : 1, inflorescence and leaf; 2, vexillum; 3, ala; 4, carina petal; 5, calyx; 6, seed, all

X 0,8. Erythrina acanthocarpa, 3.7—3.11, McCulloch s.n. (flowers in spirits): 7, vexillum; 8, ala; 9, carina petal; 10, calyx; 11, seed, all

x 0,8. Erythrina lysistemon , 3.12—3.17, Hennessy s.n. (flowers in spirits): 12, flower; 13, vexillum; 14, ala; L5, carina segment; 16, calyx;

17, seed, all x 0,8. Erythrina caffra. 3.18—3.24, Hennessy s.n. (flowers in spirits): 18, flower; 19, vexillum; 20, ala; 21, carina segment;

22, calyx; 23, infructescence; 24, seed, all x 0,8.

Botha] ia 21,1 (1991)

9

(1966); Hennessy: t. 1, 5 (1972); Van Wyk: 245 (1972); Van Rensburg:

11, 12, 13 (1982).

Tree up to 10 m tall; juvenile parts pubescent. Leaves

hysteranthous, chartaceous; leaflets broadly ovate, base

cuneate, apex acute, obtuse or acuminate; lateral leaflets

50—130 x 45-95 mm; terminal leaflet 50—130 x 45—130

mm; stipels glandular. Inflorescences ± horizontal, pre-

cocious; flowers deflexed at anthesis, not gaping. Calyx

brown; tube ± campanulate, splitting laterally to become

shallowly bilabiate at anthesis, 10-15 mm long; lobes

obsolescent, the thickened abaxial lobe prognathous in

bud. Corolla : vexillum scarlet, or more rarely, flesh pink

or off-white, enfolding inner floral parts at anthesis, 45-78

X 20—35 mm, subglabrous; alae exceeding carina; carina

segments partly connate abaxially, 9—15 x 4-8 mm.

Stamens diadelphous with vexillary stamen free or partly

coherent. Fruit subligneous, falcate, moniliform, smooth.

Seeds scarlet or vermilion red, 8—10 x 5—6 mm; hilum

oval, depressed, blackish, ±4x2 mm (Figure 3.12 —

3.17).

This winter/early spring-flowering species is distin-

guished from E. caffra, with which it has often been

confused, by its smaller stature; its narrower, chevron-

shaped inflorescences; the scarlet (as opposed to vermilion

red) colour of its vexilla; its longer, narrower, less strongly

arcuate vexilla which do not spread or reflex at maturity

but enfold and conceal the inner floral parts. The fruit and

seed of E. lysistemon and E. caffra are alike. These species

have not yet achieved perfect reproductive isolation. Where

their ranges overlap they hybridise. The hybrid, E. x

dyeri, is fertile.

Erythrina lysistemon is tolerant of a wider range of

climatic and soil types than is E. caffra and has a wider

distribution range (Figure 4). North of the Flora region

it occurs in Mozambique, Zimbabwe, Zambia, Malawi and

Tanzania.

Vouchers: Codd 7987 (GRA, K, PRE); Dyer 5748 (K, PRE); He mm

155 (PRE); Prosser 1038 (K, NBG, PRE); Ward 4274 (PRE).

6. E. caffra Thunb. , Prodromus plantarum capensium

2: 559 (1823); DC.: 412 (1825); Spreng.: 243 (1826); E.

Mey.: 149 (1836); Eckl. & Zeyh.: 259 (1836); Harv.: 236

(1862), pro parte; Sim: 201 (1907); Marloth: 81 (1925),

pro parte; Collett: 223 (1941), pro parte; Codd: 508 (1955);

Batten & Bokelmann: 78 (1966); Hennessy: 15 (1972);

Palmer & Pitman: 955 (1972); Krukoff & Bameby: 405

(1974); Hennessy: t. 1709 (1976); Coates Palgrave: 329

(1977); Jacot Guill. et al.\ 522 (1979); Van Rensburg: 2

(1982) Batten: 162 (1986). Duchassaingia caffra (Thunb.)

Walp.: 741 (1850). Corallodendron caffrum (Thunb.)

Kuntze: 172 (1891). Type: South Africa, ‘e Cap. b. Spei’,

Thunberg s.n. (UPS, holo.; K!, MO, PRE! UDW!,-

photo.).

£ fissa Presl: 69 (1832). Chirocalyx pubescens Walp. : 741 (1850), nom.

superfl. Type: a cultivated plant from Palermo Botanical Garden

(PRAHA, holo ).

£ viarum Todaro: 62 (1861). £ insignis Todaro: 66 (1861). Type: a

cultivated plant from Palermo Botanical Garden (K, lecto.!).

£ constantiana Micheli: 542 (1896). Type: ‘a cultivated plant in a

garden at Golf Jouan near Cannes. Villa Niobe, owned by Mr A.

Constant, FI. April.’ 1896, Micheli s.n. (K, holo.!).

Icqnes: Presl: t. 46 (1832); Micheli: facing p. 524 (1869); Sim: t. 53

(1907); Codd: t. 1, 2 (1955); Batten & Bokelmann: t. 66.1,2 (1966);

Hennessy: t. 4 (1972); Palmer & Pitman: 954 (1972); Hennessy: t. 1709

(1976); Van Rensburg: 2 (1982); Batten: 163 (1986).

Tree, 10—18 m tall, juvenile parts pubescent. Leaves

hysteranthous, chartaceous; slightly larger than those of

E. lysistemon ; stipels glandular. Inflorescences ± horizon-

tal or deflexed, precocious; flowers horizontal or

ascending, gaping. Calyx brown; tube campanulate,

splitting laterally to become shallowly bilabiate at anthesis,

17-20 mm long; lobes obsolescent, the thickened abaxial

lobe prognathous in bud. Corolla', vexillum vermilion red,

orange, or, rarely, creamy-white, spread and reflexed at

anthesis, exposing the inner floral parts, 42 -70 x 27-40

mm, subglabrous; alae slightly exceeding carina; carina

segments partly connate abaxially. Stamens monadelphous

with filament tube split right or left of vexillary stamen

or, rarely, diadelphous with vexillary filament free or

partly coherent. Fruit and seeds like those of E. lysistemon.

This winter/early spring-flowering species is limited in

its distribution to the eastern Cape and southern Natal

coastbelt with outlying populations in forest in northern

Zululand and on Inhaca Island off Maputo which may or

may not be natural (Figure 5). It usually occurs in coastal

and streambank forests in deep sandy soils. It differs from

other southern African members of subgenus Erythrina

in having a widely gaping flower (Figure 3.8—3.24).

Vouchers: Archibald 5906 (PRE); Bauer 249 (K, SAM); Flanagan

319 (BOL, PRE, SAM); Fourcade 4466 (BOL, STE); Ward 3014 (K, NU).

7. E. humeana Spreng., Caroli Linnaei systema

vegetabilium 3: 243 (1826); Sim: 202 (1907); Sim: 43

(1909); Phillips: t. 112 (1923); Marloth: 81 (1925); Collett:

225 (1941); Codd: 72 (1951); Batten & Bokelmann: 77

(1966); Hennessy: 21 (1972); Palmer & Pitman: 961 (1972);

Codd: 269 (1974); Krukoff & Barneby: 408 (1974);

Compton: 277 (1976); Coates Palgrave: 331 (1977); Jacot

Guill. et al.\ 522 (1979); Van Rensburg: 5 (1982). Type:

Bot. Reg. 9, t. 736A (1823) (lecto.! designated by Codd,

1974).

£ princeps A. Dietr.: 305 (1834). Corallodendron princeps (A. Dietr.)

Kuntze: 173 (1891). Type: Field Museum Nat. Hist., Chicago, negative

no. 2375 (neo.l, designated by Krukoff & Bamebv ,074>

FIGURE 4.— Distribution of Erythrina lysistemon.

10

Bothalia 21,1 (1991)

E. humei E. Mey.: 150 (1836); Harv.: 237 (1862); Bak.: 182 (1871);

Bak. f.: 370 (1929). Corallodendron humei (E. Mey.) Kuntze: 173 (1891).

Types: South Africa, ‘between Kovi and Kaprivier’, Drege s.n. (B, syn.

t); ‘between Keiskamma and Basche R., 1836’, Drege s.n. (B, syn. f;

K, isosyn.!).

E. raja Meisn.: 96 (1843). E. humei var. raja (Meisn.) Harv.: 237

(1862). Type: South Africa, Umlaas, Port Natal, Nov. 1839, Krauss 62

(K, holo.!).

E. hastifolia Bertol. f. : 568, t. 38 (1850). E. humei var. hastifolia

(Bertol. f.) Bak. f. . 370 (1929). Type: Bertol. f. , in Mem. Acc. Sc.

Bologna 2: 568, t.38 (1850). (icono.).

E. caffra sensu Ker-Gawl.: t. 736 (1823); sensu Sims: t. 2431 (1823);

sensu DC.: 412 (1825), pro parte; sensu Reichb.: t. 312 (1836), non

Thunb.

leones: Ker-Gawl.: t. 736 (1823); Sims: t. 2431 (1823); Reichb.:

t. 312 (1836); Marloth: t. 29 (1925); Batten & Bokelmann: t. 66.2

(1966); Hennessy: t. 6 (1972); Van Rensburg: 5, 6 (1982); Batten: 167

(1986).

Shrub 0,5-3, 0 m tall; juvenile parts thinly pubescent.

Leaves synanthous, chartaceous; petiole, rhachis and

sometimes veins aculeate; leaflet shape variable, oblate,

ovate-subacute, ovate-long-acuminate, or hastate; lateral

leaflets 45—130 x 30—90 mm, terminal leaflet 45—130

X 45—120 mm; stipels glandular, obtuse or truncate.

Inflorescences lax, contemporary with the leaves, distally

erect; peduncle and rhachis progressively elongating to

± 500 mm; flowers deflexed at anthesis, not gaping. Calyx

red; tube narrowly campanulate, subtruncate; teeth

spreading, the abaxial tooth oblong, others acute, + 2 mm

long. Corolla-, vexillum scarlet, deflexed, obtuse, 35-50

x 14—21 mm, subglabrous; alae oblong-arcuate, 6—12 x

3 mm; carina segments partly connate abaxially or, rarely,

free, quadrangular-unguiculate, 6—8 x 3-4 mm. Stamens

included, diadelphous with vexillary filament partly

coherent. Intrastaminal nectariferous disc with 10 rounded

lobes, white. Fruit subligneous, falcate, moniliform,

blackish, smooth. Seeds scarlet, 6,0— 7,0 x 5, 0-6,2 mm;

hilum oval, depressed, pale, 3,5 - 4,0 x 2,0 mm (Figure

3. 1-3.6).

The amount of variation in leaflet shape in this species

is noteworthy. Specimens with broadly ovate leaflets occur

mainly in the southern part of the distribution range and

those with long, narrowly hastate leaflets further north.

Between the two extremes intermediates occur. Erythrina

humeana is distinguished from E. zeyheri mainly by its

habit; its smaller stipules; smaller, chartaceous, sparsely

armed or unarmed leaflets; smooth unridged peduncle;

spreading calyx teeth and smaller fruit and seeds. Although

there is some overlap in the distribution ranges of these

two species, (Figures 6 & 7), E. humeana is the less

cold-tolerant and does not occur at high altitudes. Its range

extends northwards into Mozambique and Zimbabwe.

Summer-flowering.

Vouchers: Codd 6413 (K, PRE); Comins 1435 (PRE); Dyer 2236

(GRA); Venter 7890 (PRE); Weame 80 (NU, PRE).

8. E. zeyheri Harv., Flora capensis: 236 (1862);

Marloth: 82 (1925); Collett: 227 (1941); Dyer: t. 1011

(1947); Hennessy: 23 (1972); Krukoff & Bameby: 408

(1974); Jacot Guill. et al. \ 525 (1979); Van Rensburg: 15

(1982). Types: South Africa, ‘Betchuanaland, 1841’, Zeyher

531 (K, syn.!; PRE, SAM, isosyn.!) and ‘inter Mooye river

et Macalisberg, 1841’, Burke 69 (K, syn.!).

leones: Dyer: t. 1011 (1947); Hennessy: t. 7 (1972); Van Rensburg:

15 (1982).

Suffrutex with branched, corky, perennial subterranean

stems and enlarged rootstock; aerial stems annual, semi-

FIGURE 7.— Distribution of Erythrina zeyheri.

Bothalia 21,1 (1991)

11

erect or decumbent, 0,3— 0,5 m; juvenile parts pubescent.

Leaves synanthous, coriaceous, aculeate on petiole, rhachis

and veins; stipules ovate or oblong, chartaceous, 10—20

mm long; lateral leaflets 40—150 x 20—170 mm; terminal

leaflet 55-320 x 35—250 mm; stipels coriaceous,

triangular. Inflorescences lax, contemporary with the

leaves, erect; peduncle and rhachis progressively

elongating, up to 490 mm long, longitudinally ribbed or

terete. Flowers deflexed at anthesis, not gaping. Calyx red

or green flushed with pink; tube narrowly campanulate,

subtruncate; teeth clasping not spreading, the abaxial tooth

fleshy, oblong; others acute, ± 2 mm long. Corolla :

vexillum scarlet, deflexed, obtuse, 24-44 x 14-22 mm,

subglabrous; alae oblong-arcuate, 9—15 x 3-5 mm;

carina segments free, oblong with a proximal triangular

lobe and 1 or 2 short triangular lobes abaxially near the

truncate apex, 7—10 x 4-6 mm. Stamens included,

diadelphous with vexillary filament free or shortly

coherent. Intrastaminal nectariferous disc with 10 rounded

or truncate lobes. Fruit subligneous, falcate, moniliform,

blackish, smooth. Seeds scarlet, 10—17 x ± 10mm;hilum

oval, depressed, pale, 4—5 x 2-3 mm.

The suffrutescent habit in Erythrina is considered

(Krukoff 1974) to be advanced. Eight species, six of them

placed in subgenus Erythrina , are suffrutescent. These are

E. montana Rose & Standley, E. leptorhiza DC., E.

horrida DC. (section Leptorhizae) which inhabit temperate

habitats at high elevations in Mexico; E. herbacea L.

subsp. herbacea (section Erythrina) at the northern,

temperate limit of its range in North Carolina; E.

resupinata Roxb. (section Suberosae) which occurs at

high elevations in north India and E. zeyheri (section

Humeanae) which occurs in damp areas at higher

elevations than any other southern African species (Figure

7). In subgenus Chirocalyx, section Chirocalyx, E. baumii

Harms (including E. pygmaea Torre) and E. mendesii

Torre occur in damp depressions at elevations between 900

and 1 450 m. It would be an oversimplification to state

that dwarfing in Erythrina is an adaptation for high altitude

survival alone. Several arborescent species occur at high

elevations in the high rainfall regions of the tropics, there-

fore it is probable that a combination of high altitude, low

rainfall and edaphic factors have resulted in the

development of the suffrutescent habit in taxa occurring

towards the extremes of the range of the genus. Summer-

flowering.

Vouchers: Acocks 18843 (PRE); Codd 9406 (K, PRE); Dieterlen 547

(NH, PRE, SAM); Schrire 632 (NH); K. Smith 27 (BOL).

9. E. acanthocarpa E. Mey. , Commentariorum de

plantis africae australioris: 151 (1836); Harv. : 237 (1862);

Sim: 202 (1907); Marloth: 81 (1925); Phillips: t. 203

(1926a); Collett: 225 (1941); Batten & Bokelmann: 78

(1966); Hennessy: 25 (1972); Krukoff & Barneby: 409

(1974); Jacot Guill. et al.: 521 (1979); Van Rensburg:

17 (1982). Corallodendron acanthocarpum (E. Mey.)

Kuntze: 172 (1891). Type: South Africa, ‘in vallibus inter

Klipplatrivier et Zwartkey, alt. 3000—4000 ped., 1836’,

Drege s.n. (B, holo. t); ‘zwischen Windvogelberg und

Zwartkey auf einer Grasflache, 3000—4000 ped., F. Nov.’,

Drege s.n. (BM, lecto.-K, photo.!, designated by Krukoff

& Barneby 1974).

leones: Sim: t. 52 (19(17); Marloth: t. 25 (1925); Phillips: t. 203 (

1926a); Batten & Bokelmann: t. 65.3 (1966); Hennessy: t. 8 (1972); Van

Rensburg: 17 (1982).

Shrub, 1—2 m tall with enlarged, succulent rootstock

and much-branched, perennial aerial stems; juvenile parts

pubescent. Leaves synanthous, chartaceous, aculeate on

petiole, rhachis and veins; leaflets broadly ovate or trans-

versely elliptic, apiculate or subacute; lateral leaflets

25—47 x 20—45 mm; terminal leaflet 30—48 x 25—55

mm. Inflorescences lax or compact, erect, contemporary

with leaves; flowers horizontal or ascending at anthesis,

not gaping. Calyx green with purple flush; tube broadly

campanulate, sometimes obscurely bilabiate, shortly

toothed, the abaxial tooth obtuse, the others acute, 1-3

mm long. Corolla-, vexillum tricoloured, red proximally

grading to yellow distally with green tip, horizontal or

ascending, obtuse, 42-55 X 22-32 mm, minutely

velutinous; alae rhomboid-acute, 4,0— 7,0 x 1,5— 2,0 mm;

carina segments free, exceeding alae, lanceolate long-

acuminate, 7—15 x 1,5— 4,0 mm, with tips exserted.

Stamens exserted, diadelphous. Intrastaminal nectary

discoid or obscurely 10-lobed. Fruit subligneous, slightly

or not constricted between the seeds, brown, aculeate.

Seeds brown, 16 x 10-12 mm; hilum oval, depressed,

pale, 2, 0-3,0 x 1, 5-2,0 mm (Figure 3.7—3.11).

The profusely branched, perennial aerial system of this

species makes it, by definition, a shrub, yet the greatly

enlarged subterranean rootstock is a modification best

developed elsewhere in the genus in the suffrutescent

species. This species is isolated in the genus by the

combination of habit, small leaflets, unique colouration

of the vexilla, unusual shape and proportions of alae and

carina, aculeate fruit and brown seeds. Some specimens

have been shown to be tetraploid (Darlington & Wiley

1955) with n = 42. It is not known whether any individuals

in this endemic taxon (Figure 8) are diploid. Spring/

summer-flowering.

Vouchers: Acocks 18843 (PRE); Galpin 1653 (BOL, GRA, NU, PRE);

McCabe & Atherstone 41 (K); Tyson s.n. (BOL, K).

NATURAL HYBRIDS

10. E. x coddii Barneby & Krukoff in Lloydia 37: 443

(1974). Type: South Africa, Natal, Zululand, Hlabisa Dis-

trict, Hluhluwe Game Reserve, Deane 7 (K, holo.!).

FIGURE 8. — Distribution of Erythrina acanthocarpa, ♦; four natural

hybrids, Erythrina x coddii, O; Erythrina X johnsoniae, □;

Erythrina X dyeri, •; Erythrina X hennessyae, ▲.

12

Bothalia 21,1 (1991)

Tree with stout bole, juvenile parts densely or sparsely

hirsute. Leaves hysteranthous, variable, intermediate in

texture, size, shape and indumentum between those of the

putative parents, E. latissima and E. lysistemon. Inflores-

cences precocious, ± horizontal, compact; flowers slightly

gaping at anthesis. Calyx dull reddish-brown; tube

spathaceous splitting abaxially to the base; lobes distinct,

of varying lengths. Corolla-, vexillum deep scarlet,

somewhat spreading, narrower than that of E. latissima ;

alae exceeding carina; carina segments partly connate as

in E. lysistemon, or free as in E. latissima. Stamens

diadelphous vexillary filament partly coherent. Fruit

unknown.

Specimens of this apparently sterile hybrid taxon are at

present known from the Hluhluwe Game Reserve in

northern Natal, from Ndwedwe in Natal, from the

northern Transvaal and outside the Flora region from

Harare in Zimbabwe (Figure 8). The putative parents,

E. latissima and E. lysistemon are sympatric in these areas

and the hybrid shows characters which are intermediate

between those of these two species. Spring-flowering.

Vouchers: Lugg s.n. sub NH 40477 (NH); Mugwedi 19 (J); Ward 2954

(NH).

11. E. x johnsoniae Hennessy in Flowering Plants

of Africa 48, t. 1911 (1985). Type: South Africa, Natal,

Mitchell’s Farm between Ramsgate and Southbroom, Codd

9403 (PRE, holo. ! ; NH, iso.!).

Icon: Hennessy: t. 1911 (1985).

Tree, juvenile parts densely or sparsely hirsute. Leaves

hysteranthous, resembling those of E. latissima, but

smaller. Inflorescences precocious, ± horizontal,

compact; flowers gaping at anthesis. Calyx olive green;

tube spathaceous, splitting + abaxially to the base; lobes

distinct, to 5 mm long, or obsolete. Corolla: vexillum

bright scarlet, spread and partly reflexed, smaller than

those of E. latissima and E. caffra : alae exceeding carina;

carina segments partly connate as in E. caffra. Stamens

diadelphous or vexillary stamen absent. Fruit unknown.

This apparently sterile hybrid taxon is known only from

the Port Shepstone District in southern Natal (Figure 8)

where the putative parents, E. latissima and E. caffra are

sympatric. One tree in the garden of the Natal Herbarium

in Durban was established from a truncheon planted in

1957. Like E. x coddii this hybrid is the product of a cross

between parents belonging to different subgenera and

shows some characters of each subgenus. It differs from

E. X coddii mainly in its consistently more coriaceous

leaflets; more nearly horizontal flowers; more widely

gaping flowers and brighter red vexilla. Spring-flowering.

Vouchers: S.M.Johnson 1332 (K, NH, PRE); Nichols 779 (NH);

Schrire 1854 (NH).

12. E. x dyeri Hennessy in Bothalia 16: 48 (1986).

Type: Natal, Durban, Hennessy 445 (UDW, holo.!; NH,

PRE, iso.!).

Icon: Hennessy: 50 (1986).

Tree, intermediate between its putative parents E.

lysistemon and E. caffra, therefore impossible to dis-

tinguish accurately from either in the vegetative phase.

Inflorescences compact, + horizontal, precocious; flowers

gaping, not deflexed, resembling those of E. caffra. Calyx

tube campanulate, bilabiate at anthesis, brown; lobes

obsolete except the abaxial lobe prognathous in bud.

Corolla: vexillum bright scarlet like that of E. lysistemon,

spread and reflexed like that of E. caffra, shorter than that

of either parent; alae and carina intermediate in size and

shape. Stamens intermediate. Fruit and seeds indistinguish-

able from those of either parent.

This fertile hybrid of the two members of section Caffrae

occurs in areas where the parents are sympatric (Figure

8) or grow together in cultivation. At maturity the hybrid

is a slightly smaller tree than E. caffra. The inflorescence

shape, attributable to the gaping, + horizontal flowers,

is like that of E. caffra, but the colour is like that of E.

lysistemon. The extent to which introgression occurs is

not known. Winter-flowering.

Vouchers: Codd 7983 (PRE); Codd 7999 (K, PRE).

13. E. x hennessyae Bameby & Krukoff in Lloydia

37: 448 (1974). Type: South Africa, Transvaal, Waterberg,

Shangri La Guest Farm near Nylstroom, Meeuse 9335 (K,

holo.!).

leones: none known.

Shrub up to 3 m tall. Leaves sparsely aculeate; leaflets

long-acuminate from a broadly ovate base, like those of

some forms of E. humeana. Inflorescences compact, +

horizontal, contemporary with the leaves; flowers deflexed,

not gaping. Calyx russet, narrowly campanulate, shortly

bilabiate at anthesis; lobes obsolete. Corolla: vexillum red,

subacute like that of E. lysistemon-, alae, carina and sta-

mens similar to E. lysistemon. Fruit unknown.

The first recorded gathering of this rare (Figure 8),

apparently sterile hybrid was made at Rustenburg in 1911.

Subsequently a second specimen was discovered on a

koppie near Nylstroom and several plants were established

in the gardens of Shangri La Guest Farm from cuttings

taken from the wild specimen. In stature, foliage and the

synanthous condition, this hybrid resembles E. humeana-,

in flower morphology it closely resembles E. lysistemon.

Its sterility confirms the validity of the separation of

sections Caffrae and Humeanae. Late winter/spring-

flowering.

Vouchers: McGregor s.n. sub Moss 2628 (J); O. Nation s.n. (K).

EXOTIC SPECIES OF ERYTHRINA COMMONLY IN CULTIVATION

IN SOUTHERN AFRICA

Synoptic key to exotic species

(the hybrid taxon, aff. E. x sykesii is not included

in this key)

la Calyx dehiscing apically at anthesis, lacking true limb:

2a Calyx tube shallowly campanulate, broader than long;

carina falcate, at least half as long as vexillum; filaments

connate for ± three-quarters of their length, shortly

exserted from carina; seeds brown or blackish, some

times mottled (Subgenus Micropteryx):

Bothalia 21,1 (1991)

13

3a Abaxial calyx lobe distinct, fleshy; vexillum rhombic-

ovate, long-unguiculate, fully reflexed at anthesis,

yellowish-orange; alae well developed, ± half as long

as carina; carina obliquely ovate, connate; pod sub-

ligneous, slightly constricted between the seeds; seeds

mottled, brown (Section Duchassaingia) 14. E. fusca

3b Abaxial calyx lobe not as above; vexillum ovate, shortly

clawed, spread but not fully reflexed at anthesis; alae

minute; carina lanceolate:

4a Inflorescence lax, terminal or lateral; hypanthium short,

not or scarcely differentiated from calyx tube; vexil-

lum deep red or scarlet; alae truncate; pod sub-

ligneous, not or slightly constricted between the

seeds; seeds brown, mottled (Section Cristae-galliy.

5a Inflorescences terminal, leafy; fruit dorso-ventrally

compressed with interseminal constrictions, ± 12 mm

broad; funicle not persistent 15. E. crista-galli

5b Inflorescences lateral, leafless; fruit laterally compressed,

without interseminal constrictions, ± 20 mm broad;

funicle persistent 16. E. falcata

4b Inflorescence compact, secund, lateral; hypanthium

stipe-like, clearly differentiated from the turbinate

calyx tube; vexillum orange-scarlet; alae obtuse; pod

chartaceous, without interseminal constrictions; seeds

brown, usually mottled (Section Micropteryx)

17. E. poeppigiana

2b Calyx tube tubular or narrowly campanulate, longer

than broad; carina ± straight, half, or less than half

as long as vexillum; filaments connate for ± two-thirds

of their length, long-exserted from carina; seeds usually

red or partly red (Subgenus Erythrina):

6a Vexillum scarlet, enfolding inner floral parts; carina

petals free, linear-oblong, half as long as vexillum;

pod subligneous, without interseminal constrictions;

seeds brown, mottled (Section Stenotropis)

20. E. speciosa

6b Vexillum pink, enfolding inner floral parts; carina petals

free, obtusely spathulate, a quarter as long as vexil-

lum; pod subligneous, with interseminal constrictions;

seeds particoloured red and black (Section Erythrina )

21. E. pallida

lb Calyx dehiscing longitudinally along median abaxial line to

the base at anthesis, becoming spathaceous (Subgenus

Chirocalyx ):

7a Leaflets ovate; inflorescence slender, lax, subsecund; vexil-

lum red, enfolding inner floral parts; carina segments

connate, toothed; pod subligneous, with interseminal

constrictions; seeds red (Section Macrocymbium) ....

18. E. senegalensis

7b Leaflets palmately lobed; inflorescence stout, compact;

vexillum red, spread and partly reflexed at anthesis;

carina petals free, obtuse; pod subligneous, with well-

defined interseminal constrictions; seeds orange-red

(Section Chirocalyx) 19. E. livingstoniana

Section Duchassaingia (Walp .) Krukoff in Lloydia 37:

340 (1974). Erythrina ‘group’ Fuscae Krukoff: 226 (1939b).

Duchassaingia Walp.: 741 (1850). Type species: Duchas-

saingia glauca Walp. = Erythrina fusca Lour.

14. E. fusca Lour., Flora cochinchinensis 427 (1790);

Merrill: 209 (1935); Krukoff: 229 (1939b); Verde.: 547

(1971); Krukoff & Barneby: 340 (1974); Hennessy:

23 (1975); Hennessy: t. 1754 (1977). Types: Cochin

China (Vietnam), Louriero s.n. (syn. , not found); Gelala

aquatica Rumph., Herb. Amboin. 2: 235, t. 78 (1750)

(icono.).

E. glauca Willd.: 428 (1801). Type: Venezuela, Caracas, based on notes

by F. Bredemeyer (fide Krukoff).

E. ovalifolia Roxb. : 254 (1832); Sprague: 198 (1909). Type: India,

Calcutta, Roxburgh drawing 972 (K, lecto. , designated by Verde. 1971).

leones: Krukoff & Barneby: 341 (1974); Hennessy: t. 1754 (1977).

Tree up to 24 m tall. Leaflets ovate-elliptic, silvery

beneath; lateral leaflets 105—165 x 70—105 mm; terminal

leaflet 130—190 x 110—115 mm. Inflorescences precocious;

flowers widely gaping. Calyx broadly campanulate with

fleshy abaxial lobe forming a solid spur 2—4 mm long.

Corolla-, vexillum long-clawed, reflexed ± 90° at anthesis,

pale yellowish or pale orange; alae rhomboid, exposed,

bicoloured red/cream or deep orange, shorter than carina;

carina segments partly connate, obliquely ovate, paler than

alae. Stamens diadelphous. Fruit subligneous, slightly

constricted between the seeds, blackish. Seeds dark brown,

mottled, 12-18 x 5-8 mm.

This species is cultivated on the Natal coastbelt. Until

1984 no fruit set had been observed in the Natal plants

which were propagated from cuttings. Viable seed was

produced in 1984 by trees in the Natal Provincial Adminis-

tration garden at Mayville in Durban, which suggests that

some of the trees have been ‘adopted’ by an effective

pollinator. Several species of birds visit the trees, including

Indian mynahs ( Acridotheres tristis). Late winter/spring-

flowering.

Erythrina fusca has a wider distribution range than any

other species of Erythrina , occurring naturally on three

continents, Africa, Asia and South America in the littoral

and sublittoral zones, and on many tropical islands. Its

seeds are reputed to be distributed by ocean currents and

in river water. It was one of the first tree species to

recolonise the island of Krakatau after the cataclysmic

volcanic eruption of 1883.

Vouchers: Hennessy 385, 387 (UDW).

Section Cristae-galli Krukoff in Lloydia 37: 342

(1974). Erythrina ‘group’ Crista-galli Krukoff: 227 (1939b).

Type species: Erythrina crista-galli L.

15. E. crista-galli L., Mantissa plantarum 99 (1767);

DC.: 413 (1825); Krukoff: 228 (1939b); Verde.: 543 (1971);

Hennessy: 37 (1972); Krukoff & Barneby: 342 (1974);

Hennessy: 23 (1975); Van Rensburg: 22 (1982). Type:

Herb. Linn. Cat. no. 888.4 (lecto.).

leones: Hennessy: t. 14 (1972): Krukoff & Barneby: 343 (1974); Van

Rensburg: 21, 22 (1982).

Tree up to + 5 m tall locally. Leaves synanthous; leaflets

ovate, glabrous or glabrescent, ± 60—90 x 30—60 mm.

Inflorescences terminal, leafy, usually with 3 flowers in

axil of each leaf; flowers widely gaping, resupinate. Calyx

broadly campanulate, + 10 x 24 mm; abaxial lobe

acuminate, + 5 mm long. Corolla: vexillum ovate, shortly

clawed, ± 50 x 35 mm, deep crimson, scarlet or pale

red; alae minute, obtuse; carina segments partly connate,

lanceolate-acute, falcate, + 40 x 11 mm. Stamens shortly

exserted. Fruit subligneous, dorso-ventrally compressed,

constricted between the seeds, ± 12 mm broad, brown.

Seeds dark brown, mottled, ± 12 x 10 mm; funicle not

persistent.

This, the most widely cultivated species of Erythrina ,

in South Africa and elsewhere, is indigenous in eastern

Bolivia, Paraguay, northern Argentina and Uruguay.

Production of an annual crop of flower-bearing leafy

branches from each woody branch, only the strongest of

which persist to continue growth while the remainder

wither, die and drop off, results in mature trees which

14

Bothalia 21,1 (1991)

appear to have been pollarded, and are of relatively short

stature. Spring/summer-flowering.

The common name ‘Cock’s comb’ coral tree from the

Latin crista-gal li (cock’s comb), is an allusion to the short

fan-shaped free portion of the stamens which is exserted

beyond the keel.

Characters which separate E. crista-galli from E. falcata

are summarised in the key to exotic species.

Vouchers: Atkinson s. n. sub J 31397 (J); Dahlstrand 2382 (J); Eliovson

s.n. sub J 26210 (J).

16. E. falcata Benth. in Martius, Flora brasiliensis

15: 172 (1859); Krukoff: 232 (1939a); Krukoff: 684 (1941);

Krukoff & Barneby : 344 (1974). Corallodendron falcatum

(Benth.) Kuntze: 172 (1891). Type: Brazil, Martius s.n.

(M, holo.).

leones: Krukoff: t. 1, 2a, 2b (1941); Krukoff & Barneby: 344 (1974).

Tree up to 10 m tall locally. Leaves hysteranthous;

leaflets ovate, glabrescent, 20—60 x 18—32 mm. Inflores-

cences lateral, crowded towards tips of branches, lax,

leafless; flowers widely gaping. Calyx broadly campanu-

late, + 6 x 11 mm. Corolla: vexillum short-clawed, ovate-

orbicular, + 35 x 32 mm, vermilion red; alae minute,

truncate; carina segments partly connate, lanceolate,

obtuse, falcate, ± 25 x 7 mm. Stamens hardly exserted.

Fruit subligneous, not or hardly constricted between seeds,

+ 20 mm broad, brown. Seeds dark brown, mottled, ±

15 X 10 mm; funicle persistent, ± 3 mm long.

Indigenous in subAndean southern Peru, Bolivia, eastern

Brazil, Paraguay and northern Argentina, this species is

cultivated as an ornamental street tree in parts of South

America, in Australia and in South Africa; several trees

are known in Johannesburg. Fruit and seeds have been

collected from one of the Johannesburg plants ( Herdman

s.n. sub J 48618). All the trees in our area are red-

flowered, but the occasional white-flowered form has been

recorded in Bolivia. Spring/summer-flowering.

Vouchers: Herdman s.n. sub J 48618 (J); Jack s.n. sub J 36293 (J).

Section Micropteryx Krukoff in Lloydia 37: 344

(1974). Erythrina ‘group’ Vernae Krukoff: 234 (1939a).

Type species: Micropteryx poeppigiana Walp. = Erythrina

poeppigiana (Walp.) O.F. Cook.

17. E. poeppigiana (Walp.) O.F. Cook in Bulletin of

the United States Department of Agriculture, Botany 25:

57 (1901); Krukoff: 235 (1939a); Krukoff & Barneby: 347

(1974); Hennessy: 23 (1975). Micropteryx poeppigiana

Walp.: 740 (1850). Erythrina micropteryx Poeppig in

Urban: 327 (1899). Type: Peru, Poeppig s.n. (F, holo.;

GH, NY —photo.).

Icon: Krukoff & Barneby: 347 (1974).

Tree up to 20 m tall locally. Leaves hysteranthous;

leaflets broadly ovate-deltate; lateral leaflets subtended by

a pair of cupulate stipels. Inflorescences lateral, crowded

towards tips of branches, ± compact, secund; flowers

gaping. Calyx turbinate, expanded distally, ± 10 x 7 mm.

Corolla : vexillum short-clawed, ovate, acute, ± 40 x 15

mm, bright vermilion red; alae minute, suborbicular;

carina segments partly connate, lanceolate, acute, falcate,

± 35 x 8 mm. Stamens shortly exserted. Fruit charta-

ceous, not constricted between the seeds. Seeds brown,

immaculate or mottled.

Widely cultivated in the tropics as an ornamental tree

and frequently grown for shade in coffee and cacao planta-

tions, this species is indigenous in Venezuela, Panama,

subAndean Colombia, Ecuador, Peru and Bolivia. Mature

specimens exist in the Durban area where, although fruit

set is initiated, the follicular pods abort early and no

mature seeds have been observed. E. poeppigiana is easily

recognisable as its cup-shaped stipels are unique in the

genus. Spring-flowering.

Voucher: Hennessy 399 (UDW).

Section Macrocymbium (Walp.) Barneby & Krukoff

in Lloydia 37: 414 (1974). Macrocymbium Walp.: 149

(1850). Type species: Macrocymbium vogelii (Hook, f.)

Walp. = Erythrina vogelii Hook. f.

18. E. senegalensis DC., Prodromus systematis

naturalis 2: 413 (1825); Guill. & Perr.: 224 (1832); Bak.:

181 (1871); Aubrev.: 296, t. 118A (1936); Hutch. & Dalz.:

406 (1927); Hepper: 562 (1965); Krukoff & Barneby: 414

(1974); Hennessy: t.1786 (1979). Duchassaingia senegalen-

sis (DC.) Hassk.: 194 (1858). Type: Senegal, Roussillon

s.n. (not traced).

E. latifolia Schumach.: 333 (1827). Chirocalyx latifolius (Schumach.

& Thonn.) Walp.: 148 (1853). Type: Guinee, collector undesignated (K,

iso.).

E. guineensis G. Don: 371 (1832). Type: Sierra Leone, collector

undesignated (not traced).

leones: Aubrev.: t. 118 A (1936); Krukoff & Barneby: 415 (1974);

Hennessy: t. 1786 (1979).

Tree up to 5 m tall. Leaves synanthous or hysteranthous;

leaflets ovate, glabrous, 50-120 x 30—75 mm. Inflores-

cences lateral, crowded towards tips of branches, lax,

subsecund; flowers deflexed, not gaping. Calyx tube

spathaceous with oblique mouth, splitting abaxially, ± 12

x 6 mm; lobes obsolescent, represented by 5 black

papillae. Corolla: vexillum ovate, subobtuse, short-clawed,

+ 35 x 18 mm, scarlet; alae oblong, ± 12 x 5 mm;

carina segments partly connate, quadrangular-unguiculate

with distal margin toothed abaxially, ± 7 x 10 mm.

Stamens ± included. Fruit subligneous, moniliform, ±

12 mm broad. Seeds orange to vermilion red, 6,0— 7,5 x

5, 0-5, 5 mm.

Cultivated specimens of E. senegalensis, which is

indigenous in West Central Africa and the Sudan, have

been established in tropical Australia and in South Africa

in the Durban area. The specimens in Australia produce

fruit and seeds, but to date those in Durban have failed

to do so. Spring/early summer-flowering.

Voucher: Hennessy 393 (UDW).

19. E. livingstoniana Bak. in Flora of tropical Afri-

ca 2: 182 (1871); Sim: 43 (1909); Bak. f.: 370 (1929);

Hennessy: 39 (1972); Krukoff & Barneby: 421 (1974);

Coates Palgrave: 331 (1977); Hennessy: t. 1737 (1977).

Bothalia 21,1 (1991)

15

Type: Malawi, ‘14—19° S. lat. , 60 miles up river Shire’,

Kirk 154, with fig. (K, holo.!).

leones: Sim: t. 48 (1909); Hennessy: t. 15 (1972); Hennessy: t. 1737

(1977).

Tree up to 20 m tall, heavily armed with antrorse

prickles. Leaves hysteranthous; leaflets 90-200 x 85-190

mm, trilobed or sub 5-palmatilobed, terminal lobe deltate,

subacute or obtuse, lateral lobes subquadrangular or

sub-bilobed. Inflorescences lateral, crowded towards tips

of branches, compact; flowers horizontal, gaping. Calyx

spathaceous, splitting abaxially; lobes obsolete. Corolla :

vexillum ovate, obtuse, shortly clawed, arcuate, spreading,

± 50 x 45 mm, scarlet; alae cochleariform, ± 20 x 10

mm; carina segments free, suborbicular, + 17 x 15mm.

Stamens exposed. Fruit ligneous, moniliform, ± 30 mm

broad, grey-buff. Seeds bright orange, 12—15 x 9-11 mm;

hilum depressed, white.

An African species indigenous in parts of Malawi,

Mozambique and possibly in the Triangle region of

Zimbabwe, now in cultivation in South Africa in the

Durban area where it was introduced by Dr Vincent Wager

in 1951. Flowering of the Durban trees is erratic, occurring

in late summer with not all the trees flowering in a single

year. Fruit and seed development occur and the number

of plants grown from seed is rapidly increasing. The

formidable armament of the stems makes propagation from

truncheons difficult. This is a spectacular species which

requires a lot of water. The heavy armament and palmati-

lobed leaflets make this species easily recognisable.

Vouchers: Hennessy 389 (UDW); 390 (K, PRE. UDW); Sim 21134

(NU).

Section Stenotropis (Hassk.) Krukoff in Lloydia 37:

359 (1974). Erythrina subgenus Stenotropis (Flassk.) Bak.:

188 (1876). Erythrina ‘group’ Speciosae Krukoff: 243

(1939a). Stenotropis Hassk.: 183 (1855). Type species:

Stenotropis berteroi Hassk., nom. illeg. = Erythrina

poianthes Brot. = E. speciosa Andrews.

20. E. speciosa Andr. , The botanist’s repository 7,

t. 443 (1806); Krukoff: 243 (1939a); Krukoff: t. 1, figs.

5a, b (1941); Krukoff & Barneby: 360 (1974); Hennessy:

21 (1975). Type: Bot. Rep. t. 443 (lecto., designated by

Krukoff 1974).

E. poianthes Brot.: 342 (1824). Type: a cultivated plant of unknown

origin.

E. reticulata Presl: 22 (1834). Micropteryx reticulata (Presl) Walp.:

741 (1850a). Corallodendron reticulatum (Presl) Kuntze: 173 (1891). Type:

a cultivated plant of unknown origin.

Stenotropis berteroi Hassk.: 183 (1855) (nom. illegit.)

leones: Andrews: t. 443 (1806); Krukoff & Barneby: 360 (1974);

Hennessy: 21 (1975).

Small tree. Leaves hysteranthous or synanthous; leaflets

broadly ovate or parabolic, acute or acuminate, ± 150 x

120—180 mm. Inflorescences lateral, compact, erect;

flowers horizontal-ascending, cylindrical, not gaping.

Calyx campanulate, truncate, with triangular-subulate

abaxial tooth 1,5— 2,0 mm long. Corolla : vexillum narrow-

ly elliptic-oblong, acute, shortly clawed, distally falcate,

± 60 x 15 mm, scarlet; alae ± 9,0 x 2,5 mm; carina

segments free or connate for ± 6 mm, linear-oblong.

obtuse, ± 27,0 x 4,5 mm. Stamens included. Fruit

subl igneous with slight interseminal constrictions. Seeds

dark brown (not seen locally).

Indigenous in south-eastern Brazil and cultivated in

tropical South America, New Zealand, Australia and in

South Africa in the Durban area where flowering occurs

in late winter or early spring contemporary with E. lysiste-

mon and E. cajfra. The sterile hybrid taxon, E. X sykesii

Barneby & Krukoff ( E . hybrida of horticultural literature)

which is cultivated in parts of southern Africa may have

as one of its parents E. speciosa.

Vouchers: Hennessy 395, 444 (UDW).

Section Erythrina Krukoff & Barneby: 365 (1974),

including sect. Corallodendra Krukoff: 394 (1974). Type

species: E. corallodendrum L.

21. E. pallida Britton & Rose in Bulletin of the Torrey

Botanical Club 48: 332 (1922); Krukoff: 263 (1939a);

Krukoff & Barneby: 399 (1974). Type: Trinidad, Britton

2656 (NY, holo.; GH, US, iso.).

E. corallodendron sensu Hennessy: 35 (1972), non L.

leones: Hennessy: 1. 13 (1972); Krukoff & Barneby: 399 (1974).

Tree up to 10 m tall. Leaves hysteranthous; leaflets

broadly ovate or parabolic, acute or obtuse, 70-100 x

60—90 mm. Inflorescences lateral, compact, erect; flowers

horizontal, cylindrical, not gaping. Calyx campanulate,

truncate with obtuse abaxial lobe ± 2 mm long, deep

purple. Corolla', vexillum linear-oblong, obtuse, shortly

clawed, minutely adpressed pilose distally, ± 70 x 20

mm, pink; alae oblong, obtuse, ± 12 x 6 mm; carina

segments free, obtusely spathulate, ±9x5 mm. Stamens

included. Fruit subligneous, moniliform, black. Seeds

particoloured red/black, ± 10 x 7 mm.

The unusual combination of deep purple calyx and pink

corolla make E. pallida easily recognisable. The specific

name, pallida refers to the pale vexilla. Indigenous in

several West Indian islands and Venezuela, this species

is widely cultivated in South America, Australia, tropical

east Africa and South Africa. The cultivated plants were

initially misidentified as E. corallodendrum L., a closely

related, red-flowered species with glabrous vexilla. Spring-

flowering.

Vouchers: Davidson 2240 (J); Hennessy 447 (UDW); Hilliard 4710

(UDW).

22. Erythrina aff. E. x sykesii Barneby & Krukoff

in Lloydia 37: 447 (1974). Type: C.T. White 9073 (NY,

holo.).

Icon: Krukoff & Barneby: 447 (1974).

Tree up to 10 m tall. Leaves hysteranthous; leaflets

resembling those of section Caffrae. Inflorescences lateral,

compact, semi-erect or ± horizontal; flowers horizontally

ascending, gaping. Calyx campanulate, unilaterally or

bilaterally asymmetrically cleft at anthesis, abaxial tooth

triangular-subulate. Corolla : vexillum ovate, acute, shortly

clawed, slightly spreading, 50—60 x ± 25 mm, bright

scarlet; alae linear-oblong, obtuse, ± 20 x 5 mm; carina

16

Bothalia 21,1 (1991)

segments partly connate, oblong-ovate, ± 25 x 10 mm.

Stamens exposed. Fruit and seeds unknown.

The history of the introduction of this hardy taxon into

southern Africa in unknown. It is widely cultivated in the

western Cape, southern Cape, parts of Namibia and at least

one specimen (purchased in the western Cape) is estab-

lished in Pietermaritzburg, Natal. The plants agree with

the description of E. x sykesii except that the carina is

invariably partly connate in locally cultivated plants. E.

x sykesii was described from plants cultivated in New

Zealand and Australia and in photographs of the

Australasian plants made by Dr Ian Whitton, the habit,

inflorescence and flower structure appear identical to those

of the specimens grown in southern Africa.

The parentage of the hybrid is unknown. In inflores-

cence shape and attitude; the well-developed triangular-

subulate abaxial calyx tooth; the scarlet colour of the

vexillum and in shape and relative proportions of alae and

carina, the hybrid resembles E. speciosa. In shape of

leaflets; the asymmetrically bilabiate calyx; the gaping

flower with exposed stamens and in connation of the

relatively large carina segments, the hybrid is like E.

cajfra. Both E. speciosa and E. caffra are cultivated in

Australasia and their flowering times overlap. Although

Krukoff & Barneby (1974) have suggested E. lysistemon

as a putative parent because the hybrid possesses a calyx

of the type known only in section Cajfrae, this seems

unlikely as E. lysistemon has a closed flower with included

stamens. The stature of the mature trees more closely

resembles that of E. caffra than that of E. lysistemon. Until

the hybrid is successfully resynthesised, the parentage of

this taxon remains speculative. Winter/spring-flowering.

Vouchers: Coetzees.n. sub NBG 16949 (NBG); Hennessy 461 (UDW).

SPECIMENS EXAMINED (ERYTHRINA)

The specimens are listed alphabetically according to the

name of the collector. The figures in parentheses refer to

the number of the taxon in the text. The herbaria in which

the specimens examined are housed, are indicated by the

letter codes of Holmgren & Keuken, Index herbariorum

(1974), except that of the University of Durban-Westville,

UDW, which is as yet unlisted.

Abbott 1263 (5) NH. Acocks 10233 (7) NH, PRE; 10733 (8) NH, PRE;

13163 (2) K; 18843 (9) PRE. Ahrens 14 (7) NU. Allsopp 1009 (7) NH,

NU, PRE. Archibald 5906 (6) PRE. Atkinson s.n. sub J 31397 (15) J.

Badri s.n. (5) UDW. Balkwill 361 (7) J. Barber 786 (8) K. Barbour s.n.

(9) K. Barnard s.n. sub SAM 33302 (3) SAM. Barrett 375 (7) PRE. Baur

249 (6) K, SAM. Bavuri s.n. (7) NBG. A. Bayer 1389 (5) NU. F. Bayer

s.n. sub NH 21765 (8) NH. Bayliss BR1-B-114 (6) GRA; 2079 (7) NBG;

2983 (9) NBG; 7046 (5) K. Bennie s.n. (7) GRA. Beswick s.n. sub Galpin

7370 (8) BOL. Bolus 6298 (8) BOL, K; 7733 (5) BOL; 8886 (9) BOL;

11831 (7) BOL; 11832 (2) BOL. Bokelmann s.n. (5) NBG. Botha s.n.

(7) K, PRE. Bourquin 6 (8) NU; 259 (5) NU. Bowker s.n. (7) K.

Brayshaw 11 (5) NU; 106 (8) NU. Bredenkamp 356 (8) PRE; 1480 (7)

PRE. Breyer s.n. sub TM 19416 (7) PRE; s.n. sub TM 24377 (5) PRE.

Brown & Shapiro 380 (2) K. Burchell 3701, 3920 (6) K. Burke 69 (8)

K, SAM; s.n. (8) K; s.n. (5) K. Burn Davy 615 (5) BOL; 999(8) NH;

4108 (8) BOL; 15024 (8) J, K.

Carnegie 1490/31 (2) NBG. Chase 5800 (5) K. Chetty 19 (7) UDW.

dinning s.n. (4) WIND. Codd 1896 (2) PRE; 1978 (8) PRE; 4418 (2)

K, PRE; 6042 (2) PRE; 6149(5) PRE; 6413 (7) K, PRE; 6547 ( 5) PRE;

7807 (7) K; 7817 ( 5) PRE; 7982 (5) GRA, K, PRE; 7983 (12) K; 7987

(5) GRA, K, PRE; 7988 (6) K; 7990 (6) K; 7991 (5) K; 7993 (5) GRA,

PRE; 7994 (5) K; 7998 (5) NH, PRE; 7999 (12) K, PRE; 8000 (6) K;

8002 (5) NH, PRE; 8003 (5) K; 8005 (5) GRA, PRE; 8006 (5) GRA;

8008 (5) NH; 9297 (7) GRA, K, PRE; 9403 (11) NH, PRE; 9406 (8)

K, PRE; 9744 (2) K, NBG, PRE; 10080 (8) K, PRE. Codd & Dyer 4636

(7) K, PRE. Coetzee 254 (5) PRE; s.n. sub NBG 16949 (22) NBG.

Collector un-named s.n. sub A 3 (7) GRA, PRE; s.n. sub NH 14466

(2) NH; s.n. sub Wood 13078 (7) NH. Collins s.n. sub J 32003 (5) J.

Colvin s.n. sub Moss 23024 (5) J. Comins 411 (7) NU; 440 (5) NU; 1435

(7) PRE. Compton 17745 (2) NBG; 19105 (7) BOL, NBG; 19794 (5)

BOL, NBG; 26643 (7) K, NBG; 26977 (5) K, NBG, PRE; 27446 (2)

NBG; 27903 (5) NBG; 27947 (2) NBG, PRE; 29774 (7) NBG; 31134

(5) NBG. Conrath 245 (8) K. Cooper 204 (6) K; 267 (9) BOL, K; 992

(8) K. Crewe 45 (7) K, NU. Culverwell 908 (5) PRE.

Dahlstrand 2382 (15) J. Davidson 2240 (21) J; 2276 (7) J; 22975 (5) J.

Davies s.n. sub A 7143 (7) GRA. Deane 7 (10) K, PRE. Dersley s.n.

sub J 30513 (8) J. De Winter 395 (5) PRE; 3745 (4) K, PRE, WIND;

5927; 4110 (4) PRE. De Winter <& Leistner 5601 (3) WIND. Dieterlen

547(8) NH, PRE, SAM. Dinter 769 (3) SAM. Dohse & Lindahl 106

(7) NH. Drege 1691 (6) BOL; s.n. 1836 (i), 1836 (ii), 1837 ( 9) K. Du

Plessis 15 (4) PRE. Du Toil 547 (8) NH, PRE, SAM.

Ecklon 1836a (6) K; 1836b (9) K. Ecklon & Zeyher 1692 (9) SAM.

Edwards 765 (7) NU; 1026 (8) NU. Eliovson s.n. sub J 26210 (15) J;

s.n. sub J 26293 (8) J. Eyles 5065 (10) K.

Flanagan 125 (7) BOL, NU, SAM; 319 (6) PRE; 1827 ( 8) SAM. Fiddes

5 (8) NU; 12 (7) NU. Fisher 412, 413 (8) NU. Forbes 350 (5) NH.

Fourcade 1406 (6) STE; 4466 (6) BOL, STE. Francois 89 (8) NU.

Frankish 8 (7) NU.

Galpin 67 (6) GRA; 494 (5) BOL, GRA; 1653 (9) BOL, GRA, NU,

PRE; 3388 (7) PRE; 11906 (7) K; 12989 (8) BOL, K, PRE; 14734 (7)

BOL, K, NH, PRE. Geldenhuis 270 (6) PRE. Germishuizen 350 (5)

PRE; 1632 (7) PRE; 3870 (8) PRE. Gerrard 134 (8) K. Gerstner 3356

(5) PRE; 4292 (8) NH; 5920 (7) PRE. Getliffe 157(2) NU. Gibson 25,

225 (3) WIND. Giffen 1043 (7) PRE. Giess 11515 (3) WIND. Gillett 2903

(5) K. Gilliland 3005, s.n. sub J 26231 (8) J. Glen 268 (7) J; 446 (5)

J. Goodman 889 (7) NU; 1110 (2) NU. Goossens 810 (8) K. J.L.

Gordon-Gray s.n. (6) NU. K.D. Gordon-Gray s.n. (8) NU. Green 14

(5) J. Gueinzius s.n. sub SAM 15586 (7) SAM. Guy & Jarman 249 ( 5) NU.

Haagner s.n. sub A 7145 (8) GRA. Hafstrom & Acocks 2309 (7) PRE.

Hall s.n. sub J 46746 (7) J. Hanekom 2321 (5) K, PRE; 2454 (8) K,

PRE. Harris s.n. sub NH 28844 (2) NH. Harrison 260 (5) PRE; 262

(7) NH. Hattingh 1 (5) PRE. Helner 356 (9) PRE. Hemm 155 (5) J,

PRE; 851 (7) J. Henderson 1503 (7) NBG; 1949 (8) NBG. Hennessy

264 (7) UDW; 385, 386 (1) UDW; 387 (14) UDW; 389, 390 (19) UDW;

393 (18) UDW; 395; 444 (20) UDW; 399 (17) UDW; 445 (12) UDW;

447 (21) UDW; 461 (22) UDW; s.n. (8) UDW; s.n. (11) UDW. Henrici

1210 (8) PRE. Henriques 238 (4) K. Herbst s.n. (8) NBG. Herdman

s.n. sub J 48616 (16) J. Hillary 388 (5) NU. Hillary & Robertson 570

(5) PRE. Hilliard 1339 (7) NU; 2191 (8) NU; 4710 (21) UDW. Hinges

3706 (3) PRE. Hojfe 16 (8) PRE. Holland s.n. sub NBG 306/26 (7) BOL.

Holt 344 (2) PRE. Hornby 2722 (19) K. Hosken 16 (1) NU. Howlett s.n.

(5) NH. B. Huntley 697 (1) NU. K.D. Huntley 223 (2) NU. Hutchinson

2531 (7) BOL. Hutchinson & Gillett 4155 (5) K.

Jack s.n. sub J 36293 (16) J. Jacobsen 2602 (7) PRE. Jacot Guillarmod

8502 (6) K. Jankowitz 574 (3) WIND. Jarman & Guy 36 (7) NU. Jenkins

s.n. sub TM 9294 (8) PRE. Jenkinson s.n. sub TM 7339(5) PRE. Johnson

1332 (11) K, NH, PRE.

Keerath 11 (5) UDW. Keet 1614 (3) WIND. Kemp 21689 (5) J. Kerfoot

7379 (7) J. Killick 304 (7) NU. Kirk 154 (19) K. Kluge 1386 (5) PRE.

Krauss 62 (7) K; 263 (2) K, PRE; 286 (5) K. Krupko 25826, 27258 (7) J.

Langham s.n. sub NH 12129 (8) NH. Lansdell s.n. sub NH 16114 (2)

NH. Larson 82 (5) J. Lawn 1012 (2) NH. Lawson 285 (7) NH. Leach

8463 (7) K. Leendertz 217 (5) PRE; 693 (8) PRE; 953 (5) PRE; 2681,

3899 (8) PRE. Leith s.n. (8) PRE. Le Roux 216 (4) PRE, WIND. Uengme

453 (7) PRE. Louw 541 (8) NH, PRE. Lucas 32 (8) J. Lugg s.n. sub

NH 40477 (10) NH.

Macnae 1582 (8) J. MacOwan 1216 (1) BOL, K, SAM; 1413 (6) BOL,

K; 1435 (9) K; s.n. (7) GRA. Maguire s.n. sub Moss 22208 (5) J. Mal-

comess s.n. (6) NBG. E. Marais s.n. (5) PRE. W. Marais 476 (6) PRE;

1331 (8) K, PRE. Markotter s.n. sub STE 31572 (2) STE. Martin s.n.

(6) NU. Martindale s.n. (5) NU. McCabe & Alherstone 41 (9) K. McClean

6 Ogilvie s.n. sub NH 27899 (5) NH. McDonald 178 (5) NU;

Bothalia 21,1 (1991)

17

254 (2) K, NU, PRE. McGregor s.n. sub Moss 2628 (13) J. McKen s.n.

(7) K. Meeuse 9335 (13) K. Melville 101276 (5) K. Merxmuller & Giess

1241 (3) WIND. Methven 156 (5) K. Miller 10 (5) NU. Mogg 18222,

23431, 33387 (5) J; 18372, 23360, 25235 (8) J; 21368 (7) J. S B. Mogg

s.n. sub Mogg 18147 (8) J. Mohle 398 (5) PRE. Moll 1241-, s.n. (9) NH.

Moll & Strey 3887 (7) K, NH, PRE. Molyneux s.n. (6) K, NH. Morris

225 (7) NU; 534 (7) K, NH, NU, PRE. Moss 4548, 4551 (5) J; 4549,

8606, 10623, 18231 (8) J; 4552 (7) J; 4553 (2) J. Moss & Rogers 1134

(8) J. Mugwedi 19 (10) J; 1299 (2) PRE; 1520 (2) J. D. 11 Muller 1052

(8) PRE. M. Muller 3502 (3) WIND. Munro 21638 (5) J.

D.B. Naidoo 13 (5) UDW. V. Naidoo 4 (5) UDW. Nation s.n. (13) K.

Neethling s.n. (8) K. Nel 177 (2) NH. Nelson 84 (7) K; 223, 522 (8)

K. Netschisaulu 2200(2) PRE. Netshiungani 851 (7) PRE. Nichols 779

(11) NH. Nicholson 2 (5) NH. Noel s.n. sub A 1415 (5) GRA.

Onderstall 740 (2) PRE.

Pascoe & Waugh 6 (7) NH. Patel 1 (5) UDW. Pegler 89 (6) GRA, K,

PRE, SAM; 235 (2) BOL, GRA, K, SAM; 1134 (6) BOL; 1179(1) BOL,

GRA, SAM. Peitch & Son s.n. (9) K. Pelletier s.n. sub J 30478 (5) J.

Pennefather 81 (2) NU. Phelps 283 (19) K. Phillips s.n. (5) PRE. Pole

Evans 13212 (8) K; s.n. (9) PRE. Pont 530 (8) PRE. Poole y 815 (7) NU.

Porter 326 (1) J. Porter & Ward 279 (7) NH. Pott s.n. sub TM 15084

(8) PRE; s.n. sub J 37663 (2) J; s.n. sub J 37870 (5) J. Preller 135 (8)

PRE. Pretorius s.n. (8) NBG. Pringle s.n. (9) NBG. Prior 26 (5) K,

PRE; 359 (2) K, PRE. Prosser 1038 (5) J, K, NBG, PRE; 1155 (8) J,

K, NBG, PRE; 1356 (7) J, K, NBG. Purchase 2 (5) J; 57 (7) J.

Ramsay s.n. sub Wilson s.n. (8) PRE. Rehmann 6282 (2) K. Reynolds

1606 (9) K, PRE; 5770 (8) K; 9673 (2) NBG. Roberts 2887 (8)’ PRE.

Rodda 6 (5) NU. Rodin 3976 (1) BOL, K. Rogers 1629 (7) J; 4634 (7)

K; 14814 (8) J, K; 23010 (2) J; s.n. sub A 7142 (9) GRA. Rose-Innes

47(5) J, K, PRE. Ross 1282 (5) NU; 2123 (5) PRE. Ross & Moll 5053

(2) K, PRE; 5069 (6) NH. Rudatis 23 (7) STE; 268 (8) STE; 412, 781

(5) K. Rudd 4 (5) NU. Rump s.n. (2) NU. Rusch f. 7958 (4) BOL, K.

Rycroft 2980 (8) NBG.

Sanderson 44 (2) K; 863 (8) NH. Saunders s.n. sub NH 3440 (8) NH.

Scharedfezen 1/300 (3) WIND. Scheepers 429 (5) PRE; U71 (2) K, PRE.

Schlieben 7009 (5) K. Schrire 479 (7) NH; 632 (8) NH; 1854 (11) NH.

Seydel 369 (3) K; 4048 (3) WIND. Shirley s.n. (8) NU. Short s.n. sub

Mogg 25887 (5) J. Sim 1254, 1255 (7) BOL, NU, PRE; 1256 (6) NU;

1257(9) NU; 1258 (2) NU; 1362 (2) BOL; 21134 (19) NU. Singh 13 (5)

UDW. Smellie s.n. sub J 22490 (8) J. C.A. Smith 308 (5) PRE; 7051

(2) PRE. K. Smith 27(8) BOL. PA. Smith 1872 (4) K, PRE; 2928 (1)

K, PRE. Smuts 66 (5) K, PRE; s.n. (5) PRE. Somers s.n. (7) NH.

Standard V11D1 (8) PRE. Stephen 703 (1) K, PRE. Steyn 810(8) NBG.

Stirton 8206 (5) NH. Store s.n. (8) K. Story 1528 (5) PRE. Steinke s.n.

sub UDW 9291 (14) UDW. Strey 3614 (7) K, PRE; 3994 (7) PRE; 5698

(1) NU, UDW; 6663 (6) NH, PRE; 9018 (5) K, NH; 9069 (2) NH; 9134

(2) PRE; 10574, 10575, 10579(5) NH; 11157(1) K, NH, NU. Suther-

land s.n. (2) K. Swierstra s.n. sub T.M. 6231 (8) PRE.

L. E. Taylor 5453 (7) NBG. R. Taylor 26 (7) NH. Tinley 756 (7) NU.

Thakersee 2 (5) UDW. Theron 1472 (5) PRE. Thode s.n. sub A 2799

(8) PRE; s.n. sub STE 31574 (2) STE; s.n. sub STE 31575/6 (8) STE;

s.n. sub STE 31577 (7) STE; s.n. sub STE 31578 (6) STE; s.n. sub STE

31580/1 (5) STE; s.n. sub STE 31582 (8) STE. Thunberg s.n. (6) (photo.)

K. Trauseld 870 (8) NU. Turner 16 (5) K. Tyson 1053 (2) BOL, SAM;

1761, 3124 (1) SAM; s.n. (9) BOL, K.

Vahrmeijer & Du Preez 2555 (3) PRE. Van der Bijl s.n. sub STE 31579

(5) STE. Van der Schijff 13 (5) PRE; 402 (5) PRE; 1126 (2) PRE. Van

Vuuren 1225 (5) PRE. Van Wyk 1656 (5) PRE. Venter 7890 (7) PRE.

Venter & Vorster 167 (5) PRE. Verdoorn s.n. sub PRE 22839 (8) PRE.

Von Koenen 582 (3) WIND.

Wade s.n. (5) NU. Walter 305, 614 (3) WIND. Ward 481 (5) NU; 1135

(1) NU; 1511 (5) NH, NU; 2232 (1) NU; 2688 (2) K, NH, NU, PRE;

2954 (10) NH; 3014 (6) K, NU; 4274 (5) PRE; 4306 (6) NH, NU; 5779

(6) NH, NU, UDW. M.C. Ward 192 (7) NU. Watt s.n. (4) WIND; s.n.

sub J 25519 (8) J. Weame 80 (1) NU, PRE. Wells 1465 (5) NU; 1547

(2) NU; 2114 (7) K. Werdermann & Oberdieck 1934 (7) K ex B, PRE;

1226 (8) K ex B. Werger 176 (8) K. West 133 (7) K, NH; 2134 (2) NH;

3276 (4) K. Wiss 372 (3) PRE. Wood 4076 (8) K, NH; 9776 (7) NH;

10016 (5) BOL, NH; s.n. sub Galpin 3388 (7) PRE. R. Wood 164 (7)

NU. Wylie s.n. sub Wood 9224 (7) NH; s.n. sub NH 22344 (7) NH, PRE.

Young 1140 (8) J.

Zeyher 531 (8) K, PRE, SAM; s.n. sub SAM 32895 (7) SAM.

MUCUNA

Mucuna Adans., Families des Plantes 2: 325 (1763),

nom. cons.; DC.: 404 (1825); Benth.: 533 (1865); Bak.:

184 (1871); Bak.: 185 (1876); Taubert: 266 (1894); Taubert:

194 (1897); Prain: 404 (1897); Phillips: 337 (1926b);

Hutch.: 405 (1927); Bak. f.: 378 (1929); Phillips: 423

(1951); Hauman: 126 (1954); Hauman: 98 (1955); Hepper:

561 (1965); White: 160 (1962); Hutch.: 433 (1964); Torre:

250 (1966); Verde.: 287 (1970); Verde.: 561 (1971); Dyer:

269 (1975); Compton: 278 (1976); Verde.: 743 (1980); Lack-

ey: 312 (1981); Troupin: 319 (1982). Type species: Mucu-

na urens (L.) DC. (conserved type).

Stizolobium P. Br.: 290 (1756) nom. rej. Type species:

Stizolobium pruriens (L.) Medik. = Dolichos pruriens

L. = Mucuna pruriens (L.) DC.

Zoopthalmum P. Br.: 295 (1756) nom. rej. Type species:

Dolichos urens L. = Mucuna urens (L.) Medik.

Cacuvallum Medik.: 392 (1787). Type species: Cacuval-

lum altissimum (Jacq.) Medik. = Dolichos altissimus

Jacq. = Mucuna altissima (Jacq.) DC.

Citta Lour.: 456 (1790). Type species: Citta nigricans

Lour. = Mucuna nigricans (Lour.) Steud.

Marcanthus Lour.: 460 (1790). Type species: Marcan-

thus cochinchinensis Lour. = Mucuna utilis Wall, ex

Wight = M. pruriens (L.) DC. var. utilis (Wall, ex Wight)

Bak. ex Burck.

Homera Neck.: 43 (1790). Type species: Homera urens

(L.) Neck. = Dolichos urens L. = Mucuna urens (L.)

Medik.

Negretia Ruiz & Pav.: (1794). Type species: Negretia

urens (L.) Ruiz & Pav. = Dolichos urens L. = Mucuna

urens (L.) Medik.

Labradia Swed.: 164 (1801). Type species: Labradia

urens (L.) Swed. = Dolichos urens L. = Mucuna urens

(L.) Medik.

Carpopogon Roxb. : 54 (1814); Roxb. : 283 (1825). Type

species: Carpopogon giganteum (Willd.) Roxb. =

Dolichos giganteus Willd. = Mucuna gigantea (Willd.)

DC.

Macroceratides Raddi: fig. 5 (1820). Type species:

Macroceratides pseudo- stizolobium Raddi = Mucuna

macroceratides DC.

Pillera Endl.: 91 (1833). Type species: Pillera macro-

carpa (Wall.) Endl. = Mucuna macrocarpa Wall.

Woody lianes or herbaceous climbers; hirsute or

glabrescent. Leaves stipulate, stipellate, pinnately trifolio-

late. Inflorescences axillary, paniculate, pseudoracemose

or subumbellate; bracts and bracteoles deciduous; flowers

18

Bothalia 21,1 (1991)

maroon, cardinal red, lilac or green. Calyx tube broadly

campanulate, limb bilabiate, adaxial lip of two + connate

lobes, abaxial lip 3-lobed. Corolla : vexillum shorter

than alae and carina, ovate-orbicular, shortly clawed,

proximally auriculate; alae slightly shorter than or

equalling carina; carina rostrate, beak horny. Stamens 10,

diadelphous; anthers dimorphic, 5 large sub-basifixed

alternating with 5 shorter versatile or dorsifixed, anthers

barbate in subgenus Mucuna, glabrous in Stizolobium.

Ovary sessile or stipitate, few-ovuled; style filiform,

incurved, proximally pubescent; stigma capitate, small.

Fruit ovoid or oblong, valves thick, with an indumentum

of irritant bristles; sutures wingless or winged. Seeds

biconvex lenticular, exarillate, with elongated hilum

girdling more than half the circumference (subgenus

Mucuna) or ellipsoid-oblong with conspicuous rim aril and

short linear hilum (subgenus Stizolobium). 2n = 22.

A genus of + 100 circumtropical and subtropical

species, three of which are indigenous in the FSA region.

Pending revision of the genus on a world basis, Mucuna

is retained in its broad sense, with two well-defined sub-

genera, Mucuna and Stizolobium (P. Br.) Prain. Both

subgenera are represented in southern Africa.

POLLINATION AND ECONOMIC IMPORTANCE

Red-flowered species of Mucuna are mostly bird-

pollinated; green or pale-flowered species mainly

bat-pollinated and Kenneally (in Verdcourt 1980) has

suggested that at least one Australian species, M. reptans

Verde., a prostrate plant with musk-scented, mottled

blackish purple, red, yellow, or green flowers, is rodent-

pollinated.

Despite the attractiveness of the flowers of many species

the genus has little economic value because the irritant

bristles make the material difficult, even dangerous to

handle. Exceptions are M. nova-guineensis Scheff. , ‘red

jade vine’, a woody liane cultivated for its spectacular

flowers and M. pruriens (L.) DC. var. utilis (Wall, ex

Wight) Bak. ex Burck, ‘Florida velvet bean’, an innocuous,

large-seeded cultivar of unknown origin cultivated as a

fodder crop. Both these taxa are grown in the FSA region.

Ingestion of irritant bristles of many species induces

severe diarrhoea. In primitive medicine, potions containing

irritant bristles derived mainly from pods were employed

as vermifuges. Severe dermatitis and inflammation of the

eyes result from contact with the urticating bristles, and

inflammation of nasal and buccal mucosa, laryngitis and

gastro-intestinal disturbance from inhalation and/or

ingestion of the bristles. Cases of dementia and even death

induced by accidental exposure of domestic animals to

Mucuna bristles have been reported (Burtt Davy 1932).

A pruritogenic proteolytic enzyme, mucunain, and a

histamine-releasing substance, 5-hydroxytryptamine

(serotonin) are toxic principles isolated from the irritant

bristles of Mucuna spp. (Watt & Breyer-Brandwijk 1962).

Vernacular names of the two indigenous species of

subgenus Stizolobium are hell-fire bean, brandboontjie,

jeukpeul and isifefeta.

I. Subgenus Mucuna: Verde.: 561 (1971). Zoop-

thalmum P. Br. : 295 (1756). Subgenus Zoopthalmum (P.

Br.) Prain: 406 (1897). Type species: Mucuna urens (L.)

Medik.

Woody lianes; anthers dimorphic, barbate; fruit with

winged sutures; seeds biconvex lenticular, exarillate; hilum

elongated girdling more than half the circumference of the

seed.

II. Subgenus Stizolobium (P. Br.) Prain in Journal

of the Asiatic Society, Bengal 66: 406 (1897); Verde.: 561

(1971). Stizolobium P. Br.: 290 (1756). Type species:

Mucuna pruriens (L.) DC.

Herbaceous climbers; anthers dimorphic, beardless;

fruit wingless; seeds ellipsoid-oblong with conspicuous

rim aril; hilum short, linear.

KEY TO INDIGENOUS SPECIES OF MUCUNA

la Woody liane; flowers green or greenish white; anthers hirsute;

fruit 80-150 x 40—60 mm, each suture bordered longi-

tudinally by two wings; seeds biconvex lenticular, ± 25

mm diameter; hilum girdling, ± 50 mm long

1. M. gigantea

lb Herbaceous climbers; flowers dark red, purple or lilac;

anthers glabrous; fruit 50 -90 x 10—15 (20) mm, sutures

wingless; seeds ellipsoid-oblong, 10-18 x 7-12 mm;

hilum straight, ± 5 mm long:

2a Leaflets chartaceous, acute or acuminate, not blackening

when dried, grey pilose; calyx silvery grey pubescent

with or without admixed orange irritant bristles; fruit

with or without irritant bristles; rim aril cream:

3a Fruit thickly invested with golden orange or red-brown

irritant bristles; pericarp with longitudinal wrinkles

obscured by indumentum; seeds pinkish brown with

darker mottling or dark blackish brown, ± 10 x

8 mm 2a. M. pruriens var. pruriens

3b Fruit lacking irritant bristles, glabrous or velutinous;

pericarp with obvious longitudinal wrinkles; seeds

white, mottled or black, 12—18 x 10—12 mm

2b. M. pruriens var. utilis

2b Leaflets coriaceous, obtuse, blackening when dried, grey

velutinous or grey pubescent; calyx indumentum of

admixed adpressed golden irritant bristles and golden

pubescence; fruit thickly invested with golden orange

irritant bristles; pericarp lacking longitudinal wrinkles;

seeds pinkish brown or mottled, ±8x7 mm; rim

aril black 3. M. coriacea

1. M. gigantea (Willd.) DC ., Prodromus systematis

naturalis 2: 405 (1825); Verde.: 546 (1971); Hennessy: t.

1998 (1989). Dolichos giganteus (Willd.: 1041 (1802).

Carpopogon giganteum (Willd.) Roxb.: 54 (1814). Stizolo-

bium giganteum (Willd.) Spreng.: 281 (1827). Type: India,

Malabar; Rheede, Hort. Malab. 8: 63, t. 36 (1688)

(icono.).

M. quadrialata Bak.: 186 (1871); Bak. f.: 379 (1929). M. gigantea

(Willd.) DC. subsp. quadrialata (Bak.) Verde.: 287 (1970); Verde.: 564

(1971). Types: Mozambique, Zambezi River, Kirks.n. (K, syn. !); Zanzibar

Is., Kirk s.n. (K, syn.!).

M. longipedicillata Hauman: 98 (1955). Type: Congo, Kivu,

Ghesquiere 3771 (BR, holo.; K, iso.!).

M. flagellipes sensu Bak. f. : 379 (1929), quoad Bagshawe 1312, non

Hook. f.

leones: Rheede: t. 36 (1688); Verde.: fig. 82.2 (1971); Wilmot-Dear:

fig. 2A,B,C (1984); Hennessy: t. 1988 (1989).

Bothalia 21,1 (1991)

19

Woody liane ± 30 m long, ± 70 mm diameter at base;

branches glabrous. Petiole pulvinate, 50-100 mm long;

rhachis 20-35 mm long; stipels filiform, + 3 mm long;

petiolules ± 6 mm long; leaflets coriaceous, ovate or

elliptic, the laterals asymmetrical, 40-120 x 20-75 mm,

acuminate and apiculate, adpressed pale-hirsute at first,

later glabrous. Inflorescences borne on woody stems 2-3

years old; axillary, paniculate, pendent; peduncle (80— )130

-200 mm long, floriferous in distal 25-30 mm;

secondary branches 5—11, ± 10 mm long, each (2-)

3-flowered; flowers resupinate or not; pedicels 12-22 mm

long, hirsute; bracts oblong, 7,0 x 2,5 mm, caducous,

hirsute, green becoming black; bracteoles paired at

apex of pedicel, oblong, 12 x 5 mm, caducous, hirsute,

blackening. Calyx green (R.H.S. Agathia green 60),

velutinous with admixed golden stinging bristles; tube

broadly cup shaped, ± 10 x 15 mm; lobes 5, unequal,

broadly triangular, subacute, 4, 5-5,0 x 5,5-10,0 mm, the

upper pair connate forming an emarginate lip. Corolla :

vexillum green (R.H.S. pod green 061), glabrous or hirsute

near proximal margins, suborbicular with obtusely bilobed

apex, ± 30 x 30 mm; alae green (R.H.S. pod green

061/2), proximally hirsute, lanceolate, subacute, ± 50 x

15 mm; carina partly connate, green (R.H.S. pod green

061/3), upcurved ± 90° in free, thickened, distal 5 mm

with incurved horny apiculus ± 1,5 mm long, + 47 x

10 mm. Stamens diadelphous, + 45 mm long; anthers

included, dimorphic alternating, adaxial versatile, ovoid,

densely barbate; abaxial sub-basifixed, obelliptic, sparse-

ly hirsute. Intrastaminal nectariferous disc 10-lobed, ±

1,5 mm high. Ovary subsessile, densely hirsute, 3-ovuled;

style proximally hirsute, flexed midway, distally glabrous,

upcurved; stigma capitate, shaggy proximally. Fruit (2-)3-

seeded, oblong or elliptic-oblong, 80-150 x 40-60 mm,

each suture 2-winged; indumentum dense, of golden

irritant bristles. Seeds brown, mottled, biconvex lenticular,

25—29 X 21—25 mm, exarillate; hilum girdling, 50 mm

long (Figure 9).

Recorded in the FSA region from two localities,

Mdloti River estuary and Sihadla, Kosi River system

in Natal (Figure 10), from Mozambique, Uganda,

Tanzania, Kenya, Zaire, Zanzibar Is, Mascarene Is,

Seychelles Is, and also India, Malesia, China, Taiwan,

Japan, Australia and Polynesia. Over much of its range

its habitat is coastal, near water, on forest margins, but

in East Africa it has been recorded inland from altitudes

up to 1 140 m.

The Mdloti population is rooted on the landward side

of an estuarine fringe of Barringtonia racemosa (Lecy-

thidaceae), with the vine-like stems extending into the

upper canopy. The pendent, green-flowered, pseudo-

umbellate inflorescences are in the canopy and very

difficult to discern. Anthesis occurs simultaneously in an

inflorescence. Nectar production is copious. Olive sun-

birds (Cyanomitra olivacea ) visit the flowers, yet no

seed-set has been observed in this population, therefore

it is assumed that the sunbirds are not effective pollinating

agents for this species. This assumption is strengthened

by the observation that fallen flowers are untripped, with

anthers and stigma included in the carina. It is thought

that M. gigantea is chiropteriphilous and that the more

southerly local population may lie south of the normal

range of the effective pollinator(s).

Vouchers: Gillett 2I573/B (K); Mogg 28907 (K, J); Muir 4769 (K);

Vaughan 2124 (BM); Ward & M.C. Ward 40 (K, UDW); Ward, M.C. Ward

& R. Kyle 92 (UDW).

2. M. pruriens (L.) DC., Prodromus systematis

naturalis 2: 405 (1825); Bak.: 187 (1871); Bak.: 187 (1876);

Hutch.: 405 (1927-28); Bak. f.: 380 (1929); Hauman: 128

(1954); Hepper: 561 (1954); White: 160 (1962); Torre:

252 (1966); Verde.: 566 (1970). Dolichos pruriens L. in

Stickman: 23 (1754b); L.: 1162 (1759). Stizolobium

pruriens (L.) Medik.: 399 (1787). Type: Indonesia,

Rumphius Herb. Amb. 5, t. 142 (1750) (icono.!).

2a. var. pruriens

leones: Rumphius: t. 142 (1750); Browne: t. 31.4 (1756); Hauman: fig.

2A, B, C (1954); Verde.: fig. 82.7 (1970).

2b. var. utilis (Wall, ex Wight) Bak. ex Burck in

Annales du Jardin botanique de Buitenzorg 11: 187 (1893).

M. utilis Wall, ex Wight: 280 (1840). Type: not designated.

Marcamhus cochinchinensis Lour.: 461 (1790). Type: Cochin China,

Louriero s.n. (BM, holo.).

Carpopogon niveus Roxb.: 54 (1814). M. nivea (Roxb.) Wight & Am.:

255 (1834). Type: Roxburgh drawing no. 1601 (K, lecto., designated by

Wilmot-Dear 1984).

C. capitatus Roxb.: 284 (1832). Mucuna capitata (Roxb.) Wight &

Arn.: 255 (1834). Type: Roxburgh drawing no. 285 (K, lecto., desig-

nated by Wilmot-Dear 1984)

M. velutina Hassk.: (1842). Type: specimens cultivated at Bogor.

Stizolobium deeringianum Bort: 31 (1909). Mucuna deeringiana (Bort)

Merrill: 118 (1910). Type: Argo, Florida, 1890, S.C. Carleton (not located).

leones: Bort: 31.2,3 (1909); Hauman: fig. 2D.

Twining herb up to 3 m long. Stems hirsute becoming

glabrous. Petiole 70-260 mm long; stipels subulate, 4-5

mm long; leaflets chartaceous; terminal leaflet ovate,

obovate or elliptic, (55— )85 — 140( — 160) x (45— )80-100

(—115) mm, acute, acuminate or, rarely, obtuse, apiculate;

lateral leaflets markedly asymmetrical, (55— )70— 190 x

(45— )80— 190 mm, with sparse to dense adpressed whitish

indumentum. Inflorescence axillary, paniculate, deflexed

or pendent; peduncle (20— )30-40 mm long, silvery

pubescent; secondary branches 5 mm long, 2-3-flowered;

bracts and bracteoles caducous; pedicels 2—4 mm long.

Calyx green with adpressed whitish indumentum; tube

cupuliform, 6 x 10 mm; lobes 3—9 mm long, abaxial lobe

narrowly triangular, laterals broadly triangular, the upper

pair connate to form an acute lip. Corolla-, vexillum

maroon or pale lilac, ovate, 17—22 x 14—15 mm; alae

26-35 x 12 mm; carina 27- 28 mm long with sharply

upcurved apical homy beak. Stamens diadelphous; anthers

dimorphic, glabrous. Ovary and style-base densely hirsute;

style flexed midway; stigma capitate, sparsely fringed.

Fruit 4— 5(-6)-seeded, oblong or sigmoid, 40—110 X

13 — 15( — 20) mm, longitudinally wrinkled, with dense

indumentum of brown or rufous stinging bristles (var.

pruriens) or, in cultivated forms (var. utilis), glabrescent

or velutinous. Seeds black or mottled brown, oblong-

ellipsoid, compressed, 10 x 7,5 mm (var. pruriens) or,

in cultivated forms (var. utilis), white, brown, mottled or

black, 12-19 x 10-13 mm; rim aril creamy white (Figure

11).

20

Bothalia 21,1 (1991)

FIGURE 9. — Mucuna gigantea: 1—12, Ward & Hennessy s.n. (flowers in spirits); 14, Gillett 21573/B\ 15, Muir 4769. 1, inflorescence; 2, juvenile

shoot and young leaf; 3, fully open flower; 4, vexillum; 5, ala; 6, carina segment; 7, gynoecium, all X 0,8; 8, basifixed anther, face view

after dehiscence; 9, basifixed anther, dorsal view; 10, versatile anther, face view after dehiscence; 11, versatile anther, lateral view prior

to dehiscence, all x 5,3; 12, stigma, X 8; 13, lobed nectary, x 2,3; 14, fruit, x 0,8; 15, seed, x 0,8.

Bothalia 21,1 (1991)

21

FIGURE 10. — Distribution of Mucuna gigantea , ■; M. pruriens var.

pruriens. A; and M. coriacea , #

The wild form, M. pruriens var. pruriens has been

recorded from only two localities in the FSA region to date

(Figure 10). Elsewhere widely distributed in tropical

Africa, Madagascar, Asia and tropical America. The

innocuous cultivated form, var. utilis, which is larger in

all its dimensions than var. pruriens and is of unknown

origin, is widely used as a fodder crop and as green

manure.

M. pruriens var. pruriens is distinguished from M.

coriacea mainly by its thinner, larger, usually acute or

acuminate leaflets; the silvery calyx indumentum; the

longitudinally wrinkled pericarp and the white aril.

Vouchers: var. pruriens : Codd 6545 (K, PRE); Glen s.n. sub TM 11460

(PRE); Glen s.n. sub TM 11554 (PRE). var. utilis: Du Plessis s.n. sub

PRE 53353 (PRE); Schrire 908 (NH).

with emarginate apex, ± 35 x 22 mm; alae maroon with

a zone of golden hairs, ± 40 X 14 mm; carina ± 40 mm

long with sharply upcurved horny beak. Stamens diadel-

phous; anthers dimorphic, glabrous. Ovary and style-base

densely hirsute; style flexed midway; stigma capitate,

sparsely fringed. Fruit 4-5-seeded, oblong, curved,

50—80 X 10-15 mm, without longitudinal wrinkles, with

dense indumentum of golden orange irritant bristles. Seeds

brown or mottled brown, elliptic-oblong, compressed,

8-9 x 7—7,5 mm; rim aril black (Figure 11).

Material of this African species from Tanzania, Zambia,

northern Mozambique and Malawi has been placed in

subsp. coriacea and larger-flowered forms from Tanzania,

Uganda, Zaire, Zimbabwe, Angola, southern Mozambique

and southern Africa in subsp. irritans (Figure 10). The

indumentum of the pericarp of subsp. irritans is heavier

than that of subsp. coriacea, whereas that of the leaves

is more dense in subsp. coriacea. The merit of the separa-

tion of two subspecies is questionable as intermediates

exist.

Characters which distinguish M. coriacea from M.

pruriens are mentioned under M. pruriens.

Vouchers: Compton 27664 (K); Crawford 465 (PRE); Hemm 903 (J);

Netshiungani 903 (PRE); Stirton 8682 (PRE).

SPECIMENS EXAMINED ( MUCUNA )

The specimens are listed alphabetically according to the

name of the collector. The figures in parentheses refer to

the number of the taxon in the text. The Herbaria in which

the specimens examined are housed, are indicated by the

letter codes of Holmgren & Keuken, Index herbariorum

(1974), except that of the University of Durban-Westville,

UDW, which is as yet unlisted.

3. M. coriacea Bak. in Flora of tropical Africa 2;

187 (1871); Bak. f.: 380 (1929); Verde.: 288, 291, 292

(1970); Verde.: 567 (1971); Compton: 378 (1976). Types:

Mozambique, Chupanga, Kirk s.n. (K, syn.!); Malawi,

Manganja Hills, Metier s.n. (K, syn.!).

subsp. irritans (Burtt Da\y) Verde, in Kew Bulletin

24: 292 (1970); Verde.: 568 (1971); Onderstall: 116 (1984).

M. irritans Burtt Davy : 414 (1932). Type: Transvaal,

Nelspruit, Rogers 20255 (K, holo.!; J, iso.!).

M. coriacea sensu Hauman: 131 (1954), non Bak. sensu stricto.

Icon: Onderstall: p. 117 (1984).

Twining herb ± 3 m long. Stems with adpressed fulvous

hairs. Petiole 30—60 mm long; stipels subulate; leaflets

coriaceous, elliptic-ovate, obtuse and apiculate, 40—90 x

30—70 mm, the laterals asymmetrical, with adpressed

fulvous hairs, becoming subglabrous. Inflorescence axil-

lary, paniculate, deflexed or pendent; peduncle (12—)

25 — 70(— 110) mm long, fulvous pubescent; secondary

branches ± 5 mm long, 2— 3-flowered; bracts and

bracteoles caducous; pedicels 4-10 mm long. Calyx green

with dense indumentum of mixed fulvous pubescence and

adpressed tawny stinging bristles; tube cupuliform, 6 x

10 mm; lobes broadly triangular, the upper pair connate

to form an emarginate lip. Corolla : vexillum maroon, ovate

Bagshawe 1312 (1) BM; 1404 (3) BM. Balsinhas & Marrime 381 (2a)

BM. Bayliss 2719 (3) NBG. Bingham 208 (2a) (BM). Bruyns-Haylett

131 (3) NU. Burger 29 (3) PRE. Bunt 2202 (3) BM. Bunt Davy 2578

(3) K. Button s.n. sub PRE 56597 (3) PRE. Chase 888 (2a) BM. N.C.

Chase s.n. (3) NU. Codd 6545 (2a) K, PRE; 7813 (3) K, PRE; 7900

(3) PRE. Compton 27664 (3) NBG, PRE. Crawford 465 (3) PRE. Cross

22 (3) NBG.

Du Plessis s.n. sub PRE 53353 (2b) PRE. Du Toil 2423 (2b) NH. Dyer

3161 (3) K, PRE.

Eyles 361 (3) BOL, SAM.

Faulkner 250 (3) BM.

Galpin 9647(3) PRE; 9670(3) K, PRE. Gillett 2173/B (1) K. Glen s.n.

sub TM 11460 (2a) PRE; s.n. sub TM 11554 (2a) PRE. Gossweiler 5656,

6538 (2a) BM. Grobbelaar 579, 1096 ( 3) PRE.

Hemm 903 (3) J. Holland s.n. (3) BOL. Holt 248 (3) PRE.

Leach 12735 (3) K, PRE. Leeuwenberg 11024 (3) PRE. Legat s.n. sub

PRE 150 (3) PRE. Lemos & Marrime 324 (2a) BM. Liebenberg 2453

(3) PRE.

Mogg 28907 (1) J; 30431. s.n. sub J 36208, s.n. sub J 35333 (3) J. Mogg

& Pedro 3877 (3) PRE. Moss 18449 (3) J. Moss & Rogers 196 (3) J.

Muir 4769 (1) K. Mundy 2799 (3) BM.

Netshiungani 903 (3) PRE.

Obermeyer 29216 (3) PRE.

Pearson 2168 (2a) BOL. Pierce 19 (3) PRE.

Repton 883 (3) PRE. Rodin 4138 (3) BOL, K, PRE. Rogers 20225 (3)

J, K; 20829 (3) PRE; 23850 (3) K; 23857 (3) J.

22

Bothalia 21,1 (1991)

FIGURE 11. — Mucuna coriacea subsp. irritans: 14.1—14.9, Grobbelaar 579: 14.10, 14.11, Stirton 8682. 1, part of flowering shoot; 2, vexillum;

3, ala; 4, carina segment, all x 0,8; 5, 6, versatile anther, side and dorsal views; x 5,4; 7, 8, basifixed anther, face and dorsal views,

X 5,4; 9, stigma, X 9,6; 10, infructescence, x 0,8; II, seed X 0,8. Mucuna pruriens var. pruriens, Glen s.n. sub TM 11554 : 12, fruit,

X 0,8. Mucuna pruriens var. utilis, Du Plessis s.n. sub PRE 53353 : 13, fruit, X 0,8; 14, seed, X 0,8.

Bothalia 21,1 (1991)

23

Sampson s.n. (3) K; s.n. sub PRE 4412 , s.n. sub PRE 53348 (3) PRE.

Saunders 166 (3) K, NH. Scheepers 123, 218 (3) PRE. Schlechter 4552

(3) BOL. Schlieben 6281 (3) BM. Schrire 908 (2b) NH. Sim 20716 (3)

NU. Smuts s.n. sub PRE 53390 (3) PRE. Stephen 210, 297 (3) PRE.

Steinacker 3544, s.n. sub PRE 172 (3) PRE. Slirton 1753 A (3) K; 8682,

8739 (3) PRE. Swynnerton 450 (3) BM.

Teague 495 (3) BOL, STE. Thorncroft 406 (3) NH. Torre 198 (3) BM.

Van Dam s.n. sub TM 211, s.n. sub TM 22929 (3) PRE. Van der Schijff

2653 (3) K, PRE. Vaughan 2124 (1) BM.

Ward & Hennessy s.n. (1) UDW (in spirits). Ward & M.C. Ward 40 (1)

K, UDW. Ward, M.C. Ward & R. Kyle 92 (1) UDW. Watt 2340 (3) J.

Wright 574 (3) NU.

Young 1500 (3) J.

ACKNOWLEDGEMENTS

I thank the Keepers and Curators of the herbaria listed

for the loan of material or for permitting me to work in

their herbaria; the Council of the University of Durban-

Westville for granting me study leave; the CSIR Founda-

tion for Research Development for providing funds for

travelling and for running expenses; Mr B.D. Schrire,

former South African Liaison Officer, Royal Botanic

Gardens, Kew, for invaluable assistance in obtaining ar-

chaic literature and for his hospitality; Mrs Jean Brenan,

Kew, and my son Mr T.D.J. Hennessy for their hospitality;

the staff of the National Botanical Institute for their as-

sistance; my colleagues at the Universities of Natal

(Pietermaritzburg) and Durban-Westville for collecting

fresh material, for advice and for companionship,

particularly Mr C.J. Ward who discovered both known

populations of Mucuna gigantea in southern Africa, led

me to the more southerly of the two, climbed into the

canopy to photograph inflorescences in situ and collected

material for me; and Mrs R. Bunsee for assistance in

typing the manuscript. I also thank my spouse. Lieutenant

Commander B.J. Hennessy for his support and com-

panionship.

REFERENCES

ADANSON, M. 1763. Families des Plantes 2 : 324 — 327. Vincent, Paris.

ANDREWS, H.C. 1806. The botanist's repository 7: t. 443. London.

AUBREVILLE, A. 1936. Flore Forestiere Cote d'Ivoire 296: t. 118A.

BAKER. E G. 1929. The Leguminosae of tropical Africa 2: 366—381.

Unitas, Ostend.

BAKER, E.G. 1938. Notes from the British Museum Herbarium. The

Journal of Botany 76: 238.

BAKER, J.G. 1871. Erythrina, Mucuna. In D. Oliver. Flora of tropical

Africa 2,1: 181—188. Reeve, London.

BAKER, J.G. 1876. Mucuna, Erythrina. In J.D. Hooker, Flora of British

India 2,1: 185-190. Reeve, London.

BAKER, H.G. & BAKER, 1. 1983. Floral nectar constituents in rela-

tion to pollinator type. In C.E. Jones & R.J. Little, Handbook

of experimental pollination biology: 117—141. Scientific and

Academic Editions, Van Nostrand & Reinhold, New York.

BATTEN, A. 1986. Flowers of southern Africa: 162. Frandsen, Sand-

ton, Johannesburg.

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

Cape Province. Books of Africa, Cape Town.

BENTHAM, G. 1837. Commentationes de leguminosarum generibus.

Annalen des Naturhistorischen Museum in Wien 2: 113. Sollinger,

Wien.

BENTHAM, G. 1859. Erythrinineae. In C.F.P. von Martius, Flora

brasiliensis 15: 171—175.

BENTHAM, G. 1865. Erythrina, Mucuna. In G. Bentham & J.D. Hooker,

Genera plantarum 1: 531—533. Reeve, London.

BERTOLONI, G. 1850. Erythrina hastifolia. Memorie della R. Accademia

della scienze dell’Istituto di Bologna 2: 568, t. 38.

BORT 1909. Stizolobium deeringianum. United States Department of

Agriculture, Bureau of Plant Industry Bulletin No. 141: 31, t. 2, 3.

BRITTON, N.L. & ROSE, J.N. 1922. Erythrina pallida. Bulletin of the

Torrey Botanical Club 48: 332.

BROTERO, F. DE A. 1824, Descriptions of two new species of Erythri-

na. Transactions of the Linnean Society 14: 342.

BROWNE, P. 1756. The civil and natural history of Jamaica: 290—296.

BURCK, W. 1893. Mucuna pruriens var. utilis. Annales du Jardin

botanique de Buitenzorg 11: 187.

BURTT DAVY, J. 1932. A manual of the flowering plants and ferns of

the Transvaal with Swaziland 2: 414. Longmans, Green, London.

COATES PALGRAVE, K. 1977. Trees of southern Africa: 328—332.

Struik, Cape Town.

CODD, L.E.W. 1951. Trees and shrubs of the Kruger National Park.

Botanical Survey Memoir No. 26: 72.

CODD, L.E.W. 1955. Notes on certain South African Erythrina species.

Bothalia 6 : 507— 511.

CODD, L.E.W. 1963. Erythrina baumii. The Flowering Plants of Africa

36: t. 1412.

CODD, L.E.W. 1974. The identity of Erythrina princeps. Bothalia 11:

269-271.

CODD, L.E.W. 1977. Erythrina abyssinica. The Flowering Plants of Africa

44: t. 1738.

COOK, O.F. 1901. Erythrina poeppigiana. Bulletin of the United States

Department of Agriculture, Botany 25: 57, t. 13.

COLLETT, D.G. 1941. The South African species of Erythrina. Bothalia

4: 219-228.

COMPTON, R.H. 1976. The Flora of Swaziland. Journal of South Afri-

can Botany Supplementary Vol. No. 11: 277, 278.

DARLINGTON, C.D. & WILEY, A.P. 1955. Chromosome atlas of

flowering plants: 172. George Allen & Unwin, London.

DE CANDOLLE, A.P. 1825. Prodromus systematis naturalis regni

vegetabilis 2: 404-416. Treuttel & Wurtz, Paris.

DE WILDEMAN, E. 1920. Erythrina bequaertii. Revue de zoologique

africaine 8, Supplement botanique B15— B17.

DIETRICH, A.G. 1834. Erythrina princeps . Allgemeine Gartenzeitung

2: 305.

DON, G. 1832. A general system of gardening and botany 2: 371.

Rivington, London.

DYER, R.A., 1931. Mucuna gigantea (Willd.) DC. Kew Bulletin 1931:

464.

DYER, R.A. 1947. Erythrina zeyheri. The Flowering Plants of Africa

26: t. 1011.

DYER, R.A. 1975. The genera of southern African flowering plants. Vol.

1: 269, 270. Department of Agricultural Technical Services,

Pretoria.

ECKLON, C.F. & ZEYHER, K.L.P. 1836. Enumeratio plantarum africae

australis extratropicae: 243. Hamburg.

ENDLICHER, S.L. 1833. Prodromus florae norfolkiae: 91. Wien.

GILLETT, J.B. 1962. The fruit and seeds of Erythrina brucei and the

identity of E. abyssinica. Kew Bulletin 15: 425—429.

GUILLEMIN, J.A., PERROTTET, G.S. & RICHARD, A. 1832. Horae

senegambiae tentamen 6: 224. Paris.

HARMS, H.A.G. 1913. Leguminosae africanae VI. Erythrina decora.

Botanische Jahrbiicher fur Systematik, Pflanzengeschichte und

Pflanzengeographie 49: 441, 442.

HARMS, H.A.G. 1915. Erythrina. In H.G.A. Engler & C.G.O. Drude,

Die Vegetation der Erde 9: Die Pflanzenwelt Afrikas: 656—659.

Engelmann, Leipzig.

HARVEY, W.H. 1859. Thesaurus capensis 1: 39, t. 61, 62.

HARVEY, W.H. 1862. Erythrina. In W.H. Harvey & O.W. Sonder, Hora

capensis 2,1: 236—238. Reeve, London.

HASSKARL, J.C. 1842. Mucuna velutina. Flora 25: II.

HASSKARL, J.C. 1855. Stenotropis. Retzia 1: 183, 184.

HASSKARL, J.C. 1858. Hortus bogoriensis descriptus: 197.

HAUMAN, L. 1954. Mucuna. Flore du Congo beige et du Ruanda-

Urundi 6: 126—138.

HAUMAN, L. 1955. Mucuna. Bulletin du Jardin botanique de I’Etat

d Bruxelles 25: 98, 99.

HEMSLEY, A.J. & FERGUSON, I.K. 1985. Pollen morphology of the

genus Erythrina (Leguminosae: Papilionoideae) in relation to

floral structure and pollinators. Annals of the Missouri Botanical

Garden 72 : 570 -590.

HENKEL, J.S. 1934. A field book of the woody plants of Natal and

Zululand: 199. 200. Robinson, Durban.

HENNESSY, E.F. 1972. South African erythrinas. Natal Branch, Wildlife

Protection and Conservation Society of South Africa, Durban.

HENNESSY, E.F. 1975. Erythrina in southern Africa. Veld & Hora 61,3:

20-23.

HENNESSY, E.F. 1976. Erythrina cajfra. The Flowering Plants of Africa

43: t. 1709.

24

Bothalia 21,1 (1991)

HENNESSY, E.F. 1976. Erythrina kitissima. The Flowering Plants of

Africa 43: t. 1710.

HENNESSY, E.F. 1977. Erythrina livingstoniana. The Flowering Plants

of Africa 44: t. 1737.

HENNESSY, E.F. 1977. Erythrina fusca. The Flowering Plants of Africa

44: t. 1754.

HENNESSY, E.F. 1979. Erythrina senegalensis. The Flowering Plants

of Africa 45: t. 1786.

HENNESSY. E.F. 1985. Erythrina x johnsoniae. The Flowering Plants

of Africa 48: t. 1911.

HENNESSY, E.F. 1986. A fourth natural Erythrina hybrid from South

Africa. Bothalia 16: 48—51.

HENNESSY, E.F. 1989. Mucuna gigantea. The Flowering Plants of Africa

50: t. 1998.

HEPPER, F.N. 1954. Papilionaceae. In J. Hutchinson & J.M. Dalziel,

Flora of West tropical Africa, edn 2, revised by R.W.J. Keay 1:

561-563. Crown Agents for Overseas Governments, London.

HOCHSTETTER, C.F. 1846. Chirocalyx abyssinicus. Flora 29: 600.

HOLMGREN, P.K. & KEUKEN, W. 1974. Index he rbariorum. Bohn,

Scheltema & Holkema, Utrecht.

HUA, H. 1898. Erythrina comosa. Bulletin mensuel de la Societe

Linneenne de Paris n.s. 1: 52.

HUTCHINSON, J. 1927. Mucuna, Erythrina. In J. Hutchinson & J.M.

Dalziel, Flora of West tropical Africa, edn 1: 405, 406. Reeve,

London.

HUTCHINSON, J. 1933. Erythrina lysistemon. Kew Bulletin 1933: 422.

HUTCHINSON, J. 1964. The genera of flowering plants 1: 431—434.

Oxford University Press, Oxford.

HUTCHINSON, J. & DALZIEL, J.M. 1927. Flora of West tropical Africa,

edn 1 Crown Agents, London.

JACOT GUILLARMOD, A., JUBB, R. A. & SK.EAD. C.J. 1979. Field

studies of six southern African species of Erythrina. Annals of

the Missouri Botanical Garden 66 : 521—527.

KER-GAWLER, J.B. 1823. Erythrina caffra. The Botanical Register 9:

t. 736.

KRUKOFF, B.A. 1939a. The American species of Erythrina. Brittonia

3 : 205 - 337.

KRUKOFF, B.A. 1939b. Preliminary notes on Asiatic-Polynesian species

of Erythrina. Journal of the Arnold Arboretum 22: 225—233.

KRUKOFF, B.A. 1941. Supplementary notes on the American species

of Erythrina. American Journal of Botany 28: 683-691.

KRUKOFF, B.A. & BARNEBY, R.C. 1974. Conspectus of species of

the genus Erythrina. Lloydia 37: 332 —459.

KUNTZE, C.E.O. 1891. Revisio generum plantarum vascularium I: 172,

173. Felix, Leipzig.

LACKEY, J.A. 1981. Erythrinineae. In R.M. Polhill & PH. Raven,

Advances in legume systematic s 1: 312. Royal Botanic Gardens,

Kew, Richmond.

LAMARCK, J.B. A. P.M. DE 1786. Encyclopedic methodique 2: 392.

Paris.

LETTY, C.L. 1962. Wild flowers of the Transvaal : 169, t. 84.3. Division

of Botany, Department of Agriculture, Pretoria.

LINNAEUS, C. 1753. Species plantarum (Facsimile edn 1959) 2 : 706.

London.

LINNAEUS, C 1754a. Genera plantarum, edn 5: 316. Stockholm.

LINNAEUS, C. 1754b. Dolichos pruriens. In Olavus Stickman,

Dissertatio herbarium amboinense : 23. Uppsala.

LINNAEUS, C. 1759. Systema naturae, edn 10,2: 1162, 1163. Stockholm.

LINNAEUS, C. 1767. Mantissa plantarum: 99. Salvi, Stockholm.

LOUIS, J. 1935. Revision des especes congolaises du genre Erythrina

L. Bulletin du Jardin botanique de TEtat a Bruxelles 13: 295—319.

LOURIERO, J. DE 1790. Flora cochinchinensis : 471, 456, 457, 460, 461.

MAJOT ROCHEZ, R. & DUVIGNEAUD, P. 1954. Erythrina. Flora

du Congo beige et du Ruanda-Urundi 6: 113—125.

MARLOTH, R. 1925. The Flora of South Africa 2, 1: 81, 82, t.29. Darter,

Cape Town.

MEDIKUS, F.K. 1787. Cacuvallum. Vorlesungen der Churpfalzischen

physicalisch-bconomischen Gesellschaft 2: 392. Mannheim.

MEISNER, C.F. 1843. Contributions towards a flora of South Africa:

Erythrina, Chirocalyx. The London Journal of Botany 2: 96—99.

MERRILL, E.D. 1910. Mucuna deeringiana. Philippine Journal of

Science 5: 118.

MERRILL, E.D. 1935. Erythrina fusca. Transactions of the American

Philosophical Society 24: 209.

MEYER, E.H.F. 1836. Commentariorum de plantis africae australioris

1: 149—151. Leipzig.

MICHELI, M. 1896. Erythrina constantiana. Revue horticole 68: 524,

525.

MILLER, P. 1754. The gardener ’.v dictionary, edn 4 : 372 — 374. London.

NECKER, N.J. DE 1790. Elementa botanica 3: 43. Neuwied.

ONDERSTALL, J. 1984. South African wild flower guide 4: Transvaal

Lowveld and Escarpment. Botanical Society of South Africa, Cape

Town.

OSBECK, P. 1754. Dagbok dfwer en Ostindisk resa: 93. Stockholm.

PALMER, E.M. & PITMAN, N. 1972. Trees of southern Africa 2:

953—962. Balkema, Cape Town.

PHILLIPS, E.P. 1922. Erythrina caffra. The Flowering Plants of South

Africa 2: t. 59.

PHILLIPS, E.P. 1923. Erythrina humeana. The Flowering Plants of South

Africa 3: t. 112.

PHILLIPS, E.P. 1926a. Erythrina acanthocarpa. The Flowering Plants

of South Africa 6: t. 203.

PHILLIPS, E.P. , 1926b. The genera of South African flowering plants

1. Botanical Survey Memoirs No. 10: 336, 337.

PHILLIPS, E.P. 1951. The genera of South African flowering plants 2.

Botanical Survey Memoirs No. 25: 422, 423.

POEPPIG, E.F. 1899. Erythrina micropteryx. In I. Urban, Symbolae

antillanae 1: 327.

PRAIN, D. 1897. Some Additional Leguminosae: Mucuna. Journal of

the Asiatic Society, Bengal 66: 404-408.

PRESL, K.B. 1832. Symbolae botanicae 1: 69, t. 46. Prague.

PRESL, K B. 1834. Symbolae botanicae 2: 22, t. 68.

RAFINESQUE-SCHMALTZ, C.S. 1812. Xy phanthus. Florula ludo-

viciana: 103. Wiley, New York.

RADDI, G. 1820. Macroceratides. Memorie di matematica e di fisica

della Societa italiana delle scienze, Modena : 13, 14, fig. 5.

REICHENBACH, H.G.L. 1836. Erythrina caffra. Flora exotica 5: t.312.

Leipzig.

RHEEDE, H.A. VAN 1688. Hortus indicus malabaricus 8: 63, t.

36.

RICHARD, A. 1847. Tentamen florae abyssinicae 1: 213, 214. Paris.

ROXBURGH, W. 1814. Hortus bengalensis 54. Serampore.

ROXBURGH, W. 1825. Flora indica 3 : 2 83-287. Serampore.

ROXBURGH, W. 1832. Flora indica, edn 2: 254. Serampore.

RUIZ, L.H. & PAVON, J.A. 1794. Florae peruvianae et chilensis prodro-

mus 98: t. 21. Madrid.

RUMPHIUS, G.E. 1750. Herbarium amboinense 6: 393, 394, t. 142.

Utrecht.

SALT, H. 1814. List of new and rare plants collected in Abyssinia during

the years 1805 and 1810. A voyage to Abyssinia, Appendix 4: 63,

64. Rivington, London.

SCHREIBER, A. 1970. Fabaceae. In H. Merxmiiller, Prodromus einer

Flora von Sudwestafrika 60: 38. Cramer, Lehre.

SCHUMACHER, C.F. 1827. Beskrivelse af Guineeiske planter: 333.

Copenhagen.

SCHUMACHER, C.F. & THONNING, T. 1853. Chirocalyx latifolius.

In C.F. Schumacher, Beskrivelse af Guineeiske planter : 333,

Copenhagen.

SIM, T. R. 1907. The forests and forest flora of the Colony of the Cape

of Good Hope: 201, 202. Taylor & Henderson, Aberdeen.

SIM, T.R. 1909. Forest flora and forest resources of Portuguese East

Africa: 43, 44. Taylor & Henderson, Aberdeen.

SIMS, J. 1823. Erythrina caffra. Curtis’s Botanical Magazine 51: t.

2431.

SPRAGUE, T A. 1909. Erythrina ovalifolia. Kew Bulletin 1909: 198, 199.

SPRENGEL, A. 1827. Caroli Linnaei systema vegetabilium curae

posteriores 5: 281. Gottingen.

SPRENGEL, K.P.J. 1826. Caroli Linnaei systema vegetabilium 3: 243.

Gottingen.

STAFLEU, F.A.(ed.) 1976. Taxonomic literature. Regnum vegetabile 52.

International Bureau for Plant Taxonomy and Nomenclature.

Utrecht.

SWEDIAUR, 1801. Labradia. Arzneimittel-Forschung 1: 164.

TAUBERT, P.H.W. 1894. Erythrinineae. Die Naturlichen Fflanzenfamilien

3: 363-366.

TAUBERT, P.H.W. 1897. Beitrage zur Flora von Afrika XII. Mucuna.

Botanische Jahrbiicher 23: 194, 195.

THUNBERG, C.P. 1800. Prodromus plantarum capensium 2: 121.

Edman, Uppsala.

THUNBERG, C.P. 1823. Flora capensis, edn 2: 559. Cotae, Stuttgart.

TODARO, A. 1861. Nuovi generi e nuovi specie di plante coltivate net

rel orto botanica di Palermo: 66 —70.

TORRE, A.R. DA 1965. Taxa angolensi a nova vel minus cognita — IV.

Erythrina mendesii. Boletim da Sociedade Broteriana ser. 2, 39:

212, 213, t. 9.

TORRE, A.R. DA 1966. Erythrininae. Conspectus Florae Angolensis

3: 247-253, t. 23.

TRIMEN, H. 1894. A handbook to the flora of Ceylon 2: 60—64. Dulau,

London.

Bothalia 21,1 (1991)

25

TROUPIN, G.M.D.J. 1982. Flore des plantes ligneuses du Rwanda. 306,

319, fig. 105. Musee Royal de l’Afrique Central-Tervuren,

Belgique.

VAN RENSBURG, T.F.J. 1982. Coral tree — tree of the year 1982.

Directorate of Forestry Pamphlet No. 298. Pretoria.

VAN WYK, P. 1972. Trees of the Kruger National Park 1: 223 - 227.

Purnell, Cape Town.

VERDCOURT, B. 1970. Studies in the Leguminosae — Papilionoideae

for the Flora of tropical East Africa II. Kew Bulletin 24: 235—307.

VERDCOURT, B. 1971. Erythrina, Mucuna. Flora of tropical East Africa,

Leguminosae (4) — Papilionoideae (2): 541—571. Crown Agents

for Overseas Governments. London.

VERDCOURT, B. 1981. New taxa of Mucuna (Leguminosae—

Phaseoleae) from East Africa and Australia. Kew Bulletin 35:

743 -752.

WALPERS, W.G. 1850. Micropteryx, Duchassaingia. Linnaea 23:

739 -742.

WALPERS, W.G. 1853. Noch ein paar Worte iiber Erythrina L. Flora

36: 148, 149.

WATT, J.M. & BREYER-BRANDWIJK, M. G. 1962. The medicinal

and poisonous plants of southern and eastern Africa: 631—634.

Livingstone, Edinburgh & London.

WHITE, F. 1962. Forest flora of Northern Rhodesia: 152, 160, 433.

Oxford University Press.

WIGHT, R. 1840. leones plantarum Indiae orientalis: 1: 280. Madras.

WIGHT, R. & ARNOTT, G.A.W. 1834. Prodromus florae peninsulae

Indiae orientalis 1: 253—255. Parbury & Allen, London.

WILLDENOW, C.L. 1801. Erythrina glauca. Magazin fur die neusten

Entdeckungen in der gesammten Naturkunde der Gesellschaft

Naturforschender Freunde zu Berlin 3: 428.

WILLDENOW, C.L. 1802. Dolichos giganteus. Species plantarum 3:

1041. Berlin.

WILMOT-DEAR, M. 1984. A revision of Mucuna( (Leguminosae —

Papilionoideae) in China and Japan. Kew Bulletin 39: 23—65.

WOOD, J.M., 1906. Erythrina tomentosa. In J.M. Wood & M. Evans,

Natal plants 4: t. 384, 385. Bennett & Davis, Durban.

WOOD, J.M. 1912. Erythrina cajfra. In J.M. Wood & M. Evans, Natal

plants 6: t. 542. Bennett & Davis, Durban.

.

.

Bothalia 21,1: 27-34 (1991)

A new species of Merxmuellera (Arundineae, Poaceae) from South

Africa

N.P. BARKER* and R.P. ELLIS**

ABSTRACT

Merxmuellera setacea N.P Barker from the south-western Cape, South Africa, is formally described. Descriptions of

the morphology and leaf anatomy, in transectional and surface view, as observed under the light- and scanning electron

microscope are given. Comparisons between this taxon and other southern African arundinoid species are made on the

basis of morphological and anatomical observations. While the morphology of M. setacea is in keeping with the generic

limits of Merxmuellera Conert, the leaf blade anatomy appears to be intermediate between Merxmuellera, Pentameris Beauv.

and Pentaschistis (Nees) Spach.

UITTREKSEL

Merxmuellera setacea N.P. Barker uit die Suidwes-Kaap, Suid-Afrika, word formeel beskryf. Beskrywings van die morfologie

en blaaranatomie, in deursnee- en oppervlakaansig, soos met die ligmikroskoop en aftaselektronmikroskoop waargeneem,

word gegee. Hierdie takson word op grond van morfologiese en anatomiese waarnemings met ander arundinoiede spesies

van suidelike Afrika vergelyk. Die morfologie van M. setacea strook met die genusafbakening van Merxmuellera Conert,

maar die anatomic van die blaarskyf is skynbaar intermedier tussen Merxmuellera, Pentameris Beauv. en Pentaschistis (Nees)

Spach.

Merxmuellera setacea N.P Barker, sp. nov.

Merxmuellera sp. (Ellis 5500) in Gibbs Russell et al.\ 218

(1990).

Vaginae foliorum glabrae, basin versus ventricosae;

laminae folium usque ad 150 mm longae, setaceae,

pungentes; lemma praeter caespem unum submarginalem

pilorum alborum in utraque lato, in circa tertia parte

longitudinis corporis lemmae situm, glabra.

A caespitose, perennial grass. Rhizomes producing

closely spaced buds which develop into bulb-like structures

as they become covered in thickened, swollen leaf sheath

bases. Flowering culms erect, 0,73—1,0 m tall, usually with

three nodes. Leaves : sheaths glabrous to somewhat glossy

when young, bearded at the mouth, basally swollen,

thickened and somewhat pithy; ligule a fringe of hairs

up to 0,75 mm long; leaf blades permanently infolded,

setaceous, glabrous below, up to 150 mm long with

pungent apices. Inflorescence contracted, shortly

branched, lanceolate panicle up to 150 mm long, with

50—100 (or more) spikelets. Spikelets 3(— 4)-flowered,

12—15 mm long (excluding awns), 4—6 mm wide, upper-

most floret sometimes rudimentary; glumes 8—13 mm

long, as long as or longer than the body of the proximal

floret, 1-nerved, straw-coloured to pale greenish yellow

to olive, with keel sometimes tinged dark purple, glabrous

and thinly papery; lemma 8,5—13,0 mm long, bilobed,

lobes attenuating into a 5 mm long lateral bristle, glabrous

except for one submarginal tuft of hairs on each side

situated one third of the way up the lemma body; central

awn geniculate, 11—15 mm long, yellow to brown, loosely

twisted basally, kneed about 3 mm above point of insertion;

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

address: Bolus Herbarium, University of Cape Town, Private Bag,

Rondebosch 7700.

** Grassland Research Centre, Private Bag X05, Lynn East, Pretoria 0039.

MS. received: 1990-06-22.

palea 4,5— 5,5 mm long, narrowly obovate, pubescent in

the mid region between the keel and the margin, glabrous

between the keels; lodicules 2, up to 1,8 mm long, with

a distinct waist, extending around and beyond the ovary,

apically ciliolate; anthers up to 2,75 mm long; ovary ellip-

soid, glabrous.

TYPE. — Cape, 3319 (Worcester): Groot Winterhoek

Wilderness area, NW of Groen Mt., Suurvlakte plateau,

(-AA), 17-XI-1987, Ellis 5500 (PRE, holo.!). Figure 1.

First collected in 1987, M. setacea is so far known from

only two localities (Figure 2).

Specimens examined

CAPE.— 3219 (Wuppertal): Skurweberge (—CD), Blinkberg Pass, Ellis

5520. 3319 (Worcester): Groot Winterhoek (— AA), Suurvlakte plateau,

Ellis 5499, 5500.

MORPHOLOGY OF MERXMUELLERA SETACEA

Merxmuellera setacea is quite distinct from all other

members of the genus in that it has basally thickened and

swollen sheath bases covering the rhizome and axillary

buds (Figures 1; 3A, B). The internal tissue of the

thickened sheath is pithy in nature and the abaxial surface

is glabrous to glossy. The short leaf blades (up to 150 mm

in length) are permanently infolded and setaceous, hence

the specific epithet. Three other species of Merxmuellera

have swollen basal parts: M. rufa (Nees) Conert, M.

lupulina (Thunb.) Conert and M. decora (Nees) Conert.

However, in these three species, the basal sheaths are

densely villous, and unthickened.

The unusual growth form of this species, a linear series

of bulb-like structures, appears to occur as a result

of the rhizomes producing numerous closely spaced

shoots. These shoots develop beneath the soil surface,

producing leaves and occasionally flowering culms. The

28

Bothalia 21,1 (1991)

FIGURE 1. — Holotype of Merxmuellera setacea (PRE): short, basal,

setacous leaves, swollen culm base and rhizome covered in

glabrous papery scales. This specimen bears a label with the words

‘Voucher specimen for TAXON PRECIS 3 1988’ on it, indicating

it was cited as a voucher specimen in Grasses of southern Africa

(Gibbs Russell et al. 1990).

basally thickened leaf sheaths overlap each other,

producing a bulb-like structure unique to this species. The

close proximity of. growth points along the rhizome, all

developing as described above, produce a line of ‘bulbs’

beneath the soil surface, and a crest- or hedge-like growth

of leaves above the soil surface.

M. setacea also has a distinct floral morphology. The

lemma has one tuft of long, white, submarginal hairs

on each side of the lemma body (Figure 4C). The only

other species in the genus with a somewhat similar

arrangement of hairs is M. dura (Stapf) Conert, which can

be distinguished from M. setacea not only by the mor-

phology of the basal leaf sheaths but also by differences

in other floral characters, such as glume venation (glumes

3— 5-nerved in M. dura , 1-nerved in M. setacea ), and the

lemma indumentum (M. dura has an additional tuft or

fringe of hairs around the central awn base, particularly

visible on the inner side of the lemma, which is not present

in M. setacea ).

From observations made in the field, it appears that M.

setacea only grows in very damp to waterlogged soil in

seepage areas and streambanks. This waterlogging appears

to be seasonal, as all sites where the plant was collected

had a very shallow soil layer overlying bedrock, and would

dry out rapidly between rainfall events. This species, there-

fore, probably functions most of the time as a xerophyte

in a lithophytic community. The sclerophyllous anatomy

of the leaf blade, which is permanently infolded and

setaceous, is in keeping with a xerophyte rather than a

hydrophyte, which would have soft, spongy, mesophytic

leaves with lacunae.

It appears that this species, like several other Cape

fynbos grass taxa, survives underground as a geophyte

during the protracted interfire periods. During these

periods, which may endure for four to 40 years with a

mean of 15 years (Van Wilgen 1987), the woodier elements

of the vegetation exclude the herbaceous taxa, including

the grasses.

This particular interfire survival strategy is typical of

a number of Cape fynbos grass taxa, which have all

developed swollen, geophytic basal parts (Linder & Ellis

1990b). The geophytic base may also protect the innova-

tion bud against fire and desiccation. In the period

subsequent to a burn, this geophytic base regenerates

vigorously from the rhizomes, producing the growth form

described above.

The generic placing of the new species posed problems:

the only other southern African arundinoid genera which

have species with tufts of hairs on the lemma, although

not in the pattern reported here for M. setacea , are species

of Dregeochloa Conert, Karroochloa Conert & Tiirpe,

Schismus Beauv. and Centropodia Reichenb.

A position for this new taxon in Karroochloa was

rejected as the spikelets of M. setacea are too large, far

exceeding the 4—7 mm described in Karroochloa (Conert

& Tiirpe 1969); the glumes of species in Karroochloa are

described as being 3— 5(— 7)-nerved, those of M. setacea

are 1-nerved. Species of the genus Schismus do not have

lemmas with a central, geniculate awn (Conert 1971); the

spikelet size and glume venation differ markedly, and

consequently this genus was also rejected for the new

taxon.

32°

34°

FIGURE 2.— The known distribution of Merxmuellera setacea.

Bothalia 21,1 (1991)

setaceous, pungent leaves arising from swollen culm base, covered

in glabrous, papery scales, X 0,7; B, outline of a transverse section

through thickened, swollen basal region of leaf sheath, x 1,35.

Species of Dregeochloa, particularly D. calviniensis

Conert, bear a superficial resemblance to M. setacea in

a number of characters: the basal parts are swollen in both

taxa (although the rhizome is covered by papery scales and

not by swollen sheath bases in D. calviniensis)', the

spikelets are of comparable size, and the leaf blades are

short and setacous in both taxa. However, the differences

in glume venation (3— 5-nerved in D. calviniensis ); ques-

tionable homology between the basal structures of these

two species and marked differences in the leaf anatomy

of Dregeochloa (Ellis 1977) and M. setacea, made the plac-

ing of M. setacea in Dregeochloa extremely doubtful.

Centropodia, too, was deemed to be a very poor position

for the new taxon, because the lemmas of the taxa of this

genus have a transverse row of tufts of hairs across the

lemma backs and wide, flat or rolled leaf blades. In

addition to this, anatomical evidence of any similarity

between M. setacea and Centropodia is negligible, as

Centropodia has C4 leaf blade anatomy, whereas all other

29

South African representatives of the Arundineae have C,

anatomy (Ellis 1984, 1989).

Because of the presence of tufts of hairs on the lemma

of the new species, neither Pentaschistis nor Pentameris

were considered, despite the fact that both these genera

have species with 1-nerved glumes, and all taxa in these

FIGURE 4. — Merxmuellera setacea. A, inflorescence, X 1; B, view

of exploded spikelet showing the 1-nerved glumes and three florets,

X 4; C, lemma showing the lateral bristles, geniculate central

awn and submarginal tufts of hairs, X 4; D, palea showing pubes-

cence between margins and keels, X 4; E, diagram of lodicules,

showing pronounced ‘waist’ and ciliolate apex. Lodicules cover

and extend beyond ovary, the outline of which is indicated by the

regular dotted line, x 40.

30

Bothalia 21,1 (1991)

two genera have 1— 2-flowered spikelets (Linder &

Ellis 1990a; Gibbs Russell et al. 1990). A position in

Merxmuellera was chosen as this genus has spikelets with

3—10 florets, variously nerved glumes and lemmas which

are irregularly pilose or with hairs arranged in tufts, and

a central awn which is usually geniculate, 8—20 mm in

length (Conert 1970, 1971).

LEAF ANATOMY OF MERXMUELLERA SETACEA

The leaf blade anatomy of all the South African species

of Merxmuellera has been described (e.g. Ellis 1981a, b).

In order to obtain further evidence relating to the taxo-

nomic position of the new taxon, a detailed anatomical

study of M. setacea was carried out for comparison with

that of other species of Merxmuellera.

Leaf blade in transverse section

Outline : permanently infolded, setaceous blade with a

reduced U- or elliptical shape. Adaxial channel deep and

narrow. Two halves of leaf lamina not symmetrical about

the median vascular bundle (Figure 5A, B) but differ in

length and shape of the margins. Ribs and furrows: well

developed adaxial ribs and furrows present; furrows deep,

cleft-like and located between all larger vascular bundles

but occurring irregularly over some of the smaller bundles

(Figure 5C— F); ribs rounded, often with flattened tops,

and tending to the massive type, present over all larger

vascular bundles. No abaxial ribs and furrows.

Median vascular bundle: not structurally distinct from

other first order bundles; located off centre in the blade

due to asymmetrical nature of laminae (Figure 5A, B).

Vascular bundle arrangement: 5 first order bundles alter-

nate with single third order bundles; pattern variable in

margin where one first order bundle is much reduced in

size and an additional third order bundle may be present

on one side of median bundle (Figure 5B); this results in

asymmetrical laminae. First order bundles centrally lo-

cated in the blade but the third order bundles are situated

much closer to the abaxial surface (Figure 5C— F).

FIGURE 5. — Transectional leaf blade anatomy of Merxmuellera setacea. A, permanently infolded blade with asymmetrical laminae; B, asym-

metrical outline without discernible midrib; C, abrupt, truncated margin of blade; D, other margin of same blade as in C, tapering and not

truncated; note prickles on adaxial surface. E-F, leaf blade viewed with different filters: E, green filter showing darkly stained, lignified

supporting tissue, note the continuous hypodermal sclerenchyma layer connected to the vascular bundles by girders, and the darkly

staining, lignified inner and outer bundle sheath cells associated with the first order vascular bundles; F, red filter showing compact mesophyll

tissue consisting of small, angular chlorenchyma cells. Ellis vouchers: A, E, F, 5520; B, C, D, 5499. A, B, X 100; C, D, E, F, X 250.

Bothalia 21,1 (1991)

31

FIGURE 6, — Abaxial epidermis of Merxmuellera setacea. A, epidermal pattern at low magnification showing absence of zonation; B, alternating

files of slightly fusiform long cells separated by cork-silica cell pairs, circular silica bodies closely associated with tall cork cell; C,

very uniform arrangement of costal long and short cells; D, consistent pattern of epidermal cell arrangement and form. Ellis vouchers:

A, B, 5520; C, 5499; D, 5500. A, x 160; B, C, D, x 250.

Vascular bundle description : third order bundles elliptical

with xylem and phloem tissue indistinguishable. First

order bundles elliptical with narrow metaxylem vessels

with unthickened secondary walls. Sclerosed phloem

divided by intrusive fibres present (Figure 5C— F).

Vascular bundle sheaths : first order bundle outer sheaths

elliptical with wide adaxial and abaxial interruptions; cells

elliptical, small (same size as the innner sheath cells) with

thin walls (Figure 5C, D) but often with distinctly

thickened walls (Figure 5E, F) which then structurally

resemble mestome sheath cells. Inner sheath complete,

consisting of relatively large cells with heavily but

uniformly thickened walls. Third order bundles without

adaxial interruptions and without thickened outer bundle

sheath cells (Figure 5E, F).

Sclerenchyma : adaxial girders associated with the first

order vascular bundles inversely anchor-shaped with long,

tall and multiseriate stems; abaxial girders well developed

and trapezoidal in shape; associated with first and third

order bundle; laterally fused, forming continuous abaxial

hypodermal fibrous layer. All fibres with very thick,

lignified secondary walls and no sclerenchyma tissue

staining blue-green with fast green (Figure 5F). Chloren-

chyma : irregular arrangement, W-shaped mesophyll tissue

occupying the sides and bases of furrows; tissue consisting

of small, regular isodiametric cells which are tightly

packed with very small intracellular air spaces (Figure 5F).

Colourless cells : absent. Adaxial epidermal cells : bulli-

form cells very small, located at bases of furrows. No

macro-hairs or papillae; prickle hairs with long barbs

common, these without bulbous bases. Abaxial epidermal

cells: no bulliform cells or appendages. Continuous, thick

cuticle present over uniform, but small epidermal cells.

Abaxial epidermis in surface view

Zonation: costal and intercostal zones not distinguishable

(Figure 6A). Long cells: elongated, rectangular with side

walls parallel or slightly bowed, slightly thickened, not

undulate (Figure 7A, B). Nuclei present (Figure 6B— D).

Cell shape and size consistent throughout epidermis

(Figure 6A). Short cells: cork-silica cell pairs alternate

with all long cells in all files. Silica cells closely associated

with tall cork cell (Figure 6D). Stomata: absent. Papillae:

absent. Prickles: absent. Microhairs: absent. Macrohairs:

absent. Silica bodies: round to slightly elliptical (Figure

6B— D; 7B); present throughout the epidermis; same width

as long cells.

Adaxial epidermis in surface view

Long cells: elongate, rectangular without sinuous walls

(Figure 7C). Short cells: appear to be absent. Stomata:

low dome-shaped, flush with the epidermal surface (Figure

7E, F); restricted to the bases and sides of the adaxial

furrows. Papillae: absent. Prickles: common with long

barbs and without swollen bases (Figure 7C, D); prickles

on top of adaxial ribs with blunt apices, whereas those

overlying the furrows interlocking with sharp, long barbs

(Figure 7C). Microhairs: elongate, at least 3 times the

length of the stomata; basal cell very long and narrow with

minute distal cell only */s the length of the hair (Figure

7E, F); only present in adaxial furrows. Macrohairs:

absent. Silica bodies: appear to be absent.

The leaf anatomy of M. setacea is unusual in several

respects and does not closely resemble that of any other

species of Merxmuellera. Certain anatomical similarities

32

Bothalia 21,1 (1991)

with some species of Pentaschistis and Pentameris are,

however, evident.

The asymmetrical leaf outline of M. setacea, together

with the off-centre location of the median vascular bundle,

is an unusual feature, but one that is not unique in the tribe.

Similar outlines rarely occur in M. aureocephala (J.G.

Anders.) Conert (Ellis 1981b), Pentaschistis pyrophila

Linder and P. eriostoma (Nees) Stapf (Ellis & Linder in

prep.). This feature in isolation does not appear to be

taxonomically important but it is always associated with

other anatomical attributes which appear to link M. setacea

to these other taxa and their close relatives.

One such feature is the alternating arrangement of first

and third order vascular bundles across the leaf width. This

pattern is not found in M. stricta (Schrad.) Conert and

its allies (Ellis 1981b) where lateral first order bundles are

not separated by smaller bundles. M. dura also has this

M. stricta type of vascular bundle arrangement. Thus the

similarities between M. setacea and M. dura as suggested

by the lemma vesture are not supported by the vascular

bundle arrangement observed in these two species. M.

disticha (Nees) Conert, M. davyi (C.E. Hubb.) Conert,

M. macowanii (Stapf) Conert and sometimes M. aureo-

cephala, however, share the alternating vascular bundle

arrangement with M. setacea.

FIGURE 7. — Epidermal ultrastructure of Merxmuellera setacea. A, B, abaxial epidermis: A, uniform arrangement of rectangular long cells

alternating with paired silica bodies and cork cells. This pattern is consistent along all epidermal cell fdes, note the absence of epidermal

appendages; B, detail of rectangular to slightly fusiform long cells with inflated anticlinal walls and thickened periclinal walls, silica

bodies oval to round in shape. C, D, adaxial epidermis: C, elongated prickle hairs common, interlocking over adaxial groove and

resembling asperites on adaxial ribs; D, detail of prickles showing barbed prickles on sides of adaxial rib, whereas those on the top of the

rib tend to be domed. E, F, adaxial microhairs located on the sides and bases of the cleft-like furrows: E, elongated microhair with basal

cell considerably longer than the small cap cell; F, apical cell only 9s the length of the elongated basal cell, note stomata located at the

base of the furrow beneath the interlocking prickles. Ellis vouchers: A, B, E, 5499\ C, D, F, 5500. A, C, X 200; B, D, E, F, X 650.

Bothalia 21,1 (1991)

33

A similar alternating arrangement of vascular bundles

occurs in most species of Pentameris, particularly P.

macrocalycina (Steud.) Schweick. and Pentameris sp. 1

sensu Barker in Gibbs Russell etal. (1990) [=R obtusifolia

(Hochst.) Schweick. sensu Ellis 1985)] as well as many

species of Pentaschistis (Ellis & Linder in prep.). Relevant

examples are P pyrophila, P. eriostoma and P. curvifolia

(Schrad.) Stapf and its allies.

The sclerosed phloem in M. setacea occurs in M.

macowanii, M. davyi and M. aureocephala (Ellis 1981b)

but not in M. disticha. This feature also occurs in M.

stricta (very rarely), M. drakensbergensis (Schweick.)

Conert and M. stereophylla (J.G. Anders.) Conert (Ellis

1981a), but these latter species do not have the same

vascular bundle arrangement as M. setacea. Intrusive

fibres also do not occur in either Pentameris or

Pentaschistis. This feature is, therefore, not congruent with

the evidence from other anatomical characters, but does

support the placement of M. setacea in Merxmuellera

rather than either Pentameris or Pentaschistis.

The secondary thickening of the walls of the outer

bundle sheath cells of M. setacea appears to be a unique

feature in South African arundinoid species, but was only

observed in the first order vascular bundles of the type

specimen, Ellis 5500 (Figure 5E, F). Other specimens

from the same locality ( Ellis 5499), as well as from

elsewhere, lacked this feature. Secondary wall thickening

of outer bundle sheath cells is therefore, not a consistent

feature of this species and merits little taxonomic

significance.

The lignified fibres of the sclerenchyma of M. setacea,

together with the continuous hypodermal sclerenchyma

layer, are another feature this species shares with those

species also possessing the same type of bundle arrange-

ment. However, similar sclerenchyma also occurs in M.

drakensbergensis and M. stereophylla (Ellis 1981a) which

have the M. stricta type of bundle arrangement.

The unusual epidermal features of adaxial prickles and

the extremely elongate microhairs of M. setacea appear

to be very significant in the classification of this species.

These features are closely correlated with many of the

features of the transectional anatomy discussed above. M.

stricta and its allies have a papillate adaxial epidermis and

the microhairs are short with basal and distal cell equal

in length. This type of hair may also occur on the abaxial

surface. Even where adaxial prickles are present (as in

M. drakensbergensis and M. stereophylla) these differ

considerably from those of M. setacea in that they are

closely surrounded by raised inflated papillae. M. setacea

therefore differs considerably from M. stricta, M. guil-

larmodiae Conert, M. drakensbergensis, M. stereophylla,

and even M. dura which resembles this group of species

very closely. This type of papillate epidermis and short

microhair in turn also occurs in Pentaschistis colorata

(Steud.) Stapf and closely allied Pentaschistis species (Ellis

& Linder in prep.).

Arundinoid species which also have elongated adaxial

microhairs with minute distal cells and adaxial prickles

are: Pentameris macrocalycina, Pentameris sp. 1 (sensu

Barker in Gibbs Russell et al. 1990), P thuarii Beauv. and

P. longiglumis (Nees) Stapf as well as Pentaschistis

pyrophila Linder, P. eriostoma (Nees) Stapf and P curvi-

folia (Schrad.) Stapf. and its close allies (P. pungens

Linder, P. caulescens Linder, P. scandens Linder and P.

acinosa Stapf).

CONCLUSIONS

The inclusion of M. setacea in the Arundinoideae is

fully supported by the leaf anatomical evidence although

this subfamily lacks reliable diagnostic features (Ellis

1986). The presence of microhairs precludes pooid

relationships (Watson et al. 1985). This is supported by

the dome-shaped stomata with guard cells flush with the

epidermis and the absence of oblong or nodular silica

bodies (Ellis 1986). The non-Kranz anatomy with compact

chlorenchyma, simple midrib, absence of papillae and the

elongated, finger-like microhairs, in combination, also

suggest arundinoid affinities. Within this subfamily this

suite of anatomical characters also supports the classifi-

cation of M. setacea in the tribe Arundineae and excludes

its inclusion in any of the other tribes.

The lack of congruence between anatomical characters,

such as the asymmetrical leaf outline, the arrangement of

first and third order vascular bundles and various adaxial

epidermal features, and the generic classification of these

taxa appears to be a reflection of the unsatisfactory

delimitation of the arundinoid genera Merxmuellera,

Pentaschistis and Pentameris.

Anatomical evidence indicates affinities for M. setacea

with certain species of Merxmuellera, together with other

species presently included in both Pentameris and

Pentaschistis, and the new taxon appears to be truly

intermediate in its leaf anatomy, making it particularly

valuable in gaining a better understanding of the complex

character distribution patterns exhibited by these taxa.

Individual anatomical characters show clear disjunctions

but, in combination, no reliable diagnostic differences

appear to separate the various taxa displaying resemblances

to M. setacea.

Before the taxonomic significance of these indications

can be fully appreciated, it will be necessary to under-

take an extensive evaluation of as many characters as

possible in order to arrive at a more natural delimitation

of genera in this poorly understood tribe. The correct

taxonomic placement of this new species is therefore

extremely difficult. Thus, although the true affinities of

this species are obscure, its identification and description

will undoubtedly contribute to a better understanding of

the complex interrelationships between these arundinoid

genera. It therefore seems reasonable to follow the spikelet

indications and include this new species in Merxmuellera

until such time as generic delimitations in this tribe become

clearer.

ACKNOWLEDGEMENTS

We would like to thank Ms G. Condy for the pen and

ink art work, Mrs S.M. Perold for assistance with the SEM

work, Mrs A. Romanowski for the preparation of the

photographic figures and Dr H.F. Glen for the Latin

translation.

34

Bothalia 21,1 (1991)

REFERENCES

CONERT, H.J. 1970. Merxmuellera , eine neue Gattung der Gramineen

(Poaceae, Arundinoideae). Senckenbergiana Biologica 51:

129-133.

CONERT, H.J. 1971. The genus Danthonia in Africa. Mitteilungen der

Botanischen Staatssamrnlung, Munchen 10: 299—308.

CONERT. H.J. & TURPE, A M. 1969. Karroochloa, eine neue Gattung

der Gramineen (Poaceae, Arundinoideae, Danthonieae). Sencken-

bergiana Biologica 50: 289—318.

ELLIS, R.P. 1977. Leaf anatomy of the South African Danthonieae

(Poaceae). I. The genus Dregeochloa. Bothalia 12: 209—213.

ELLIS, R.P. 1981a. Leaf anatomy of the South African Danthonieae

(Poaceae). IV. Merxmuellera drakensbergensis and M. stereophyl-

la. Bothalia 13: 487-491.

ELLIS, R.P. 1981b. Leaf anatomy of the South African Danthonieae

(Poaceae). V. Merxmuellera macowanii, M. davyi and M. aureo-

cephala. Bothalia 13: 493—500.

ELLIS, R.P. 1984. Leaf anatomy of the South African Danthonieae

(Poaceae). IX. Asthenatherum glaucum. Bothalia 15: 153—159.

ELLIS, R.P. 1985. Leaf anatomy in the South African Danthonieae

(Poaceae). XIII. Pentameris macrocalycina and P. obtusifolia.

Bothalia 15 : 579 - 585.

ELLIS, R.P. 1986. A review of comparative leaf blade anatomy in the

systematics of the Poaceae: the past twenty-five years. In T.R.

Soderstrom el al. , Grass systematics and evolution. Smithsonian

Institution Press, Washington D.C.

ELLIS, R.P. 1989. Leaf anatomy of the South African Danthonieae

(Poaceae). XVIII. Centropodia mossamedensis. Bothalia : 19:

41-44.

ELLIS, R.P. & LINDER, H.P. in prep. An atlas of leaf anatomy in the

genus Pentaschistis (Poaceae: Arundinoideae: Arundineae).

Memoirs of the Botanical Survey of South Africa.

GIBBS RUSSELL, G.E., WATSON, L., KOEKEMOER, M., SMOOK,

L., BARKER, N.P., ANDERSON, H.M. & DALLWITZ, M.J.

1990. Grasses of southern Africa. Memoirs of the Botanical Survey

of South Africa No. 58: 1—437.

LINDER, H.P. & ELLIS, R.P. 1990a. A revision of Pentaschistis

(Arundineae, Poaceae). Contributions from the Bolus Herbarium

No. 12.

LINDER, H.P. & ELLIS, R.P. 1990b. Vegetative morphology and

interfire survival strategies in the Cape Fynbos grasses. Bothalia

20: 91-103.

VAN WILGEN, B.W. 1987. Fire regimes in the fynbos biome. In R.M.

Cowling et al. , Disturbance and the dynamics of fynbos biome

communities. South African National Scientific Programmes

Report No. 135. CSIR, Pretoria.

WATSON, L., CLIFFORD, H.T. & DALLWITZ, M.J. 1985. The

classification of Poaceae: subfamilies and supertribes. Australian

Journal of Botany 33: 433—484.

Bothalia 21.1: 35-49 (1991)

Synopsis of the genera Nesaea and Ammannia (Lythraceae) in southern

Africa

K.L. IMMELMAN*

Keywords: Ammannia , Lythraceae, Nesaea, southern Africa, taxonomy

ABSTRACT

The genera Nesaea Commers. ex H.B.K. and Ammannia L. in southern Africa, which comprises Namibia, Botswana,

South Africa, Swaziland and Lesotho, were revised. In Nesaea, Tl species (seven new), four varieties and one new form

were recognized. Ammannia was revised using the DECORANA computer program, and four species were recognized.

Keys to the taxa are presented, as well as synonymy (restricted to the region), diagnoses and descriptions of new taxa, and

notes on distribution, taxonomy and nomenclature. The following taxa are new: Nesaea alata Immelman, N. angustifolia

Immelman, N. cymosa Immelman, N. minima Immelman, N. sagittifolia (Sond.) Koehne var. ericiformis Koehne forma

swaziensis Immelman, N. saluta Immelman, N. wardii Immelman and N. zambatidis Immelman.

UITTREKSEL

Die genusse Nesaea Commers. ex H.B.K. en Ammannia L. in suidelike Afrika, wat uit Namibie, Botswana, Suid-Afrika,

Swaziland en Lesotho bestaan, is hersien. In Nesaea word 27 spesies (sewe nuut), vier varieteite en een nuwe vorm erken.

Ammannia is met behulp van die DECORANA-rekenaarprogram hersien, en vier spesies word erken. Sleutels tot die taksons,

asook sinonimie (tot die streek beperk), diagnoses en beskry wings van nuwe taksons, en aantekeninge oor verspreiding,

taksonomie en nomenklatuur word gegee. Die volgende taksons is nuut: Nesaea alata Immelman, N. angustifolia Immelman,

N. cymosa Immelman, N. minima Immelman, N. sagittifolia (Sond.) Koehne var. ericiformis Koehne forma swaziensis

Immelman, N. saluta Immelman, N. wardii Immelman en N. zambatidis Immelman.

INTRODUCTION

This synopsis of Nesaea and Ammannia in southern

Africa was written during the preparation of an account

of the Lythraceae for the Flora of southern Africa ( FSA ).

A combined key to the taxa of the two genera in the area

is provided. Synonomy is restricted to those names based

on southern African material, or names which have been

used by various authors within the area. Distribution and

habitat are recorded. Notes on taxonomy, nomenclature

and typification are provided in some cases. Eight new

taxa are described.

Nesaea Commers. ex H.B.K.

The first person to give a monographic account of

Nesaea was Koehne (1882, 1903), who dealt with the

whole of the Lythraceae. He described a number of new

species and transferred many more from the ‘holdall’ genus

Ammannia. The two genera were distinguished for the first

time on the basis of capsule dehiscence (see key to genera

below). Koehne divided Nesaea into five sections and a

number of subsections and series.

A number of partial treatments of the genus have

followed, dealing with specific aspects such as pollen or

with the genus within a limited region, e.g. those of

Panigrahi (1976, 1979, 1980a,b, 1986), Keay (1954), Fohnert

& Roessler (1966) and Fernandes (1970, 1978).

Nesaea is a widespread genus of 50—70 species, most

of which occur in Africa and Madagascar with a few

* Grassland Research Centre, Department of Agricultural Development,

Private Bag X05, Lynn East, Pretoria 0039.

MS. received: 1990-01-02.

species in tropical Asia, Australia, North and Central

America. Twenty-seven species, four varieties and one

form are recorded from the Flora of southern Africa

region. These are all erect to prostrate herbs or small

shrubs occurring either on temporarily or permanently

damp open soil, or occasionally submerged.

Four of Koehne’s five sections are represented in

southern Africa, only the monospecific sect. Crinipedium

being absent. Only a few species and infraspecific taxa

are endemic to southern Africa, however, with the majority

of species being recorded from the northern part of the

region, and clearly tropical African in origin. The genus

is not represented in the winter rainfall region of southern

Africa.

In contrast to Ammannia (see below), the boundaries

of species of Nesaea are distinct, and it is my opinion that

there is little, if any, hybridisation in the southern African

species of the genus.

Dimorphic flowers occur in the closely related species

N. rigidula (Sond.) Koehne and TV. alata Immelman.

Trimorphic flowers are encountered in TV. sagittifolia

(Sond.) Koehne and TV. schinzii Koehne, also closely

related to each other (Koehne 1882, 1903).

Table 1 and Figure 1 illustrate the relative lengths of

the style and stamen filaments in pin and thrum flowers

of the above species (excluding TV. alata). There are twice

as many stamens as calyx lobes in these species, the

filaments occurring in two different lengths. The length

of the capitate stigma is not included in the measurements

of style length.

36

Bothalia 21,1 (1991)

TABLE 1. — Style and stamen filament lengths (mm) of three species

of Nesaea

predominantly autogamous (rarely cleistogamous), but

with at least a low level of outcrossing. Species do

apparently sometimes hybridise, and a well-established

American taxon is probably of hybrid origin, indicating

that some species are not exclusively autogamous. Graham

also points out that some species have large brightly-

coloured petals and basal nectaries, which would not be

expected in an entirely self-fertile species.

A few species of Nesaea are very similar to Amman-

nia , and these add to the difficulties of determination. With

some specimens it may not be possible to establish the

type of capsule dehiscence. The species of the two genera

are therefore treated in a combined key.

Ammannia L.

The genus was first described by Linnaeus (1753), who

placed three species in Ammannia , one of which, A.

baccifera L., is included here. The genus was much

confused with the subsequently described genera Rotala

and Nesaea. Their boundaries were first clarified by

Koehne (1880) using characters of capsule dehiscence.

Subsequent treatments of Ammannia , as in Nesaea , have

dealt with specific aspects of the genus, or with the genus

in a restricted region. Fernandes (1978) recognized five

species and two subspecies in the Flora zambesiaca area.

Graham (1985) revised the five species occurring in the

western hemisphere, of which two also occur in our area.

She also provided a summary of the biology of the genus.

Ammannia comprises + 25 species, widespread in

tropical, subtropical and temperate regions, with 16 species

recorded from Africa. Four species occur in the Flora of

southern Africa region, all herbaceous and all growing in

standing water or marshy areas in the summer rainfall

areas. One has been recorded as a weed in rice paddies,

but does not appear to be a serious problem in our region.

None of the species of Ammannia is endemic to the

southern African region, and all are primarily tropical

African (or even more widespread), extending into

southern Africa at their southern limits only.

Di- or tri-stylous flowers such as seen in a few species

of Nesaea, do not occur in Ammannia. The pollen of

Ammannia, as in Nesaea, is prolate and tricolpate, with

six well-defined pseudocolpi (Panigrahi 1979).

Despite the revisions mentioned above of some species

of Ammannia, many specimens cannot be identified, and

appear to be intermediate between the accepted species.

Although some characters, e.g. style length, peduncle

length and petals, seem to have parallel variation, they

occasionally vary in different combinations. These

exceptional specimens have made it difficult to decide

where to draw the boundaries between taxa, as the position

of the boundary may differ according to which characters

are used as diagnostic.

Possible reasons for the great variability in Ammannia

are that the species are autogamous, or that they

may hybridise. According to Graham (1985) they are

METHODS

Material from all the major and some minor South

African and Namibian herbaria was examined. In Nesaea

the traditional taxonomic methods proved sufficient to

1.2

FIGURE 1. — Dimorphic and trimorphic flowers in Nesaea, showing

relative lengths of style and one pair of stamens: measurements

used are medians. 1.1, N. rigidula; 1.2, N. schinzii; 1.3, N. sagitti-

folia subsp. sagittifolia.

Bothalia 21,1 (1991)

37

TABLE 2.— Eigenvalues for specimens of Ammannia (first two axes

only)

distinguish taxa, but in Ammannia no clear pattern

emerged. A computer program package was therefore

used.

A suitable computerised method of analysing the

variablity encountered is the PHYTOTAB package, which

includes DECORANA (detrended correspondence

analysis). These programs were written by Westfall & Ded-

nam (PHYTOTAB) and Hill (DECORANA) to ordinate

and determine discontinuities in ecological data, thus de-

termining community limits (Hill 1979). It was thought that

DECORANA could be equally useful in helping to eluci-

date species limits in taxonomy.

Fifty-two specimens from PRE and WIND of the

southern African Ammannia species were examined and

scored for eight important characters. Characters used

were those chosen by previous workers as useful in keys,

i.e. shape of leaf base, peduncle and pedicel length,

presence or absence of petals, length of the calyx, stamen

filaments and style, and the presence or absence of calyx

appendages. Before being included in the data set, the

character states were aggregated into classes.

DECORANA was then used to group the specimens

according to their overall similarity. This resulted in four

sets of eigenvalues, of which the first two sets (Table 2)

were used as the X and Y axes of a scatter plot (Figure

2). Five of the eight characters used were then individually

superimposed on the common scatter plot to make a set

of pictorialised scatter plots (Figures 3-7).

RESULTS

Twenty-seven species, four varieties and one form were

distinguished in Nesaea.

In Ammannia , on the basis of results obtained from

DECORANA alone, there are two distinct groups present,

each of which can be divided into two further, if less

clearcut, groups.

When the basic scatter plot was used as the basis for

drawing up pictorialised scatter plots, i.e. using overall

similarity of specimens combined with single characters,

the boundaries between the four possible taxa became far

clearer. The scatter plots which most clearly showed these

boundaries proved to be those where petals (absent, small

or large), style length, peduncle length, leaf bases and

calyx appendages (absent, minute or clearly defined) were

used. This technique also showed up a few specimens

which did not fit into any group, emphasizing the

variability of the taxa. Much of the confusion surrounding

species delimitation probably arises from the description

of intermediate specimens as formal, named taxa.

The four taxa are here recognized at species rather than

subspecific rank. Synonymy is restricted to those names

based on southern African material, and names which have

been used by various authors within the area. Notes are

provided where necessary, and distribution and habitat

described.

Key to the genera

Capsule dehiscing irregularly; flowers always monomorphic,

in dense or lax cymes and subtended by large or small

subulate bracts, never aggregated into heads; calyx

lobes 4(5); stamens (2)4( — 8) 1. Ammannia

Capsule dehiscing first by a small apical operculum, then

irregularly; flowers mono-, di- or trimorphic; some

species with flowers aggregated into heads which are

subtended by enlarged, often folded, bracts; calyx lobes

4(5)6; stamens 4(5) — 8 — L2 2. Nesaea

Combined key to species of Nesaea and Ammannia

la Calyx lobes regularly 5; stamens regularly 5 2.9. N. rautanenii

lb Calyx lobes 4(5)6; stamens 4(5)6—12:

2a Calyx lobes 6; stamens 12:

3a Primary bracts up to 8 mm long, folded along midline; calyx appendages 0,7— 1.5 mm long; flowers homo-

morphic 2.10. N. radicans

3b Primary bracts + 1 mm long, flat; calyx appendages very short to (usually) absent; flowers dimorphic or

homomorphic:

4a Soft, probably annual herbs without woody taproot; petals pink or purple; inflorescence subsessile;

style of pin flowers 3,6— 4,3 mm long 2.16. N. alata

4b Perennial herbs with stems woody at base, arising from a stout woody taproot; petals orange; inflorescence

on peduncle 2—15 mm long; style of pin flowers 5,0— 8,6 mm long:

5a Leaf bases cuneate to obtuse; open flowers on primary peduncles 10 mm or longer (in southern African

specimens); flowers dimorphic; N & E Transvaal, Swaziland 2.18. N. heptamera

5b Leaf bases cordate to sagittate; open flowers on primary peduncles 2—5 mm long (in southern African

specimens); flowers homomorphic; Namibia, Botswana and N & W Transvaal 2.17. N. rigidula

38

FIGURE 2.— Scatter diagram of eigenvalues for Ammannia , first two

axes only. Possible boundaries between species, .

FIGURE 3. — Scatter diagram of eigenvalues for Ammannia , plotted

against shape of leaf bases. Leaves: all auriculate, #; cuneate

and auriculate on same plant, ©; all cuneate, O.

FIGURE 4. — Scatter diagram of eigenvalues for Ammannia , plotted

against length of primary peduncles (mm). <0,4: O; 0,5— 0,9:

®; 1,0-1, 9: •; 2, 0-2, 9: 4; 3, 0-3,9: 4"; 4, 0-5, 9: 4^;

6-14: -f-.

Bothalia 21,1 (1991)

FIGURE 5. — Scatter diagram of eigenvalues for Ammannia , plotted

against petals. Petals: large and brightly coloured, #; small and

white, C>; absent, O.

FIGURE 6. — Scatter diagram of eigenvalues for Ammannia , plotted

against style length (mm). 0,12—0,2: O; 0,3— 0,8: (*) ; 0,9— 1,0:

• ; 1,1— 1,6: 4; 1, 7-1,8: 41; 1,9-2, 0: 2, 1-2,2:

FIGURE 7. — Scatter diagram of eigenvalues for Ammannia , plo'tted

.gainst calyx appendages. Appendages: large, 0; small, ©;

absent, O.

Bothalia 21.1 (1991)

39

2b Calyx lobes 4( — 5); stamens 4-8:

6a Flowers in a ± hemispherical head subtended by large conduplicate bracts much longer than flowers, base

of bracts often whitish with darker veining, apices abruptly acuminate

7h Sy‘,e Z? 0,3 '"7 l0ng; pett abSent 2.14. N. schlechteri

7b style 0,65 mm or longer; petals present:

8a Style 2,5 mm or longer; robust herb up to 0,6 m high; inflorescence usually sessile 2 15 N tolypobotrys

8b Style up to 2,0 mm long; plant up to 0,2 m high; inflorescence pedunculate:

9a Style 1, 2-2,0 mm long; calyx appendages 0, 4-1,0 mm long 2.11. N. cordaia

9b Style 0.65-0,85 mm long; calyx appendages up to 0,2 mm long 2.12. N. angustifolia

6b Flowers, if in a hemispherical head, not enclosed by large conduplicate bracts, bracts shorter to somewhat longer

than flowers not white with darker veining at base, concolorous or may be darker along midline with

whitish membranous margins, apices obtuse, acute or attenuate, not abruptly acuminate:

10a Plants minute, up to 15 mm high, with ± 2 pairs of leaves and a solitary terminal flower 2.13. N. minima

10b Plants at least 100 mm high, with numerous pairs of leaves and numerous flowers:

11a Primary bracts half as long to longer than calyx, usually with dark centres and whitish membranous margins'

12a Style longer than 0,8 mm; stamens exserted:

13a Petals absent or cream-coloured; stamens 4; leaves usually spiral on at least some branches densely

hirsute; SE Botswana and E Transvaal 2.19. N. passerinoides

13b Petals present, bright purple; stamens 8; leaves always opposite, glabrous; Namibia Transvaal

Natal and E Cape:

14a Style 0,8-2, 3 mm long; calyx appendages 0,3-0,8 mm long; bract 0,9-l,7 mm long; N Namibia

2.6. N. teixeirae

14b Style 2.6-4.0 mm long; calyx appendages 0,7-l,4 mm long; bract 4-8 mm long; Transvaal,

Natal and E Cape 2.15. N. tolypobotrys

12b Style up to 0.5 mm long; stamens included:

15a Calyx lobes and stamens 5 2.8. N. sarcophylla

15b Calyx lobes 4; stamens 4—8:

16a Stamens 8; Zululand (Natal); stigma sessile 2.5. N. wardii

16b Stamens 4—8; Namibia, Botswana, Transvaal, OFS, N Cape; if stamens 8 or plants in Natal, then

style 0,15—0,5 mm long:

17a Style 0,25-0,5 mm long; stamen filaments 1,25-2,0 mm long:

18a Stamens 8; small pink petals present 2.27. N. drummondii

18b Stamens 4; petals absent 2.3. N. aspera

17b Style up to 0,25 mm long; stamen filaments up to 1,2 mm long:

19a Stamens 8; small pink petals present 2.27. N. drummondii

19b Stamens 4; petals absent:

20a Style sessile; plant asperous or glabrous; primary bracts 1,0-1, 5 x ± 0,3 mm; N Namibia

2.2. N. ondongana

20b Style ± 0,25 mm long; plant glabrous; bracts 2, 2-3, 6 X 0,65-1,45 mm; Transvaal and

N Cape 2.4. N. cymosa

lib Primary bracts less than half as long as calyx, usually of one colour:

21a Style 0,12—1,0 mm long (—1,8 mm in N. zambatidis)', stamens included or shortly exserted, filaments up

to 2,0(— 4,0) mm long; base of larger leaves usually cuneate or cordate (except in N. zambatidis and

Ammannia spp ):

22a Stamens 8; petals present:

23a Stems quadrangular with well-defined asperous ribs; style 0,2 -0,4 mm long, included; mouth

of calyx very slightly constricted, tube ± cylindrical; Namibia, Botswana, N Transvaal and

N Cape 2.27. N. drummondii

23b Stems terete, glabrous; style 0,65-1,8 mm long, shortly exserted; calyx widest at mouth, tube

broadly campanulate; N & E Transvaal and Swaziland 2.24. N. zambatidis

22b Stamens 4; petals present or absent:

24a Stems usually terete, may be angled, soft, often decumbent and rooting at nodes; leaves always

flat:

25a Leaf bases usually all auriculate, rarely auriculate and cuneate on same plant or all cuneate;

pedicels (0.5— )1, 5— 3,0 mm long; petals absent 1.2. A. senegalensis

25b Leaf bases all cuneate or upper leaves auriculate, rarely all auriculate; pedicels 0,5(— 3,0) mm

long; small white or large pink petals often present:

26a Pedicels 0,5(— 2,0) mm long; petals either absent or small and white 1.1. A. baccifera

26b Pedicels 2—5 mm long; petals either absent or large and pink:

27a Large pink petals present; pedicels 2—5 mm long 2.1. N. crassicaulis

27b Petals absent; pedicels 2—3 mm long 2.2. N. ondongana

24b Stems always quadrangular and sharply angled, often winged, plants erect and wiry, not rooting at nodes;

leaves often with rolled margins, especially when young:

28a Style up to 0,25 mm long; plant up to 0,5 m high; Zululand and N Natal 2.22. N. woodii

28b Style 0,35—1,0 mm long; plant up to 0,2(— 0,5) m high; Namibia, Botswana, Transvaal, OFS

and N Cape:

29a Petals absent; calyx appendages long, acute and spreading; inflorescence sessile or peduncles

and pedicels each up to 1 mm long; wings on stem usually asperous 2.7. N. anagalloides

29b Small pink petals present, at least in bud; appendages absent; inflorescence subsessile or

peduncles up to 6 mm long, pedicels up to 4 mm long; stem glabrous 2.21. N. saluta

21b Style 1,3— 7,0 mm long; stamens exserted, filaments 3,0 mm or longer (shorter in Ammannia auriculata );

base of larger leaves sagittate:

30a Style up to 3 mm long; stamens 4—8, filaments up to 4 mm long:

31a Stamens 8; calyx without appendages 2.24. N. zambatidis

31b Stamens 4; calyx with or without appendages:

32a Petals absent; styles 1,3— 1,6 mm long; stems and leaves densely hirsute over whole surface;

leaf margins rolled, leaves often spirally arranged on at least some stems 2.19. N. passerinoides

32b Petals usually present (may be small); styles 0,5— 2,6 mm long; stems not hirsute but may be asperous

on wings only; leaves flat, always opposite:

33a Styles 0,5—0,75 mm (—1,3 mm in tropical Africa) long; calyx 0,9— 1,1 mm long 1.3. A. prieuriana

33b Styles (1 — )1,45 — 2,6 mm long; calyx 1, 1-2,0 mm long:

40

Bothalia 21,1 (1991)

34a Peduncles up to 6 mm long; style 1,45-2,6 mm long 2.20. N. dinteri

34b Peduncles (3— )6— 14 mm long; style (1— )1,3 — 1,5 mm long 1.4. A. auriculata

30b Style (in at least some flowers in inflorescence) longer than 3,5 mm; stamens 8, at least some longer

than 5 mm;

35a Leaves spirally arranged; stems terete 2.26. N. sagittifolia

35b Leaves opposite or, if spirally arranged, then stems quadrangular:

36a Calyx 1,0-2, 5( —5,0) mm long; flowers trimorphic; styles 1,0-4,75 mm long; pin flowers with the

short stamens usually O3 to 2i 3 the length of the long stamens; widespread but rare in Namibia

2.25. N. schinzii

36b Calyx 2,5— 5,0 mm long; flowers homomorphic, all with long styles; styles (4,25— )5,25 — 6,75 mm

long; short stamens usually 2 13 to equal the length of the long stamens; restricted to Namibia

2.23. N. luederitzii

1 Ammannia L., Species plantarum 1: 119 (1753).

Type species: Ammannia latifolia L. (fide Britton &

Brown: 577 (1913)).

Annual (possibly short-lived perennial) herbs. Leaves

opposite-decussate, sessile, entire. Inflorescence pedun-

culate or sessile, lax or condensed dichasial cymes of

(1)3 -numerous (lowers; bracts 2 below each flower, small,

subulate. Flowers 4( — 5)-merous, never heterostylous.

Calyx tube broadly to narrowly campanulate, becoming

globose in fruit, usually 4-lobed, lobes alternating with

conical appendages or appendages absent, appendages

very short to nearly as long as lobes. Petals 1—4 or absent,

caducous. Stamens (2 — )4( — 8), inserted on calyx tube,

included or exserted, subequal. Ovary incompletely

2— 4(5)-locular, sometimes unilocular, upper portion of

septa incomplete; style absent to 1,5 mm long. Capsule

globose or ellipsoid, dehiscing irregularly; seeds numerous

and small, concave-convex, often with a float of thin-walled

cells on the concave side, often with hairs on the convex

side and margin which evaginate in moisture.

1.1 A. bacciferaL., Species plantarum 1: 120(1753);

Hiern: 478 (1871); Koehne: 259 (1880); Koehne: 53, fig.

5M (1903); Burtt Davy: 198 (1926); Keay: 165 (1954);

Pohnert & Roessler: 2 (1966); Webb & Moore: 302 (1968);

Fernandes: 177 (1970); Fernandes: 308 (1978); Graham:

405, fig. 2, map 1 (1985). Type: China, Savage H. (LINN

156/4, lecto. Graham 1985; microfiche in PRE, IDC

177.99: III. 4!).

A. aegyptiaca Willd. : 6, t. 6 (1803); DC.: 78 (1828); Fernandes: 309

(1978). A. baccifera L. subsp. aegyptiaca (Willd.) Koehne: 260 (1880);

Fernandes: 309 (1978). Iconotype: Willdenow in Flort. Berol. 1: 1.6 (1803).

(B-WILLD; photo, in PRE!).

A. wormskioldii Fisch., C.A. Mey. & Ave-Lallemant: 42 (1840);

Koehne: 256 (1880) as A. wormskjdldii\ Koehne: 53, fig. 51 (1903);

Pohnert & Roessler: 3 (1966); Fernandes: 177 (1970); Fernandes: 308

(1978). Type: Brazil, hb. Berol. (Bf).

A. salicifolia auct. non Monti: sensu Fliern: 478 (1871).

A. wormskioldii var. alata Koehne: 48 (1908). Type: Namibia,

Okahandja, 1 200 m, July, Dinter 253 (HBG!, SAM!).

The species occurs in N Namibia, NE Botswana, the

E, central and S Transvaal, the Orange Free State and the

N Cape. It was also seen by the author from Burundi,

Tanzania, Kenya, Zimbabwe and Angola. Other authors

have recorded it from Africa both south and north of the

Sahara, possibly Madagascar, Mauritius, the Middle East,

India and Ceylon, Java, the Phillipines, Russia, China,

Japan and Australia (Koehne 1880). It has been recorded

from Italy, Guadaloupe and Jamaica as an adventive

(Graham 1985). The species is found growing in moist

soil near water bodies, or in water.

The iconotype of Ammannia aegyptiaca is poor, but

shows the relevant features. Leaf bases are attenuate, the

flowers are + sessile, there are no petals, small calyx

appendages appear to be present, and the stigma is

subsessile. In the description these characters are repeated,

and the description of the calyx appears, with some

difficulty in the interpretation, to indicate the presence of

appendages.

The description of A. wormskioldii is short, but indicates

a plant with 4-merous flowers having 4 small white petals,

the flowers being in sessile axillary cymes. Although the

type has not been seen, the description is adequate to

identify it.

1.2 A. senegalensis Lam. ex Poir. in Lam., Ency-

clopedic Methodique, Botanique 1,1: 130 (1783); Lam.:

tab. 77 (1791); Lam.: 328 (1810); DC.: 77 (1828); Hiern:

477 (1871) p.p. excl. syn. A. prieuriana\ Koehne: 255

(1880); Koehne: 52 (1903); Burtt Davy: 198 (1926); Keay:

165 (1954); Fernandes: 176 (1970). Type: E Senegal,

Roussillon s.n. (P, microfiche no. 238/18, photo, in PRE!).

The species occurs in N Namibia and N Botswana, the

N and E Transvaal, Swaziland and N Natal. It was also

seen by the author from Tanzania, Zambia, Zimbabwe and

Angola; it is recorded by other authors from Egypt

(Koehne 1880), and from West Africa and the Sudan (Keay

1954). It occurs in or near water.

The primary peduncles of the type specimen are short

and the calyx appendages present, but styles are not visible

in the photograph. The styles are sessile according to

Koehne (1880) and in the illustration in Koehne (1903).

According to Graham, A. senegalensis is a synonym of

A. auriculata, but she has also seen only a photograph

of the type.

1.3 A. prieuriana Guill. & Perr. in Guill., Perr. &

A. Rich., Florae senegambiae tentamen 1: 303 (1833);

Koehne: 248 (1880); Koehne: 48 (1903); Keay: 164 (1954);

Fernandes: 175 (1970); Fernandes: 306 (1978). Holotype:

Gambia, marshy places around Albreda, Leprieur s.n. (G!)

A. pusilla Son<J.: 40 (1850). Type: wet places near Sandriver, May,

Zeyher , Fib. Sonder (S; SAM, mixed specimen! with Nesaea anagalloides

and a Bergia sp.).

The species occurs in E and central Transvaal, and

Natal. It was seen by the author from Burundi, Tanzania,

Kenya, Malawi, Zambia and Mozambique, and is also

recorded from West Africa (Koehne 1880; Keay 1954).

The species grows near or in water.

Bothalia 21,1 (1991)

41

The type of the species has styles 0,94-1,1 mm long,

the leaves auriculate and the primary peduncles short. The

calyx appendages are present but there are no petals.

The type of A. pusilla is given in Graham (1985) as

'Senegal or Nigeria, swampy places near Sand-rivier,

Zeyher 541 (Bt).’ However, 'Senegal or Nigeria’ is not in

the original citation in Sonder, and his article is titled (in

German) ‘Additions to the flora of South Africa’. Zeyher

did not collect in Senegal or Nigeria, and the specimen

is certainly from a South African locality. The specimen

has peduncles + 2 mm long, the calyx 1,5 mm long with

small but well-defined appendages, there are no petals and

the style is + 1 mm long.

1.4 A. auriculata Willd. in Hortus berdlinensis 1: 7,

t. 7 (1803); DC.: 80 (1828); Koehne: 244 (1880); Britton

& Brown: 469 (1879); Koehne: 45, fig. 5B (1903); Keay:

164 (1954); Pohnert & Roessler: 2 (1966); Webb & Moore:

302 (1968); Fernandes: 175 (1970); Fernandes: 305 (1978);

Graham: 403, map 1 (1985) p.p. excl. syn. A. pusilla.

Iconotype: Willdenow in Hort. Berol. 1: t. 7 (1803), (B-

WILLD; photo, in PRE!).

A. senegalensis var. auriculata (Willd.) Hiern: 477 (1871).

The species occurs in N Namibia, N Botswana and the

E Transvaal. It was also seen by the author from Burundi,

the Congo, Tanzania, Zambia, Zimbabwe, Mozambique

and Angola. Other authors record it from West Africa

(Keay 1954), the Middle East, India, China and Australia

and as introduced in Central and North America and the

Caribbean, and possibly also in South America (Graham

1985). It is found growing in vleis, moist places in

grassveld, near waterfalls, etc.

This species may be confused with Nesaea saluta , but

can be differentiated by a number of characters. The

peduncles are (3— )6— 14 mm long, and also have a

tendency to curve upwards, especially the older ones, while

those of the Nesaea are up to 6 mm long (but usually much

shorter) and quite straight. The calyx and style are more

than 1 mm long and large pink petals are present.

This species, according to Graham (1985), is one of the

most important weeds of rice in India, and is controlled

by raising the water level high enough to prevent it from

establishing itself. It has been introduced into the New

World, where the earliest record was in the 1930’s from

Guadaloupe. It has since been found in Jamaica, but

appears to be of limited distribution.

The numerous varieties and forms of this species which

were described by Koehne, Graham considers

meaningless, as the species is known to be highly variable.

The iconotype of the species is very poor but, together

with the description, can be interpreted as follows: leaves

with bases auriculate-cordate, inflorescences with 3( — 5)

flowers on primary peduncles + 5 mm long, calyx without

appendages (this is not certain), petals present, style

shorter than stamens.

2 Nesaea Commerson ex H.B.K., Nova genera et

species plantarum 6 (quarto): 191 (1824). Type species:

N. triflora (L.f.) H.B.K.

Annual or perennial herbs, shrublets, rarely shrubs.

Leaves opposite, ± decussate, rarely ternate or alternate

(spiral), sessile or shortly petiolate, entire. Inflorescence

basically a cyme, this variously condensed, reduced or

aggregated, often capituliform and subtended by large

bracts at base, sometimes flowers solitary. Flowers

bisexual, sometimes distylous or tristylous, usually 4-, 5-,

or 6-merous (in FSA species), bibracteolate. Calyx tube

variously-shaped, lobed, lobes alternating with conical

calyx appendages, these may be absent or up to as long

as lobes. Petals 0 or as many as calyx lobes, caducous.

Stamens 4—12 (in FSA species), inserted on calyx tube,

in 1—2 series, subequal or long and short stamens alter-

nating, included or exserted. Ovary sessile, 2-5-locular;

style nearly absent to much longer than calyx. Capsule

globose to ellipsoid, opening by an apical operculum,

lower part dehiscing irregularly later; seeds numerous,

small, concave-convex, without inflating wing.

2.1 N. crassicaulis (Guill. & Perr.) Koehne in

Botanische Jahrbiicher 3: 324 (1882); Keay: 166 (1954);

Pohnert & Roessler: 6 (1966); Fernandes: 182 (1970);

Fernandes: 287 (1978). Types: Senegal, Cape Verde,

marshy places near Khann, marked ‘herbarium Richard’

and ‘Herbier E. Drake’, March, 1834, Perrottet (P, lecto.!,

here designated); Senegal, by springs at Ghielcouil, Cayor,

May, 1827, Leprieur s.n. (G! , P!) .

Ammannia crassicaulis Guill. & Perr.: 303 (1833); Hiern: 479 (1871).

The species occurs in Namibia (Caprivi) and N

Botswana (Okavango Swamps). It has also been seen by

the author from Portuguese Guinea, and recorded by others

from Angola, Zambia, Zimbabwe and Mozambique

(Fernandes 1970, 1978) and West Africa, Zanzibar and

Madagascar (Keay 1954). It grows in standing or running

water.

2.2 N. ondongana Koehne in Botanische Jahrbiicher

: 165 (1900a); Koehne: 78 (1900b); Koehne: 225 (1903);

Pohnert & Roessler: 7 (1966); Fernandes: 182 (1970);

Fernandes: 285, tab. 69 (1978). Type: Namibia, Ambo-

land, Ondongana, Jan., 1886, Rautanen 206 (Z, lecto.,

here designated!; H!).

Two varieties are recognized:

Calyx 1,2— 2,0 mm long; plant usually asperous, usually erect

2.2a. N. ondongana subsp. ondongana var. ondongana

Calyx up to 1,2 mm long; plant usually glabrous, usually erect

or may be decumbent

2.2b. N. ondongana subsp. ondongana var. evansiana

2.2a N. ondongana subsp. ondongana var. ondon-

gana.

The typical variety has been seen from the Kaokoveld

and on the banks of the Kundne River (N Namibia). It

has been seen by the author from Zimbabwe, and is also

42

Bothalia 21,1 (1991)

recorded by other authors from Botswana, Zambia and

Mozambique (Fernandes 1978). It grows in damp places

in sand or in pans.

The species is very similar to Ammannia baccifera but

the capsule is definitely that of a Nesaea.

2.2b N. ondongana subsp. ondongana var. evan-

siana (A. Fernandes & Diniz) A. Fernandes in Boletim

Sociedade Broteriana Ser. 2 , 48: 115 (1974); Fernandes:

285 (1978). Ammannia evansiana A. Fernandes & Diniz:

155 (1957). Type: Botswana, Kachikau area, swampy areas

by Chobe River [± 36 miles N of Kachikau on road to

Kazane], Pole Evans 4187 (SRGH, holo. ; PRE, iso.!).

The variety has been seen from Namibia (Caprivi),

Botswana and Swaziland. It has also been seen by the

author from Mozambique, Zambia and Malawi. It grows

on the edge of water bodies, dried-up pans and seasonal

floodplains; it may be partially submerged in still shallow

water, or mat-forming.

2.3. N. aspera (Guill. & Perr.) Koehne in Botanische

Jahrbiicher 3: 327 (1882); Koehne: 226 (1903); Keay: 166

(1954); Pohnert & Roessler: 5 (1966); Fernandes: 289

(1978). Holotype: (Senegambia), Senegal, Cape Verde,

moist sandy areas around Kounoun, March, collector

probably either Perrottet or Leprieur (not found in P).

Ammannia aspera Guill. & Perr.: 304 (1833); Hiern: 480 (1871).

The species occurs in central and N Namibia, Botswana

and N Natal. It has been seen by the author from Mozam-

bique, and has also been recorded from Angola, Zimbabwe

and tropical Africa generally (Keay 1954; Fernandes 1978).

It grows near pans and springs.

The type has not been seen, but our material matches

the type description well.

2.4 N. cymosa Immelman, sp. nov. , N. asperae

proxima, sed cymis pedunculatis, bracteis integris et stylo

breviore differt.

The species is most similar to N. aspera , but differs in

having pedunculate cymes, entire bracts and a shorter style.

TYPE. — Cape Province, 2822 (Glen Lyon): Hay

division, Paardekloof [Perdekloof], desiccating pool in

kloof, sandy soil, 5-1942 (—DA), Cooke s.n. (PRE, holo.),

44378 (BOL), 6448 (KMG).

Slender annual herb, branching freely from base or

along (?prostrate) stems, rooting at nodes; stems + 0,08

m long, probably quadrangular, glabrous, up to 1,5 mm

in diameter. Leaves subsessile, glabrous, lanceolate, up

to 17 x 4 mm, apex broadly acute, base cuneate (lower

nodes) or shallowly cordate. Inflorescence a lax or

moderately dense axillary cyme of + 5 flowers, this

subtended by large but not enclosing bracts, subsessile or

older inflorescences with primary peduncle up to 3 mm

long. Bracts glabrous, longer than flowers, 2,2— 3,6 x

0,65—1,45 mm, broadly lanceolate, apex acute, base

cuneate, without solid keel, margins whitish. Calyx 4( — 6)-

lobed, campanulate, glabrous, ± 1,2 mm long, append-

ages large, erect, 0,4— 0,6 mm long. Petals absent. Stamens

4, opposite calyx lobes, included, filaments + 1 mm long.

Style ± 0,23 mm long, included, seeds produced early.

The species is found in the N Cape and E Transvaal,

in moist sand or on rock sheets by water.

NAMIBIA. — SWA, no date given, Schoenfelder S891 (PRE).

TRANSVAAL.— 2530 (Lydenburg): Lowveld Botanic Garden, on rock

sheet in spray of Nels River waterfall, full sun, SE slope, 640 m, 21-5-1980

(— BD), Kluge 2288.

CAPE. — 2724 (Taung): Vryburg, Zoetvlei Farm (P.O. Box 2188,

Vryburg), 1 500 m, where water has stood in Windmill ca" ■>, 1988

(— AA), Speedy s.n. B (cf. 15/18) (PRE).

2.5 N. wardii Immelman , sp. nov., N. asperae

proxima, sed staminibus 8, caulibus quadrialatis et

stigmatibus sessilibus differt.

This species is most similar to N. aspera but differs in

having eight stamens, 4-winged stems and sessile stigmas.

TYPE.— Natal, 2831 (Nkandla): Umfolozi Game

Reserve, in moist mud at edge of pan, 130 m, 7-12-1954

(— B), Ward 2456 (PRE, holo.; NU, iso.).

Robust decumbent herb, probably annual, glabrous,

unbranched; stems rooting at nodes, ± 0,35 m high,

4-angled and strongly 4-winged, up to 3,5 mm in diameter.

Leaves glabrous, lanceolate, 30—40 x 5-11 mm, base

cordate-sagittate, apex obtuse. Inflorescence an axillary

cluster of 4—5 flowers, primary peduncles very short,

pedicels 1—3 mm long. Bracts lanceolate, up to 2,2 x 0,3

mm but usually less. Calyx 4-lobed, glabrous, globular

in fruit, up to 2,5 mm long, with sharply acute, long,

erect appendages up to 1 mm long, calyx lobes almost

obsolete. Petals absent. Stamens 8, included, fugaceous;

filaments ± 0,7 mm long. Stigma sessile. Seeds produced

in bud.

Two specimens have been seen, both collected from

Zululand, and growing on the edge of a pan.

NATAL.— 2732 (Ubombo): Mkuzi Game Reserve, margin of Bube

Pan, 107 m, 25 1-1960 (— CA), Tmley 3 (PRE).

This Zululand species is named after Mr C.J. Ward, in

recognition of his years of work and extensive collecting

in that region.

2.6 N. teixeirae A. Fernandes in Boletim Sociedade

Broteriana Ser. 2, 43: 141 (1969). Type: Angola, Huilla,

Namuculungo, 1 100 m, 25-6-1957, Teixeira 2538 (LUA,

holo.; LISC, iso.).

Within the FSA area there are a few records from NW

Namibia. The species has also been seen by the author

from Angola. It grows in marshy areas.

Although the type has not been seen, the PRE material

has been compared with a specimen from K ( Pearson

2542 ) authenticated by Fernandes. It also keys to this

species in Fernandes (1970).

Bothalia 21,1 (1991)

43

2.7 N. anagalloides (Sond.) Koehne in Botanische

Jahrbiicher 3 : 327 (1882); Koehne: 228 (1903); Burtt Davy:

199 (1926). Type: Rhinosterkop, marshy places by Vaal

River, May, Zeyher 541 (SAM, lecto.!, here designated).

Ammannia anagalloides Sond.: 40 (1850); Sond.: 515 (1868).

N. loandensis auct. non (Hiern) Koehne: Pohnert & Roessler: 6 (1966).

The species has been recorded from scattered localities

in the Transvaal and N Cape, growing in rock pools, moist

areas and pans in grassveld, and on the margins of streams.

Koehne (1882) distinguished between N. loandensis and

N. anagalloides as follows:

Bracteoles shorter than pedicels N. loandensis

Bracteoles longer than pedicels N. anagalloides

The name N. loandensis is used by Pohnert & Roessler

(1966) for the material in Namibia, and a single voucher

is cited. A duplicate of this voucher specimen is in PRE,

and agrees well with my concept of N. anagalloides. None

of the material seen by me from southern African herbaria,

and identified as W. loandensis \ matches the type (BM!),

which is a soft herb with terete, prostrate stems and short

included styles.

Our material rather matches N. anagalloides as

described by Koehne, except in having the bracts shorter,

not as long as the pedicels. The lectotype in SAM, how-

ever, has short bracts, so that this character is presumably

variable in the species.

2.8 N. sarcophylla {Hiern) Koehne in Botanische

Jahrbiicher 3: 328 (1882); Koehne: 228, fig. 43E (1903);

Fernandes & Diniz: 104 (1958); Fernandes: 177 (1970);

Fernandes: 284 (1978). Types: Angola, Mossamedes, Bero

near Cavalleiros, ‘sparsim in arenosis humidis juxta ripam

fluminum,’ 7-1859, Welwitsch 2365 (FISU, lecto. fide

Fernandes (1970); BM!).

Ammannia sarcophylla Hiern: 479 (1871).

N. sarcophylla has been recorded twice from north and

central Namibia, as well as from Angola (see type) and

Mozambique. It grows in wet areas in riverbeds.

2.9 N. rautanenii Koehne in Bulletin de l'Herbier

Boissier 6: 750 (1898a); Koehne: 165 (1900a); Koehne: 231,

fig. 45B (1903); Pohnert & Roessler: 7 (1966); Fernandes:

298 (1978). Type: Namibia, Amboland, Wasserstellen,

12-6-1898, Rautanen 218 (K!).

One specimen was seen from the Caprivi (Namibia).

The species is also recorded from Zambia (Fernandes

1978). It is found growing in vleis.

This is the only species of Nesaea in the region which

regularly has 5-partite flowers.

2.10 N. radicans Guill. & Perr ., in Guillemin,

Perrottet & Richard, Florae senegambiae tentamen 8: 306,

tab. 70 (1830-33); Hiern: 474 (1871); Koehne: 231 (1903);

Keay: 166 (1954); Fernandes: 188 (1970); Fernandes: 298,

299 (1978). Syntypes: Cape Verde, Khann, marshy places;

Cape Verde, N’Batal; Cape Verde, Kounoun; Senegal, near

N’Boro, Cayor, Perrottet or Guillemin (not seen).

Two varieties are recognized:

Plant glabrous or almost so 2.10a. var. radicans

Plant with ± dense crispate hairs 2.10b. var. floribunda

2.10a var. radicans

A single specimen was seen from the Flora area, from

Mochudi in Botswana. It has been seen by the author from

Zimbabwe, Mozambique, Zaire, Tanzania, Senegal and

Zanzibar, and is also recorded from Zambia (Fernandes

1978). The ecology is as for var. floribunda.

2.10b var. floribunda (Sond.) A. Fernandes in Bole-

tim Sociedade Broteriana Ser. 2, 48: 117 (1974); Fernandes:

299 (1978). Type: Natal, Omblasriver, near Port Natal

[Umlaas River, near Durban], April, Dr'ege s.n. (S!).

N. floribunda Sond.: 517 (1868); Hiern: 474 (1871); Burtt Davy: 199

(1926); Pohnert & Roessler: 6 (1966); Fernandes: 189 (1970).

This is the variety commonly encountered in the Flora

area, having been recorded from N Namibia, N and SE

Botswana, Transvaal, Swaziland, Natal and E Cape. It has

been seen by the author from Mozambique and Tanzania,

and is also recorded from Zambia, Malawi and Zanzibar

(Fernandes 1970, 1978). It grows in moist ground near

dams, vleis and streams, often among boulders or low

vegetation.

2.11 N. cordata Hiern in Flora of tropical Africa 2:

475 (1871); Koehne: 166 (1900a); Koehne: 232 (1903); Burtt

Davy: 199 (1926); Keay: 166 (1954); Pohnert & Roessler:

6 (1966); Fernandes: 187 (1970); Fernandes: 302 (1978).

Syntypes: Sudan, Madi, Speke & Grant s.n. ; Niger, Barter

s.n. ; Angola, Pungo Andongo, 2400—3800 ft., Welwitsch

2J27(BM, lecto. fide Panigrahi (1976); PRE, isolecto.!).

N. cordata occurs in the northern half of Namibia, SE

Botswana, Transvaal and Swaziland. It has also been seen

from Mozambique, Zimbabwe, Tanzania, Nigeria and

Ghana by the author, and by other authors from Angola,

Zambia and Malawi (Fernandes 1970, 1978) and from West

Africa (Keay 1954). It grows in seepage areas, pans and

by streams or springs, usually in grassveld on their banks

but occasionally submerged in shallow standing or slow-

flowing water.

2.12 N. angustifolia Immelman, sp. nov. , N. erectae

proxima, sed calycis appendicis brevissimis, stylo breviori

differt.

Most similar to N. erecta , but differs in having very

short calyx appendages and a shorter style.

TYPE.— Transvaal, 2327 (Ellisras): Ellisras, in shallow

seepage water, frequent, 25-5-1967 (—DA), Louw 4145

(PRE).

44

Bothalia 21,1 (1991)

Slender annual herb, 0,06—0,17 m high, glabrous or

sparsely pubescent with spreading, relatively long

hairs, unbranched or branching from base, branches

probably quadrangular when young, 0,5— 1,0 mm in

diameter. Leaves glabrous or pubescent, linear to linear-

lanceolate, 7,0—20,0 x 1,0— 1,5 mm, apex broadly acute,

base cuneate. Inflorescence of pedunculate axillary heads,

the (1 — )3 — 13 flowers enclosed by a pair of large bracts,

primary peduncle 3—23 mm long, flowers ± sessile in

head. Bracts glabrous or hirsute, lanceolate to ovate, ±

2,5— 5,8 x 0,8— 3,0 mm, folded along midline with a solid

keel, without a pale base or dark veining, apices acuminate

and strongly recurved. Calyx 4-lobed, campanulate,

1,8— 2,5 mm long, appendages small, up to 0,2 mm long.

Petals ± 0,7 mm long, pink or magenta. Stamens 4—8,

included to shortly exserted, of two different lengths (when

8), filaments 0,8— 1,1 and 1,6— 1,8 mm long. Style exserted,

0,65—0,95 mm long.

The species is recorded from the E Transvaal lowveld

and central Transvaal, growing in moist places in bushveld.

TRANSVAAL. — 2431 (Acomhoek): Kruger National Park. Satara,

Pumbe Pan, damp soil next to pan, 3-1967 (-BB), Van Wyk 4788 (PRE);

Manyeleti Game Reserve, bushveld, 420 m. 18-3-1976 (-DA),

Bredenkamp 1585 (PRE). 2528 (Pretoria): Transvaal, Sandfontein,

19-1-1894 (— AB), Schlechter 4243 (BOL).

2.13 N. minima Immelman , sp. nov. , N. cordatae

proxima, sed bracteis longioribus, petalis nullis, floribus

solitariis differt.

Similar to TV. cordata , but differs in having the bracts

longer, the petals absent and the flowers solitary.

TYPE. — Botswana, 1824 (Kachikau): Zwezwe ‘flats’

(receive the overspill of the Ngwezumba River in Feb.,

March), heavy white sand, open grassveld which is moist

below the surface, together with sedges, 18-5-1977, (— CB)

Smith 2037 (PRE).

Annual herb, glabrous, unbranched; stems very slender,

almost filamentous, 0,015 m high, less than 0,5 mm in

diameter. Leaves in 1 or 2 pairs per plant, glabrous, elliptic

to ovate-oblong, 4x1 mm long, base cuneate, apex

obtuse. Inflorescence of solitary terminal flowers, primary

peduncles very short, pedicels ± 0,5 mm long. Bracts

glabrous except for a few papillae along margin, broadly

lanceolate to ovate, longer than calyx, ±2x1 mm long,

with a solid keel but not folded along midline, apex obtuse.

Calyx 4-lobed, glabrous except for a few hairs on

appendages, broadly campanulate (only specimens in fruit

seen), ± 1,5 mm long; with well-defined conical

appendages, 0,3 -0,4 mm long. Petals absent. Stamens 4,

included, attached near base of tube, ± 1,5 mm long. Style

not seen. Seeds produced in bud (flowers cleistogamous?),

with evaginating hairs.

There is only a single record of this distinctive species,

from N Botswana, growing in the moist grassveld of a

floodplain.

2.14 N. schlechteri A. Fernandes in Garcia de Orta

4,2: 189, t. 1 (1980). Type: Transvaal, by Sandfontein,

humid depression, 1 330 m, 19-1-1894, Schlechter 4243

(MO, holo.; PRE, iso.!).

Within the FSA area there is one record from the

Ruacana Falls (Namibia), and one from the Transvaal (type

specimen). The Namibian specimen was growing on a wet

boulder near the falls.

2.15 N. tolypobotrys Koehne in Botanische

Jahrbiicher 22: 151 (1895); Koehne: 166 (1900a); Koehne:

232 (1903); Burtt Davy: 199 (1926); Fernandes: 297 (1978).

Type: Natal, on fields by Umhlatuzaan River, 100 ft., Med-

ley Wood 334 (BM).

Recorded from the SE Transvaal, Natal and E Cape, as

well as from Mozambique (Fernandes 1978). It grows in

wet places, often in sandy soil or on rock sheets.

2.16 N. alata Immelman sp. nov., TV. rigidulae

proxima, sed floribus roseis vel purpureis, inflorescentia

subsessili, stylis brevioribus differt.

The species is most similar to TV. rigidula, but differs

in having flowers pink to purple, the inflorescence sub-

sessile and the styles shorter.

TYPE. — Transvaal, 2331 (Phalaborwa): Kruger

National Park, Mtomene Pan, wet soil near bank of pan,

5-1977 (—AC), Gertenbach 7006 (PRE, holo.; KNP, iso.).

Erect annual herb, glabrous, ± 0,45 m high, unbranched

or with a few branches, young stems quadrangular,

strongly winged, wings may be minutely asperous, up to

2 mm in diameter. Leaves glabrous, lanceolate to nearly

linear, 25—55 x 2—7 mm, base shallowly sagittate, apex

broadly acute. Inflorescence an axillary cluster of (1 — )3 — 5

flowers, subsessile or peduncles up to 2 mm long. Bracts

glabrous, lanceolate, ± 1,5 x 0,3— 0,5 mm. Flowers

dimorphic. Calyx 6-lobed, glabrous, broadly campanulate,

2-3 mm long, appendages very small or absent, with 12

dark green stripes. Petals ± 1,5 mm long, purple. Stamens

12, of two lengths in the flower. Pin flowers: stamen

filaments 3,9 and 4,3— 4,7 mm long, exserted; style

3,6— 4,3 mm long, exserted. Thrum flowers : stamen

filaments ± 1,5 mm long, included; style 1,5— 4,0 mm

long, included.

Two specimens have been seen, one from the Transvaal

lowveld and one from Swaziland, in or next to still water.

SWAZILAND. — 2631 (Mbabane): Lubombo Mountains, 17 km N of

Siteki on road to Mhlumbe, between Farms Groenpan and Cyrildene,

in shallow pool in semi-shade, 500 m, 8-5-1976 (— BD), Culverwell 59

(PRE).

The opening mechanism of the capsule was examined

and is definitely that of a Nesaea.

2.17 N. rigidula (Sond.) Koehne in Botanische

Jahrbiicher 3: 333 (1882); Koehne: 166 (1900a); Koehne:

235 (1903); Burtt Davy: 199 (1926); Pohnert & Roessler:

7 (1966); Fernandes: 184 (1970). Type: Transvaal, Aapies-

river, Oct., Zeyher 542 (SAM!)

Lythrum rigidulum Sond.: 42 (1850); Sond.: 516 (1868).

N. nuicronata Koehne: 249 (1888); Koehne: 236 (1903). Type: Namibia,

Amboland, Omatope, Feb. 1886, ?Schinz (not seen).

Bothalia 21.1 (1991)

45

The species occurs in NE Namibia and the Transvaal

(type specimen). It has been seen by the author from

Zimbabwe, and has also been recorded from Botswana,

Angola and Zambia (Fernandes 1970, 1978). It grows in

moist open places such as margins of pans and in vleis.

It has a thick woody taproot and probably produces

annual deciduous shoots.

N. mucronata is given as a synonym of N. rigidula by

Pohnert & Roessler (1966), and by Fernandes (1978). I have

not seen the type, but accept their placing the name here.

2.18 N. heptamera Hiern in Flora of tropical Africa

2 : 472 (1871). Holotype: Mozambique, Zomba, and E end

of Lake Schirwa, 10-1861. Metier s.n. (K! ).

There are scattered records from the N and E Transvaal,

and from Swaziland. The species is, however, more

common in east tropical Africa, having been seen by the

author from Zimbabwe and recorded by other authors from

Zambia, Malawi, Mozambique, Zaire and Tanzania

(Fernandes 1978). It is found in moist places in grassveld,

often in saline areas.

Like N. rigidula , it has a thick woody taproot that

probably produces annual deciduous shoots: this would

also enable the two species to survive grass fires.

2.19 N. passerinoides (Hiern) Koehne in Botanische

Jahrbiicher 3: 338 (1882); Koehne: 237 (1903); Fernandes:

296 (1978); Fernandes: 186 (1970). Type: Angola, Huilla,

March, Welwitsch s.n. (probably 2336) (PRE!).

Ammannia passerinoides Hiern: 480 (1871).

There are a few scattered records from SE Botswana

and SE Transvaal. The species has been seen by the author

from Zimbabwe; it is also recorded by other authors from

Angola (Fernandes 1978). It grows beside water, but little

has been recorded about its habitat in our region.

Fernandes (1978) gives the habitat as ’grasslands, margins

of vleis and termite mounds, on black clayey or sandy

soils’.

N. passerinoides is closely related to N. sagittifolia but

can be distinguished from it by the absence of petals and

the homomorphic, short-styled flowers.

2.20 N. dinteri Koehne in Botanische Jahrbiicher: 166

(1900a); Koehne: 25 (1900b); Koehne: 68 (1902); Koehne:

237 (1903); Pohnert & Roessler: 6 (1966); Fernandes: 294

(1978). Type: Namibia, Hereroland. Okaruse, April,

Dinter 606 (Bt?, not traced elsewhere).

2.20a subsp. elata A. Fernandes in Boletim Sociedade

Broteriana Ser. 2, 48: 122, tab. 9 (1974); Fernandes: 294,

tab. 73 (1978). Type: Zambia, Mumbwa, Chunga, Kafue

National Park, Mitchell 18/50 (LISC, holo.; COL PRE!,

SRGH).

N. transvaalica A. Fernandes: 121. tab. 8 (1974). Type: Transvaal. Brits.

Assen area, on Farm Welgevonden. marshy area, 10-4-1936, Obermeyer

s.n. (SRGH. holo.!: PRE, iso.!).

The subspecies occurs in N Namibia and N Botswana,

central and E Transvaal, and Swaziland. It has been seen

by the author from Zimbabwe and Tanzania, and it has

also been recorded from Zambia and Malawi (Fernandes

1978). It occurs in or near water.

Superficially this species is very similar to Nesaea

saluta, but it can be distinguished by the longer styles and

the well-defined calyx appendages.

The styles of the type specimen of N. dinteri subsp. elata

are 2,2— 2,6 mm long, the calyx 1,85 mm, and the four

stamen filaments are ±2,4 mm long and opposite the

calyx lobes. N. transvaalica , on the other hand, has the

calyx 1,6 mm long, with large appendages and well-defined

asperous ribs and large, bright pink petals. The stamens

are 1,85 mm long and opposite the calyx lobes, and the

style is 1,25—1,6 mm long. The specimen is a small

unbranched asperous herb. Except for the small size and

short styles, this specimen is typical of N. dinteri ; it is

probably either very young or depauperate.

2.21 N. saluta Immelman, sp. nov,. N. dinteri subsp.

elatae proxima, sed stylis brevioribus, calycis appendici-

bus nullis differt.

The species is most similar to Nesaea dinteri subsp.

elata but differs in having shorter styles and no calyx

appendages.

TYPE.— Namibia, 1918 (Grootfontein): Grootfontein,

27-4-1934 (-CA), Dinter 7347 (PRE, holo.; BOL, HBG,

WIND, iso.).

Slender annual herb up to 0,2 m high, simple or

branching freely; stems quadrangular, glabrous, up to 1,5

mm in diameter. Leaves sessile, glabrous, oblong-

lanceolate, 8—18 x 2—4 mm, apex broadly acute, base

truncate to (usually) shallowly cordate, to slightly hastate,

sometimes discolorous. Inflorescence very variable with

age, of simple few-flowered axillary cymes on short

primary peduncles becoming lax and much-branched with

up to 20 flowers in fruit, primary peduncles up to 6 mm

long but usually shorter. Bracts narrowly lanceolate, entire,

± 0,5 mm long. Calyx 4-lobed, narrowly campanulate be-

coming globular in fruit, glabrous, 0,8-l,0 mm long,

without appendages, buds often apiculate at apex where

calyx lobes join. Petals small, soon caducous (sometimes

entirely absent?), pale pink or purplish brown. Stamens

4, opposite calyx lobes, shortly exserted, filaments 0,5 -1,0

mm long. Style exserted, 0, 4-0,8 mm long.

Within the Flora region the species was recorded from

Namibia and the E Transvaal; it was also seen from

Angola. It is found growing in moist, often sandy places.

NAMIBIA.— 1917 (Tsumeb): track between Tsumeb and Abenab, 9

km from main road, small vlei, 25-4-1963 (— BD), Kers 424 (PRE,

WIND, mixed specimen). 1918 (Grootfontein): Grootfontein, Gumab,

5-5-1934 (— CA), Schoenfelder 4419 (BOL, HBG. PRE); Farm

Oliewenhof, GR 215. palm flats, grey clay soil. 8-3-1974 (— CB). Merx-

mtiller & Giess 30129 (PRE, WIND).

2.22 N. woodii Koehne in Pflanzenreich 17 (IV, 216):

238 (1903). Types: Natal, Zululand, by Tugela River, Wylie

46

Bothalia 21,1 (1991)

sub Wood 5689a (PRE, lecto.! here designated; BOL!,

SAM!); Natal, Zululand, by Tugela River, Wylie sub Wood

8599 (NH!, marked as Lythrum rivulare).

N. woodii occurs in N Natal; it has also been seen from

Zimbabwe. It grows in moist sand on pan margins, stream

banks and sand dunes.

The species is closely related to N. schinzii, but differs

in having homomorphic flowers with 4 included stamens,

consistently shorter styles, and often longer and broader

bracts.

2.23 N. luederitzii Koehne in Verhandlungen des

Botanischen Vereins der Provinz Brandenburg 30:

251 (1888); Koehne: 238 (1903); Pohnert & Roessler: 6

(1966). Type: Namibia, Damaraland, Ganahomtes, Tsoa

Xaubthal, 26-9-1885, Schenk 420 (Z!).

Two varieties are recognized:

Leaves all opposite; pedicel of median flower 5—8 mm long; style

slightly longer than stamens; plant usually glabrous ..

2.23a. var. luederitzii

Leaves partly opposite, partly alternate; pedicel of median

flower 1—1,5 mm long; style much longer than stamens;

plant often hispidulous 2.23b. var. hereroensis

2.23a var. luederitzii

N. straminea Koehne; 167 (1900a); Koehne; 26 (1900b); Koehne: 239

(1903). Type: Namibia, Hereroland, Tsoachaubsand, Salem, September,

Dinter 139 (Bt).

The variety is scattered throughout Namibia, and is

probably the most common species there, if herbarium

records are taken as representative. It is found growing

in dry riverbeds and on streambanks.

N. luederitzii is closely related to N. schinzii, from

which it can be distinguished by the characters given in

the key. Sterile plants can often be distinguished from that

species by the yellowish stems and exfoliating bark at the

base of older stems of N. luederitzii. N. schinzii is rare

in Namibia.

From the type description I consider N. straminea a

synonym of N. luederitzii var. luederitzii. It is so treated

by Pohnert & Roessler (1966).

2.23b var. hereroensis Koehne in Botanische Jahr-

biicher 29: 167 (1900a); Pohnert & Roessler: 6 (1966).

Type: Namibia, Hereroland, Otjinene, on limestone, 13

April, 1899, Dinter 613 (Z!).

This rather rare variety is recorded from central and NE

Namibia. It grows in dry riverbeds.

2.24. N. zambatidis Immelman, sp. nov. , N. schinzii

proxima, sed floribus homomorphis, caulibus crassis

teretibus differt.

The species is most similar to N. schinzii, but differs

in the homomorphic flowers and in the stems being thick

and terete.

TYPE. — Transvaal, 2229 (Waterpoort): near Water-

poort, east of Pylkop Station, north of the Soutpansberg,

Farm Driehoek, in rocky outcrop in Mopane veld, growing

in pools of water in rocks, 700 m, 9-5-1984 (— DD);

Balkwill 1651 (PRE, holo.; NU, iso.).

Robust, probably annual herb; stems thick, tough and

spongy, up to 5 mm in diameter, glabrous, few-branched,

may or may not root at nodes, ± 0,3 m high. Leaves

glabrous, crowded, lanceolate, 37-65 x 6-8 mm, bases

sagittate, apex broadly acute to obtuse. Inflorescence a

dense subsessile axillary cluster of ± 5 flowers. Bracts

glabrous, narrowly lanceolate, 0,5— 0,7 x 0,1 mm. Calyx

4(5)-lobed, glabrous, broadly campanulate, 1,5 -3,0 mm

long, without appendages. Petals ± 4 mm long, pink.

Stamens twice as many as calyx lobes, included or shortly

exserted; filaments of two lengths with longest opposite

calyx lobes, 1,1— 1,8 and 1,8— 3,9 mm long. Style shortly

exserted, 0,65—1,8 mm long.

Specimens have been seen from the N and E Transvaal,

and Swaziland; the species was also seen from Zimbabwe.

It has been recorded as growing in standing water of rock

pools and seasonal pans.

TRANSVAAL. — 2431 (Acornhoek): Timbavati Private Reserve, Farm

Kempiana 90 KU, seasonal pan, in waterlogged clay, rare, 27-2-1984

(—AD), Zambatis 1636 (PRE). 2531 (Komatipoort): Kruger National

Park, Klapalamkwenya, pools in rock sheets, 700 m, 25-2-1954; (— AB),

Van der Schijff 3057 (KNP. PRE); Kruger National Park, Klapalam-

kwenya, in water, 1 350 m, 8-1-1955, Van der Schijff 4181 (KNP, PRE);

Kruger National Park, no precise locality given, 3-1960, Brynard &

Pienaar 4472 (KNP).

SWAZILAND. —2631 (Mbabane): Mlawula Nature Reserve, below

dam at rhino pools, in water, aquatic, 229 m, 16-3-1987 (— BD), Alward

91 (PRE). 2632 (Bela Vista): Lubombo Mountains, W of Umbeluzi Poort

and S of Mlawula Station, Farm Mlawula, pool below airstrip, on

Nkumbane Stream near junction with Mlawula River, in deep still pool,

in shallow water, 170 m, 22-5-1976 (— AA), Culverwell 94 (PRE).

The capsules are typical of Nesaea, though in many

respects the species resembles Ammannia baccifera or A.

senegalensis . It can be distinguished from these, however,

by the longer styles, the presence of pink petals and the

8, not 4 stamens.

The species is named after Mr N Zambatis who has

done much collecting in the Transvaal Lowveld, including

one of the quoted specimens of this new species.

2.25 N. schinzii Koehne in Verhandlungen des

Botanischen Vereins der Provinz Brandenburg 30: 250

(1888); Koehne: 239 (1903); Pohnert & Roessler: 7 (1966);

Fernandes: 186 (1970); Fernandes: 292 (1978). Type: Cape

Province, Upington, Oshando, March 1886, Schinz 517

(BOL!).

N. schinzii var. fleckii Koehne: 409 (1895). Type: Namibia, Hereroland,

Gansberg, Fleck 231a (not seen).

N. schinzii var. rehmannii Koehne: 151 (1895); Burtt Davy: 199 (1926);

Fernandes: 186 (1970). Syntypes: Transvaal, near Trichardtsfontein,

highveld, Rchmann 6679: Transvaal, Bronkhorstspruit, Rehmann 6567:

Transvaal, Pretoria, Rehmann 4749 (BOL!).

Bothalia 21,1 (1991)

47

N. kuntzei Koehne in Kuntze: 97 (1898b); Koehne: 167 (1900a); Koehne:

69 (1902); Koehne: 238 (1903). Holotype: Natal, Ladysmith, 1 100 m,

Kuntze 31; Natal, Ladysmith, 1 100 m, Kuntze 32 (NY; photo, in BOL!;

scrap and photo, in PRE!).

Lythrum rivulare Wood & Evans ex Wood: 12 (1901a); Wood & Evans:

172 (1901b). N. rivularis (Wood & Evans ex Wood) Koehne: 68 (1902);

Koehne: 238 (1903). Type: Natal, Zululand, near Tugela River, Wylie

sub Wood 5689 (NHI).

This very common species, probably the most common

one of the genus in southern Africa, is distributed over

Namibia, NW Botswana, the Transvaal and the E OFS.

It has been seen by the author from Zimbabwe, and by

other authors from Angola and tropical east Africa as

far north as Zaire and Kenya. It grows in damp open

grassveld.

The species has a stout woody taproot that produces

annual shoots, and which probably enables the plants to

survive grass fires. N. schinzii may be confused with N.

luederitzii in the Namibian region. It is, however, rather

rare in Namibia and can be distinguished by the trimorphic

flowers. Also the styles of the long-styled (pin) flowers

are shorter than the styles of flowers of N. luederitzii , and

proportions of the short to the long stamens in one flower

are different (see key to species).

N. schinzii may also sometimes be confused with N.

sagittifolia as these two species are closely related, and

their distributions overlap in the Transvaal and OFS. The

following distinguishing characters are considered most

reliable and easy to use:

N. sagittifolia

Stems terete

Leaves spiral

Leaves sessile, imbricate, except

in var. ericiformis

Leaves with sagittate clasping

bases except in var. erici-

formis

A scrap of the type specimen of N. kuntzei is in PRE.

The only significant difference seen between it and

specimens of N. schinzii with mid-styled flowers is the

four rather than eight stamens; this is also the only

difference given by Koehne (1903) in his key. All other

differences seen are differences only of size. I consider

it to be very probably a depauperate specimen of N.

schinzii with mid-styled flowers. The reduction in number

of stamens may also be due to the depauperate state. The

gynoecium is definitely that of a Nesaea.

N. rivulare also had four stamens in Koehne’s account:

otherwise there seems to be no fundamental difference

between it and N. schinzii . According to Koehne it is

differentiated from N. woodii by the much longer leaves.

The type of Nesaea schinzii var. fleckii was not seen,

but on the basis of the description I consider it a synonym

of N. schinzii.

2.26 N. sagittifolia (Sond.) Koehne in Botanische

Jahrbiicher 3: 339 (1882); Koehne: 240 (1903); Burtt Davy:

199 (1926). Type: Transvaal, boggy places near

Magaliesberg, Zeyher 543 (BM, lecto. fide Panigrahi

(1976); D, K, S!).

Lythrum sagittifolium Sond.: 41 (1850); Sond.: 516 (1868).

This attractive species appears to be endemic to the

eastern part of South Africa. Two varieties and two forms

are recognized:

la Calyx 3-4 mm long; inflorescence usually subsessile ....

2.26a. N. sagittifolia var. sagittifolia

lb Calyx 1—2 mm long; inflorescence subsessile or pedunculate:

2a Leaves erect and imbricate, sometimes spreading, rather

narrower than usual for the species; pedicels and

primary peduncles less than 2 mm long; Swaziland

with one record from Barberton (E Transvaal)

.. 2.26ba. N. sagittifolia var. ericiformis forma swaziensis

2b Leaves spreading, small; whole plant wiry and much-

branched; primary peduncles and pedicels 2—3 mm

long; East Griqualand, E Cape, Transkei, and Natal

.. 2.26bb. N. sagittifolia var. ericiformis forma ericiformis

2.26a var. sagittifolia

Koehne: 152 (1895a); Koehne: 240 (1903).

Nesaea sagittifolia var. glabrescens Koehne: 152 (1895); Koehne: 240

(1903); Burn Davy: 199 (1926). Type: Transvaal, near Standerton,

highveld, Rehmann 6796 (Z!) .

Nesaea sagittifolia var. salicarioides Koehne: 152 (1895): Koehne: 241

(1903); Burtt Davy: 199 (1926). Type: Natal, Schulze 1887-40 (not seen).

The typical variety occurs in the Transvaal, Natal and

E OFS. It is found growing in damp open grassveld.

Like N. schinzii, it has a stout woody taproot which

produces annual shoots, and which is important in enabling

the plants to survive grass fires.

Koehne described a number of varieties of N. sagitti-

folia, but I have maintained only var. ericiformis. The

calyx of the type specimen of N. sagittifolia var.

glabrescens is + 3 mm long. Although the leaves are clear-

ly of the sagittifolia type, on the young stems they are

spreading and not at all imbricate, and are often flat with

less deeply sagittate bases, as is found in var. ericiformis.

However, the large calyces as well as the distribution, dis-

tinguish it from that variety.

2.26b var. ericiformis Koehne in Botanische Jahr-

bucher 22: 152 (1895); Koehne: 241 (1903); Burtt Davy:

199 (1926). Type: Natal, Pietermaritzburg, Rehmann 7599

(Z!).

Var. ericiformis occurs in two separate areas: Swaziland,

with one record from Barberton; and E Griqualand,

Transkei and the E Cape, with one record from the Natal

midlands (the type specimen).

The populations from these areas are morphologically

partly distinct, though the distinction between the two

forms is not clearcut. The type of the variety, for instance,

matches some aspects of each. Differences can be seen

in the leaves, primary peduncles and pedicels. Neither of

these forms can be hybrids of N. sagittifolia with N.

schinzii, though they do resemble the latter species in many

N. schinzii

Stems quadrangular

Leaves usually opposite, some-

times spiral

Leaves shortly petiolate,

spreading

Leaves with base hastate to

cuneate, not clasping stem

48

Bothalia 21,1 (1991)

respects, as N. schinzii does not occur in these areas. The

typical variety also is absent from these areas, with the

exception of Barberton.

2.26bb forma ericiformis

Stems slender and wiry. Leaves spreading, 7,0—12,0 X

1.5— 2,5 mm. Inflorescence : primary peduncle 1,0— 2,5

mm long, pedicel 1,0-2, 5 mm long. Calyx glabrous or

minutely puberulous on ribs and at apices on calyx-lobes,

1.5— 2,0 mm long. Petals 1,5— 3,0 mm long, pink. Thrum

flowers : filaments 3,0-5,0 and 2,0 mm; style 0,7— 1,0 mm.

Mid-style flowers: filaments 4 and 2,0— 2,5 mm; style

3,0— 3,5 mm. Pin flowers: filaments 4,0— 5,5 and 1,5 mm;

style 5,0— 5,5 mm. Otherwise as for forma swaziensis.

Occurs in E Cape, E Griqualand, Natal and the

Transkei. It grows in damp places in the mountains.

Occasionally a plant of var. sagittifolia will be found

having 1 or 2 weak side branches showing the slender

stems and loosely arranged leaves typical of var. erici-

formis forma ericiformis. They can easily be distinguished,

however, by the difference in flower size, and by the fact

that the stems and leaves of forma ericiformis are consis-

tently as given in the description.

2.26ba forma swaziensis Immelman, forma nov. ,

formae ericiformae simillima, sed foliis erectis imbricatis,

pedunculis et pedicellis brevioribus differt.

Similar to forma ericiformis but differs in the erect

imbricate leaves and shorter peduncles and pedicels.

TYPE. — Swaziland, 2631 (Mbabane): Manzini

District, Mpisi Government Farm, swamp, 500 m,

22-6-1962 (— BC), Karsten s.n. (49640 in PRE, holo. ;

NBG, iso.).

Stems slender, often wiry, may be minutely striate under

the microscope, hairs dense and retrorse. Leaves may be

spreading near base of branches, otherwise imbricate, hairs

dense, 7,0—12,0 x 1,5— 2,5 mm. Inflorescence: primary

peduncle 1,5 mm long, pedicels + 1,0 mm long. Bracts

puberulous, lanceolate, 0,7— 1,5 x 0,2— 0,4 mm. Calyx

usually glabrous, sometimes minutely puberulous on ribs

and at apices of calyx-lobes, rarely hirsute, 1,5 -2,0 mm

long. Petals 1,5— 3,0 mm long, pink or magenta. Thrum

flowers: filaments 4, 5-6,0 and 2,0— 3,5 mm; style 1 mm.

Mid-style flowers: 5,5 and 1,5 mm; style 4 mm. Pin

flowers: filaments 5,0— 5,5 and 2,0 mm; style 5,5— 6,5 mm.

Occurs in SE Transvaal and Swaziland. It grows in vleis

and damp places in open grassveld.

TRANSVAAL.— 2531 (Komatipoort): Barberton, Concession Creek,

830 m, 18-1-1891 (-CC), Galpin 1273 (PRE).

SWAZILAND. — 2630 (Carolina): Mbabane District, Little Usutu

Valley, swamp, 10-4-1955, (-AC/CB) Compton 25099 (NBG, PRE);

Mbabane District, Little Usutu River, swamp, ± I 000 m, 10-3-1961,

Compton 30583 (NBG). 2631 (Mbabane): Mbabane District, Kirkhill,

moist ground, I 170 m, 9-3-1956 (—AC), Compton 25753 (NBG, PRE).

2.27 N. drummondii A. Fernandes in Boletim

Sociedade Broteriana 48; 124, t. 10 (1974); Fernandes: 304

(1978). Type: Botswana, Namibian border, 16 km west of

Knau Knau, Chadum Valley, wetter patch in flat valley-

bottom, 14-3-1965, Wild & Drummond 7019 (SRGH,

holo.!; COI, K!).

The species is distributed from central to N Namibia,

Botswana, N Transvaal and N Cape. It has been seen by

the author from Zimbabwe and is also recorded from

Mozambique (Fernandes 1978). It grows near water, on

dam margins and moist areas in grassveld.

Species incertae sedis:

Ammannia crassissima Koehne in Botanische Jahr-

biicher 4: 391 (1883); Koehne: 53 (1903); Burtt Davy: 198

(1926). Type: S Abyssinia, Delhi-Dikeno, collector

unknown (Bt). Burtt Davy’s concept of the name could

fit A. baccifera. Type not seen.

Ammannia multiflora Roxb. in Carey, Flora Indica 1:

447 (1820), non Fernandes & Diniz (1956); DC.: 79 (1828);

Koehne: 247 (1880); Pohnert & Roessler: 3 (1966). A.

senegalensis forma multiflora (Roxb.) Hiern: 477 (1871).

A. senegalensis var. multiflora (Roxb.) Koehne: 48 (1903).

Type: open lands around Calcutta, Roxburgh s.n., two

specimens from Herbs. Forsyth and Rottler respectively

(K!). The type could not be matched with any species of

Ammannia or Nesaea in the FSA region, and it seems

probable that Pohnert & Roessler misapplied the name.

ACKNOWLEDGEMENTS

I thank Dr Taciana B. Cavalcanti of the University of

Sao Paulo in Brazil and Dr Shirley A. Graham of the Kent

State University in Ohio, for critically reading the MS;

and Mr G.B. Deall and Mr M. Panagos, who gave much

help in making DECORANA work and in interpreting the

results.

REFERENCES

BRITTON, N. & BROWN, A. 1913. An illustrated flora of the northern

United States, Canada and the British possessions edn 2, vol.

2: 577. Scribner, New York.

BURTT DAVY, J. 1926. Lythraceae. A manual of the flowering plants

and ferns of the Transvaal, with Swaziland, South Africa 1:

196-199. Longmans Green, London.

DE CANDOLLE, A.P. 1828. Lythrarieae. Prodromus systematis naturalis

regni vegetabilis 3 : 75 —94. Treuttel & Wurtz, Paris.

FERNANDES, A. 1969. Nesaea (Sect. Ammanniastrum ) teixeirae , sp.

nov. Boletim Sociedade Broteriana Ser. 2, 43: 141.

FERNANDES, A. 1970. Lythraceae. In A.W. Exell, A. Fernandes, &

E.J. Mendes, Conspectus florae angolensis 4: 165—189. Junta de

Investigates do Ultramar, Lisbon.

FERNANDES, A. 1974. Lythraceae africanae novae vel minus cognitae

I. Boletim Sociedade Broteriana Ser. 2, 48: 115—168, tabs. 1—18.

FERNANDES, A. 1978. Lythraceae. In E. Launert, Flora zambesiaca

4 : 276 - 323. Crown Agents for Overseas Governments and

Administrations, London.

FERNANDES, A. 1980. Lythraceae africanae novae vel minus cognitae

IV. Garcia de Orta 4,2: 189—191, tab. 1.

FERNANDES, A. & DINIZ, M.A. 1956. Contribucao parao Lythraceae

de Mozambique. Garcia de Orta Ser. 1,3: 385—408.

FERNANDES, A. & DINIZ, M.A. 1957. Lythraceae africanae novae

II. Boletim Sociedade Broteriana Ser. 2,31: 151-160, tabs. 1—11.

FERNANDES, A. & DINIZ, M.A. 1958. Contribuzao para o con-

hecimento das Lythraceae de Angola. Garcia de Orta 6,1: 87-

BO.

Bothalia 21,1 (1991)

49

FISCHER, F.E.L. VON, MEYER, C.A. & AVE-LALLEMANT, J.L.E.

1840. Animadversiones Botanicae. Index Seminum, quae hortus

botanicus imperiatis Petropolitanum pro mutua commutatione

offert. Acceduni animadversiones botanicae nonnullae 7: 41-59.

Academiae Caesarea Petropolitanae, St Petersburg.

GRAHAM, S.A. 1985. A revision of Ammannia (Lythraceae) in the

western hemisphere. Journal of the Arnold Arboretum 66,4:

395-420.

GUILLEMIN, J. A. & PERROTTET, S. 1830-1833. In J.A. Guillemin,

S. Perrottet & A. Richard, Florae senegambiae tentamen 8: 306,

tab. 70. Treuttel & Wurtz, Paris.

HIERN, W.P 1871. Lythraceae. In D. Oliver, Flora of tropical Africa

2: 464—486. Reeve, London.

HILL, M .O. 1979. DECORANA: a FORTRAN program for correspon-

dence analysis and reciprocal averaging. Cornell University,

Ithaca, New York.

HUMBOLDT, F.W.H.A. VON, BONPLAND, A.J. & KUNTH, C.S.

1824. Nova genera et species plantarum 6 (quarto): 191. Lutetiae

Parisiorum, Paris.

HUTCHINSON, J. & DALZIEL, J.M. 1927. Lythraceae. Flora of West

tropical Africa 1,1: 142—145. Crown Agents, London.

KEAY, R.W.J. 1954. Ammannia, Nesaea. In J. Hutchinson, & J.M.

Dalziel, Flora of west tropical Africa, edn 2, 1,1: 164-166. Crown

Agents, London.

KOEHNE, E. 1880. Lythraceae monographice describuntur II: Amtnan-

nia. Botanische Jahrbucher fur Systematik, Pflanzengeschichte

und Pflanzengeographie 1: 240—262.

KOEHNE, E. 1882. Lythraceae monographica describuntur: Tribus II

Nesaeae. Botanische Jahrbucher fur Systematik, Pflanzen-

geschichte und Pflanzengeographie 3: 319—352.

KOEHNE, E. 1883. Lythraceae monographice describuntur, addenda

et corrigenda. Botanische Jahrbucher fur Systematik, Pflanzen-

geschichte und Pflanzengeographie 4: 386—408.

KOEHNE, E. 1888. In H. Schinz, Beitrage zur Kenntnis der Flora von

Deutsch-Siidwest-Afrika und der angrenzenden Gebiete II:

Lythraceae. Verhandlungen des Botanischen Vereins der Provinz

Brandenburg 30: 248—252.

KOEHNE, E. 1895. Lythraceae africanae. Botanische Jahrbucher fur

Systematik, Pflanzengeschichte und Pflanzengeographie 22:

149-152.

KOEHNE, E. 1898a. In H. Schinz, Beitrage zur Kenntnis der

afrikanischen Flora: Lythraceae. Bulletin de I'Herbier Boissier

6: 750.

KOEHNE, E. 1898b. Lythraceae. In O. Kuntze, Revisio generum

plantarum 3,2 : 96 —97. Felix, Leipzig.

KOEHNE, E. 1900a. Lythraceae novae. Botanische Jahrbucher fur Sys-

tematik, Pflanzengeschichte und Pflanzengeographie 29: 154—168.

KOEHNE, E. 1900b. In H. Schinz, Beitrage zur Kenntnis der

afrikanischen Flora 12: Lythraceae. Memoires de I'Herbier

Boissier 20 : 24 - 26, 78 -79.

KOEHNE, E. 1902. Lythrum rivulare. Journal of Botany, British and

Foreign 40: 68—69.

KOEHNE, E. 1903. Lythraceae. In A. Engler, Das Pflanzenreich: regni

vegetabilis conspectus 17 (IV, 216): 1-325. Engelmann,

Weinheim.

KOEHNE, E. 1908. Lythraceae: Nachtrage II. Botanische Jahrbucher

fur Systematik, Pflanzengeschichte und Pflanzengeographie 42,

Beibl. 97: 47-53.

LAMARCK, J.B.P.A.M. DE. 1783. Ammane. Encyclopedic methodique

1,1: 130—131. Panckoucke, Paris.

LAMARCK, J.B.P.A.M. DE. 1791. Recueil de Planches de Botanique

1: tab. 77. Agasse, Paris.

LAMARCK, J.B.P.A.M. DE. 1810. Tableau Encyclopedic Methodique

des trois regnes de la nature: Botanique, Suppl. 1: 328—329.

Agasse, Paris.

LINNAEUS, C. 1753. Species plantarum 1: 119. Salvii, Stockholm.

PANIGRAHI, S.G. 1976. Studies on generic delimitations on the four

genera Rotala, Ammannia, Nesaea and Hionanthera (Lythraceae)

— a historical survey. Bulletin of the Botanical Survey of India

18: 178-193.

PANIGRAHI, S.G. 1979. A contribution to the palynotaxonomy of four

herbaceous genera of Lythraceae. Fifth International Palynological

Conference, Lucknow (1976-77) 1: 422—431.

PANIGRAHI, S.G. 1980a. Studies on the cuticle and stomata of

Ammannia, Rotala, Nesaea and Hionanthera (Lythraceae).

Journal of the Orissa Botanical Society 2: 13—18.

PANIGRAHI, S.G. 1980b. Contribution of anatomy to the systematics

of Ammannia. Phytomorphology 30,4: 320—330.

PANIGRAHI, S.G. 1986. Seed morphology of Rotala L., Ammannia L.,

Nesaea Kunth. and Hionanthera Fernandes & Diniz (Lythraceae).

Botanical Journal of the PJnnean Society 93: 389—403.

POHNERT, H. & ROESSLER, H. 1966. Lythraceae. In H. Merxmiiller,

Prodromus einer Flora von Siidwestafrika 95: 1—9. Cramer,

Lehre.

ROXBURGH, W. 1820. Ammannia. In W. Carey, Flora of India 1:

446-449. Mission Press, Serampore.

SAINT-HILAIRE, A.F.C.P. DE. 1833. Flora brasiliae meridionalis III:

135 (110), t. 187. Belin, Paris.

SONDER, O.W. 1850. Lythrarieae. Linnaea 23: 40-42.

SONDER, O.W. 1868. Lythrarieae. In W.H. Harvey & O.W. Sonder, Flora

capensis 2: 514-517. Reeve, London.

WEBB, D.A. & MOORE, D.M. 1968. Lythraceae. In T.G. Tutin et al. ,

Flora europaea 2: 300-303. University Press, Cambridge.

WILLDENOW, C.L. 1803. Hortus berolinensis 1: 6—7, tab. 6—7.

Schiippel, Berlin.

WOOD, J.M. & EVANS, M.S. ex WOOD, J.M. 1901a. Colonial

herbarium report for the year 1900: new Natal plants. Reports

of the Natal Botanic Gardens and Colonial Herbarium : 8—12.

WOOD, J.M. & EVANS, M.S. 1901b. New Natal plants. Journal of

Botany, British and Foreign 39: 169-172.

Bothalia 21,1: 51-54 (1991)

Studies in the genus Riccia (Marchantiales) from southern Africa. 22.

R. rubricollis , now validated, typified and described

S. M. PEROLD*

Keywords: Marchantiales, Riccia rubricollis, southern Africa, taxonomy, typification

ABSTRACT

Riccia rubricollis is a very rare endemic species, only known from a few collections by Duthie at Knysna. Unfortunately

she and Garside did not publish anything on it, and Arnell’s (1963) description has neither a Latin diagnosis nor was a

type specimen cited. These omissions have now been rectified here, following the recent rediscovery of Duthie’s collections

at BOL.

The species is characterised by conspicuous, mostly dark purple antheridial necks in the male plant and the spores have

large, deepset areolae on the distal face. Although the species has some features typical of section Ricciella of subgenus

Ricciella, it is not placed in this section because the sporangia do not bulge ventrally.

UITTREKSEL

Riccia rubricollis is n baie skaars endemiese spesie, slegs bekend van n paar Duthie-versamelings in Knysna. Ongeluk-

kig het sy en Garside nooit iets daaroor gepubliseer nie, en Arnell (1963) se beskrywing is sonder ’n Latynse diagnose,

en 'n tipe eksemplaar word ook nie aangehaal nie. Dit word nou hier reggestel. nadat Duthie se versamelings onlangs weer

by BOL opgespoor is.

Die spesie word gekenmerk deur opvallende donker pers anteridiele nekke in die manlike plant, en die spore het groot,

ingesinkte areolas op die distale kant. Alhoewel die spesie enkele kenmerke besit wat tipies is van seksie Ricciella van subge-

nus Ricciella, word dit nie onder hierdie seksie geplaas nie aangesien die sporangia nie ventraal uitbult nie.

Riccia rubricollis Garside <£ Duthie ex Perold

Thallus dioicus, ?perennis, in turmis gregariis, raniis

interdum superpositis, usque ad 12 mm longis, 2,8— 3,0

mm latis, oblongis vel linearibus, apicem versus

angustatis; stramineis, partis senioribus dorsaliter foveatis.

Contextus assimilans sectione ± 12 cavernulis trans latitu-

dinem thalli, lateraliter devexis apicemque versus dilatatis.

Squamae parvae, hyalinae, ventrales solum apicem versus.

Antheridia collis conspicuis, atropurpurea, in plantis

masculis in seria media secundum ramos; propter hoc

signum species facile distinguenda. Sporulae 92-105 pm

diametro, triangulo-globulares, porphyreae, vel (atro)brun-

neae, alatae; superficie distali areolis 5 vel 6 diametro

transversus, (10 — )20 — 25 /xm latis, parietibus ± 4 pm

crassis, 6 jxm altis; superficie proximali nota triradiali

distincta, areolis plerumque incompletis. Chromosoma-

tum numerus ignotus, quod plantae vivae non suppetentes.

Species rarissima, solum e loco vulgo dicto Knysna nota.

TYPE. — Cape, 3423 (Knysna): Knysna, Belvidere, on

turf in shady places, not far from lagoon (— AA),

Sept. /Oct. 1929, Duthie 5014 (BOL, lecto.; PRE, isolecto.)

Riccia rubricollis is known only from Knysna in the

southern Cape, where it was collected on a few occasions

by Duthie (Figure 1). It has so far not been found else-

where in southern Africa. It grows at low altitude, in damp,

shady places, with R. purpurascens Lehm. & Lindenb.,

Fossombronia sp. and Pleuridium sp. The rainfall is

600—800 mm per year and may occur all year round.

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

MS. received: 1990-07-02.

Thallus dioicous, ?perennial, in gregarious patches

(Figure 3A) sometimes with branches overlying, or

scattered, not in rosettes, yellowish green, occasionally

with some purple blotches; branches once (Figure 2A) or

twice symmetrically or asymmetrically furcate, rarely

simple, and then apically bilobed, moderately to widely

divergent, up to 12,0 mm long, segments 2,5— 6,0 x

2,8— 3,0 mm, 0,7— 1,0 mm thick medianly, thinner toward

winged margins, in section 2-4 times wider than thick;

oblong or somewhat linear as in species of section Ric-

ciella, tapered toward apex (Figure 3C); groove deep only

at apex, soon shallow and wide; margins subacute, rather

irregularly undulating, winged; flanks sloping obliquely

to very obliquely, yellowish; ventrally rounded or median-

ly keeled; when dry, margins not indexed, somewhat

FIGURE 1.— Distribution of R. rubricollis in southern Africa.

52

Bothalia 21,1 (1991)

FIGURE 2. — Riccia rubricollis. Morphology and anatomy. A, female

thallus; B, male thallus; C, epidermal cells and air pores (hatched),

overlying air chambers, as seen from above; D, air pore (hatched),

with surrounding cells; E, cross section of part of thallus show-

ing air chambers; F, cross section of female thallus. A-F, Dulhie

5014. Scale bars on A, B, F = 1 mm; C, E = 100 ,xm; D = 50

ftm.

raised, straw-coloured to light brown, dorsally pitted in

older parts.

Anatomy, dorsal epidermis forming slightly domed to

Hat covering over large, wide air chambers, from above

cells oblong-hexagonal or 5-sided, 62-80 x 50 /am, walls

up to 10 /am deep; air pores small toward apex (Figure

3D), only + 30 /am wide, surrounded by slightly curved,

narrow, elongated cells, soon disintegrating, proximally

air pores rapidly enlarging (Figure 2D), leaving air

chambers exposed; assimilation tissue 400—750 /am thick,

occupying more than '/2 to most of thickness of thallus,

air chambers 5- or 6-sided (Figure 2C), elongated, mostly

in one layer, ± 12 across width of thallus (Figure 2F),

sloping laterally and widening toward top, ± 200—250

pm wide, enclosed by unistratose, chlorophyllose cell

plates (Figure 2E); storage tissue + 300 /am thick, less

than '/2 the thickness of thallus, cells transparent,

angular, 30—70 /im wide, smaller below; rhizoids arising

medianly from ventral epidermal cells, mostly tuberculate,

17—22 wide. Scales small, hyaline, ventrally situated

toward apex only.

Antheridia in a row along midline of male thalli, when

mature bulging dorsally, with conspicuous, mostly dark

purple necks (Figure 2B), up to 300 X 25 /am. Archegonia

3 or 4 serially arranged toward base, necks purple, thin,

hidden. Sporangia up to 1100 /am wide, deeply imbedded,

not bulging dorsally or ventrally, but eventually opening

to upper surface, containing ± 620 spores each. Spores

92— 100(— 105) /am in diameter, triangular-globular, polar,

reddish brown to deep russet-brown, semitransparent to

opaque; wing 5 /am wide, wider and perforated at marginal

angles (Figure 4E), and often perforated elsewhere too,

margin finely crenulate, slightly undulating; ornamenta-

tion completely or incompletely reticulate, rather different

on 2 spore faces: distal face (Figure 4C, F) with 5 or 6

areolae across diameter, central ones larger, 20—25 ^m

wide, toward margin smaller, 10 pm wide, walls + 4 /im

thick and 6 pm deep, shallower laterally (Figure 4D), not

raised at nodes (Figure 4E); proximal face with triradiate

mark prominent (Figure 4A), widening toward marginal

angles at junction with wing (Figure 4B), areolae up to

12 pm wide, mostly incomplete, walls slightly raised at

nodes. Chromosome number: unknown, as living material

not available.

This species is characterized by ± linear, apically

tapering branches, which, in the male plants, bear a single

median row of conspicuous, deep purple antheridial necks.

Stolons are confined to the thickened perennating tips of

some branches (Figure 3B). Although more robust and

fleshy, R. rubricollis is clearly related to R. purpurascens

Lehm. & Lindenb., because of its somewhat linear habit,

but it is not classified together with it and R. stricta

(Lindenb.) Perold in section Ricciella (subgenus Ricciella ),

as the sporangia do not conspicuously bulge ventrally.

Na-Thalang (1980) regards the Australian species R.

collata , as having the closest affinity to R. rubricollis ,

although the latter plant is larger and on the distal spore

face, the areolae are wider.

The specific epithet, R. rubraosteolata, in Duthie’s

handwriting, appears on the labels of her collections, one

of which she kept in cultivation for some time at

Stellenbosch, but unfortunately she and Garside did not

publish a description of this rare, endemic species. AmeH’s

(1963) illustrations are reported to be after Garside, but

the originals have not been traced. Arnell eventually

described the species under the name, R. rubricollis , but

did not give a Latin diagnosis and cited no specimens,

Bothalia 21,1 (1991)

53

FIGURE 3. — Riccia rubricollis. Mor-

phology of the thallus. A,

crowded thalli; B, thallus with

apical stolon; C, thallus, show-

ing tapering apex and groove;

D, air pores. A-D, Duthie

5014 A, by A. Romanowski;

B-D, SEM micrographs by

S.M. Perold. Scale bars on

A— C = 1 mm; D = 50 /zm.

FIGURE 4. — Riccia rubricollis.

Spores. A, proximal face; B,

proximal face, side view; C, F,

distal face; D, distal face, side

view; E, much enlarged areo-

lae and perforations on distal

face. A— F, Duthie 5014. A-E,

SEM micrographs; F, LM pho-

tograph by S.M. Perold. Scale

bars on A-E = 50 p.m; di-

ameter of spore on F, + 100

/im.

omissions which render it invalidly published under

Articles 36.1 and 37.3 respectively, of the ICBN (Greuter

et al. 1988). Arnell must, however, have examined the

specimens, to have written the descriptions. Duthie’s

collections have only recently become available to me for

study, although I had previously specifically asked about

their whereabouts in a letter to the late Prof. E.A. Schelpe.

He had, however, assured me that there were no such

specimens held at BOL. In my study (Perold 1989) of the

spore wall ornamentation as an aid in identifying southern

African species of Riccia , I therefore mistakenly stated

that there are no examples of R. rubricollis.

The largest collection of this species, Duthie 5014 , con-

sists of two packets; one of them is selected here as the

lectotype and is held at BOL; the duplicate specimen, the

isolectotype, is held at PRE.

54

Bothalia 21,1 (1991)

SPECIMENS EXAMINED

CAPE. — 3423 (Knysna): Knysna, Belvidere, on turf in shady places,

not far from lagoon (— AA), Sept. /Oct. 1929, Duthie 5014 (BOL, lec-

to.; PRE, isolecto.); Knysna, Eastford Tower (— AA), Duthie 5406 (BOL).

ACKNOWLEDGEMENTS

The author wishes to express her sincere gratitude to

the Curator for the loan and to Mesdames Lorber and

Thompson, all of BOL, for finding and for sending the

specimens to PRE on loan. Dr H.F. Glen is deeply thanked

for the Latin diagnosis, and so are Mrs A. Romanowski for

the photographic work, Mrs J. Mulvenna for typing the

manuscript, and Ms J. Kimpton for the drawings.

REFERENCES

ARNELL, S. 1963. Hepaticae of South Africa, pp. 441. Swedish Natural

Science Council, Stockholm.

GREUTER, W. et al. 1988. International Code of Botanical Nomen-

clature. pp. 328. Koeltz Scientific Books, Konigstein.

NA-THALANG, O. 1980. A revision of the genus Riccia (Hepaticae)

in Australia. Brunonia 3: 61—140.

PEROLD, S.M. 1989. Spore wall ornamentation as an aid in identifying

the southern African species of Riccia (Hepaticae). Journal of

the Hattori Botanical Laboratory 67: 109—201.

Bothalia 21,1: 55-66 (1991)

Notes on African plants

VARIOUS AUTHORS

POACEAE

AIRA PRAECOX , A NEW RECORD FROM SOUTHERN AFRICA

Aira praecox L. (Aveneae: Pooideae) is a cosmopolitan

grass. Originally from Europe, where it is widespread,

it has become naturalized in N America, S America and

Australia (Hubbard 1984), with no records from sub-

saharan Africa till now (Gibbs Russell et al. 1985; Clayton

1970, 1972). The species has recently been found on Table

Mountain, Cape Peninsula, along the footpath from

Window Stream to Maclear’s Beacon, where it grows in

disturbed acidic sand, especially in shallow sand on rocks,

and flowers from September to November. This habitat

agrees closely with that described by Hubbard (1984).

There is one previous collection of the species, predictably

made by Ms Esterhuysen in 1977 from the same locality.

This specimen was, however, misidentified as Aira caryo-

phyllea L.

The genus Aira L. only includes annual grasses (Clayton

& Renvoize 1986), and its members can be recognized by

the two florets of which the upper is sessile or subsessile.

The rhachilla between the florets is absent or very short,

and indeed there is no rhachilla extension either. Aira

includes eight species, of which two or three are known

from southern Africa. This depends on whether A.

cupaniana Guss. and A. caryophylla L. are both recog-

nized locally (Sokutu in Gibbs Russell et al. 1990).

However, A. praecox is readily distinguished from both

these species by its compact, spikelike inflorescences.

Aira praecox L.

Plants delicate, annual; stems solitary to several in a tine

cluster, to 180 mm tall. Nodes three, swollen, nitid, the

culms directly below the nodes coarsely deflexed-scabrid.

Leaves two per culm; sheaths 15—30 mm long, smooth

to finely scaberulous; ligules membranous, to 3 mm long,

the apex lacerated; blades to 30 x 1 mm, V-shaped, mar-

gins scabrid, the apex rounded. Inflorescence a spike-like

panicle, pale green, 10—25 x 3—5 mm, the rhachis

generally obscured by the spikelets; nodes pale, nitid;

rhachis and pedicels scaberulous; pedicels shorter than

the spikelets. Spikelets 3 mm long, with two flowers

inserted practically at the same level, i.e. there is no

visible rhachis and no rhachis extension. Glumes 3 mm

long, 1-veined, taller than the florets, acute, scaberulous

in the apical third. Lemmas 5-veined, 2,5 mm long, shortly

bilobed, rounded on the backs, the lower half smooth

and the upper half scaberulous, with a small tuft of

hairs at the base; awned from '/4 above the base, awns

3,5— 3,7 mm long, with a short column. Palea 2 mm

long, bikeeled, bilobed. Lodicules two, fleshy, ovate with

a small basal tooth. Anthers yellow, 0,2 mm long. Ovary

glabrous.

Specimens examined

CAPE — 3318 Cape Town: Table Mountain, along path from Win-

dow Stream to Maclear’s Beacon (-CD), Esterhuysen 34816 (BOL);

Under 5045 (BOL, PRE).

REFERENCES

CLAYTON, W.D. 1970. Gramineae (Part 1). In E. Milne-Redhead &

R.M. Polhill, Flora of tropical East Africa: 1—177. Crown Agents

for Overseas Governments and Administrations, London.

CLAYTON, W.D. 1972. Gramineae. In F.N. Hepper, Flora of West

tropical Africa: 349-574. Crown Agents for Overseas Govern-

ments and Administrations, London.

CLAYTON, W.D. & RENVOIZE, S.A. 1986. Genera graminum — grass-

es of the World. Her Majesty’s Stationary Office, London.

GIBBS RUSSELL, G.E., REID, C., VAN ROOY, J. & SMOOK, L.

1985. List of species of southern African plants, edn 2, part 1.

Memoirs of the Botanical Survey of South Africa No. 51: 1—152.

GIBBS RUSSELL, G.E., WATSON, L.', KOEKEMOER, M., SMOOK,

L., BARKER, N.P. ANDERSON, H.M. & DALLWITZ, M.J.

1990. Grasses of southern Africa. Memoirs of the Botanical Survey

of South Africa No. 58.

HUBBARD, C.E. 1984. Grasses: a guide to their structure, identification,

uses and distribution in the British Isles, 3rd edn. Penguin Books,

England.

H P LINDER

* Bolus Herbarium, University of Cape Town, Rondebosch 7700.

MS. received: 1990-03-28.

ZYGOPHYLLACEAE

A NEW SPECIES OF ZYGOPHYLLUM FROM THE CAPE PROVINCE

Zygophyllum chrysopteron Retief sp. nov., Z. retrofrac-

tum Thunb. affinis, sed fructibus 5-alatis non fusiformibus;

ramis minus implexis; distributione dissimili differt.

TYPE.— Cape Province, 3021 (Vanwyksvlei): 1 km out

of Vanwyksvlei, roadside (— BD), Smook & Harding 782

(PRE, holo.) Figure 1.

56

Bothalia 21,1 (1991)

KSS HERBARIUM PRETORIA

3o;’i hd ~ iFr.M ■ ^apc

Zjrgopftyllum retrofractum tfionb,

woody, fruits tinned pink

JL

FIGURE 1. — Holotype of Zygophyllum chrysopteron (Smook & Harding

782).

A much-branched woody shrub up to 1 m high; branch-

lets ascending, reflexed or secund, striate, internodes short,

0,5— 3,0 mm. Leaves bipinnate; lamina broadly obovate

to orbicular, 1,5— 2,0 X 1,5— 2,0 mm, sometimes very

shortly petiolate with only the two leaflets visible against

the stems. Stipules ovate, very small. Flowers solitary in

the axils of the leaves; 5-partite; pedicels 1 mm long. Calyx

with lobes adnate at the base; lobes oblong, 2,5 x 1,0 mm.

Corolla with petals spathulate, 3 mm long, white. Stamens

FIGURE 2. — Distribution of Zygophyllum chrysopteron, #; Z. retrofrac-

tum, O.

10, adnate to a disc; filaments 2,5 mm long, appendages

deeply divided, margins entire; anthers 1 mm long. Style

1,5 mm long. Ovary glabrous. Fruit a 5-winged capsule,

subglobose in outline, 3—4 x 2—3 mm. Seeds trigonous

to oblong, 0,5— 1,5 mm long.

CAPE PROVINCE. — 2920 (Boomrivier): 5 miles [8 km] W of

Loogkolk (—AC), Pole Evans 2244 (PRE). 2921 (Kenhardt): Jagbult,

40 miles [64 km] W of Marydale (—DA), Story 1108 (PRE). 2922

(Prieska): Prieska (-DA), Bryant, J. 293 (PRE); Karabee (-DB), Roux

131 (PRE). 3020 (Brandvlei): Rietkolk (-BC), Le Roux & Lloyd 230

(PRE); Kroon (— CB), Dean 645 (PRE). 3021 (Vanwyksvlei): 1 km from

Vanwyksvlei (— BD), Smook & Harding 782 (PRE). 3022 (Carnarvon):

Rhenosterkolk (— CA), Acocks 1719 (PRE). 3120 (Williston): Williston

Station (— BD), Smith 2442 (PRE). 3221 (Merweville): Layton (— BB),

Shearing 1379 (PRE). 3222 (Beaufort West); Nieuweveld escarpment

(—BA), Adamson D155 (PRE); Aardoorns (— DD), Retief & Reid 92

(PRE). 3223 (Rietbron): Rooidam (— CA), Retief & Reid 45 (PRE);

Kalkdam (—CD), Retief & Reid 349 (PRE); 1 m [1,6 km) S of Kiwietskuil

(-DA), Acocks 14657 (PRE). 3320 (Montagu): Touws River (—AC),

Van Breda 206 (PRE); 17 km N of Matjiesfontein (—BA), Van Wyk 524

(PRE). 3322 (Oudtshoorn): 5 km from the Beaufort West/Laingsburg

road on the road to Oudtshoorn (— BC), Van Wyk 1850 (PRE).

Zygophyllum chrysopteron is endemic in the Cape

Province (Figure 2). It occurs in various veld types of the

Karoo. The veld types are characterised by dwarf trees

and shrubs, grass and succulents. The rainfall is very low

in these areas, up to 200 mm a year. Z. chrysopteron can

be found growing on red or grey sandy soil; on stony,

gravelly soil; limestone and shaly slopes. The species

flowers in October and November and bears fruit till July.

Z. chrysopteron is often heavily grazed.

In the past, dwarf shrubs, much-branched with re-

curved, tangled and secund branches have been identified

as Zygophyllum retrofractum Thunb. A study of the species

revealed that two different fruit types occur within

specimens placed under this name, known as Z. retrofrac-

tum. In February 1937, Acocks however, collected two

specimens at Rhenosterkolk, Carnarvon District, and

noted on one of the labels that his numbers 1719 & 1731

did not represent the same entity, which is indeed the case.

The winged fruits of Z. chrysopteron (Figure 3), together

with a more erect habit and a different distribution pattern

(Figure 2) separate this species from its nearest relative,

Z. retrofractum. This latter species has spindle-shaped

fruits with ridges (Figure 3), a conspicuously spreading

habit and occurs from Namibia through Namaqualand as

far south as Barrydale (Figure 2). In the Williston,

Carnarvon and Prieska areas, both species are found

FIGURE 3. —Fruit of Zygophyllum: A, retrofractum , X 8; B, chrysopte-

ron, X 8.

Bothalia 21,1 (1991)

57

growing in the same region, as Aeocks’s specimens also

revealed.

Besides Z. chrysopteron, a number of other species of

Zygophyllum (e.g. Z. morgsana , Z. sujfruticosum and Z.

microcarpum) have winged fruits. The fruits of these

species are larger in size and the structure of their leaves

is not the same as those of Z. chrysopteron , which are

bipinnate with broadly obovate to orbicular blades, very

small in size.

The specific epithet chrysopteron refers to the golden

yellow colour of the fruits. According to the label of Smook

& Harding 782, the fruits can be tinged with pink.

E. RETIEF

MS. received: 1990-07-30.

VERBENACEAE

STACHYTARPHETA SPECIES IN SOUTHERN AFRICA

The genus Stachytarpheta , comprising some 70-100

species, is found mainly in tropical America. Some species

have become widely dispersed as weeds in the tropics and

a few are cultivated as ornamentals. Up till now, only two

species have been recorded for the FSA region and both

were presumably introduced. They have been variously,

and often incorrectly referred to as S. indica , S. jamai-

censis or S. urticifolia (Ross 1972; Wells et al. 1986; Wel-

man in Gibbs Russell et al. 1987). Verdcourt is currently

revising the genus for the FTEA (in press). Using an adapt-

ed version of his key, the species found in the FSA region

can be distinguished as follows;

Corolla-limb 10-20 mm wide; inflorescence-rachis coarse,

over 5 mm wide when dry (up to 10 mm wide after

flowering); leaves slightly discolorous, densely velvety

pubescent on lower surface 5. mutabilis

Corolla-limb usually under 10 mm wide; inflorescence-rachis

more slender, about 3 mm wide (up to 7 mm wide

after flowering); leaves glabrous S. urticifolia

S. mutabilis (Jacq.) Vahl is cultivated and occurs as an

escape from cultivation. It has so far only been recorded

from the northern and eastern Transvaal.

TRANSVAAL.— 2330 (Tzaneen): Westfalia (-CA), Van Wyk 4301

(PRE). 2430 (Pilgrim’s Rest): Lekgalameetse Nature Reserve (-AB),

Stalrnans 754 (PRE). 2530 (Lydenburg): Nelspruit, Westacres suburb

(-BD), Clarke s.n. PRE 733686 (PRE). 2531 (Komatipoort): Witrivier

(-AC), Stirton 7277 (PRE).

S. urticifolia (Salisb.) Sims occurs as a naturalized weed

in the eastern Transvaal and Natal. This taxon was up till

now referred to as S. indica (L.) Vahl (Ross 1972; 300;

Welman 1987; 169) but that species does not seem to occur

as far south.

TRANSVAAL. — 2530 (Lydenburg): Nelspruit, Westacres suburb

(— BD), Clarke s.n. (PRE); Mataffin (— BD), Durant LN 1180 (PRE).

NATAL. — 2931 (Stanger): lower Tugela District, Groutville (—AD),

Acocks 11823 (PRE). 3030 (Port Shepstone): Southbroom (— CB), Marais

1149.

REFERENCES

ROSS, J.H. 1972. Flora of Natal. Memoirs of the Botanical Survey of

South Africa No. 39.

VERDCOURT, B. in press. Flora of tropical East Africa.

WELLS, M.J., BALSINHAS, A. A., JOFFE, H.. ENGELBRECHT,

V.M., HARDING, G. & STIRTON, C.H. 1986. A catalogue of

problem plants in southern Africa. Memoirs of the Botanical

Survey of South Africa No. 53.

WELMAN, W.G. 1987. Verbenaceae. In G.E. Gibbs Russell et al..

Memoirs of the Botanical Survey of South Africa No. 56, edn 2,

part 2.

P.P.J. HERMAN

MS. received: 1990-08-27.

SELAGINACEAE

M1CRODON BRACTEATUS— THE CORRECT NAME FOR M. LUC1DUS

Thunberg (1794) named one of his collections from

southern Africa Selago bracteata Thunb. Ventenat (1804)

described and illustrated other material of the same species

and named it Selago lucidus Vent. When Choisy (1823)

described his new genus Microdon Choisy, he recognized

that the plant that Ventenat had drawn should be included

in Microdon and therefore made the combination Microdon

lucidus (Vent.) Choisy. Taxonomists, such as Walpers

(1844) and Rolfe (1884), followed Choisy and retained his

combination. The earliest epithet, however, is that of Thun-

berg and thus the correct combination is made below.

Microdon bracteatus (Thunb.) l.H. Hartley comb,

nov. Type: Cape Province, without precise locality, Thun-

berg s.n. (microfiche IDC 13904).

Selago bracteata Thunb.: 100 (1794); Thunb.: 465 (1823).

Selago lucidus Vent.: t. 26 (1804). Microdon lucidus (Vent.) Choisy:

97 (1823); E. Mey.: 253 (1835); Drege: 76, 77, 202 (1843); Choisy: 22

(1848); Walpers: 168 (1844); Rolfe: 354 (1884); Rolfe: 175 (1901). Icono-

type: Ventenat, Jardin de la Malmaison: t. 26 (1804).

REFERENCES

CHOISY, J.D. 1823. Memoire sur lafamille de Selaginees, lu a la Societe

de Physique et d'Histoire naturelle de Geneve le janvier 1822.

Paschoud, Geneve.

CHOISY, J.D. 1848. Selaginaceae. In A.P. De Candolle, Prodromus

systematis naturalis regni vegetabilis 12: 1—26. Victoris Masson,

Paris.

DREGE, J.F. 1843. Zwei pflanzengeographische Documente. Leipzig.

MEYER, E.H.F. 1835. Commentariorum de plantis africae australioris.

Leopoldum Voss, Leipzig.

58

Bothalia 21,1 (1991)

FIGURE 4 .—Amphithalea rostrata (R1-RI0) and A. tomentosa (Tl— T9) showing the differences between the two species: R1 & Tl, flowers in

lateral view (note the cucullate standard petal and squarrose wing petal in A. rostrata)-, R2 & T2, leaves in abaxial view (note the revolute

margin and pilose vestiture in A. rostrata)-, R3 & T3, calyces opened out with the upper lobes to the left, vestiture not shown; R4 & T4,.

standard petals; R5 & T5, wing petals; R6 & T6, keel petals; R7 & T7, stamens; R8 & T8, pistils; R9 & T9, bracts; RIO, flowering branch.

Rl— RIO from Oliver 8752 in STE; Tl— T9 from Schutte 327 in JRAU; scale in mm.

Bothalia 21,1 (1991)

59

ROLFE, R.A. 1884. On the Selagineae described by Linnaeus, Bergius,

Linnaeus fil. and Thunberg. Journal of the Linnean Society

(Botany) 20: 338-358.

ROLFE, R.A. 1901. Selagineae. In W.T. Thiselton-Dyer, Flora capensis

5,1: 175-177. Lovell Reeve, London.

THUNBERG, C.P. 1794. Prodromus plantarum capensium. Edman,

Uppsala.

THUNBERG, C.R 1823. Flora capensis, edn Schultes. J.G. Cottae,

Stuttgardt.

VENTENAT, E.P. 1804. Jardin de la Malmaison 1,5. Du Limprimerie

de Crapelet, Paris.

WALPERS, W.G. 1844. Repertorium botanices systemalicae 3,4.

Hinrichs, Leipzig.

I.H. HARTLEY*

* C.E. Moss Herbarium, Botany Department, University of the

Witwatersrand. P.O. Wits 2050.

MS. received: 1990-07-09.

FABACEAE

A NEW SPECIES OF AMPHITHALEA (LIPARIEAE)

In a recent revision of the genus Amphithalea Eckl. &

Zeyh., Granby (1985) recognized twenty species and four

subspecies. A distinct new species has been discovered

since.

Amphithalea rostrata A.L. Schutte & B-E. van Wyk,

species insignis carina rostrata, vexillo cucullato, aliis

squarrosis, flore vinoso et foliis revolutis pilosis, a

speciebus nobis notis bene distincta.

TYPE. — Cape, 3419 (Caledon): Carruthers Hill, SEof

Pearly Beach (—DA), 10.06.1985, Oliver 8752 (STE, holo.;

PRE, iso.).

A diffuse shrublet, up to 0,6 m tall. Branches sparsely

leafy with scars left from fallen leaves, ± glabrous; young

twigs thin and slender, densely leafy, pilose. Leaves

alternate, simple, ovate to broadly ovate with strongly

revolute margins; blade (4,5 — )7,0— 10,0 x (2,0— )3,5— 5,0

mm; petiole very short, less than 0,5 mm long; apex acute;

base rounded or somewhat cordate; densely pilose

abaxially, sparsely so adaxially. Stipules absent. Inflores-

cences single-flowered, axillary, condensed to 1— 2(— 4)-

flowered terminal nodding capitula. Bracts linear, 2,5 -3,0

mm long. Bracteoles absent. Flowers subsessile, 7—8 mm

long, dark maroon-red. Calyx 5,5 mm long, purplish, with

the upper two lobes fused much higher up; lateral and

lower sinuses nearly of equal depth, the lateral ones

slightly deeper; lobes triangular, acuminate to somewhat

caudate; densely pilose. Standard 8,0— 8,5 x 4,0— 4,5 mm,

ovate with the upper part cucullate; apex mucronate; base

cordate; claw very short, less than 0,5 mm long. Wing

petals shorter than the keel, 6,0— 6,5 mm long, oblong to

narrowly oblong, auriculate, distinctly pocketed, apical

part of the lamina curved downwards; sculpturing upper

basal, 3 rows of transcostal lamellae; claw 1,5— 2,0 mm

long. Keel petals 7,0 -7,5 mm long, semicircular, with the

distal part hidden in the upper part of the standard,

auriculate, pocketed; apex distinctly rostrate; claw 1,0— 1,5

mm long. Stamens 8,0— 8,5 mm long; diadelphous,

vexillary stamen free, the others fused for ± half the

length; anthers uniform in size. Pistil 5,5 mm long, sessile;

style strongly curved upwards, glabrous; ovary with a

single ovule, densely sericeous. Fruit and seed unknown.

Figure 4.

The flower structure of A. rostrata is comparable only

to that of A. tomentosa. These two species are the only

ones with a rostrate keel, a mucronate standard and oblong

wing petals. A. rostrata differs in the strongly revolute leaf

margins (leaves flat in A. tomentosa), the pilose vestiture

(silky-canescent in A. tomentosa), the maroon-coloured

flowers (yellow in A. tomentosa), the cordate standard

(oblong in A. tomentosa) and the absence of protuberances

at the apices of the keel and wing petals (see Figure 4).

Unlike all other species of Amphithalea, the upper part

of the standard petal is cucullate (not recurved or straight)

and the wing petals are distinctly squarrose (not straight)

and shorter than the keel (not as long or longer than the

keel).

A. rostrata was discovered by Mr E.G.H. Oliver of the

Stellenbosch Herbarium. We thank him for directing us

to the only known locality, which is at Carruthers Hill,

south-east of Pearly Beach (Figure 5). The population

occurs amongst fynbos on a sandy and stony south-facing

slope, at 120 m above sea level.

FIGURE 5. — The known geographical distribution of Amphithalea

rostrata.

CAPE.— 3419 (Caledon): Carruthers Hill, SE of Pearly Beach (—DA),

10.06.1985, Oliver 8752 (STE. holo.; PRE, iso.), 13.11.1989, Van Wyk2896

(JRAU).

REFERENCE

GRANBY, R. 1985. Revision of the genus Amphithalea (Liparieae—

Fabaceae). Opera Botanica 80: 1—34.

A.L. SCHUTTE* and B-E. VAN WYK*

* Botany Department, Rand Afrikaans University, P.O. Box 524,

Johannesburg 2000.

MS. received: 1990-07-06.

60

Bothalia 21,1 (1991)

ROSACEA E

A NEW SPECIES OF CLIFFORTIA FROM THE SOUTH-WESTERN CAPE

Cliffortia burgersii Oliver & Fellingham, sp. nov. ,

sectione Alatae, C. alatae N.E. Br. affinis, sed differt

fructu grandiori roseo cum alis 3-4, staminibus

10— 11(— 13), foliis parvis flavovirentibus muticis glabris,

habitu frutescenti ramoso compacto. Crescit in planitie

arenosa calcarea dehoopensi pone mare.

Frutex ramosissimiis ad 1,5 m altus. Rami indumento

crispo albo, glabrescentes brunnescentes cortice irregu-

lariter fidenti; internodia ± 10 mm longa. Folia trifoliolata;

vagina 0,8 mm longa, cristis longitudinalibus tribus;

stipulae deltoideae, 0,3 mm longae; petiolus 0,3— 0,5 mm

longus; folioli recti ad falcati, 3,0— 7,0 x 0,4— 0,5 mm,

ericoidei sulcati glabri, obtusi ad acuti. Flores masculi :

bracteolae ovatae, 1,3 mm longae, scariosae ciliatae

puberulae; pedicellus receptaculumque 1,5 mm longus

glaber; sepala 3—4, elliptica acuta, 3,0— 3,5 x 2,1— 2,2

mm, viridescentes, caespite subapicali pilorum crisporum

adaxiali; stamina 10 — 11( — 13); filamenta 2,0— 3,0 mm longa

glabra; antherae 0,6— 0,8 mm longae. Flores feminei :

bracteolae rhomboideo-ovatae, 1,2 mm longae, ciliatae,

interne villosae, externe puberulae; pedicellus 1,5 mm

longus in frutice ad 3,7 mm, puberulus pilis retrorsis;

sepala 3—4, late elliptica acuta mucronata, 2,0 x 1,3 mm,

glabra sed pilosa basi abaxiali caespite subapicali papil-

larum; ovanwmapicebasiqueemarginatum, 1,7 x 1,8 mm,

villosum pilis retrorsis; stylus solitarius, 0,2— 0,5 mm

longus, viridescens glaber; stigma expansum fimbriatum

atrorubens 1,7 mm latum. Fructus a latere visus orbicu-

laris, (8)9 X 10(11) mm, alis prominentibus, apice basique

emarginatus, parte centrali ellipsoidea viridescenti lanugi-

nosa, spinis parvis vel cristis pectinatis; alae (2)3-4, 3(4)

mm latae, rosae glabrae vel sparse pilosae pilis crispis.

TYPE. — Cape, 3420 (Bredasdorp): De Hoop Reserve,

Hats NE of the Reserve Centre, 30 m, 7 June 1989, Oliver

& Fellingham 9148a (female) (STE, holo. ; BOL, K, MO,

PRE).

A much-branched erect shrub to 1,5 m tall, mostly

shorter and compact through grazing, dioecious but with

an occasional fertile female flower on a male plant.

Branches greenish yellow with a white crisped indumen-

tum when young, becoming glabrous and brown with age,

the bark then dull grey and splitting irregularly; intemodes

about 10 mm long. Leaves trifoliolate; vagina 0,8 mm long

with 3 longitudinal ridges; stipules deltoid, 0,3 mm long;

petiole 0,3— 0,5 mm long; leaflets 3—7 x 0,4— 0,5 mm,

straight to falcate, ericoid, glabrous, obtuse to acute some-

times with a pale sharp apex, sulcate below with the groove

hairy. Male flowers : bracteoles ovate, 1,3 mm long,

scarious, ciliate, puberulous; pedicel and receptacle 1,5

mm long, glabrous but for a ring of hairs below the sepals;

sepals 3—4, 3,0— 3,5 x 2,0— 2,2 mm, greenish, some with

an occasional maroon-red longitudinal line, elliptic, acute,

with a subapical tuft of crisped hairs adaxially; stamens

10 — 11( — 13), filaments 2, 0-3,0 mm long, maroon-red,

glabrous; anthers 0,6— 0,8 mm long, maroon-red with a

white connective; an occasional male flower with a rudi-

mentary ovary and well developed style and stigma. Female

flowers', bracteoles rhomboid-ovate, 1,2 mm long.

ciliate, villous inside, puberulous outside, the lower half

scarious soon turning brown; pedicel 1,5 mm long

lengthening rapidly to 3,7 mm in the fruit, puberulous with

retrorse hairs; sepals 3-4, broadly elliptic, 2,0 x 1,3 mm,

acute, mucronate, glabrous but pilose near the base

abaxially and with a subapical tuft of long papillae soon

shrivelling, sepals persisting in the fruit; ovary emarginate

at both ends, 1,7 x 1,8 mm, villous with retrorse hairs

longer near the base, 3 -4-winged; style solitary, 0,2 -0,5

mm long, green, glabrous, sunken into a tube formed by

a ring of short red-brown staminodes; stigma formed by

the style expanding into a broad, deep red fimbriate

pompom-like structure 1,7 mm across. Fruit: (8)9 x 10(11)

mm, circular in outline, emarginate at the apex and base,

prominently winged; the central portion ellipsoid, green,

white woolly, equipped with irregular, hard, horny, deep

red short spines and/or a narrow sturdy to more

wing-like comb in place of the fourth wing (see Figure

6J); wings (2)3-4, somewhat crescent-shaped, 3(4) mm

broad, reddish pink, undulate or flat, entire, glabrous or

with a few short crisped hairs, and with numerous fine

radiating ridges. Figure 6.

Diagnostic features

C. burgersii (Figure 6A— J) is similar to C. alata (Figure

7A — E), but differs in the more branched compact growth

form, the smaller yellow-green blunt glabrous leaves

(bluish grey-green, distinctly red mucronate and crisped

puberulous all over in C. alata)\ the larger number of

stamens, 10 — 11( — 13) (6-8 in C. alata)', the size of the

fruit, (8)9 x 10(11) mm [(5)6 X (6)7 mm in C. alata ];

the colour of the fruit, green and reddish pink (red all over

in C. alata) and the number of wings in the fruit, 3—4

(2—3 in C. alata).

Distribution and habitat

C. burgersii is known only from the De Hoop Nature

Reserve on open flats at almost sea level where it grows

in Limestone Fynbos on sand over limestone sheets (Figure

8). This is in striking contrast to the distribution and habitat

of C. alata. Originally C. alata was recorded only from

Muiskraal on the northern slopes of the Langeberg at 420

m. Two more recent collections, Kruger 1345 and Van der

Merwe 293, have extended the distribution further west

along the Langeberg to the northern slopes in the Groot-

vadersbosch State Forest area, but at 620 m. The habitat

at Muiskraal is mainly Dry Mountain Fynbos on sand

derived from sandstone of the Nardouw Subgroup, but also

Central Mountain Renosterveld on clays. At Grootvaders-

bosch it is probably a drier form of Mesic Mountain

Fynbos, or perhaps also Dry Mountain Fynbos, again on

the Nardouw Formation.

In situ, C. burgersii is fairly conspicuous due to the very

noticeable reddish pink fruits which cover the ultimate

branches of the plants. These fruits remain on the plant

for several months during the winter period gradually

drying and turning brown by summer.

Bothalia 21,1 (1991)

61

FIGURE 6. — Cliffortia burgersii : A, branch, natural size: B, leaf, X 5: C, apex of leaflet, x 20; D, female flower in side view, x 5; E, male

flower from above, x 5; F. fruit in side view, note persistent sepals, x 5; G-J, fruits as seen from above, variations in fruit wings, x

5. All drawn from Oliver & Fellingham 9148a & b (STE).

Discussion

In C. burgersii five different patterns can be seen in the

arrangement of the fruit wings. Two opposite wings

interspersed by two opposite combs occur very rarely. A

Y-shaped arrangement of three wings is more frequent

(Figure 6G). Equally frequent is the arrangement of three

wings, where two are opposite and the third at right angles

to these (Figure 6H). Four equally spaced wings occur

more commonly (Figure 61). Most commonly the fourth

wing is replaced by a comb (Figure 6J). In C. alata the

two-winged form is the most prevalent, the three-winged

form occurring only occasionally (Figure 7D & E). Other

winged species in the Section Alatae are C. teretifolia L.f.

and C. semiteres Weim. These species have small, incon-

spicuous, narrowly winged, dull coloured fruits quite

unlike those of C. alata and C. burgersii. The only other

species with narrow straight wings is C. phyllanthoides

Schltr. in the Section Bifoliolae. Other winged species

occur in Section Inflexae , but there the wings are all curved

and the fruits small and hidden in contrast to the showy

exposed fruits of C. alata and C. burgersii.

This distinct new species is named after Chris Burgers

of the Cape Department of Nature and Environmental

Conservation who is making a special study of the flora

of the De Hoop area and of rare and endangered species

in general.

FIGURE 7. — Cliffortia alata: A, leaf, X 5; B, apex of leaflet, x 20; C, fruit, x 5; D & E, fruits as seen from above, variations in fruit wings,

x 5. All drawn from Oliver & Fellingham 9143 (STE).

Specimens examined

C. burgersii sp. nov.

CAPE.— 3420 (Bredasdorp): De Hoop Nature Reserve; De Hoop flats,

along public road, 28-7-1979, (-AD), Burgers 2061 (PRE, STE); along

public road to Potberg, on De Hoop, 30-1-1985, (-AD), Fellingham 926

(NBG, PRE, STE); 1 km from gate of Nature Reserve on public road

towards Potberg, 20-3-1985, (—AD), Fellingham 984 (BOL, K, PRE,

STE); flats next to old public road about 1 km before turn-off to the

Opstal, 11-3-1989, (—AD), Fellingham 1444 (BOL, K, MO, PRE, STE);

Oats NE of the Reserve Centre, female plants, 7-6-1989, (—AD), Oliver

& Fellingham 9184a (BOL, K, MO, PRE, STE); ibid., male plants, Oliver

& Fellingham 9184b (BOL, K, MO, PRE, STE).

FIGURE 8. — Known distribution of Cliffortia burgersii, •; and C.

alata, ▲ .

C. alata N.E. Br. in Kew Bulletin 15: 121 (1901);

Weim.:80(1934). Type:Cape Colony, Riversdale Division;

on the Karoo plains at Muis Kraal, near Garcia’s Pass,

alt. 1000 ft, Galpin 3925 (BOL!, K!, male & female

marked as type by N.E. Brown).

CAPE. — 3320 (Montagu): Langeberg at Moeras rest huts, Grootvaders-

bosch State Forest, 19-6-1971, (— DD), Kruger 1345 (STE); Grootvaders-

bosch State Forest, near Doom River huts, 630 m, 29-1-1989, (— DD),

Van tier Merwe 293 (STE). 3321 (Ladismith): Muiskraal, near foothills

in dip next to dam, NW aspect, 370 m, 10-8-1983, (— CC), Bohnen 8238

(PRE, STE); Muiskraal, 10-1904, (-CC), Bolus 11269 (BOL, K, STE);

3-10-1897, Galpin 3925 (BOL, K); 7-1927, Levyns 2118 (BOL); 11-8-1948,

Levyns 9008 (BOL); 15-9-1981, Fellingham 110 (K, PRE, STE);

Muiskraal, S of the farmstead, 420 m, 7-6-1989, (— CC), Oliver &

Fellingham 9143 (K, MO, PRE, STE); Garcia’s Pass, near the top,

13-10-1981, (-CC), Mauve & Hugo 221 (K, PRE, STE); 20-7-1937, Salter

6772 (BOL, K); 14-9-1937, Salter 6904 (BOL, K).

REFERENCES

BROWN, N.E. 1901. Cliffortia alata. Kew Bulletin 15: 121.

WEIMARCK, H. 1934. Monograph of the genus Cliffortia. Gleerups-

ka Univ. Bokhandeln, Lund.

E.G.H. OLIVER* and A.C. FELLINGHAM*

* Stellenbosch Herbarium, National Botanical Institute, P.O. Box 471,

Stellenbosch 7600.

MS. received: 1990-07-05.

CUPRESSACEAE

TETRACUN1S ARTICULATA, A HITHERTO UNRECORDED NATURALISED ALIEN CONIFER IN SOUTH AFRICA

INTRODUCTION

Tetraclinis articulata, a monotypic Mediterranean genus

of the Cupressaceae allied to Widdringtonia and Callitris ,

is a native of Morocco, Algeria and Malta with a small

outlying population near Cartagena in south-east Spain

(Dallimore & Jackson 1974; Kriissman 1972). While

inspecting various exotic trees in the historic naval

cemetery at Simonstown with the late Prof. E.A. Schelpe

in 1984, several randomly planted mature specimens as

well as naturalised saplings were observed which were

later positively identified by us as Tetraclinis articulata.

Subsequently, two additional groups were found further

up Simonstown Mountain flanking Runciman Drive, above

the cemetery. The latter groups consisted of approximately

18 mature specimens planted at regular intervals in rows

and measuring between 10 and 12 metres in height with

the trunk diameter at chest height ranging from 250 to 400

mm. (Figure 9). Moreover, saplings as well as young

seedlings in various stages of development were also noted

on adjacent undeveloped plots and on the surrounding

mountainside above Simonstown, competing with other

alien trees such as various Eucalyptus and Acacia species.

It appears that Tetraclinis is in the process of becoming

naturalised locally in the environs of Simonstown,

especially in disturbed areas, although at this stage it shows

no indication of becoming a rampant arborescent pest

plant in fynbos. In late summer (February, March) the

cones open spontaneously releasing their seeds, which

have two laterally placed wings to assist in their dispersal

(Figure 10). Under windy conditions Tetraclinis seeds are

probably distributed several hundred metres from the

parent .plants. When felled or injured the trees coppice

extensively.

Bothalia 21,1 (1991)

63

FIGURE 9.— A, mature group of

Tetraclinis articulata on

Runciman Drive, Simonstown;

B, naturalised T. articulata

seedling on Simonstown

Mountain in association with

Elytropappus rhinocerotis and

Acacia cyclops.

ESTABLISHMENT AND DATE OF INTRODUCTION TO

SOUTH AFRICA

With the exception of a single record of a specimen

cultivated at The Wilds, Johannesburg in the 1950’s, the

presence of Tetraclinis articulata has not been noted in

South Africa as an ornamental, a commercial forest

species, or as an alien adventive either in local herbaria,

or in publications listing conifers cultivated in this country

(Poynton 1984; Poynton pers. comm.; Wells et al. 1986).

The Simonstown plantings and adjacent naturalised

occurrences of this species have apparently hitherto been

overlooked and may well be the only major occurrence

of the species in our region. Since the Royal Navy formerly

maintained important naval bases at Simonstown and on

Malta, and as Tetraclinis is indigenous to Malta, it seems

plausible to assume that the Simonstown plantings were

introduced either as seed or seedlings from Malta by Royal

Navy personnel at some stage in the past. All the mature

Tetraclinis trees at Simonstown are of comparable age.

Judging from their size they appear to be fully mature.

A photograph in the Simonstown Museum depicts a formal

Empire Day (May 24th) tree-planting ceremony at which

the Tetraclinis trees on Runciman Drive were planted

out as half metre seedlings from paraffin-tin containers.

Museum staff have dated this photograph as having been

taken between 1904 and 1910. Additional confirmation of

the date of this event appears in a brochure entitled

‘ Souvenier of Simonstown 1910' (Anon. 1910) which also

records that the town council had spent £795 over the previ-

ous five years (i.e. 1905—1910) planting trees in the Simon-

stown Municipal area. How and why the seed or seedlings

of T. articulata were brought to South Africa is unclear

but it appears to have been a single isolated

B

FIGURE 10. — A, dehisced female

cones of Tetraclinis articulata,

dorsal & ventral views; B,

seeds showing prominent later-

al wings; C, fruiting branch

bearing immature female

cones. Each scale division =

2 mm.

64

Bothalia 21,1 (1991)

event instigated by an enthusiastic plant introducer. These

trees have evidently been producing both male and female

cones for several decades as indicated by the size (3—4

m tall) of the largest self-sown saplings.

NATURAL OCCURRENCE AND USES

Extensive forests of Tetraclinis cover some 350 000 ha

in Morocco, where this species is an important source

of a dark, fragrant timber (Howes 1949). It is among

the ecologically dominant tree species of Mediterranean

woodland in North Africa. White (1983) recognises

Tetraclinis articulata forest as a distinct plant community

within the broader classification of Mediterranean sclero-

phyllous forest occurring from southern Morocco to

Tunisia in the oceanic and maritime semi-arid zones

(rainfall 500—700 mm per year), between sea level

and 1 500 metres, on both calcareous and siliceous

soils. Well-grown Tetraclinis forest is from 12 m to 15 m

tall (White 1983). The hard, short-grained timber has

been esteemed since Roman times and may be finely

figured like birds-eye maple (Dallimore & Jackson 1974).

Apart from producing a commercially valuable timber,

this species is also the source of a resin which exudes

from the trunk known as African or Mogador Sandarac

and was used in the manufacture of clear varnishes (Howes

1949).

POTENTIAL USES IN SOUTH AFRICA

Tetraclinis articulata is clearly a very suitable tree for

semi-arid Mediterranean-type climatic conditions, having

an ability to withstand considerable periods of drought

(Dallimore & Jackson 1974).

It would probably be worth cultivating in the more arid

western areas of the winter rainfall region as a street tree

or an ornamental and might also possibly serve as a

commercial source of timber. However, in the moister

regions of the western Cape, e.g. the Cape Peninsula, there

are indications that this species could invade natural fynbos

on a limited scale.

No mention of Tetraclinis was made in the Flora of the

Cape Peninsula (Adamson & Salter 1950). This may

indicate that the spread of seedlings into areas surrounding

the original plantings has only become noticeable in the

last 40 years and that prior to 1950, seedling spread was

not especially evident due perhaps to the fact that the

original plantings had only recently reached sexual

maturity. The group at Simonstown now produces an abun-

dant seed crop annually, the cones dehiscing mainly in

February and March.

DESCRIPTION

Tetraclinis articulata (Vahl) Masters in Journal of the

Royal Horticultural Society: 250 (1892); Masters: 14

(1893); Tutin: 38 (1964); Dallimore & Jackson: 602 (1974);

Kriissmann: 318 (1972).

Thuja articulata Vahl: 96 (1794).

A monoecious, broadly conical evergreen tree 12—15

m tall; trunk up to 0,5 m in diam. Bark reticulately fissured

scaly, grey-brown. Branchlets laterally flattened, dichoto-

mous, glabrous, dark green, with two longitudinal

striations on each surface. Leaves in fours, scale-like,

adnate-decurrent, the tv/o laterals slightly larger than, and

enclosing, the other pair. Cones terminal, solitary. Female

cones stalked, ovoid, 4-valved, glaucous, 10-15mm in

diam., one pair of cone scales cordate-triangular, the

opposite pair smaller, ovate-truncate. Male cones sessile,

8-10 x 3 mm, sporophylls opposite and decussate,

papyraceous. Seeds conic, 15-18 mm across with two

prominent, upwardly deflected lateral wings.

Specimens examined

CAPE.— 3418 (Simonstown): Simonstown, mountainside above town

(-AB), Jan., Rourke 1824 (NBG, PRE, PRF).

TRANSVAAL. — 2628 (Johannesburg): The Wilds, cultivated, Sept.

1952, H. Friede 3 (PRE).

ACKNOWLEDGEMENTS

I am indebted to Mr P. Salter for assistance in the field

and to Miss C. Salter of the Simonstown Museum for

providing historical records.

REFERENCES

ADAMSON, R.S. & SALTER, T.M. 1950. The Flora of the Cape

Peninsula. Juta, Cape Town.

ANON. 1910. Souvenier of Simonstown 1910 : 46. Simonstown Munici-

pality. (Document in Simonstown Museum.)

DALLIMORE, W. & JACKSON, A.B. 1974. A handbook ofConiferae

and Ginkgoa'ceae , edn 4, revised by S.G. Harrison. Edward

Arnold, London.

HOWES, F.N. 1949. Vegetable gums and resins. Chronica Botanica,

Waltham, Mass.

KRUSSMANN, G. 1972. Handbuch der Nadelgeholze. Paul Parry, Berlin.

MASTERS, M.T. 1892. List of conifers and taxads. Journal of the Royal

Horticultural Society 14: 250.

MASTERS, M.T. 1893. Notes on the genera of Taxaceae and Coniferae.

Journal of the Linnean Society 30: 14, 15.

POYNTON, R .J. 1984. Characteristics and uses of selected trees and

shrubs cultivated in South Africa , edn 4. Directorate of Forestry,

Pretoria.

TUTIN, T.G. 1964. Cupressaceae. In T.G. Tutin et al.. Flora europaea

1. Cambridge University Press, Cambridge.

VAHL, M. 1794. Symbolae botanicae sive plantarum 3: 96.

WELLS, M.J., BALSINHAS, A. A., JOFFE, H., ENGELBRECHT,

V.M., HARDING, G. & ST1RTON, C.H. 1986. A catalogue of

problem plants in southern Africa. Memoirs of the Botanical

Survey of South Africa No. 53.

WHITE, F. 1983. The vegetation of Africa. A descriptive memoir to

accompany the Unesco/AETFAT/UNSO vegetation map of Africa.

Natural Resources Research 20. UNESCO, Paris.

J.P. ROURKE*

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

7735.

MS. received: 1990-10-26.

Bothalia 21,1 (1991)

65

PTERIDOPHYTA

A UNIQUE LOCALITY FOR OLEANDRA DISTENTA , THE FIRST RECORDED FOR THE ORANGE FREE STATE

During a phytosociological investigation of the Koranna-

berg (2827CC Excelsior and 2 827CD Clocolan) in the

south-eastern Orange Free State, a colony of Oleandra

distenta Kunze was discovered. This is the first record of

this species for the Orange Free State (Anthony & Schelpe

1985; Burrows 1990; Jacobsen 1983; Schelpe & Anthony

1986; Roux 1986). It is also the first time that the species

is recorded from an area with an average annual rainfall

as low as 617 mm (Weather Bureau 1954). According to

Jacobsen (1983) O. distenta Kunze occurs almost exclu-

sively in high rainfall areas that receive a rainfall of more

than 1 000 mm per annum. According to Burrows (1990);

Jacobsen (1983) and Schelpe & Anthony (1986) O. dis-

tenta Kunze also occurs in the Magaliesberg, where it was

first collected by Zeyher (Sim 1915) and where the annual

rainfall is 703 mm (Weather Bureau 1954). Plants in that

area were described by Jacobsen (1983) as relicts of an

earlier, wider distribution.

This fern species also occurs from the Transkei, through

Natal, the eastern and northern Transvaal into eastern

Zimbabwe, Mozambique (Figure 11), Malawi, Zambia and

throughout tropical Africa. It is also found on the

Seychelles, Madagascar, Reunion, Mauritius and the

Comoro Islands (Burrows 1990).

It grows at very low altitudes in Natal but reaches

altitudes well over 2 000 m above sea level in the tropics

(Burrows 1990; Jacobsen 1983; Schelpe & Anthony

1986). This lithophyte forms extensive, tangled colonies

over exposed rocky outcrops or scrambles over boulders

in light shade (Burrows 1990; Schelpe & Anthony

1986). In tropical Africa it can occur as a high-level

epiphyte on forest trees (Jacobsen 1983; Schelpe & An-

thony 1986).

At the Korannaberg it was found at an altitude of 1 720

m above sea level on a north-easterly aspect. The fern’s

FIGURE 11. — Oleandra distenta Kunze. Presently known distribution,

• ; new locality at Korannaberg, ♦.

widely creeping rhizomes are attached to cracks at the

base of a sandstone cliff of the Clarens Formating (Figure

12).

During summer water constantly seeps through the cracks

in the sandstone. During the first part of the day the colony

is fully exposed to sunlight, but according to Schelpe &

Anthony (1986) and Burrows (1990), the species rarely

grows in full sun. During the dry season the plant is

frequently deciduous, dropping most of its fronds (Burrows

1990; Jacobsen 1983; Schelpe & Anthony 1986).

ACKNOWLEDGEMENTS

I am indebted to the directors and Council of the

National Museum, Bloemfontein, for financing this

research and the financial support of the Free State branch

of the Wildlife Society is also gratefully acknowledged.

FIGURE 12.— Typical habitat of

Oleandra distenta Kunze at

Korannaberg. A, fern colonies;

B, dolorite sill; C, cliff face.

66

Bothalia 21,1 (1991)

REFERENCES

ANTHONY, N.C. & SCHELPE, E.A.C.L.E. 1985. A checklist of the

Pteridophytes of the Flora of southern Africa region. Bothalia

15: 541-544.

BURROWS, J.E. 1990. Southern African ferns and fern allies. Frandsen

Publishers, Sandton.

JACOBSEN, W.B.G. 1983. The ferns and fern allies of southern Africa.

Butterworths, Durban.

ROUX, J.P. 1986. A checklist of Pteridophyta of the north-eastern Orange

Free State. Bothalia 16: 83—85.

SCHELPE, E.A.C.L.E. & ANTHONY, N.C. 1986. Pteridophyta. In O.A.

Leistner, Flora of southern Africa. Government Printer,

Pretoria.

SIM, T.R. 1915. The ferns of South Africa , 2nd edn. Cambridge University

Press, Cambridge.

WEATHER BUREAU 1954. Climate of South Africa. Climate Statistics.

Part 1. WB 40. Government Printer, Pretoria.

P.J. DU PREEZ* and PC. ZIETSMAN*

* National Museum, P.O. Box 266, Bloemfontein 9300.

MS. received: 1990-11-08.

AMARYLLIDACEAE

THE CORRECT AUTHOR CITATION FOR CLIVIA MINIATA

The authority for Clivia miniata is universally indicated

as Regel (1864: 131). The most important examples are

Baker (1888: 62; 1896: 229), Wood (1909: t. 503), Phillips

(1921: t. 13), Compton (1976: 123), and Gibbs Russell et

al. (1985: 111).

This citation is incorrect. Regel (1864: 131) himself did

not claim to be describing C. miniata as new, but cited

Vallota miniata Lindley (1854: 119) in full. The miscon-

ception may have originated in Lindley (1854: 119) writing

the name as Vallota ? miniata , creating the impression that

the name is not validly published [I.C.B.N. (1988) Art.

34.1.b: not validly published ‘when it is merely proposed

in anticipation of the further acceptance...’]. From

Lindley’s text it is quite clear that he was in no doubt about

the taxonomic acceptability of the species, and that the

question mark indicates doubt only about the correct

placement of the species in the genus Vallota. Clearly

Vallota miniata is validly published in terms of Art. 34.2

which states that a name is validly published when

published with such ‘a question mark or other indication

of taxonomic doubt, yet published and accepted by the

author’.

The correct author citation is therefore Clivia miniata

(Lindley) Regel, based on Vallota miniata Lindley.

REFERENCES

BAKER, J.G. 1888. Handbook of the Amaryllideae: 62. Bell, London.

BAKER, J.G. 1896. Amaryllidaceae. In W.T. Thiselton-Dyer, Flora

capensis 6: 171-246. Reeve, Ashford.

COMPTON, R.H. 1976. The flora of Swaziland. Journal of South African

Botany, Supplementary Volume 11: 123.

GIBBS RUSSELL, G.E. et al. 1985. List of species of southern African

plants. Edn 2, Part 1. Memoirs of the Botanical Survey of South

Africa No. 51: (11.

GREUTER, W. etal. 1988. International code of botanical nomenclature.

Regnum vegetabile 118. Koeltz, Konigstein.

LINDLEY, J. 1854. New plants. 47. Vallota ? miniata. The Gardeners’

Chronicle August 1854: 119.

PHILLIPS, E.P. 1921. Clivia miniata. The Flowering Plants of South

Africa 1: t. 13.

REGEL, E. 1864. Clivia miniata Lindl. Gartenflora 13: 131.

WOOD, J.M. 1909. Clivia miniata. Natal Plants 6: t. 503.

P. VORSTER*

* Botany Department, University of Stellenbosch, 7600 Stellenbosch,

South Africa.

MS. received: 1990-11-14.

Bothalia 21,1: 67-72 (1991)

Mixoploidy and cytotypes: a study of possible vegetative species

differentiation in stapeliads (Asclepiadaceae)

F. ALBERS* and U. MEVE*

Keywords: adventitious roots, Asclepiadaceae, cytotypes, euploidy, growth forms, mixoploidy, Stapelieae

ABSTRACT

Mixoploidy is common in proembryos and embryos as well as in meristems of radicles, primary, secondary and adventitious

roots and in innovation shoots of stapeliad species (Asclepiadaceae). The proportion of polyploid cells in the meristem of

single adventitious roots is significantly higher than in meristems of primary and secondary roots. This may lead to a complete

polyploidisation of adventitious roots. Innovation shoots display a low percentage of polyploid cells, comparable to the condition

found in primary and secondary roots. Nevertheless cells of different euploid levels are frequently found. During field studies,

however, individuals of a given population were always found to have the same chromosome number; infraspecific polyploidy

was found in only three out of 305 investigated stapeliad species. Genera with an orthotropic growth form were found to

be diploid throughout, whereas the genera of which the members spread vegetatively by means of innovation shoots, comprise

di-, tetra- and polyploid species. This phenomenon can be ascribed to ecological rather than morphological factors. The

often proposed hypothesis that new cytotypes can have a vegetative origin is not found to be acceptable.

UITTREKSEL

Miksoploldie kom algemeen voor by proembrio’s en embrio’s asook by meristeme van kiemworteltjies, primere, sekondere

en bywortels en by verjongingslote van aasblomspesies (Asclepiadaceae). Daar is in verhouding meer poliploiede selle in

die meristeem van enkelvoudige bywortels as in meristeme van primere en sekondere wortels. Dit mag tot die algehele

poliploidisering van bywortels lei. Die persentasie poliploiede selle by verjongingslote is laag en kan vergelyk word met

die toestand wat by primere en sekondere wortels aangetref word. Selle van verskillende euploiede vlakke word nietemin

dikwels aangetref. Tydens ondersoeke in die veld is bevind dat individue in ’n bepaalde bevolking altyd dieselfde

chromosoomgetal het; infraspesifteke poliploidie is by slegs drie uit 305 aasblomspesies wat ondersoek is, aangetref. Genusse

met 'n ortotropiese groeivorm was deurgaans diploied, terwyl genusse waarvan die plante spreidende verjongingslote het,

di-, tetra- en poliploiede spesies bevat. Hierdie verskynsel kan aan ekologiese eerder as morfologiese faktore toegeskryf

word. Die hipotese wat dikwels voorgestel word dat nuwe sitotipes ’n vegetatiewe oorsprong kan he, is nie aanvaarbaar be-

vind nie.

CONTENTS

Introduction 67

Material and methods 67

Results 68

1. Mixoploidy in embryos and meristematic tissue

of adult individuals 68

a. Proembryos and embryos 68

b. Radicles and secondary roots 68

c. Adventitious roots 69

d. Primordia of innovation shoots 69

2. Mixoploidy in plagiotropic and orthotropic

genera 70

3. Chromosome number and ecology of popula-

tions 70

Discussion 71

Acknowledgements 72

References 72

INTRODUCTION

Previous casual collections and the karyological study

of stapeliad species have elucidated the spectrum of

chromosome numbers of this stem-succulent group of the

Asclepiadaceae (preliminary summary by Albers 1983).

* Institut fur Botanik, Westfalische Wilhelms-Universitat, Schlossgarten

3, D-4400 Munster, Germany.

MS. received: 1990-04-17.

The mixoploid character of many root tips, present as early

as in the meristematic stage, has often hampered the

determination of the chromosome number of single taxa.

Due to this peculiarity the vegetative origin of new

cytotypes has been suggested (Reese 1973). Even though

only root tips had been studied at the time, Reese (1973)

proposed a similar cytological concept for shoot tips,

which was originally introduced by Sharma (1956), who

suggested a vegetative species differentiation, but failed

to submit convincing experimental proof. To test the

hypothesis of vegetative cytotype differentiation, wide-

ranging karyological studies were carried out on

individuals, populations and genera. Data and material

were collected during several periods of field work in

southern Africa.

MATERIAL AND METHODS

Field work was carried out during periods of several

months’ duration in southern Africa in 1983, 1986 and

1988. The material collected is cultivated in the Botanical

Garden of the University of Munster. The present study

is based mainly on the following taxa from the Cape

Province, South Africa, and Namibia:

Duvalia caespitosa (Mass.) Haw.: 2n = 44.

CAPE.— 3118 (Vanrhynsdorp): 15 km S, Meve 147. 3119 (Calvinia):

40 km N of Niewoudtville, Meve 245\ 13 km N of Niewoudtville, Meve

252.

68

Bothalia 21,1 (1991)

Duvalia pubescens N.E. Br. : 2n = 44.

CAPE.— 2816 (Oranjemund): Richtersveld, Hellskloof, Albers & Meve

33. 3017 (Hondeklipbaai): 3 km E of Kamieskroon, Meve 154. 2917

(Springbok): 8 km N of Concordia, Meve 222.

Hoodia gordonii (Mass.) Sweet: 2n = 22.

CAPE.— 3219 (Wuppertal): Biedouw Valley, Albers el al. K1301.

NAMIBIA. — 2816 (Oranjemund): Lorelei, Albers et al. K1364.

Orbea namaquensis (N.E. Br.) Leach: 2n = 22.

CAPE.— 2817 (Vioolsdrif): 50 km N of Lekkersing, Albers & Meve

15. 2917 (Springbok): 4 km N of Steinkopf, Albers & Meve 112\ 5 km

N of Concordia, Meve 163.

Pectinaria articulata (Ait.) Haw. subsp. borealis Bruyns:

2n = 22.

CAPE.— 2816 (Oranjemund): Richtersveld, Hellskloof, Albers & Meve

32.

Stapelia asterias Mass.: 2n = 22.

CAPE. — 3321 (Ladismith): N of Calitzdorp, Albers et al. KII24.

Stapelia gariepensis Pillans: 2n = 44.

CAPE. — 2816 (Oranjemund): Richtersveld, Hellskloof, Albers & Meve

23, 24\ Grootderm, Jurgens s.n..

Stapelia hirsuta L.: 2n = 22.

CAPE. — 3318 (Cape Town): Malmesbury, Albers etal. KI5I3, KI5I4,

K1515.

Trichocaulon dinteri Berger: 2n = 22.

CAPE.— (Oranjemund): Richtersveld, Beauvallon, Albers et al. K1346.

NAMIBIA — 2716 (Witputz): 8 km S of Rosh Pinah, Albers & Meve 45.

Tridentea longipes (Liickhoff) Leach: 2n = 22.

CAPE. — 2816 (Oranjemund): Richtersveld, Numees, Albers & Meve

38, 39.

NAMIBIA. — 2716 (Witputz): 38 km N Rosh Pinah. Albers & Meve

76.

Earlier chromosome counts (Reese & Kressel 1967;

Reese 1971; Albers 1974, 1975, 1976, 1981; Albers & Delfs

1983; Albers & Austmann 1987; Albers, Austmann &

Meve 1988, 1990) have also been used in the present study.

Population studies were carried out in grid squares.

Individual plants, stem parts and fruits were collected

within each plot. Seeds of fruits which had already set

in habitat could be harvested from a number of plants in

the Botanical Garden of the University of Munster. Root

tips were collected from individual plants transplanted to

the greenhouse, from rooting stems (adventitious roots)

and from germinated seeds. Furthermore, the meristems

of the stem tip, the proembryo and the embryo were

studied. Apart from the latter, mitotic equatorial plates

were generally used to count chromosome numbers.

Sections and squash preparations were produced and

stained according to standard techniques (Snow 1963). For

the embryo sections, the nuclei were measured and the

ploidy level estimated according to the different diameters

of these nuclei.

RESULTS

Within the Stapelieae, two different growth forms are

known. In southern Africa the genera Hoodia, Quaqua

and Trichocaulon are characterized by an orthotropic

growth form, while other genera, such as Duvalia,

Huernia, Orbea, Pectinaria, Piaranthus, Stapelia and

Tridentea display plagiotropic growth forms. In the latter

taxa the axis ceases to grow at an early stage. Basally,

numerous innovation shoots are formed, which are often

lying on the ground, lending a mat-like appearance to the

plant (Albers et al. 1989). These lateral shoots form

adventitious roots. In order to clarify the extent and

qualitative significance of mixoploidy, embryos as well as

root- and shoot-primordia of individuals of these two

growth form types were studied.

1 Mixoploidy in embryos and meristematic tissue of adult

individuals

a. Proembryos and embryos

In Trichocaulon dinteri, Duvalia caespitosa and D.

pubescens, single polyploid cells can be found as early

as the proembryo stage. Even though the ploidy level

cannot always be established with certainty by means of

counts in equatorial plates, the size of the nucleus and the

number of chromocentres allow such a judgement. The

chromocentres are extraordinarily prominent in

meristematic cells of the Asclepiadaceae and, as Czeika

(1956) has already pointed out, for Echidnopsis cereiformis

Hook. f. (Stapelieae), it is easy to differentiate between

diploid and polyploid cells. Polyploid cells are also found

in the tissue of mature embryos ( <5%). Whether chromo-

some numbers are exclusively doubled, or whether higher

ploidy levels occur, cannot always be determined with

certainty, judging from the number of chromocentres. The

polyploid cells are single cells evenly dispersed throughout

the whole tissue of the cotyledons, the hypocotyl and the

radicle.

b. Radicles and secondary roots

Between 1,9% and 6,3% polyploid cells occur in the

apical region of the radicle (Table 1). These figures

represent the mean of different root tips of an individual

plant; individual values, however, ranged from 0 to 8,3%.

Polyploid cells do not simply possess a double chromo-

some number. Cells with 33 chromosomes were observed

in diploid species and cells with 55, 66 and, very rarely,

77 and 88 chromosomes were seen in tetraploid species.

There are obviously no divergent aneuploid cells. Merely

one individual with 2n=22 + IB was found in the East

African Echidnopsis scutellata (Defiers) Berger. Irregulari-

ties during the anaphase were also never detected.

Bothalia 21,1 (1991)

69

TABLE 1, — Percentage of aberrant polyploid cells in the meristematic

area of the radicle

Cells with chromosome numbers above the diploid level

of a species are scattered over the whole cross section

of the apex with a slight increase in number in the

protoderm.

Species with orthotropic growth possess an allorhizal

root system. Even though examination of the secondary

roots of Trichocaulon dinteri occasionally revealed small

sectors of triploid cells in the periblem and the plerome,

the percentage of polyploid cells in the apical regions

corresponds with the lower value already found in

the radicles. The primary roots studied support this

observation.

c. Adventitious roots

In species with mat-like growth forms, the number of

shoots can increase considerably during a single growth

period. Their adventitious roots have previously been

studied by Reese (1973). In tips of adventitious roots, he

demonstrated the entire spectrum of different ploidy levels;

however, he did not compare adventitious roots with

radicles, primary and secondary roots.

Apart from single euploid nuclei, polyploid layers and

sectors can be found more frequently in adventitious roots

(Figure 1). This root type therefore displays more

cytochimaera features than the ones previously mentioned

(Reese 1973: fig. 1). Although the adventitious roots

normally contain the expected chromosome number, the

share of (aberrant) polyploid cells, which did not reach

10% in radicles and secondary roots, can rise to more than

50% in adventitious roots of Duvalia and Huemia spe-

cies. In rare cases entire adventitious roots attain a higher

ploidy level (see also Reese 1973).

Reese (1973) mentions two chromosomally aberrant

examples in which a tripling of the basic number could

be observed, apart from normal diploid values. This

euploidy has, however, been found in many species

presently studied. Root tips of diploid species can contain

cells with 2n=22, 33, 44 and 66 chromosomes (Figure

LA). Penta- to octoploid cells (Figure IB, C) can frequently

be found in tetraploid species, and hepta- and octoploid

cells in hexaploid species.

A haploid cell with 11 chromosomes was observed only

twice: in the tetraploid Duvalia vestita Meve, and in

the diploid Pachycymbium carnosum (Stent) Leach. No

explanation for this unusual karyological condition can be

offered.

The number of triploid and pentaploid cells is occa-

sionally very high; these cells may even make up the entire

root. Adventitious roots of innovation shoots, sequen-

tially branching off the same plant, do not change in their

karyological composition. Thus the proportion of

polyploid cells to diploid cells does not increase with

time.

d. Primordia of innovation shoots

In apical meristems of innovation shoots, relatively few

polyploid cells are found. Of the twelve examined

individuals belonging to the genera Duvalia , Orbea,

Pectinaria and Piaranthus, only three plants showed

FIGURE 1. —Metaphase plates in root

tip meristems. A, Duvalia ele-

gans (2n=22) with a normal

and an aberrant equatorial plate

(22 < — > 33 chromosomes);

B, D. vestita (2n=44) with 55

chromosomes; C, D. caespi-

tosa (2n=44) with 77 chromo-

somes. All x 1600.

70

Bothalia 21,1 (1991)

TABLE 2. — Mixoploidy in the apical area of innovation shoots

mixoploidy in the apical area. In these plants the

percentage of polyploid cells of different ploidy levels was

also small (Table 2). Exclusive occurrence of unequal

multiples of haploid chromosome numbers, apart from the

expected diploid numbers, is particularly conspicuous.

2 Mixoploidy in plagiotropic and orthotropic genera

The above-mentioned peculiarities have been found

predominantly in adventitious roots. Nevertheless, differ-

ences between genera of different growth forms are

smaller than expected. In orthotropic genera 7,7% of the

individuals display mixoploidy, while this number is about

9,5% in plagiotropic genera (Table 3).

3 Chromosome numbers and ecology of populations

Growth forms in the Stapelieae have been discussed

recently by Albers et al. (1989). Studies concerning

ecology and population dynamics in the tribe are not

available. However, several months of field work allow us

to recognize some clearcut tendencies. Plagiotropic species

are regularly associated with xerophytic shrubs (about

0,5 -1,0 m high), but rarely with succulent shrubs. They

grow generally at the shaded bases of these shrubs. The

fine-grained soil found in this habitat, due to reduced

erosion and the accumulation of drifting sand, furnishes

the ideal substrate for stapeliads. This preference for a

certain habitat expresses itself in the ephemeral character

of these species; and in highly variable population sizes

and densities in space as well as in time. This behaviour,

which characterises pioneer plants, is particularly con-

spicuous in disturbed areas.

One population of Duvalia caespitosa (Touwsrivier,

Karoo), initially consisting of 33 individuals (and a total

of 648 shoots) has been observed since 1983 and

considered suitable for studies of population karyology as

well as population dynamics. The high density of

individuals, which causes meshing of individual plants in

several places, is in part a consequence of vegetative

reproduction by means of innovation shoots. As demon-

strated by vegetative reproduction in cultivation, every

shoot can be the starting point for a new plant. Even so,

no individuals have been found for which chromosome

numbers in adventitious roots or in apical meristems were

higher than 2n=44 throughout, the normal chromosome

number for the tetraploid D. caespitosa.

Observations on this stand, repeated in 1986 and 1988,

demonstrate clearcut changes within the whole stapeliad

population: only a few individuals of D. caespitosa were

left in 1988, whereas another species, Stapelia pillansii

N.E. Br., was becoming more frequent. Populations of

Tridentea longipes (Grasvlakte, Richtersveld), Stapelia

hirsuta (Malmesbury) and Orbea namaquensis (N of

Lekkersing, Richtersveld) belonging to the same growth

TABLE 3. — Chromosome numbers and extent of mixoploidy in individuals and species of different growth forms

Bothalia 21,1 (1991)

71

form, also provided no evidence for the formation of new

cytotypes.

Orthotropic species in general form stable, long-lived

populations on fully sun-exposed stands. Habitat observa-

tions in populations of Trichocaulon dinteri in the Richters-

veld (Beauvallon) illustrate in addition the strong influence

of topographic and edaphic factors on orthotropi-

cally growing species. Seventeen out of 19 individuals of

a small population, restricted to a small hill (6 m2) with

very coarse and permeable substrate, grew on the

microclimatically more moderate south-western slope. In

contrast, no plant could be found on the sandy and

level area around that hill. The clearcut boundary of the

population, however, has to be attributed to the occurrence

of rather unstable quartzite sands. They create a high-risk

habitat for the slow-growing Trichocaulon plants, which

are incapable of vegetative reproduction.

Investigations of root tips of primary and secondary roots

and seedlings grown from seeds harvested from individu-

als of T. dinteri at this locality did not yield any new karyo-

logical information. Only a few single cells differed from

the diploid chromosome number in the manner discussed

earlier. Since no innovation shoots were formed and thus

no vegetative reproduction occurred, formation of different

cytotypes is considered to be impossible in this growth

form.

DISCUSSION

Endopolyploidy has been found in a vast number of

species throughout the vascular plants (John & Lewis

1968). On the other hand, a low level or absence of

endopolyploidy seems to characterize a smaller number

of species Tschermak-Woess (1956). Endomitotic poly-

ploidisation commonly occurs in the zone adjacent to the

apical meristem (Fenzl & Tschermak-Woess 1954). This

fact has been confirmed by the present observations in

radicles and both secondary and primary roots of the

Stapelieae, in which a relatively low percentage of aberrant

chromosome numbers has been found in the apical zone.

A different point of view, however, needs to be taken

for the adventitious roots widespread in the Stapelieae.

The conspicuous mixoploid character of some adventitious

roots has been mentioned in earlier studies on chromo-

some numbers in the Stapelieae (Reese 1971; Reese &

Kressel 1967). Reese (1973) comments on the different

degrees of polysomaty in some stapeliad species, and

develops his own ideas about their origin. His suggestion

that the adventitious roots, and to a lesser degree the secon-

dary roots, tend towards mixoploidy, was confirmed in the

present study. The mixoploidy level rises in the meristems

of adventitious roots. According to Reese, cells of differ-

ent polyploid levels become involved as early as the for-

mation stage of the primordia.

Endopolyploidisation commonly leads to a doubling

of chromosome numbers (Tschermak-Woess 1971). Occa-

sionally higher polyploid cells have been observed

(summary in Tischler & Wulff 1953-1963: 321-336).

Though they occur in the stapeliads in extraordinarily high

numbers, they have not been mentioned at all by Czeika

(1956) and only as an oddity by Reese (1973). They are

found in different root types as well as in shoot tips. While

valencies of euploid cells commonly exceed the diploid

value, an equatorial plate has been found in Duvalia vestita

and Pachycymbium carnosum with 11 chromosomes.

Mitotic irregularities during the anaphase have never

been noted here. It is questionable whether a nucleus

fragmentation observed by Czeika (1956) in older tissue

of Echindopsis cereiformis (Stapelieae) was responsible

for the formation of aneuploid nuclei, since amitoses are

more or less disorganized processes (Nagl 1976) and are

never followed by cell divisions (Nagl 1978).

Since the number of cells with different polyploid levels,

especially in adventitious roots of plagiotropically growing

shoots, can be considerable and since these polyploid cells

can in extreme cases constitute the whole root, this

phenomenon can be identified as a source of obviously

wrong chromosome numbers in the literature. Some

stapeliad species have been thought of as triploid and

therefore as potential hybrids (Reese & Kressel 1967).

Control counts have, however, unambiguously established

the diploid character of these species. In contrast, triploidy

has been proven in a natural hybrid (Duvalia caespitosa

2n=44 x Huernia pillansii 2n=22; Albers & Meve

unpubl.), which was collected by Bruyns (1981).

A small percentage of polyploid cells, comparable to

those found in radicles and secondary roots, has been

found in apical meristems of innovation shoots, which

enlarge the body of individual plants of plagiotropic

species. Since every shoot roots as it comes into contact

with the soil, it forms a possible starting point for a new,

genetically identical plant.

Speculative formation of new cytotypes by means of

mixoploid meristems in this vegetative process, could not

be proven in individuals of larger populations. The possi-

bility of such a process is also contradicted by results of

karyological studies in apical meristems of innovation

shoots, which have never been found to be exclusively

polyploid. The exclusive presence of polyploid cells in a

small percentage of adventitious roots had previously led

to such speculations (Reese 1973). The different mode of

formation of adventitious roots (from parenchymatic tis-

sue adjacent to the outer phloem) and of innovation shoots

(exogeneously from axillary buds) must also be taken into

account.

The fact that species of the genera Hoodia, Quaqua and

Trichocaulon (a total of 19 species was investigated) with

orthotropic growth forms, are exclusively diploid, seems

to support the hypothesis of cytotype formation, since

tetraploidy and hexaploidy are widespread in the genera

with plagiotropic growth forms. However, this clearcut dis-

tinction between the two morphologically different groups

must be attributed to habitat conditions rather than to

growth form. Hoodia, Trichocaulon and Quaqua (p.p.)

predominantly inhabit extremely dry and relatively stable

biotopes in the winter rainfall area of southern Africa and

their populations could be shown to be highly constant

during the time of our studies over five years. Most other

genera, in contrast, are confined to far more unstable

karroid areas, in which they are apparently often relatively

short-lived.

72

Bothalia 21,1 (1991)

The formation of different ploidy levels in the

plagiotropic growth form obviously is not caused by the

branching off of cytologically different innovation shoots,

but has been achieved generatively by promotion of poly-

ploid races in the conquest of new habitats (e.g. Hanelt

1966). The possible vegetative formation of new cytotypes

requires a high percentage of different infraspecific

cytotypes. Extensive investigations of species and sub-

species has so far only revealed two cytotypes each in

Orbea variegata (L.) Leach, Huernia hislopii Turill subsp.

hislopii , and Duvalia polita N.E. Br. In terms of stem and

flower morphology, each pair member is indistinguishable

from the other; the two cytotypes of the first species,

however, inhabit ecologically clearly distinct habitats. In

D. polita there are indications of mixed populations with

diploid and tetraploid cytotypes. In summary, there are

no clues for an enlarged number of cytotypes nor for a

special mode of origin for the existing ones.

Angiosperms tend to form mosaic tissues; the degree

to which mosaic tissues are produced is more or less

species-specific. This condition, which can occur as early

as in the apical meristems, however, obviously does not

lead to the formation of new cytotypes under natural

conditions, even though such a mechanism has been

suggested several times (Sharma 1956; John & Lewis 1968;

Reese 1973). This fact is surprising in the light of the

successes of plant breeding research in the production of

polyploid strains.

ACKNOWLEDGEMENTS

Our studies have been made possible by financial

support of the Deutsche Forschungsgemeinschaft, the

CSIR (RSA) and the University of Stellenbosch (RSA).

For advice on many aspects we are grateful to Mr M.B.

Bayer and Mr L. Leach, both formerly at the Karoo

Botanical Garden, Worcester, RSA. Prof. J.J.A. van der

Walt, Dept, of Botany, University of Stellenbosch, has

supported our work in various ways. Finally we wish to

thank Dr Sigrid Liede for translation of our manuscript,

and numerous other colleagues in South Africa, Namibia,

Lesotho and Zimbabwe.

REFERENCES

ALBERS, F. 1974. In A. Love, IOPB chromosome number reports XLIV.

Taxon 23 : 373, 374.

ALBERS, F. 1975. In A. Love, IOPB chromosome number reports XLDC

Taxon 24 : 507, 508.

ALBERS, F. 1976. In A. Love, IOPB chromosome number reports LI.

Taxon 25: 161-163.

ALBERS, F. 1977. In A. Love, IOPB chromosome number reports LVHI.

Taxon 26: 557, 558.

ALBERS, F. 1981. In A. Love, IOPB chromosome number reports LXX.

Taxon 30: 76, 77.

ALBERS, F. 1983. Cytotaxonomic studies in African Asclepiadaceae.

Bothalia 14 : 795 -798.

ALBERS, F. & AUSTMANN, M. 1987. In A. Love, IOPB chromosome

number reports XCV. Taxon 36: 494—496.

ALBERS, F. & DELFS, W. 1983. In A. Love, IOPB chromosome number

reports LXXXI. Taxon 32: 667, 668.

ALBERS, F., AUSTMANN, M. & MEVE, U. 1988. Chromosome

number report. IOPB Newsletter 11: 10-12.

ALBERS, F., AUSTMANN, M. & MEVE, U. 1990. Chromosome

number report. IOPB Newsletter 15: 11—14.

ALBERS, F., DELFS, W., KUSCH, G. & MEVE, U. 1989. Lebens-

formen der Ceropegieae und Stapelieae (Asclepiadaceae) in ariden

Zonen Afrikas. Beitrage zur Biologie der Pflanzen 64: 59—74.

BRUYNS, P.V. 1981. Field hybridisation — a further comment. Asklepios

24: 53, 54.

CZEIKA, G. 1956. Strukturveranderungen endopolyploider Ruhekeme

im Zusammenhang mit wechselnder Bundelung der Tochter-

chromosomen und karyologisch-anatomische Untersuchungen an

Sukkulenten. Osterreichische Botanische Zeitschrift 103: 536—

566.

FENZL, E. & TSCHERMAK-WOESS, E. 1954. Untersuchungen zur

karyologischen Anatomie der Achse der Angiospermen. Oster-

reichische Botanische Zeitschrift 101: 140—164.

HANLET, P. 1966. Polyploidie-Frequenz und geographische Verbreitung

bei hoheren Pflanzen. Biologische Rundschau 4: 183—196.

JOHN, B. & LEWIS, K.R. 1968. The chromosome complement.

Protoplasmatologia. Handbuch der Protop lasmaforschung. Wien,

New York.

NAGL, W. 1976. Zellkern und Zellzyklen. Stuttgart.

NAGL, W. 1978. Endopolyploidy and polyteny in differentiation and

evolution. Amsterdam, New York, Oxford.

REESE, G. 1971. Untersuchungen iiber die Chromosomenzahlen der

Stapelieae II. Portugaliae Acta Biologica , Ser. A, 12: 1—23.

REESE, G. 1973. Uber einen bemerkenswerten Fall von Polysomatie in

Wurzelspitzen. Cytologia 38: 593—601.

REESE, G. & KRESSEL, H. 1967. Untersuchungen uber die Chromo-

somenzahlen der Stapelieae. Portugaliae Acta Biologica , Ser. A,

10: 33-54.

SHARMA, A.K. 1956. A new concept of a means of speciation in plants.

Caryologia 9: 93—130.

SNOW, R. 1963. Alcoholic hydrochloric acid-carmine as a stain for

chromosome in squash preparations. Stain Technology 38: 9—13.

TISCHLER, G. & WULFF, H.D. 1953-1963. Allgemeine Pflanzen-

karyologie. Suppl. Angewandte Pflanzenkaryologie. Handbuch

der Pflanzenanatomie 2. Berlin.

TSCHERMAK-WOESS, E. 1956. Karyologische Pflanzenanatomie.

Protoplasma 46 : 798 - 834.

TSCHERMAK-WOESS, E. 1971. Endomitose. Handbuch der Allge-

meinen Pathologie 2,2. Berlin, Heidelberg, New York.

Bothalia 21,1: 73 - 89 (1991)

Invasive alien woody plants of the Orange Free State

L. HENDERSON*

Keywords: alien. Grassland Biome, invasive plants (woody), Nama-Karoo Biome, Orange Free State, roadside survey. Savanna Biome

ABSTRACT

The frequency and abundance of invasive alien woody plants were recorded along roadsides and at watercourse crossings

in 66% (151/230) of the quarter degree squares in the study area. The survey yielded 64 species of which the most prominent

(in order of prominence) in streambank habitats were: Salix babylonica, Populus x canescens, Acacia dealbata and Salix

fragilis (fide R.D. Meikle pers. comm.). The most prominent species (in order of prominence) in roadside and veld habitats

were: Opuntia ficus-indica, Primus persica, Eucalyptus spp., Rosa eglanteria, Pyracantha angustifolia and Acacia dealbata.

Little invasion was recorded for most of the province. The greatest intensity of invasion was recorded along the perennial

rivers and rocky hillsides in the moist grassland of the eastern mountain region bordering on Lesotho and Natal.

UITTREKSEL

Die frekwensie en volopheid van uitheemse houtagtige indringerplante is langs paaie en by oorgange oor waterlope in

66% (151/230) van die kwartgradevierkante in die studiegebied aangeteken. Daar is 64 spesies aangetref waarvan die vemaamste

(in volgorde van belangrikheid) langs stroomoewers Salix babylonica, Populus X canescens. Acacia dealbata en Salix fragilis

(fide R.D. Meikle pers. meded.) was. Die vernaamste spesies (in volgorde van belangrikheid) langs paaie en in veldhabitats

was Opuntia ficus-indica, Prunus persica, Eucalyptus spp., Rosa eglanteria, Pyracantha angustifolia en Acacia dealbata.

Daar was min indringing in die grootste deel van die provinsie. Die ergste indringing is langs standhoudende riviere en

op rotsagtige heuwels in die vogtige grasveld van die oostelike berggebied langs die Lesotho- en Natalse grense aangetref.

CONTENTS

Introduction 73

Survey history and objectives 73

The study area 73

Method 74

Sampling method 74

Abundance ratings 75

Sampling level achieved 75

Data treatment — formulae used 75

Frequency 75

Prominence value 75

Mean species abundance rating in roadside and veld

habitats 76

Mean abundance of invaders per km in roadside and

veld habitats 76

Results 76

The streambank habitat 76

The whole study area 76

Analysis according to veld type 77

Analysis according to species 77

Frequency 77

Prominence 83

Roadside and veld habitats 83

The whole study area 83

Analysis according to veld type 83

Analysis according to species 83

Frequency 83

Pominence 83

Patterns of invasion 83

Discussion 83

Indigenous versus alien species composition 83

* Plant Protection Research Institute, Department of Agricultural

Development; stationed at National Botanical Institute, Private Bag X101,

Pretoria 0001.

MS. received: 1990-11-29.

Prominent and potentially important species 84

Relation of invasion to environmental factors .... 87

Conclusion 88

Acknowledgements 88

References 88

Appendix 89

INTRODUCTION

Survey history and objectives

This study of the Orange Free State (OFS) is the third

of eight regional surveys which together are designed to

reflect invasion by woody alien plants in the Republic of

South Africa as a whole. Surveys have been completed

for the Transvaal (Henderson & Musil 1984) and Natal

(Henderson 1989). The north-eastern OFS was surveyed

concurrently with north-western Natal in February 1987.

The remainder of the OFS was surveyed during October

1987, October 1988 and November 1988.

The objectives of the survey are: to produce a checklist

of the major invasive alien woody plants of streambank,

roadside and veld habitats in the study area; to determine

the pattern of alien woody invasion as a whole and for in-

dividual species; to attempt to relate distribution to

environmental factors and to determine which are the most

prominent and potentially important invaders.

The study area

The OFS occupies an area of 127 993 km2 (Depart-

ment of Foreign Affairs and Information 1983). It lies

between latitudes 26° and 31°S and longitudes 24° and

30°E (Figure 1). It is situated on the central plateau of

South Africa and consists largely of open rolling plains

74

Bothalia 21,1 (1991)

0

Intensive site

Survey route

FIGURE I. — The study area, survey routes and intensive sites.

interrupted at irregular intervals by low rocky hills. The

general altitude above sea level varies from 1 000 m in

the west to 2 000 m in the east. Near the borders of

Lesotho and Natal the landscape becomes more undulating

and high peaks up to 3 000 m occur along the Maluti and

Drakensberg Mountains near the junction of the OFS,

Lesotho and Natal borders.

Rain falls mainly in summer, the mean annual rainfall

more or less coinciding with altitude. It averages only

250—380 mm in the west and increases gradually to

635—760 mm in the east with 890 mm or more near the

Natal border (King 1951).

Summer temperatures are generally high, sometimes

exceeding 38°C during mid-summer. The resultant high

rate of evaporation causes a moisture deficit in virtually

all regions of the province, so that perennial streams and

continuously moist soils occur only in very limited areas

of the high rainfall eastern districts (Roberts 1968). Periods

of drought frequently occur in the spring and early summer

(Van Rensburg 1975).

Winters are cold and dry, except in the mountainous

eastern regions where mist and snow are common winter

phenomena (Roberts 1968). Frost is moderate to severe

over the entire province (Poynton 1972). Mean daily

minimum temperatures for July range from less than

— 2,5°C to about 2,5°C (Schulze & McGee 1978). The

highest frequency of below-freezing minimum tempera-

tures occurs in the eastern Drakensberg region (Tyson

1986).

There are three biomes in the study area. These are

Grassland, Nama-Karoo and Savanna (Rutherford &

Westfall 1986) (Figure 2). Thirteen Acocks Veld Types

(Acocks 1988) occur in the study area and have been

grouped into four veld type categories for the purposes

of this survey. These are temperate grassland, and moist

subtropical and transitional grassland in the Grassland

Biome; false karoo in the Nama-Karoo Biome; and

Kalahari thornveld in the Savanna Biome (Table 1 and

Figure 2).

Temperate grassland occupies the greatest area in the

OFS. Indigenous woody species are scarce, being confined

to occasional rocky hills. Moist subtropical grassland is

situated in the high rainfall mountain region adjacent to

the borders with Natal and northern Lesotho. Patches of

indigenous forest occur in valleys and on sheltered moun-

tain slopes. Transitional grassland links the temperate and

moist subtropical grasslands in the highest and wettest

parts of the OFS. It is undulating and broken country with

indigenous scrub occurring on hillsides and ravine forests

occurring in deep mountain valleys. Transitional and moist

subtropical grassland have been combined here since they

provide similar environmental conditions for the growth

of woody species, which are the subject of this survey.

Kalahari thornveld is limited to the extreme north-western

portion of the province and is characterized by an open

Acacia savanna, with a sparse ground cover of semi-arid

grasses. False karoo occupies the western and south-

western arid regions and is generally regarded as an

induced formation resulting from the degradation and

desertification of the original grassland (Acocks 1988). It

is characterized by a sparse population of dwarf shrubs

and mostly annual grasses. Trees and shrubs grow on the

many low rocky hills.

METHOD

Sampling method

The method used in this survey was the same as that

used in Natal (Henderson 1989). The presence and abun-

dance of all naturalized alien trees and large shrubs were

recorded for each veld type category, habitat type (road-

sides and adjoining veld, and streambanks) and quarter

TABLE I. — Veld type categories in the study area and the equivalent Acocks Veld Type groupings and Veld Type numbers

according to Henderson.

Bothalia 21,1 (1991)

75

SAVANNA BIOME

Kalahari thornveid

NAMA-KAROO BIOME

[♦. + . +. False karoo

GRASSLAND BIOME

] Temperate grassland

j jj Transitional grassland

| Moist subtropical grassland

FIGURE 2.— The biomes and broad veld type categories in the study

area, after Rutherford & Westfall (1986) and Acocks (1988).

degree square traversed by road. No naturalized climbers

were seen in this survey. Although the objective of the

survey was to record woody species, other large non-

woody and succulent species were included rather than

lose valuable information.

Recordings of roadside and veld invaders were made

from a moving vehicle whereas recordings of streambank

invaders were made while stationary at watercourse

crossings. Abundance estimates of roadside and veld

invaders were based on frequency of encounter within road

transects of five to ten kilometres in length. Abundance

estimates of streambank invaders were based on estimates

at specific sites.

a total road length of approximately 30 km (six transects

each five km long). Recordings were made at virtually

all watercourse crossings. Herbarium specimens of all

invader species which were flowering or fruiting, were

collected.

Survey routes and road transects were plotted on

1:250 000 maps (general survey area) and 1:50000 maps

(intensive sites) before a field trip was undertaken.

Wherever possible two or more road transects were plotted

per quarter degree square. As in the Natal survey

(Henderson 1989) road transects usually were not

contiguous but were separated by a distance of between

five and ten kilometres. Road transects along national roads

and other routes with heavy traffic were kept to a

minimum. Recordings were made at most bridges over

watercourses but some were omitted because of time

constraints and traffic considerations.

Abundance ratings

The abundance ratings for roadside and veld habitats

and streambank habitats are given in Table 2. Henderson

(1989) recommended that the abundance scale for stream-

bank habitats be revised or replaced with a cover-

abundance scale. Most of the field work for this survey

had already been completed when this recommendation

was made. The old abundance scale was therefore retained

in this survey but will be replaced by a cover-abundance

scale in further surveys.

Sampling level achieved

The sampling level achieved in this survey was 66% (151

of the total 230 quarter degree squares) at an average of

29 km travelled per square. An average of 18 km of road

transects were sampled per quarter degree square for

abundance estimates of roadside and veld invaders.

The veld type coverage in terms of quarter degree

squares and road transects sampled, kilometres travelled

and watercourse recordings made, is given in Table 3.

The width of road transects and length of watercourses

scanned for invaders varied according to local conditions.

Usually no more than 50 m of veld and 100 m of stream-

bank habitat were scanned on either side of the road for

invaders. Species occurring beyond these ranges and along

watercourses which were not crossed were recorded as

present in the given habitat type and veld type category

but were not included within the formal recordings.

Nine quarter degree squares were selected, using a

combination of systematic and subjective methods, for

more intensive surveying (Figure 1). Initial site selection

was systematic and on a country-wide basis, the sites being

situated approximately two degree squares apart along each

line of latitude and longitude. This ensured an even spread

of sites through the whole country and through each

biome. Further sites were subjectively selected to

incorporate representative parts of each veld type category

and geographical subregion in the study area. The inten-

sive sites may also be used at a later date for a quick

resurvey of the study area to assess any changes that may

have taken place. In each of the nine sites abundance

recordings of roadside and veld invaders were made along

Data treatment — formulae used

Frequency

The percentage frequency of occurrence of a species

x in veld type category y was calculated as follows:

no. of watercourse recordings/road transects

, in veld type y having species x .

frequency = — — — - — — x 100

total no. of watercourse recordings/road

transects in veld type y

All frequencies were expressed as percentages, even

though some sample sizes were less than 100. This was

done to facilitate comparisons with other similar statistics,

and the error terms associated with these estimates should

be borne in mind.

Prominence value

The prominence value is a combined measure of a

species’ frequency and abundance relative to that of all

other species, within a particular vegetation category.

76

Bothalia 21,1 (1991)

TABLE 2. — Abundance ratings

* Approximate numbers of individuals/groups per 10 km transect.

It has been derived from Curtis’ Importance Value

(Mueller-Dombois & Ellenberg 1974) and was first

described by Henderson (1989).

In streambank habitats the prominence value for a

species x in veld type category y was calculated as

follows:

frequency of species x in veld type y

scoring an abundance rating of 5, 6 or 7

sum frequency of all species in veld type

y scoring abundance ratings of 5, 6 or 7

prominence value = +

frequency of species x in veld type y

sum frequency of all species in veld

type y

The selection of abundance rating 5 as the cut-off point

is arbitrary but one at which a species can be regarded

as locally prominent (see definition in Table 2). A formula

using all abundance ratings would be preferable if each

rating could be converted to an absolute value.

Mean species abundance rating in roadside and veld

habitats (see Table 7)

The mean abundance rating** of a species x in veld type

category y was calculated as follows:

mean no. of

individuals

or groups

per 10 km

total no. of individuals or groups of species

x in veld type y

X 10

total distance along which species x was

rated in veld type y

Mean abundance of invaders per km in roadside and veld

habitats (see Table 5 and Figure 5)

The mean abundance of invaders per kilometre in veld

type category y/quarter degree square z was calculated as

follows:

total abundance* of all species in veld type y /

quarter degree square z

mean abundance =

total kilometres rated for abundance estimates in

veld type y/quarter degree square z

In roadside and veld habitats the prominence value for

a species x in veld type category y was calculated as

follows:

total abundance* of species x in veld

type y ^

sum of the abundances* of all species

in veld type y

prominence value = +

frequency of species x in veld type y

sum frequency of all species in veld

type y

RESULTS

The survey yielded 64 naturalized alien species. These

species are listed in the Appendix together with a further

six species which were obtained from specimen labels in

the National Herbarium. All these species with the

possible exception of Nicotiana glauca have been cultivated

in the OFS and have spread spontaneously from gardens

and plantations into the surrounding countryside. The

distributions and high abundance areas of 30 of the most

prominent species are given in Figures 6 & 7.

* each abundance rating was expressed in numbers of individuals or

groups recorded per transect (see Table 2). To be both conservative

and consistent the minimum number was used in each instance, e.g.

an abundance rating of 5 over ten kilometres = 50 and an abundance

rating of 5 over five kilometres = 25.

** mean no. of individuals or groups per 10 km converted to rating (see

Table 2).

The streambank habitat

The whole study area

Three hundred and ninety watercourse crossings were

sampled in which 42 species were recorded, with up to

Bothalia 21,1 (1991)

77

TABLE 3. — Sampling coverage in each veld type category, Biome and the study area

* This represents the distance along which abundance recordings were made. Total distance along which observations were made is approximately

one and a hall times that given; t according to Henderson; t according to Rutherford & Westfall 1986.

TABLE 4.— Statistics for streambanks in each veld type category, Biome and the study area

Veld type category* and Biome* Total no' Average no. of Max. no. of % crossings % crossings

of spp. spp. /crossing spp./crossing heavily invaded* invaded**

* one or more species scored an abundance rating of 5 or more; ** see data treatment-formulae used; f according to Henderson;

f according to Rutherford & Westfall 1986.

12 species in one sample. Invaders were present at 72,3%

of all crossings and 16,2% of all crossings were heavily

invaded (Table 4).

Analysis according to veld type

The greatest intensity of invasion was recorded in moist

subtropical and transitional grassland. Half the total

number of species and two-thirds (42/63) of all heavily

invaded watercourse crossings were recorded in this

relatively small area. There was a progressive decrease

in the intensity of invasion from east to west with the least

invasion occurring in Kalahari thornveld.

Analysis according to species

Frequency

Salix babvlonica was the most frequently recorded

invader in all veld type categories and the whole study

area (62,8%). It was most frequent in moist subtropical

and transitional grassland where it was recorded at 76%

of all watercourse crossings (Table 6).

TABLE 6. — Alien species occurring in streambank habitats

78

Bothalia 21,1 (1991)

F = % frequency of occurrence; I = % crossings heavily invaded; P = prominence value; * species occurring in the given category but not included in a formal recording at a watercourse crossing; t according to

Henderson; $ according to Rutherford & Westfall 1986.

TABLE 6. — Alien species occurring in streambank habitats (continued)

Bothalia 21,1 (1991)

79

E

E

o

<D

Q.

a

.o

"s

.00

op £ .jC

00 ^

C <3

Q£ C

3 Q- ~

Ci, fc U

£ £

F = % frequency of occurrence; I = % crossings heavily invaded; P = prominence value; * species occurring in the given category but not included in a formal recording at a watercourse crossing; t according to

Henderson; $ according to Rutherford & Westfall 1986.

TABLE 7. — Alien species occurring in roadside and veld habitats

80

Bothalia 21,1 (1991)

F = % frequency of occurrence; A = mean abundance rating; P = prominence value; * species occurring in the given category but not included in a formal recording in a road transect, t according to Henderson,

| according to Rutherford & Westfall 1986.

TABLE 7. — Alien species occurring in roadside and veld habitats (continued)

Bothalia 21,1 (1991)

81

F = % frequency of occurrence; A = mean abundance rating; P = prominence value; * species occurring in the given category but not included in a formal recording in a road transect; t according to Henderson;

f according to Rutherford & Westfall 1986.

TABLE 7. —Alien species occurring in roadside and veld habitats (continued)

82

Bothalia 21,1 (1991)

Bothalia 21,1 (1991)

83

Populus x canescens was the next most frequent invader

in the study area (16,7%) and was also most frequently

recorded in moist subtropical and transitional grassland

(33,0%). P. x canescens and S. babylonica were the only

species to be recorded at more than 10% of all watercourse

crossings in the study area.

Other species which were recorded at more than 10%

of crossings in a veld type category were: Acacia dealbata,

Salix fragilis, Prunus persica and Pyracantha angusti-

folia in moist subtropical and transitional grassland, and

Arundo donax and Schinus molle in false karoo (Table 6).

Prominence

Salix babylonica was by far the most prominent riverine

invader in the study area with a prominence value of 104,9

out of a combined total for all species of 200 (Table 6).

It was also the most prominent invader in each of the veld

type categories. Populus X canescens, Acacia dealbata

and Salix fragilis were less prominent but all formed

continuous stands along riverbanks in parts of moist

subtropical and transitional grassland.

Roadside and veld habitats

The whole study area

One hundred and fifty one quarter degree squares and

274 road transects were sampled in which 58 species were

recorded. Up to 21 species were recorded per quarter

degree square. Naturalized species were recorded in 90,1%

of all transects sampled but only 2,6% of all transects were

heavily invaded (Table 5).

Analysis according to veld type

The greatest intensity of invasion was recorded in moist

subtropical and transitional grassland (Table 5). Two-thirds

of the total number of species were recorded here as well

as most (5/7) of the heavily invaded transects. There was

a progressive decrease in the intensity of invasion from

the grassland types in the east to the savanna and karoo

types in the west.

Analysis according to species

Frequency

The most frequently recorded species in the study area

were Opuntia ficus-indica (39,8%), Eucalyptus spp.

(36,5%) and Prunus persica (31,0%). Opuntia sp. cf.

robusta cultivars, Gleditsia triacanthos, Rosa eglanteria,

Pyracantha angustifolia and Acacia dealbata were

recorded in more than 10% of all road transects (Table

7).

The highest percentage frequencies were recorded

in moist subtropical and transitional grassland where

Acacia dealbata, Prunus persica, Eucalyptus spp., Rosa

eglanteria and Pyracantha angustifolia were recorded in

more than 50% of all transects (Table 7).

Prominence

Opuntia ficus-indica scored the highest prominence

value of 24,7 in the study area and in each veld type

category with the exception of moist subtropical and

transitional grassland where Acacia dealbata and Rosa

eglanteria were the most prominent species (Table 7).

Only three species scored abundance ratings of 5 or

more in road transects (Table 2) and these were Acacia

dealbata, Rosa eglanteria and Pyracantha angustifolia.

Species which were occasionally abundant in isolated

localities were Opuntia sp. cf. robusta cultivars, O. ficus-

indica, Cupressus arizonica, Prunus persica and Juniperus

spp.

Patterns of invasion

Alien species are naturalized in streambank, roadside

and veld habitats throughout the province. However, the

greatest intensity of invasion in terms of species diversity

and abundance of invaders was recorded in the eastern

mountain region bordering on Lesotho and Natal (Figures

3, 4 & 5). Within this zone the most invasion was recorded

in the Harrismith, Ficksburg, Fouriesburg and Golden

Gate Districts.

A comparison of Figures 3 & 4, indicating the severity

of invasion in streambank and roadside and veld habitats

respectively, shows similar patterns except that in the

north-eastern region there is more severe invasion of the

streambank habitat than of roadside and veld habitats. This

pattern of streambank invasion is mainly the result of Salix

babylonica invasions.

DISCUSSION

Indigenous versus alien species composition

Only 184 indigenous species of trees and shrubs were

documented for the OFS by Venter & Joubert (1984). Alien

species therefore constitute approximately one third

(70/254) of the total (indigenous and naturalized) tree and

shrub species of the province.

Indigenous species belong to 58 families, the Anacardia-

ceae being the largest with a total of 21 species, 19 of which

are Rhus species. Naturalized alien species belong to 19

families, the largest being the Rosaceae with 16 species.

Alien species of Rosaceae outnumber the indigenous

species by at least three to one. Eleven families are

represented by only alien species. The largest families

which are alien to South Africa are the Pinaceae and

Cactaceae (i.e. assuming that Rhipsalis baccifera is an

early introduction and not indigenous).

Only two indigenous species, Podocarpus latifolius

(family Podocarpaceae) and the very rare Widdringtonia

nodiflora (family Cupressaceae) belong to the Gyrnno-

spermae whereas naturalized alien Gymnosperms number

nine or more species. These alien species belong to the

families Pinaceae (six or more species) and Cupressaceae

(three or more species).

Alien species of the family Salicaceae account for most

riverine invasion in the OFS. Only one indigenous species,

84

Bothalia 21,1 (1991)

a 10 or more species per quarter degree square

m 1 or more river crossings heavily invaded

1 or more river crossings invaded

Mean abundance per km:

[3 less than 2 individuals/groups

2-4 E3 5-8

GD 1 or more rivers invaded, but no formal recordings

FIGURE 3.— Invasion in streambank habitats in terms of the intensity

of invasion of watercourse crossings and species diversity per

quarter degree square.

Salix mucronata , is represented in this family (Immelman

1987) but alien species number five or more.

Prominent and potentially important species

Salix babylonica , the weeping willow, is the most

widespread riverine invader in the OFS. It forms

E3 1 or more road transects invaded

m 1 or more species locally abundant

| o | I or more species invasive, but no formal recordings

FIGURE 4.— Invasion in roadside and veld habitats in terms of the

intensity of invasion of road transects and species diversity per

quarter degree square.

FIGURE 5. — Invasion in roadside and veld habitats in terms of the mean

abundance of invaders per kilometre in each quarter degree square.

continuous stands, some stretching for many kilometres,

along the major perennial watercourses, particularly the

Caledon, Wilge and Vaal Rivers. Although it has been

planted at dams and along rivers for ornament, shade,

fodder and erosion control its wide distribution along

watercourses is most likely due to self (vegetative)

propagation and dispersal by floodwaters. It is probable

that S. fragilis is propagated and dispersed in the same

manner (Henderson in prep.).

There has been some confusion regarding the identity

of Salix fragilis (fide R.D. Meikle pers. comm.). It appears

that the same species was incorrectly referred to as S.

lasiandra by Henderson (1989). This matter will be dealt

with in a paper on invasive Salix species in South Africa

(Henderson in prep.).

Salix fragilis is less widespread than S. babylonica but

also forms pure stands which can stretch for several

kilometres, for example along the Wilge River near

Harrismith. Although aesthetically pleasing and possessing

many beneficial qualities, the Salix species pose a poten-

tial threat to the conservation of indigenous riparian species

and may alter the hydrology of the watercourses they invade

(Henderson in prep.).

There is no information available on water usage by the

alien Salix species, nor any other alien riparian species,

in South Africa (C. Bruwer 1989, Hydrological Research

Institute, pers. comm.; and D. Versfeld 1989, Jonkers-

hoek FRC, pers. comm.). This aspect of alien invasion

needs urgent attention. Rivers in many parts of South

Africa have been heavily invaded by alien woody species

(Macdonald et al. 1986) and although stream flow is

expected to be diminished, there is no evidence. 5.

babylonica and S. fragilis , which flourish along perennial

rivers in the important water catchment areas of the

Bothalia 21, J (1991)

85

25 26 27 28 29 25 26 27 28 29 25 26 27 28 29

27

28

29

30

Acacia baileyana

Acacia dealbata

Agave americana

Arundo donax

Cupressus arizonica

Eucalyptus spp.

27

28

29

30

Gleditsia triacanthos Juniperus spp. Melia azedarach

27

28

29

30

Opuntia ficus-indica

Opuntia sp. cf. robusta cultivars

Pinus spp.

27

28

29

30

FIGURE 6. — Distribution of the most prominent species. Highest abundance rating of 4 or less: •. Highest abundance rating of 5 or more:

roadside and veld habitats, D; streambank habitats. A; streambank, roadside and veld habitats, O.

86

Bothalia 21,1 (1991)

25 26 27 28 29

25 26 27 28 29

25 26 27 28 29

Populus x canescens

Populus nigra

Populus spp.

27

28

29

30

27

28

29

30

Prosopis spp.

Prunus persica

Pyracantha spp.

27

28

29

30

Robinia pseudoacacia

Rosa eglanteria

Salix babylonica

Salix fragilis Schinus molle Sesbania punicea

27

28

29

30

FIGURE 7.— Distribution of the most prominent species. Highest abundance rating of 4 or less: •. Highest abundance rating of 5 or more:

roadside and veld habitats, □; streambank habitats, A; streambank, roadside and veld habitats, O.

Bothalia 21,1 (1991)

87

central interior highlands (Henderson in prep.), might be

significant water users in these parts. Their effects on

stream flow would be most noticeable during times of

drought. There is some evidence that deciduous species,

like the Salix species, have greater water demands than

evergreen species in spring (D.B. Versfeld pers. comm.)

when water supplies are usually at their lowest.

Populus X canescens and Acacia dealbata form dense

stands in places along watercourses. P. x canescens has

a limited ability to spread as it only propagates vegeta-

tively, but its profuse suckering enables it to form very

dense stands. A. dealbata in contrast produces vast

quantities of seed which is efficiently dispersed down

watercourses enabling it to spread far afield; it could there-

fore become an important invader along the Caledon River.

It is most invasive in moist subtropical and transitional

grassland but has the potential to invade watercourses

throughout the grassland areas and it may even penetrate

the drier areas, as it has done in the Transvaal (Henderson

& Musil 1984) and Natal (Henderson 1989). Its seed is

relatively immobile out of water but may be dispersed in

soil by road-building activities. It was sometimes recorded

along roadsides for stretches of several kilometres.

Robinia pseudoacacia is a potentially important riverine

invader as it can form dense suckering stands along water-

courses. Its marked tendency to sucker has been used to

advantage for the reclamation of dongas (Poynton 1973).

It can also propagate from seed. According to Ridley

(1930) its pods are very light and wind dispersed, shed-

ding seed as they travel.

Rosa eglanteria, Acacia dealbata, Pyracantha angusti-

folia and Prunus persica are the most prominent invaders

of veld habitats in the eastern OFS. Rosa and Pyracantha

spp. are potentially the most important invaders of hillside

scrub and surrounding grasslands in this region.

R. eglanteria and P. angustifolia are most invasive in

the Fouriesburg, Clarens and Ficksburg Districts but

are naturalized and locally abundant in the entire

eastern region stretching from the Transvaal border in

the north to the Cape border in the south. They are

particularly abundant along rocky outcrops but also

occur along dongas, fence lines and in open grassland.

They are a potential threat to the conservation of

indigenous woody species, many of which are confined

to rocky outcrops. They may also pose a threat to grazing

lands by replacing valuable grasses with impenetrable woo-

dy thickets.

Both R. eglanteria and P. angustifolia produce an

abundance of brightly coloured fleshy fruits which are

most likely to be dispersed by birds but possibly also by

mammals. R. eglanteria which is a principle weed in New

Zealand (Holm et al. 1979) is largely spread by horses in

that country and is known to be eaten by cattle and goats

in Tasmania (Ridley 1930).

Prunus persica , another member of the Rosaceae is

widespread but appears to have a more limited ability to

spread. It is usually confined to roadsides, railway lines,

disturbed ground around habitation and at bridges over

rivers which suggests that its distribution is largely deter-

mined by the dispersal of its seed by humans. Its fruit

may also be dispersed by crows. Many large-seeded fruits,

including those of Prunus spp. are known to be dispersed

by crows (Ridley 1930).

Several gymnosperms are locally prominent on rocky

outcrops in the eastern OFS. These include Cupressus

arizonica, Juniperus sp. cf. pinchotii, J. virginiana and

Pinus elliottii. Juniperus berries are eaten by many species

of birds as well as small mammals throughout their native

range in the northern hemisphere (Ridley 1930) and it is

therefore likely that they are being dispersed in a similar

manner in the OFS. C. arizonica and P. elliottii seeds are

dispersed by wind. All these species are likely to be

confined to rocky outcrops, ridges and dongas where they

will receive some protection from fire.

Opuntia ficus-indica is one of the most widespread

species in the OFS but occurs only as widely scattered

plants. Its present distribution and abundance in South

Africa is to a large extent controlled by introduced

cactophagous insects (Zimmermann et al. 1986). It is

therefore unlikely to become more abundant in the OFS.

Opuntia sp. cf. robusta cultivars are also widespread but

not abundant.

Eucalyptus spp. have been planted throughout the OFS

for shade and shelter. Natural spread is usually confined

to the immediate vicinity of plantings but watercourses

might provide suitable conditions for their spread further

afield. They were recorded along riverbanks and also on

islands within perennial rivers. Some Eucalyptus spp. have

been described by Brown & Gubb (1986) as moderately

to highly invasive in many seminatural and natural habitats

including riverine systems in the more arid regions to the

west of the OFS.

Prosopis spp. are potentially important invaders of false

karoo. These species are highly invasive in the arid north-

western Cape where they have infested more than 200 000

ha of land (Harding 1987). They are limited to areas with

shallow ground water which are also the most fertile in

these arid regions (Harding 1987).

Relation of invasion to environmental factors

There is a general trend for more invasion in terms of

species diversity and abundance of invaders with increasing

moisture availability from the dry savanna and karoo veld

types in the west to the moist grassland types in the east.

The most invasion occurs in the moist eastern mountain

region bordering on Lesotho and Natal. Here invasion

occurs along the perennial rivers and in terrestrial habitats

particularly where rocky ridges and ravines afford some

protection against fire and frost. Approximately 62% of

the total alien species were recorded here. It has been

estimated by extrapolation from Venter & Joubert (1984)

that approximately 59% of indigenous woody species also

occur in the moist eastern mountain region.

The large representation of alien Rosaceae (about 23%

of the total alien species) may be partly explained in

terms of minimum temperature requirements for seed

germination. Many of the Rosaceae require stratification

to terminate seed dormancy. This requirement may limit

the distribution of species of Rubus, Rosa, Pyracantha and

Cotoneaster to regions where winter temperatures fall

Bothalia 21,1 (1991)

below 5°C (Dean et al. 1986). The whole of the OFS

experiences minimum winter temperatures of below 5°C

and the extreme eastern regions bordering on Lesotho and

Natal, where the alien Rosaceae are most prominent,

experience the highest frequencies of below-freezing

minimum temperatures (Tyson 1986).

Watercourses have played an important role in the

dispersal of several species. Salix babylonica and Acacia

dealbata are good examples. There are no records of S.

babylonica setting seed in South Africa. It spreads vegeta-

tively from branches torn off by floodwaters and deposited

downstream (Poynton 1973). A. dealbata , in contrast,

produces vast quantities of rather immobile seeds. These

seeds however are easily washed downstream enabling it

to spread far from plantings. Stream flow also promotes

the spread of suckering species such as Populus x

canescens , which sends up a multitude of suckers when

its roots become exposed, as well as P. nigra and Robinia

pseudo-acacia.

Animals, particularly birds and mammals, may be

important dispersal agents for many species. Seeds

destined for being swallowed by animals are mainly those

in a pulpy pericarp, being either drupaceous or baccate

(Ridley 1930). A large proportion (47%) of the naturalized

alien woody species in the OFS have fruits that are either

drupes or berries. This includes all the species belonging

to the Rosaceae. Rosa eglanteria and Pyracantha angusti-

folia are most noticeable as invaders of rocky hillside scrub

in the eastern OFS. This vegetation type provides food,

nesting sites and shelter for large numbers of birds and

small mammals (Scheepers 1975).

CONCLUSION

Alien woody invasion is not expected to become a

problem in the greater part of the OFS. The intensity of

invasion is expected to increase the most in the moist

grasslands in the eastern mountain region bordering on

Lesotho and Natal. Here the spread of Acacia dealbata,

Rosa eglanteria and Pyracantha angustifolia needs to be

controlled. In the western drier regions fewer species are

expected to become troublesome. The streambank habitat

however could be threatened by alien invaders such as

Prosopis and Eucalyptus spp.

Acacia dealbata appears to be the most aggressive

invader species in terms of its ability to spread far from

plantings and to produce a large number of individuals

in a given area. It threatens the whole of the eastern

mountain region and especially the perennial rivers. Salix

babylonica is the most widespread riverine invader in the

province and is particularly abundant along rivers in the

important water catchment areas. Water usage by this

species and other alien riparian species needs to be

investigated.

ACKNOWLEDGEMENTS

I thank Mrs H. Joffe of the National Botanical Institute

and Miss J. Rankin of the Plant Protection Research

Institute for their companionship and assistance in the

Field.

REFERENCES

ACOCKS, J.P.H. 1988. Veld types of South Africa, 3rd edn. Memoirs

of the Botanical Survey of South Africa No. 57.

BROWN, C.J. & GUBB, A. A. 1986. Invasive alien organisms in the

Namib Desert, Upper Karoo and the arid and semi-arid savannas

of western southern Africa. In I.A.W. Macdonald, FJ. Kruger

& A. A. Ferrar, The ecology and management of biological

invasions in southern Africa. Oxford University Press, Cape

Town.

DEAN, S.J., HOLMES, P.M. & WEISS, P.W. 1986. Seed biology of

invasive alien plants in South Africa and South West Africa/

Namibia. In I.A.W. Macdonald, F.J. Kruger & A. A. Ferrar, The

ecology and management of biological invasions in southern

Africa. Oxford University Press, Cape Town.

DEPARTMENT OF FOREIGN AFFAIRS AND INFORMATION 1983.

South Africa 1983. Official yearbook of the Republic of South

Africa, 9th edn. Chris van Rensburg Publications, Johannesburg.

HARDING, G.B. 1987. The status of Prosopis as a weed. Applied Plant

Science 1: 43—48.

HENDERSON, L. 1989. Invasive alien woody plants of Natal and the

north-eastern Orange Free State. Bothalia 19: 237-261.

HENDERSON, L. in prep. Alien invasive Salix spp. (willows) in the

grassland biome of South Africa. South African Forestry Journal.

HENDERSON, L. & MUSIL, K.J. 1984. Exotic woody plant invaders

of the Transvaal. Bothalia 15 : 297— 313.

HOLM, L., PANCHO, J.V., HERBERGER, J.P. & PLUCKNETT, D.L.

1979. A geographical atlas of world weeds. Wiley, New York.

IMMELMAN, K.L. 1987. Synopsis of the genus Salix (Salicaceae) in

southern Africa. Bothalia 17: 171—177.

KING, N.L. 1951. Tree-planting in South Africa. Journal of the South

African Forestry Association 21: 1—102.

MACDONALD, I.A.W., KRUGER, F.J. & FERRAR, A. A. (eds) 1986.

The ecology and management of biological invasions in southern

Africa. Oxford University Press, Cape Town.

MUELLER-DOMBOIS, D. & ELLENBERG, H. 1974. Aims and

methods of vegetation ecology. Wiley, New York.

POYNTON, R.J. 1972. Characteristics and uses of trees and shrubs.

Bulletin No. 39. Department of Forestry, Pretoria.

POYNTON, R.J. 1973. Two hundred selected indigenous and exotic

species: how to recognise and grow them. In W.F.E. Immelman,

C.L. Wicht & D P. Ackerman, Our green heritage. Tafelberg,

Cape Town.

RIDLEY, H.N. 1930. The dispersal of plants throughout the world. Reeve,

Kent.

ROBERTS, B.R. 1968. The Orange Free State. In O. Hedberg & I.

Hedberg, Conservation of vegetation in Africa south of the Sahara.

Proceedings of a symposium held at the 6th Plenary Meeting of

AETFAT in Uppsala, 1966. Acta Phytogeographica Suecica 54:

247-250.

RUTHERFORD, M.C. & WESTFALL, R.H. 1986. Biomes of southern

Africa— an objective categorization. Memoirs of the Botanical

Survey of South Africa No. 54.

SCHEEPERS, J.C. 1975. The plant ecology of the Kroonstad and

Bethlehem areas of the Highveld Agricultural Region. D.Sc. the-

sis, University of Pretoria.

SCHULZE, R.E. & MCGEE, O.S. 1978. Climatic indices and classifi-

cations in relation to the biogeography of southern Africa. In

M.J.A. Werger, Biogeography and ecology of southern Africa.

Junk, The Hague.

TYSON, P.D. 1986. Climatic change and variability in southern Afri-

ca. Oxford University Press, Cape Town.

VAN RENSBURG, A.D. 1975. Guide to tree-planting: the highveld and

adjoining districts of the Transvaal, OFS and Cape Province.

Pamphlet No. 163. Department of Forestry, Pretoria.

VENTER, H.J.T. & JOUBERT, A.M. 1984. Climbers, trees and shrubs

of the Orange Free State. De Villiers, Bloemfontein.

ZIMMERMANN, H.G., MORAN, V.C. & HOFFMANN, J.H. 1986.

Insect herbivores as determinants of the present distribution and

abundance of invasive cacti in South Africa. In I.A.W. Macdonald,

F.J. Kruger & A. A. Ferrar, The ecology and management of

biological invasions in southern Africa. Oxford University Press,

Cape Town.

Bothalia 21,1 (1991)

89

APPENDIX

The following is a species checklist of all naturalized alien trees and shrubs recorded during the survey as well as a few additional species cited

on National Herbarium specimen labels (PRE). Some non-woody species are included.

Acacia

baileyana F.J. Muell., Bailey’s wattle

cultriformis A. Cunn., knife-leaved wattle

dealbata Link, silver wattle

decurrens (J.C. Wendl.) Willd., green wattle

mearnsii De Wild., black wattle

Agave americana L., century plant

Ailanthus altissima (Mill.) Swingle, tree-of-heaven

Arundo donax L., giant reed

Atriplex sp.

Caesalpinia gilliesii (Wallich. ex Hook.) Benth., bird-of-paradise

Cedrus deodara (D. Don) G. Don, deodar

Cereus peruvianus (L.) Mill., queen of the night

Cestrum laevigatum Schlechtd. (PRE), inkberry

Cortaderia sp. , pampas grass

Cotoneaster

franchetii Bois. (PRE), orange cotoneaster

pannosus Franch.

sp., cotoneaster

Crataegus

X lavallei Herincq, Lavallee thorn

monogyna Jacq. (PRE), English hawthorn

Cupressus

arizonica Greene

var. glabra (Sudw.) Little, smooth-barked Arizona cypress

var. montana (Wiggins) Little, Arizona cypress

spp. , cypresses

Cydonia oblonga Mill., common quince

Cytisus scoparius (L.) Link (PRE), Scotch broom

Eucalyptus

tereticornis Sm.. forest red gum

spp., gums

Fraxinus americana L., American ash

Gleditsia triacanthos L., honey locust

Juniperus spp. (J. virginiana L.. red cedar; cf. J. pinchotii Sudw., red-

berry juniper and possibly other species)

Ligustrum sp., privet

Malus pumila Mill. var. paradisiaca (Medic.) C.K. Schneid., paradise

apple

Melia azedarach L., syringa

Morus alba L.. white mulberry

Nicotiana glauca R.C. Grah.. wild tobacco

Opuntia

ficus-indica (L.) Mill., sweet prickly pear

imbricata (Haw.) DC., chain-link cactus

spinulifera Salm-Dyck (PRE), large round-leaved prickly pear

sp. cf. robusta cultivars, spineless prickly pears

Pinus

elliottii Engelm., slash pine

patula Schlechtd. & Cham., patula pine

? taeda L., loblolly pine

sp. cf. P. halepensis Mill., aleppo pine

sp. cf. P. uncinata Mill, ex Mirb.

spp. , pines

Populus

x canescens (Ait.) J.E. Sm., grey poplar

nigra L. var. italica Muenchh., Lombardy poplar

spp. (cf. P. deltoides Bartr. ex Marsh., match poplar and possibly P.

wislizenii Sarg., valley match poplar)

Prosopis spp (P. glandulosa Torr. var. glandulosa, mesquite; cf. P

glandulosa Torr. var. torreyana, mesquite; P. velutina Wooton

(PRE), velvet mesquite)

Primus

armeniaca L., common apricot

persica (L.) Batsch., peach

Punica granatum L., pomegranate

Pyracantha

angustifolia (Franch.) C.K. Schneid., yellow firethorn

fortuneana (Maxim.) H.L. Li

rogersiana (A.B. Jacks.) Bean

Pyrus sp., pear tree

Quercus robur L., English oak

Robinia pseudoacacia L., black locust

Rosa eglanteria L., eglantine

Rubus sp. , bramble

Salix

babylonica L., weeping willow

fragilis L. (fide R.D. Meikle, pers. comm.), basket willow

Schinus mode L., pepper tree

Sesbania punicea (Cav.) Benth., red sesbania

Sophora sp. cf. S. davidii (Franch.) Skeels

Tamarix chinensis Lour., Chinese tamarisk

Trichocereus sp.

Yucca sp. cf. Y. aloifolia L., Spanish bayonet

Bothalia 21,1: 91-107 (1991)

Multivariate analysis of coastal grasslands at Mkambati Game Reserve,

north-eastern Pondoland, Transkei

C.M. SHACKLETON*, J.E. GRANGER** B. MCKENZIE+ and M.T. MENTIS+

Keywords: classification, grasslands, ordination, Pondoland, Transkei

ABSTRACT

The grassland vegetation on the coast of north-eastern Pondoland was analysed after sampling of 113 quadrats in Mkambati

Game Reserve. Data were summarised using TWINSPAN and DECORANA multivariate procedures. Several communities

and subcommunities are recognized and described in relation to measured abiotic variables. A primary gradient from the

shore inland is evident, corresponding with changes in altitude, soil conductivity, soil organic matter and soil texture. Time

since the last fire influences the invasion of shrubland species.

UITTREKSEL

Die grasveld aan die kus van noordoostelike Pondoland is ontleed nadat monsters van 113 monsterpersele in die Mkambati-

wildtuin geneem is. Data is d.m.v. TWINSPAN- en DECORANA-veelvoudige prosedures opgesom. Verskeie gemeenskappe

en subgemeenskappe word erken en met betrekking tot gemete abiotiese veranderlikes beskryf. Daar is 'n primere gradient

vanaf die kuslyn na die binneland wat ooreenkom met verskille in hoogte bo seevlak, grondgeleidsvermoe, organiese inhoud

en tekstuur van die grand. Tydsverloop sedert die laaste veldbrand beinvloed die indringing van struikspesies.

CONTENTS

Introduction 91

Study area 92

Location 92

Geology and geomorphology 92

Soils 92

Climate 92

Rainfall 92

Temperature 93

Vegetation 93

Fauna 93

Human impact 94

Historical 94

Present management (1978 to date) 94

Methods 94

Results 95

Classification of the grasslands 95

Ordination of the grasslands 96

Grassland communities at MGR 98

Themeda triandra—Centella asiatica Dwarf

Grassland Community 99

Tristachya leucothrix—Loudetia simplex Short

Grasssland Community 101

Tristachya l eucoth rix — Ath rixia phylicoides Short

Grassland Subcommunity 101

Festuca costata—Albuca setosa Medium Grass-

land Subcommunity 102

Stoebe vulgaris— Athanasia calva Short Shrub

Grassland (heathland) Subcommunity 102

*- Botany Dept, Univ. Transkei, Private Bag XI, Unitra 5100. Present

address: Wits Rural Facility. P.O. Box 7, Klaserie 1381.

** Botany Dept, Univ. Transkei, Private Bag XI, Unitra 5100. Present

address: Botany Dept, Univ. Natal, P.O. Box 375, Pietermaritzburg 3200.

+ Botany Dept, Univ. Transkei, Private Bag XI, Unitra 5100. Present

address: Botany Dept, Univ. Western Cape, P. Bag X17, Bell ville 7535.

t Botany Dept, University of the Witwatersrand, P.O. Wits 2050.

MS. received: 1989-11-30.

Cymbopogon validus-Digitaria natalensis

Medium rassland Community 103

Aristida junciformis—Helichrysum mixtum Short

Grassland Community 104

Discussion 104

Evaluation of the classification and ordination

results 104

Comparison with other coastal areas of Transkei 105

Comparison with other sour grasslands 105

Fire and ‘invasion’ of fynbos species 105

Acknowledgements 105

References 106

INTRODUCTION

Areas of uniform species composition and structure

commonly reflect a particular range of uniform environ-

mental variables (Mueller-Dombois & Ellenberg 1974).

Delineation of such homogeneous areas is desirable since

it is assumed that they would require the same manage-

ment intervention throughout the total uniform area to

achieve specified management goals (Edwards 1972).

Thus, description and classification of homogeneous

vegetation units forms the primary basis for delineation

of homogeneous physiographic units for management

purposes.

The main aim of the present contribution towards a

larger study at Mkambati Game Reserve (MGR), was to

describe and map the vegetation of the reserve in order

to assess the primary productivity of coastal grasslands

(Shackleton 1989), and to delineate management units. As

more than 80% of the reserve is grassland and this was

the most intensively exploited vegetation type, quantita-

tive sampling was restricted to this formation. Specific

aims of this paper were to determine the range in

physiognomy, species composition and spatial distribution

92

of the various grassland communities at MGR; and to

correlate these with selected biotic and/or abiotic factors.

STUDY AREA

Location

Mkambati Game Reserve (7 720 ha) is situated on the

coast of north-eastern Pondoland, Transkei (31° 13'— 20'S

and 29° 55'-30° 04'E) (Figure 1). It is bounded by the

Mtentu River to the north and the Msikaba River in the

south, with approximately 12 km of coastline forming the

eastern boundary. The only non-natural boundary is the

inland fence in the west (+ 300 m a.s.l.). MGR is charac-

terized by a combination of geomorphic, edaphic and

phytogeographic factors that are not found elsewhere along

the coast of southern Africa (Feely 1986).

Geology and geomorphology

The reserve is underlain by Palaeozoic pre-Karoo sedi-

ments of the Natal Group sandstone (Johnson & Meyboom

1976). Although doleritic intrusions are common in

Transkei, very few are present in this area (Du Toit 1912;

McKenzie 1984), but several small localized outcrops were

noted at MGR during the course of this study. A small

outcrop of Upper Cretaceous deposits is present at the

coast between the Mtentu River (northern boundary) and

the Mgwetyana stream within the reserve to the south

(Du Toit 1912; Geological Survey 1976). Between the rivers

and streams the land-sea interface is marked by generally

horizontal, abrupt rock outcrops of the Msikaba Formation

which is relatively resistant to weathering except along

lines of weakness (Feely 1986). Consequently, both the

FIGURE I — Location map of Mkambati Game Reserve.

Bothalia 21,1 (1991)

shore and foreland are homeostatic, i.e. neither prograding

nor degrading (Tinley 1985).

Situated at the southern end of the Natal monocline, the

terrain at MGR is a typical example of the tableland topo-

graphy described by Kruger (1983). Land facets are orien-

tated parallel to the coast except where drainage systems

have cut across them at right angles. Most of the slopes

are less than 5,0%, except on scarp surfaces, so that the

average gradient from the sea (the eastern boundary) to

the inland boundary (300 m a.s.l.) is 3,8%.

This gentle topography is the result of intense planation

since the Tertiary. It is interrupted by two steps parallel

to the coast that indicate surf-scars during intervals of past

geologic uplift (Du Toit 1912; King 1963; Tinley 1978).

These steps are evident at 85 m and 190 m and are charac-

terized by steeper surface gradients ranging between 10%

and 15%. They mark the position of past shorelines.

Soils

Being derived from Natal Group sandstones under a high

rainfall regime, the soils of MGR are acidic, dystrophic,

and sandy (Tinley 1978; Shackleton 1989). They are also

characterized by a weak structure, high permeability and

low available moisture capacity (Maud 1966).

Shackleton (1989) determined that the soil forms

covering the largest areas were Mispah (64,7%), Clovelly

(15,9%), Champagne (6,8%) and Pinedene (2,2%)

[nomenclature follows MacVicar etal. (1977)]. Other forms

encountered included Glenrosa, Hutton, Katspruit and

Kroonstad. Berea Formation sands form an irregular belt

across the reserve close to the shoreline.

Other than tfiose forms characterized by shallow sub-

soil horizons (e.g. Mispah and Glenrosa), the soils at MGR

are, on average, deep ( > 1,2 m). Ferricrete is not prevalent

and the primary limitation to rooting depth is saprolite and

wet season waterlogging. In some areas the saprolite itself

forms a deep, highly decomposed horizon ( > 1,0 m). Signs

of gleying and gley horizons are common in profiles in

the vicinity of the base of the slope (Shackleton 1989).

Climate

Mkambati Game Reserve has a humid, temperate

climate according to the Koppen classification (Schulze

1947). On a local scale, details of the major climatic

variables are given below.

Rainfall

The average annual rainfall is 1 200 mm (56 years data),

of which 61% is received during spring and summer

(September to February). A minimum of 50 mm is

expected every month. The monthly distribution of the

annual rainfall is illustrated in Figure 2. The nature of the

rain is usually cyclonic but more convectional during

summer. Thunderstorms occur, on average, twenty days

per annum, generally restricted to early spring (Tyson

1986).

Analysis of the distribution of annual rainfall revealed

that the median is close to the mean, reflecting a relative-

ly even distribution (Figure 3).

Bothalia 21,1 (1991)

93

FIGURE 2. — Walter-Lieth diagram for Mkambati Game Reserve.

Temperature

Characteristic of its coastal location, MGR has a

relatively equable temperature regime with respect to both

diurnal and seasonal variations. Data for Port St Johns

(50 km to the south) indicate that the warmest months are

January and February (see Figure 2). Minimum tempera-

tures are recorded during July and August. Frost is absent.

Diurnal temperature variations are relatively small, even

in winter. Differences in the mean temperature at 08h00

and 12h00 are less than 6°C for all months of the year

(Weather Bureau 1986).

Vegetation

Biogeographically MGR is situated in the Indian Ocean

Coastal Belt (Werger 1978), which extends along the

eastern seaboard of Africa from south-eastern Somalia to

Port Elizabeth (RSA). Due to abrupt small-scale changes

in climate and soils within this belt, it is divided into a

mosaic of various vegetation types ranging from edaphic

grasslands to tall forest (White & Moll 1978). The southern

portion, referred to as the Tongoland-Pondoland Regional

Mosaic, extends from the Limpopo River (25 °S) to Port

Elizabeth.

The vegetation of MGR is a typical example of the

edaphic grasslands that constitute part of the Tongoland-

Pondoland Mosaic. Recent work by Feely (1986) indicated

that significant portions of the coastal belt (including north-

eastern Pondoland) have been dominated by grasslands for

at least the last 2 000 years, but more probably through-

out the Holocene. Consequently, by implication, the

present disjunct, small patches of forest were not signifi-

cantly more extensive than at present for at least 10 000

years, contrary to general belief.

Productivity is high, permitting a high burning fre-

quency of two or three times per annum (White & Moll

1978; pers. obs.). This high burning frequency has

promoted the dominance of fire-tolerant species such as

Tristachya leucothrix, Trachypogon spicatus and Themeda

triandra and, according to White & Moll (1978), it also

promotes invasion and dominance by Aristida junciformis ,

if coupled with selective grazing.

On a finer scale the Tongoland-Pondoland Mosaic

includes Acocks’s (1988) veld type nos 1, 2, 3, 5, 6, 10,

23 and 24. By location, MGR should be classified as veld

type no. 1, Coastal Forest and Thornveld. However,

Shackleton (1989) argues that the predominance of

grassland and its species composition make it more akin

to Pondoland Coastal Plateau Sourveld (veld type no. 3).

Six major physiognomically different vegetation types

are clearly discernible at MGR. Most of these can be

subdivided into several communities on the basis of

physiognomy and species composition (Table 1).

The MGR lies in the sandstone region of southern

Natal/Pondoland which is significant in terms of its

conservation status through the presence of a relatively

large number of endemic angiosperms (Wildlife Society

of southern Africa 1977; Van Wyk & Schrire 1986) and

outlying taxa that exhibit no contiguity with the rest of

the taxon’s distribution. To date, one family (Rhyncho-

calycaceae), six genera (e.g. Jubaeopsis , Pseudoscolopia)

and more than 50 species have been listed as endemic to

the region, 40 or more of which have been noted in the

reserve (A.E. van Wyk pers. comm.).

Fauna

The reserve supports approximately 1 600 introduced

wild ungulates, mostly concentrate grazers (Shackleton

1989). The most abundant species are blesbok ( Damalis -

cus dorcas phillipsii ), blue wildebeest ( Connochaetes

taurinus), eland (Taurotragus oryx) and red hartebeest

(Alcelaphus buselaphus). Non-introduced species such as

reedbuck (Redunca arundinum), bushbuck ( Tragelaphus

scriptus) and duiker (Sylvicapra grimmia ) exist in small

numbers.

1 2

ANNUAL RAINFALL (x 102 mm)

FIGURE 3. —Frequency histogram of annual rainfall.

94

Bothalia 21,1 (1991)

TABLE 1. — Major communities at Mkambati Game Reserve (areas are approximate)

Human impact

Historical

One of the factors contributing to the proclamation of

Mkambati as a conservation area was its low settlement

density which minimised translocation of rural people to

alternative sites. The low density settlement of this entire

tableland region had certainly prevailed for the last 2 000

years (Feely 1986) and it is unlikely that it was any higher

before that. Possible reasons for this feature were discussed

by Bigalke (1979), Hall (1981) and Feely (1986). The

occurrence of marine shell-middens along the coast and

in several rock shelters further inland indicates exploita-

tion of intertidal resources since at least the Middle Stone

Age (Feely 1986; Prins & Strever 1987). It is also likely

that these hunters fired these grasslands on a frequent basis

(Hall 1984; McKenzie 1984), thereby maintaining a long

period of fire occurrence.

Present management (1978 to date)

The present area was stocked with wild herbivores in

1979 to be managed as a hunting concern. This was dis-

solved in 1983 and a Board of Directors was appointed

by the Government of Transkei. This body is presently

responsible for management policy and decisions.

The reserve is divided into six management ‘blocks’ of

approximately equal size (A— F). Prescriptions are that

each block is burnt on a biennial basis so that 50% (three

blocks) of the reserve is burnt each year. Due to poor fire

control Prionium serratum wetlands, swamp forests and

coastal forests are frequently burnt.

The provision of concentrated supplementary feeds

during the winter ceased in the early 1980’s. However,

localized patches characterized by a different species

composition persist at these sites.

METHODS

Data were collected in the field using the Braun-Blanquet

approach (see Mueller-Dombois & Ellenberg 1974; Werger

1974). This was adopted in preference to a more quantita-

tive sampling method (such as the wheel-point) as it pro-

vides a more comprehensive list of species (Novellie &

Strydom 1987), an important consideration since the flora

of MGR had been poorly collected before this study was

begun.

One hundred and thirteen quadrats (5 m x 5 m) were

distributed judgementally throughout the reserve. The

distribution of quadrats in each vegetation unit, provi-

sionally recognized during reconnaissance visits (on the

basis of physiognomy, dominant species composition and

abundance), was in approximate agreement with the

contribution of that community to the total area, but with

bias towards the smaller units (a minimum of three

quadrats per unit as far as possible). The following

provisional units were identified: (i) Tristachya leucothrix—

Loudetia simplex unit (assigned the prefix of ‘L’ — 78

quadrats), (ii) Cymbopogon validus—Digitaria natalensis

unit (‘C’ — 15 quadrats), (iii) Themeda triandra—

Centella asiatica unit (‘T’ — 8 quadrats), (iv) unburnt

moribund areas dominated by Stoebe vulgaris and

Athanasia calva (‘S’ — 4 quadrats), (v) several small areas

dominated by Aristida junciformis (‘A’ — 3 quadrats) or (vi)

by Festuca costata (‘F’ — 2 quadrats), and a (vii) Watsonia

sp. ( Shackleton 426) unit (‘W’ — 3 quadrats). Sampling

was carried out between November and February in areas

burnt in * the previous winter (except the Themeda

triandra— Centella asiatica unit which foiled to bum). This

period coincided with the flowering period for most

Bothalia 21,1 (1991)

95

species which greatly aided identification. As only 50%

of the reserve is burnt each year, field sampling was spread

over two growing seasons (1985/1986 and 1986/1987).

The following variables were recorded for each quadrat:

(i) all species observed and an estimate of their cover-

abundance on the Braun-Blanquet scale as modified by

Werger (1973), (ii) gradient (Abney level), (iii) altitude

(from 1:10000 ortho-photographs), (iv) aspect (compass),

(v) rockiness (estimated as a percentage of the ground

cover), (vi) soil form according to the Binomial Classifi-

cation of southern Africa (MacVicar et al. 1977) (auger

sample), (vii) soil depth (auger sample). A soil sample

from the A-horizon was taken at the centre of each quadrat.

These were analysed in the laboratory for pH (in KC1),

texture (Andressian pipette method), conductivity (con-

ductivity cell) and organic carbon (Walkley-Black method)

according to the methods given by the Fertilizer Society

of South Africa (1980).

Species data were subjected to analysis via TWINSPAN

and DECORANA (Hill 1979a, b) after conversion of the

cover-abundance values to a scale of 1—9. For descrip-

tions and summaries of the basic algorithms, see Hill &

Gauch (1980); Gauch (1982); Scotcher (1982) and Conlong

(1986). In each case default values of the programmes

yielded the most satisfactory results, after several iterations

with altered pseudospecies cutoff levels and/or unequal

weightings for species of differing abundance.

The ordination score of each quadrat on the first two

axes was correlated (product-moment correlation

coefficient) with each of the measured environmental

variables. To permit correlation of aspect with the

ordination scores, aspects greater than 180° were converted

to the mirror-image of aspects less than 180° as described

by Wikum & Wali (1974). This was done after correction

of the compass readings for the magnetic declination. For

quadrats with a recorded soil depth greater than 1,2 m,

a value of 1,8 m was used in the calculation of correlation

coefficients and mean depths per community.

It was probable that there would be non-linear associa-

tions between vegetation types and site characteristics

which would not be evident from the linear correlation

of axis score and environmental variables. This was

checked by means of Correspondence Analysis (CA)

described by Greenacre (1986). Organic matter was not

included in the CA because of missing values.

Final description of the grassland communities and sub-

communities was drawn from the interpretation of the

results of TWINSPAN and DECORANA and field notes.

Physiognomic nomenclature followed that of Phillips (1971)

rather than Edwards (1983), as the height divisions

suggested by Phillips (1971) resulted in a more meaningful

physiognomic differentiation of the various grassland com-

munities at MGR. Edwards’ limits resulted in most of the

vegetation units being classified into the same category

and there was, therefore, no distinction between units

except on floristics. To facilitate identification of com-

munities by management personnel, usually unfamiliar

with species identification, it was considered important

to be able to differentiate the vegetation units at MGR on

the basis of physiognomy, where possible. Phillips (1971)

classified grasslands as dwarf (<150 mm), short (150—

450 mm), medium (460—1 000 mm), tall (1,0— 2,0 m) and

high ( > 2 ,0 m).

The vegetation map (Figure 8) was drawn from ground

survey using 1:30 000 aerial photographs and 1:10000

ortho-photographs and results obtained from the multi-

variate analyses.

To provide more detailed data on species composition

to act as a baseline for further studies, sixteen wheel-point

samples (Tidmarsh & Havenga 1955) were divided

between the two most extensive grassland communities.

Each sample consisted of 200 points.

The plant names used are according to Gibbs Russell

et al. (1985, 1987). Specimens were identified by staff at

the University of Transkei Herbarium, the National

Botanical Institute (Pretoria), Mr B.D. Schrire, the SALO

at Kew and Dr O.M. Hilliard at the University of Natal,

Pietermaritzburg.

As a consequence of the inability to distinguish with

certainty between vegetative material of Rendlia altera,

Diheteropogon filifolius, Elionurus muticus, Koeleria

capensis and Bulbostylis schoenoides in all quadrats, they

were collectively recorded as ‘wire grasses’. Specimens

were housed at the National Botanical Institute, Pretoria,

and the Herbarium, University of Transkei.

RESULTS

Classification of the grasslands

The resultant hierarchical division of the sample plots

from the TWINSPAN analysis is given in Figure 4.

The short grassland immediately adjacent to the coast

was recognized as being very different from the remainder

of the reserve, with all eight quadrats (plus one from the

provisional Watsonia sp. vegetation unit) being separated

from the rest of the data set at the first level of division.

Indicator species were Centella asiatica and Ruellia

cordata. At the second level of division the included

Watsonia sp. quadrat was split from the eight Themeda

triandra— Centella asiatica quadrats. Further divisions of

this community were not recognizable in the field and were

therefore ignored in the final dendrogram.

The main group of 104 quadrats on the positive side of

the first division was divided into two groups of almost

equal size at the second level. Each consisted of quadrats

differing widely in floristics, physiognomy and habitat.

Thus, this level did not yield meaningful results for

classification.

On the other hand, the third level of division succeeded

in reducing each of these groups to recognizable units. In

the first instance, three quadrats that represented unburnt

grasslands invaded by woody species, primarily Stoebe vul-

garis and Athanasia calva, were separated as a unit. These

two species acted as indicators for the positive side of the

division. The fourth quadrat dominated by these two spe-

cies was not included in this group, probably because it

had a greater number of species (50 as opposed to 34,

27 and 17 for the other three quadrats), giving it more

AXIS 1 2 AXIS

96

Bothalia 21,1 (1991)

T. triandra- C. asiatica

samples

i'tr"

' 'Ll L

C. v a lid us -

D. natalensis

samples

C , LL, L'-'Ll L L LL L L L- simp

/ C-l L L L L.LL

L LLL LL Lk LL L \

T. leucothrix-

L. simplex samples

L L L . t,W

L LLL ^L 4,4. LLL LLL L ,

L L L L LL L L L1

L L L L I

LLL |

S vulgaris-

A. calva samples

AXIS 2

GURE 5. — DECORANA ordination of the coastal grassland communities of Mkambati Game Reserve (all samples): T, Themeda triandra—

Centella asiatica samples; C, Cymbopogon validus—Digitaria natalensis samples; L, Tristachya leucothrix—Loudetia simplex samples;

S, Stoebe vulgaris— Athanasia calva samples; F, Festuca costata—Albuca setosa samples; A, Aristida junciformis—Helichrysum mixtum

samples; W, Watsonia sp. samples.

A

C. validus- D. natalensis

samples

C C

/

/ -t

VA'1'

/ Lt L'

/ L L'

I L L L ,

I ' LU , t

l ' L \ L - L

1 L LL x '-,L \

\ L YwW. L \

\

L Lk‘lLLL "■

\

L L L

l LL LLL

L L L L

T. leucothrix- L L L

L simplex

samples

LL L

L L

L

L

AXIS 2

130 150

r)o

190 210

FIGURE 6. —DECORANA ordination omitting samples from the Themeda triandra— Centella asiatica and Stoebe vulgaris— Athanasia calva samples

(compare to Figure 5).

Bothalia 21,1 (1991)

97

FIGURE 4. — Dendrogram of TWINSPAN classification: 1, Themeda

triandra — Centella asiatica community; 2, Cymbopogon

validus—Digitaria natalensis community; 3, Tristachya

leucothrix — Loudetia simplex community; 4, Stoebe vulgaris —

Athansia calva subcommunity; 5, Festuca costata—Albuca seto-

sa subcommunity; 6, Tristachya leucothrix— Athrixia phylicoides

subcommunity.

species in common with the Tristachya leucothrix—

Loudetia simplex community on the other side of the

division at level two. From the field situation this is a

‘mis-classification’ as these four quadrats were the only

ones characterized by woody species (shrubs) and were

therefore very distinct from the rest of the data set.

The 45 quadrats on the negative side of this division

included samples from four of the provisional vegetation

units identified during reconnaissance visits. Successive

divisions of this group, at the fourth, fifth and sixth levels,

isolated the two quadrats dominated largely by Festuca

costata, although it was not selected as an indicator species

in the TWINSPAN analysis.

Most of the Cymbopogon validus—Digitaria natalensis

quadrats were subsequently separated from the Tristachya

leucothrix— Loudetia simplex quadrats at level four.

Divisions within the Cymbopogon validus—Digitaria

natalensis quadrats appeared to have few practical

implications beyond level four. The exception to this were

quadrats C23, C26 and C28 which were all situated in

management block A. This was of interest considering that

the T. leucothrix— L. simplex quadrats of this block were

also recognized as distinct from others of this community

at level six. Similarly, soils of areas in this block dominated

by C. validus and D. natalensis are different to other areas

of the reserve also dominated by these two species (S.E.

Shackleton 1989). The TWINSPAN results, therefore,

indicate that the same holds true for the areas not

dominated by C. validus and D. natalensis. The lower level

divisions also separated quadrats of this community

situated on Hutton (form) soils, but included these with

quadrats intended to characterize areas dominated by

Aristida junciformis.

The partitioning of the A. junciformis quadrats between

two groups at the fourth level, and the failure to isolate

them by the sixth level, leads us to the conclusion that these

areas of the reserve do not merit recognition as a separate

community for management purposes. These areas

obviously share a large number of species with other com-

munities, and are therefore not floristically distinct.

More difficult to interpret was the major division of the

Tristachya leucothrix— Loudetia simplex quadrats at the

second level. The majority of quadrats (55) form a major

group that is successively divided into smaller groups at

the lower levels. The remaining 23 quadrats were included

on the positive side at level two, along with the Cymbopo-

gon validus—Digitaria natalensis and Festuca costata

quadrats. Several of the measured environmental variables

associated with this group of quadrats were intermediate

between those of the pre-recognized C. validus— D.

natalensis and T. leucothrix— L. simplex communities (see

Table 5).

Ordination of the grasslands

From Figure 5 it is evident that the quadrats of both the

Themeda triandra— Centella asiatica and Stoebe vulgaris—

Athanasia calva samples form distinct units at the ends

of the first and second ordination axes, respectively. To

assist in the interpretation of these axes, three additional

iterations were performed successively: 1, omitting the

T. triandra- C. asiatica quadrats only; 2, omitting the

S. vulgaris -A. calva quadrats only; and 3, omitting

quadrats from both these two communities. The last case

is presented in Figure 6.

The clusters obtained for the first and second axes

tended to substantiate the groups identified in the TWIN-

SPAN classification. The Cymbopogon validus—Digitaria

natalensis quadrats were grouped together in both

ordination diagrams. Furthermore, the Tristachya

leucothrix -Loudetia simplex quadrats that were included

on the same side of the TWINSPAN dichotomy at level

3, along with the C. validus— D. natalensis quadrats, are

in an intermediate position between the two major com-

munities. During field sampling these quadrats were not

recognized as having close affinities with the C.

validus— D. natalensis community because they lacked

these two physiognomically distinctive species.

Interpretation of the first two ordination axes was

assisted through linear correlation of the ordination scores

with each of the measured environmental variables. These

have been summarized in Table 2.

These results indicated that there was a complex of

interrelated environmental variables (see Table 3)

correlated with the pattern illustrated in the ordination di-

agram. The primary axis, with an eigenvalue of 0,409, was

highly correlated with soil conductivity, altitude (negative)

and organic matter, and marginally correlated with

the amounts of sand (negative) and clay (all quadrats

98

Bothalia 21,1 (1991)

TABLE 2. — Correlation coefficients between the ordination scores and measured environmental variables

All samples (Figure 5) Excluding extreme samples (Figure 6)

considered). The first two were to be expected, as there

was an increase in altitude and distance from the sea across

the whole reserve. Such correlations have been shown

elsewhere for coastal regions (e.g. Westman 1981) and are

associated with the concomitant reduction in sodium

chloride input. The second axis (eigenvalue = 0,233)

showed correlation with the same variables and pH.

However, the degree of correlation was less for soil

conductivity, but greater for the amounts of sand and clay

(negative). This indicates some degree of interaction

between the two axes. There was a strong degree of

interaction between altitude and both soil texture and

conductivity. The third and fourth axes had eigenvalues

of 0,150 and 0,107 respectively, and were not considered

further. Omission of the Themeda triandra—Centella

asiatica and Stoebe vulgaris— Athanasia calva quadrats did

not alter the correlations except to strengthen the degree

of correlation with soil texture along the primary axis.

The results from Correspondence Analysis (CA) were

in agreement with those obtained above. They are

presented in Figure 7 and the breakdown of row and

column contributions are provided in Table 4.

Axes one and two accounted for 69,2% and 27,7% of

the inertia respectively. The greatest correlations with axis

one were soil conductivity and altitude (negative). Thus,

one extreme was characterized by the Themeda triandra—

Centella asiatica quadrats associated with sandy soils, high

soil conductivity, high slope and low altitude. At the

opposing extreme were the Cymbopogon validus—

Digitaria natalensis quadrats at higher altitudes, with clay

soils and low soil conductivity. The Tristachya leuco-

thrix-Athrixia phylicoides quadrats were intermediate

between the Tristachya leucothrix—Loudetia simplex and

Cymbopogon validus— Digitaria natalensis quadrats.

Environmental variables most closely associated with

the second axis were altitude and soil depth. The Stoebe

vulgaris— Athanasia calva quadrats were strongly asso-

ciated with pH and soil depth. This was the only feature

highlighted by CA not evident from the previous analyses.

Grassland communities at MGR

It was evident from synthesis of the above results that

several distinct grassland units were present in MGR.

TABLE 3. — Correlation coefficients between the measured environmental variables

Bothalia 21,1 (1991)

99

FIGURE 7. — Correspondence analysis scatterplot of coastal grassland

communities and subcommunities and associated abiotic variables

at Mkambati Game Reserve.

Small areas within the major communities differing with

respect to only the dominant species, or their ‘invasion’

by other species, were designated as subcommunities. The

following descriptions of each community deal with the

floristically dominant and more easily identified species.

The spatial distribution of these communities is depicted

in Figure 8. The name assigned to each community was

drawn from two species, the first being the physiognomic

dominant which was usually abundant, and the second

being an indicator species identified by the TWINSPAN

analysis.

Rue Ilia cordata. This community had many preferential

species, the most easily identifiable in the field being

Euryops leiocarpus, Stenotaphrum secundatum, Geranium

ornithopodum, Moraea spathulata and a short, slender

form (ecotype?) of Themeda triandra . The last three were