ISSN 0006 8241 = Bothalia

Bothalia

A JOURNAL OF BOTANICAL RESEARCH

Vol. 26,1 May 1996

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BOTHALIA

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Now available:

Molteno Formation (Triassic) Vol. 1. Introduction. Dicroidium, by J.M. & H.M. Anderson.

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

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BOTHALIA

A JOURNAL OF BOTANICAL RESEARCH

Volume 26,1

Scientific Editor: O.A. Leistner

Technical Editor: B.A. Momberg

NATIONAL

Botanical

INSTITUTE

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NASIONALE BOTANIESt

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1 NATIONAL BOTANICAL

INSTITUTE

ISSN 0006 8241

May 1996

Editorial Board

D.F. Cutler

B.J. Huntley

P.H. Raven

J.P. Rourke

M.J. Werger

Royal Botanic Gardens, Kew, UK

National Botanical Institute, Cape Town, RSA

Missouri Botanical Garden, St Louis, USA

Compton Herbarium, NBI, Cape Town, RSA

University of Utrecht, Utrecht, Netherlands

CONTENTS

Volume 26,1

l 1. Four new species and one new subspecies of Lachenalia (Hyacinthaceae) from arid areas of South

Africa. G.D. DUNCAN 1

/ 2. Saxicolous species of the genus Rinodina (lichenized Ascomycetes, Physciaceae) in southern

Africa. M. MATZER and H. MAYRHOFER 11

■J 3. FSA contributions 4: Agavaceae. G.F. SMITH and M. MOSSMER 31

,/ 4. FSA contributions 5: Buxaceae. H.F. GLEN 37

5. Notes on African plants:

/ Apiaceae. A new species of Centella. M.T.R. SCHUBERT and B.-E. VAN WYK 50

d Celastraceae. Correct orthography and author citation for Elaeodendron. R.H. ARCHER and

A.E. VAN WYK 41

sj1 Convolvulaceae. New records, name changes and a new combination in southern Africa. A.D.J.

MEEUSE and W.G. WELMAN 46

J Crotalarieae. Rare or extinct species of Argyrolobium . T.J. EDWARDS 42

v 6. Cytogenetic studies in some representatives of the subfamily Pooideae (Poaceae) in South Africa. 1.

The tribe Aveneae, subtribe Aveninae. J.J. SPIES, S.K. SPIES, S.M.C. VAN WYK, A.F. MALAN

and E.J.L. LIEBENBERG 53

d 7. Cytogenetic studies in some representatives of the subfamily Pooideae (Poaceae) in South Africa. 2.

The tribe Aveneae, subtribes Phalaridinae and Alopecurinae. J.J. SPIES, S.K. SPIES, S.M.C. VAN

WYK, A.F. MALAN and E.J.L. LIEBENBERG 63

\J 8. A tribute to Frank White (5th March 1927 to 12th September 1994). A. ANGUS and J.D. CHAPMAN 69

v

9. Obituaries: Arthur Koeleman (1915-1994). G.F. SMITH and A.E. VAN WYK 77

,/ Werner Bahne Georg Jacobsen (1909-1995). N.H.G. JACOBSEN and J.B.E. JACOBSEN 79

v 10. Book review 81

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Bothalia 26,1: 1-9(1996)

Four new species and one new subspecies of Lachenalia (Hyacinthaceae) from

arid areas of South Africa

G.D. DUNCAN*

Keywords: Hyacinthaceae, Lachenalia , new species, new subspecies, South Africa

ABSTRACT

Four new species of Lachenalia are described: L. karooica W.F.Barker ex G.D.Duncan from the Great Karoo and southwestern

Free State, L. perryae G.D.Duncan from the Little Karoo and southern Cape, as well as L. neilii W.F.Barker ex G.D.Duncan and L.

alba W.F.Barker ex G.D.Duncan, both from the Nieuwoudtville-Calvinia District of the Northern Cape. In addition, a new

subspecies, L. marginata W.F.Barker subsp. neglecta Schltr. ex G.D.Duncan is described from the Western Cape.

INTRODUCTION

Lachenalia Jacq. f. ex Murray is the largest genus

within the southern African Hyacinthaceae and comprises

more than 100 species (Duncan 1988, 1992). The genus

is endemic to this region where it has a very wide distri-

bution extending from the southwestern parts of Namibia,

southwards throughout Namaqualand, the Western Cape

and Great Karoo to as far inland as the southwestern Free

State, from where its probable boundary makes an arc to

the southeast down to Transkei on the east coast of the

Eastern Cape (Barker 1987). Most of the taxonomic work

on Lachenalia carried out this century has been under-

taken by the late Miss W.F. Barker, formerly curator of

the Compton Herbarium at Kirstenbosch Botanical Gar-

den. Her studies, both during her term as curator and sub-

sequent to her retirement, have resulted in the publication

of more than forty new species for the genus. A number

of new Lachenalia species have still to be formally de-

scribed and published, and this paper serves as the first

in a series to be published as a continuation of the recent

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

1984, 1987, 1989) and the current author (Duncan 1993)

towards a revision of the genus.

NEW SPECIES IN LACHENALIA

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

sp. nov. distinguitur inflorescentia spicata vel subspicata

floribus patentibus, oblongo-campanulatis virellis cum vel

sine basibus excaeruleis, segmentibus interioribus perian-

thii recurvatis, staminibus bene exsertis patentibus,

plerumque folio singulari lanceolato conduplicato venis

longitudinalibus depressis, supra maculis brunneis,

viridibus vel marroninis irregulariter dispersis.

TYPE. — Free State, 2925 (Jagersfontein): Fauresmith

veld, 0.5 km on road from Fauresmith to Koffiefontein,

(-CB), 6-8-1976, Chaplin s.n., (NBG, holo.!).

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

7735, Cape Town.

MS. received: 1995-10-09.

Deciduous, winter-growing geophyte 45-220 mm high.

Bulb globose, 10-20 mm in diam., white with thin brown

outer scales, usually produced into a neck terminating just

below ground level. Leaf usually solitary, occasionally 2,

lanceolate, conduplicate, 40-200 x 5-20 mm, glaucous

with depressed longitudinal veins on upper surface and a

coriaceous margin, and irregularly scattered brown, green

or maroon blotches mainly on upper surface; clasping leaf

base 5-30 mm long with magenta bands or blotches just

below ground level, shading to purplish maroon above

ground level. Inflorescence spicate or subspicate, few to

many-flowered, 20-150 mm long with a short sterile tip;

peduncle erect to suberect, 20-80 mm long, slender or

sturdy, shorter than length of leaf, mottled with very pale

maroon and green; rachis mottled with very pale maroon

and green; pedicels absent or up to 2 mm long; bracts

ovate, greenish white, 1-2 mm long. Flowers patent, ob-

long-campanulate, dull white to greenish white, often dis-

colouring to a dull purple on herbarium sheets; outer

perianth segments oblong, 5-7 x 2-3 mm, with or without

bluish bases, with purplish maroon or greenish brown gib-

bosities; inner perianth segments obovate, recurved, 7-10

x 3 mm, with brownish green keels. Stamens well exserted

beyond tip of perianth, spreading; filaments white, 10 mm

long; anthers pale to dark maroon prior to anthesis, yellow

at anthesis. Ovary ovoid, pale green, 2-3 mm long; style

white, 10 mm long, protruding well beyond stamens as

ovary enlarges. Capsule ovoid, membranous, 5-7 mm

long. Seed globose with an arillode with a terminal ridge

of medium height. Chromosome number: 2n = 16. Figures

1A; 2 & 3.

Etymology : named after the Great Karoo where the first

specimens were collected, and from where the largest

number of collections are known.

Diagnostic characters

L. karooica is characterised by a spicate or subspicate

inflorescence of patent, oblong-campanulate flowers, with

the inner perianth segments distinctly recurved and con-

siderably longer than the outer ones. The spreading sta-

mens are well exserted beyond the tip of the perianth, and

the anthers are conspicuously pale to dark maroon prior

2

Bothalia 26,1 (1996)

FIGURE 1. — Inflorescences of four new species and one new subspecies of Lachenalia. A, L. karooica, Chaplin s.n.; B, L. perryae, Perry s.n.; C,

L. neilii, Duncan 196; D, L. alba, Steiner 1366; E, L. marginata subsp. marginata, Duncan 197; F, L. marginata subsp. neglecta, Duncan

238. Scale bars: 10 mm.

to anthesis. The plant usually has a single lanceolate leaf

which is glaucous and distinctly conduplicate, with de-

pressed longitudinal veins on the upper surface, and ir-

regularly scattered brown, green or maroon blotches,

mainly on the upper surface.

L. karooica is related to L. bowkeri Baker, an as yet

poorly known species from the southern part of the

Eastern Cape, in that both have a usually single con-

duplicate, lanceolate leaf and oblong-campanulate, dull

white to greenish white, patent flowers which are ses-

sile or have very short pedicels. The anthers of both

species are distinctly pale to dark maroon prior to an-

thesis. L. bowkeri differs in having included, declinate

stamens, the tips of the outer perianth segments not re-

curved, and the leaf pale green and unblotched without

coriaceous margin.

Flowering time: June to September.

Distribution and habitat

Material of this very distinctive species was first col-

lected by Douglas F. Gilfillan in August 1899 in the Mid-

delburg District of the Eastern Cape. It has since been

recorded from several widely separated localities in the

Great Karoo and Northern Cape, and from one locality at

Karoopoort. It is distinguished from all other lachenalias

in having a single glaucous, conduplicate, lanceolate leaf

with a coriaceous margin, and oblong-campanulate flow-

Bothalia 26,1 (1996)

3

FIGURE 2. — Holotype of Lachenalia karooica , Chaplin s.n.

ers with spreading, exserted stamens, of which the an-

thers are distinctly pale to dark maroon prior to anthe-

sis. The largest number of specimens is recorded from

the Fauresmith District in the southwestern part of Free

State, and it is very likely that the plant occurs through-

out the Great Karoo and much of the Northern Cape;

it is no doubt due to the remote nature of this area and

the inconspicuousness of the flowers that only a small

number of scattered specimens has been recorded. L.

karooica , although growing in areas with predominantly

summer rainfall, follows the typical pattern of winter rain-

fall growth and summer dormancy characteristic of the

vast majority of species belonging to this genus.

At the Fauresmith site, plants occur singly or in small

groups on south-facing aspects on dolomite outcrops, often

growing in the shade of Rhus burchellii Sond. ex Engl. Two

other geophytes encountered at this site are the white-flow-

ered Freesia andersoniae L. Bolus and the yellow-flowered

Homeria pallida Baker, both widespread, spring-flowering

geophytes which follow the winter growth cycle.

Material examined

NORTHERN CAPE. — 2824 (Kimberley): between Campbell and

Schmidtsdrif, (-CA), Aug. 1963, Stayner s.n. (NBG); Farm Wolwefon-

tein, Barkly West, (-DA), July 1936, Acocks 540 (PRE). 2922 (Prieska):

near Prieska, (-DA), Aug. 1935, Bryant 592 (PRE); Prieska, (-DA),

June 1969, Hardy & Rauh 1591 (PRE). 3025 (Colesberg): Colesberg

commonage, (-CA), Aug. 1925, Botha s.n. (PRE). 3124 (Hanover): 12.6

km from T-junction on Nieuw Bethesda road to Middelburg, (-DC),

Sept. 1990, Van Jaarsveld 11160 (NBG).

FREE STATE. — 2925 (Jagersfontein): Fauresmith Botanical Reserve,

(-CB), Aug. 1938, Henrici 3158 (PRE); Sept. 1925, Pole Evans 1825

(PRE); Sept. 1925, Smith 404 (PRE); 0.5 km on road from Fauresmith

to Koffiefontein, (-CB), Aug. 1976, Chaplin s.n. (NBG); Aug. 1993,

Duncan 367 (NBG); Aug. 1976, Lumley s.n. (NBG).

EASTERN CAPE. — 3125 (Steynsburg): Farm Conway, Middelburg

Dist., (-CB), Aug. 1899, Gilfillan s.n. in Herb. Galpin 5578 (PRE). 3225

(Somerset East): Mountain Zebra National Park, top of Babylon’s Tower,

(-AB), Aug. 1963, Liebenberg 7248 (PRE).

WESTERN CAPE. — 3319 (Worcester): Karoopoort, (-BC), Aug.

1980, Karoo National Botanic Gardens expedition s.n. (NBG).

Lachenalia perryae G.D. Duncan, sp. nov. distin-

guitur inflorescentia racemosa floribus oblongo-campanu-

latis segmentibus exterioribus perianthii pallide azureis

declinatis, staminibus inclusis, folio singulari arcuato con-

duplicato anguste lanceolato infra fasciis perspicuis

viridibus vel magenteis.

TYPE. — Western Cape, 3319 (Worcester); Karoo Na-

tional Botanic Garden veld reserve, Worcester, (-CB),

Aug. 1985, P.L. Perry s.n. (NBG, holo.l; PRE).

Deciduous, winter-growing geophyte 120-320 mm

high. Bulb globose, 10-20 mm in diam., covered with

dark brown spongy tunics produced into a short neck ter-

minating just below ground level. Leaf usually solitary,

occasionally 2, 240-300 x 10-23 mm, arcuate, condupli-

cate, narrowly lanceolate, yellowish green, banded below

with dark green shading to maroon or magenta on the

pale white clasping leaf base. Inflorescence subspicate or

racemose, lax, few to many-flowered, up to 130 mm long

with a very short sterile tip; peduncle slender, suberect,

up to 260 mm long, pale green; pedicels 2-3 mm long;

bracts ovate to lanceolate, greenish white, 1-2 mm long.

Flowers patent or slightly cemuous, oblong-campanulate;

outer perianth segments oblong, very pale blue, 5-6 x 3

mm, with pale green or brown gibbosities; inner perianth

segments obovate, obtuse, 7-9 x 4-5 mm, protruding well

beyond outer perianth segments, pale white to whitish yel-

low with green keels and recurved tips. Stamens declinate;

4

Bothalia 26,1 (1996)

tanical Gardens at Worcester, in recognition of her valu-

able work in collecting and recording the bulbous flora

of the Little Karoo.

Diagnostic characters

Superficially, L. perryae is reminiscent of the very vari-

able and widespread L. unifolia Jacq., to which it is related

due to the combination of features of a single narrow,

banded leaf, patent or slightly cemuous, oblong-campanu-

late flowers with pale blue outer perianth segments and

whitish inner segments, and included stamens. It differs

mainly in leaf morphology, and to a lesser extent, in

pedicel length, flower shape and seed morphology. In L.

unifolia , which never has more than one leaf, the leaf

blade is linear, widening abruptly into a loosely clasping

base. L. perryae, on the other hand, usually has a single

leaf, or occasionally two, with a narrowly lanceolate blade

which does not widen abruptly. Furthermore, the flowers

of L. perryae have short pedicels no longer than 3 mm,

whereas L. unifolia usually has very long pedicels up to

12 mm long. Although both these species have oblong-

campanulate flowers, the perianth segments of L. unifolia

are usually appreciably longer and the tips of the inner

perianth segments of L. perryae considerably more flared

than in L. unifolia. In addition, the seed of L. perryae has

an arillode with a terminal ridge of medium height,

whereas in L. unifolia the arillode is almost obsolete.

Flowering time : July to September.

Distribution and habitat

FIGURE 4. — Holotype of Lachenalia perryae , Perry s.n.

filaments white, as long as or very slightly longer than

inner perianth segments. Ovary globose, pale green, 2 mm

long; style white, 8-10 mm long, protruding well beyond

stamens as ovary ripens. Capsule globose, 5-7 mm long.

Seed globose with an arillode with a terminal ridge of

medium height. Figures IB; 4 & 5.

Etymology : L. perryae is named after Miss Pauline

Perry, formerly horticulturist at the Karoo National Bo-

The first specimens of L. perryae were collected by

Miss W.F. Barker in August 1949 near Robertson in

the Little Karoo, and it is from this region that most

records of the species are known; the largest number

of collections have been made in the veld reserve of

the Karoo National Botanical Garden at Worcester. The

species is currently known from the Worcester District

in the north to Port Beaufort in the south and eastwards

to Albertinia, all in the Western Cape, with one isolated

record much further east near Kommadagga in the East-

ern Cape. It occurs in arid areas in clay or sandy soil

amongst succulents, usually on south-facing aspects.

Plants occur singly or in small groups, and are locally

common in some areas. It is quite a variable species as

FIGURE 5. — Distribution of Lache-

nalia perryae , •; and L. alba ,

★

Bothalia 26,1 (1996)

5

regards plant size, ranging from depauperate specimens

120 mm high to much taller individuals, up to 320 mm

in height.

Material examined

WESTERN CAPE. — 3319 (Worcester): top of Hex River Pass,

(-BD), Sept. 1974, Mauve 221 (NBG); Karoo National Botanical Garden

veld reserve, (-CB), Sept. 1949, Barker 5941 (NBG); Aug. 1970, Bayer

17 (NBG); Aug 1953, Compton 24118 (NBG); Aug. 1962, Olivier 59

(NBG); Aug. 1986, Perry s.n. (NBG); De Dooms Mountains on way to

Touws River, (-CB), Aug. 1984, Van Zijl 235 (NBG); near Nuy cement

factory, (-DA), Aug. 1970, Barker 10706 (NBG); 9 km beyond Nuy

Village, (-DA), Sept. 1975, Thomas s.n. (NBG); 0.3 km from Breede

River Bridge on road to Le Chasseur, (-DC), Aug. 1974, Van Reenen

s.n. (NBG); near Robertson, (-DD), Aug. 1949, Barker 5393 (NBG).

3419 (Caledon): 17 km E of Caledon, (-BA), Aug. 1969. Barker 10616

(NBG). 3420 (Bredasdorp): 20 km NW of Port Beaufort on Malagas

road, (-BD), Aug. 1962, Thomas s.n. (NBG). 3421 (Riversdale): 1.6 km

along Stilbaai road from Albertinia, (-BA), Aug. 1971, Thomas s.n. (NBG).

EASTERN CAPE. — 3325 (Port Elizabeth): road to Jansenville from

Kommadagga, (-BB), Sept. 1973, Bayliss 5919 (NBG, PRE).

Lachenalia neilii W.F. Barker ex G.D. Duncan, sp.

nov. distinguitur inflorescentia multiflora racemosa flori-

bus parvis oblongo-campanulatis pallide viridibus basibus

perspicuis obscure caeruleis, tumoribusque clare viridibus

vel brunneis, staminibus declinatis, foliis duobus lanceo-

latis glaucis suberectis innotatis supra venis longitudinali-

bus depressis annuloque bulbillorum basi bulbi.

of the Northern Cape. This is the second Lachenalia spe-

cies commemorating Mr Macgregor and his family; in

1979, the very rare L. macgregoriorum W.F.Barker was

described from the Charlies Hoek area of Nieuwoudtville.

Diagnostic characters

L. neilii is characterised by a many-flowered raceme

of greenish white oblong-campanulate flowers with dis-

tinct dull blue bases and bright green or brown gibbosities,

shortly exserted stamens, two glaucous, lanceolate leaves

and usually a ring of bulbils produced at the base of the

bulb. It is related to L. alba W.F.Barker ex G.D. Duncan

due to the shared features of two lanceolate, unmarked

leaves, the oblong-campanulate flowers with protruding

white inner perianth segments and the slightly exserted

stamens. L. alba differs in having bigger, more campanu-

late flowers which are pure white, without distinct dull

blue bases, and which are usually attached to the rachis

by very short pedicels. Furthermore, the bulb of L. alba

does not develop a ring of bulbils at its base as L. neilii

does.

Distribution and habitat

L. neilii has a restricted distribution in the Nieu-

woudtville and Calvinia Districts of the Northern Cape

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

woudtville-Calvinia road (R27), at turnoff to Rondekop, in

ditches on both sides of road, (-AC), 12-9-1985, G.D. Dun-

can 196 (NBG, holo.).

Deciduous, winter-growing geophyte 120-320 mm

high. Bulb globose, 12-25 mm in diam., surrounded by

strong, dark brown outer tunics, usually with a ring of

bulbils produced at the base. Leaves 2, lanceolate, glau-

cous, unmarked, suberect, with depressed longitudinal

veins on upper surface, 55-145 x 7-22 mm, clasping leaf

base pale yellowish green, with or without tiny maroon

speckles, 40-70 mm long. Inflorescence racemose, many-

flowered, 50-150 mm long with few sterile flowers at tip;

peduncle erect, sturdy, pale green, with or without pale

brown or maroon speckles, 40-110 mm long; pedicels

suberect, white, 2-5 mm long; bracts ovate to lanceolate,

white, 2-3 mm long. Flowers oblong-campanulate, green-

ish white fading to dull red, with distinct dull blue bases;

outer perianth segments oblong, 7-8 x 4 mm, greenish

white with distinct dull blue bases and bright green or

brown gibbosities; inner perianth segments obovate, tips

slightly recurved, protruding beyond outer perianth seg-

ments, white with green keels, 9-10 x 4-5 mm. Stamens

declinate; filaments white, exserted up to 2 mm beyond

perianth, 9-11 mm long. Ovary ovoid, bright green, 3 mm

long; style white, 8-11 mm long. Capsule ovoid, 7-8 x

5-6 mm. Seed globose, with an arillode with a terminal

ridge of short to medium height. Figures 1C; 3 & 6.

Flowering time: August to October.

Etymology. L. neilii is named after Mr Neil Macgregor

of Glen Lyon Farm, Nieuwoudtville, in recognition of the

sterling nature conservation work he has done in this area

FIGURE 6. — Holotype of Lachenalia neilii , Duncan 196.

6

Bothalia 26,1 (1996)

where it grows in heavy red clayey soil on open flats.

The first specimens were collected as far back as Septem-

ber 1930 by a National Botanic Gardens expedition in the

area known as Klipkoppies in the Nieuwoudtville area.

The plant is locally plentiful and has been recorded as

growing in association with another new species, L. alba ,

which is also published for the first time in this paper,

and which is also restricted to the Nieuwoudtville area in

the Calvinia District.

Material examined

NORTHERN CAPE.— 3119 (Calvinia): Klipkoppies, Nieu-

woudtville, (-AC), Aug. 1961, Barker 9535 (NBG); Sept. 1961, Barker

9667 (NBG); Sept. 1986, Brandham 3018 (NBG); Sept. 1930, National

Botanic Gardens expedition s.n. (BOL); Glen Lyon Farm, Nieu-

woudtville, (-AC), Oct. 1967, Barker 10544 (NBG); Oct. 1973, Barker

10892 (NBG); Glen Ridge Farm, Nieuwoudtville, (-AC), Aug. 1960,

Barker 9185 (NBG); Aug. 1961 , Barker 9435 (NBG); Aug. 1968, Thomas

s.n. (NBG); 1.6 km E of Nieuwoudtville on road to Calvinia, (-AC),

Nov. 1967, Barker 9532 (NBG); Nieuwoudtville Reserve, (-AC), Oct.

1983, Perry & Snijman 2420 (NBG); Nieuwoudtville-Calvinia road

(R27), at Rondekop turnoff, (-AC), Sept. 1985, Duncan 196 (NBG).

Without precise locality: Klipbokberg, March 1931, Herre s.n. (BOL);

Leliefontein, Sept. 1940, Leipoldt 3386 (BOL).

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

distinguitur inflorescentia subspicata vel racemosa floribus

grandibus oblongo-campanulatis candidis cum vel sine

basibus perpallide azureis cum vel sine carinis distinctis

viridibus, rubris vel caeruleis segmentibus interioribus peri-

anthii recurvatis declinatis, staminibusque breviter exsertis,

foliis duobus lanceolatis suberectis ad erectis glaucis innotatis

supra venis longitudinalibus depressis.

TYPE. — Northern Cape, 3119 (Calvinia); Charlies Hoek,

Nieuwoudtville, (-AC), 30-9-1973, W.F. Barker 10888

(NBG, holo.!).

FIGURE 7. — Holotype of Lachenalia alba, Barker 10888.

Deciduous, winter-growing geophyte 100-330 mm

high. Bulb globose, 15-20 mm in diam., white with thick

membranous dark brown outer tunics. Leaves 2, lanceo-

late, channelled above, suberect to erect, glaucous, un-

marked, margin sometimes undulate, upper leaf surface

with depressed longitudinal veins, 120-230 x 15-25 mm,

clasping leaf base 20-80 mm long, sometimes suffused

with dull maroonish brown zones. Inflorescence subspi-

cate to racemose, usually many-flowered, 40-170 mm

long with short sterile tip; peduncle suberect to erect, pale

green, 50-150 long; rachis pale green shading to white in

upper half, with very pale maroonish brown speckles;

pedicels white, 1-3 mm long; bracts white, ovate to

lanceolate, 2-3 mm long. Flowers pure white, oblong-

campanulate; outer perianth segments oblong, 7-8 x

3-4 mm, with or without very pale greenish blue bases,

gibbosities pale brownish green or dull red; inner peri-

anth segments protruding well beyond outer perianth

segments, recurved, obovate, 10-11 x 4—6 mm, with or

without pale to distinct brownish green, red or blue

keels. Stamens declinate; filaments white, 10 mm long,

protruding up to 2 mm beyond tip of perianth. Ovary

ovoid, pale green, 3-4 mm long; style white, 8 mm

long, protruding well beyond perianth as ovary matures.

Capsule ovoid, pale green, 7 mm long. Seed globose

with an arillode with a terminal ridge of medium height.

Figures ID; 5 & 7.

Flowering time : August to October.

Etymology, named alba due to the overall white col-

ouring of the flowers.

Diagnostic characters

L. alba is characterised by its many-flowered inflores-

cence of pure white, oblong-campanulate flowers with re-

curved inner perianth segments which may or may not

have distinct green, red or blue keels, and its two glau-

cous, lanceolate, suberect to erect unmarked leaves with

distinct, depressed longitudinal veins on the upper surface.

It is related to L. neilii, but the latter differs in having

smaller, less campanulate greenish white flowers with dis-

tinct dull blue bases borne on long pedicels up to 5 mm

long, whereas L. alba has short pedicels usually not longer

than 2 mm. Furthermore, the bulb of L. alba does not

develop the typical ring of bulbils at its base as L. neilii

does.

Distribution and habitat

As with L. neilii , this species has a restricted distribu-

tion in the Nieuwoudtville and Calvinia areas of the

Northern Cape, growing in colonies in heavy red clayey

Bothalia 26,1 (1996)

7

soil on open flats. The earliest record of the plant is the

collection made by W.F. Barker in September 1930 at

Klipkoppies, Nieuwoudtville. L. alba is recorded as grow-

ing in association with L. neilii and L. elegans W.F.

Barker, the latter being a very common, variable species

in the Nieuwoudtville-Calvinia area, but to my knowledge

there are no reports of interspecific natural hybrids be-

tween these species.

Material examined

NORTHERN CAPE. — 3119 (Calvinia): Klipkoppies, Nieu-

woudtville, (-AC), Sept. 1930, Barker s.n. (BOL); Sept. 1961, Barker

9540 (NBG); Sept. 1973, Barker 10889 (NBG); Sept. 1986, Brandham

3027 (NBG); 2 km E of Nieuwoudtville, (-AC), Sept. 1957, Acocks

19493 (NBG); Sept. 1961, Barker 9532 (NBG); Nieuwoudtville, (-AC),

Sept. 1938, Buhr s.n. (NBG); Grasberg, (-AC), Sept. 1943, Buhr s.n.

(NBG); 1 km W of Nieuwoudtville, (-AC) Sept. 1986, Brandham 3007

(NBG); Oorlogskloof, (-AC), Sept. 1939, Leipoldt s.n. (NBG); Oor-

logskloof road, 10.6 km S of junction with R27 at Nieuwoudtville, (-AC),

Sept. 1986, Steiner 1366 (NBG); Nieuwoudtville Reserve, (-AC), Sept.

1983, Perry & Snijman 2369 (NBG); Glen Lyon Farm, (-AC), Aug.

1972, Barker 10544 (NBG); Sept. 1973, Barker 10888 (NBG); Oct. 1974,

Barker 10921 (NBG); between Oorlogskloof and Papkuilsfontein, (-AC),

Sept. 1939, Leipoldt 4170 (BOL); near Toren Farm, (-BC), Oct. 1974,

Thomas s.n. (NBG); 8.2 km along Toren road from Calvinia-Loeriesfon-

tein road, (-BC), Sept. 1986, Brandham 3030 (NBG). Without precise

locality; between Vanrhynsdorp and Calvinia, Sept. 1934, Bishop S. Lavis

s.n. (BOL).

Lachenalia marginata W.F.Barker in Journal of

South African Botany 45: 204—207 (1979).

TYPE. — Nieuwoudtville, Barker 6463 (NBG, holo.!).

L. marginata is a very distinctive species with a dis-

tribution range extending from the fynbos on the Nieu-

woudtville escarpment to the Gifberg massif and the

Nardouw-Pakhuis-Cederberg Mountains and the low-ly-

ing areas around Clanwilliam where it usually occurs in

very sandy soil growing singly or in small groups. The

subsp. marginata, which is the most widespread of the

two subspecies, is recognised by its single ovate leaf with

a distinctive thickened coriaceous margin which is undu-

late and crisped, its heavily banded clasping base and its

long, sessile narrowly urceolate flowers with very con-

spicuous dark brown gibbosities on the outer perianth seg-

ments. The seed places the species into the group of

species having a medium-sized terminal inflated arillode

(Figures IE; 8 & 9).

Material of subsp. marginata examined

NORTHERN CAPE.— 3119 (Calvinia): Glenridge Farm, Nieu-

woudtville, (-AC), Aug. 1961, Barker 9433 (NBG); Aug. 1966, Barker

10451 (NBG); Sept. 1985, Duncan 197 (NBG); 4.8 km W of Nieu-

woudtville, (-AC). Aug. 1950, Barker 6465 (NBG); July 1948, Lewis

2408 (SAM); Aug. 1950, Lewis 3240 (SAM); 6.4 km W of Nieu-

woudtville, (-AC), Aug. 1950, Barker 6463 (NBG).

WESTERN CAPE. — 3118 (Vanrhynsdorp); Matzikamma, (-AB),

July 1940, Martin s.n. sub NBG 72364 (NBG); Klawer, (-DC), June

1950, Barker 7387 (NBG). 3218 (Clanwilliam): 10 km E of Clanwilliam

on Pakhuis Pass, (-BB), Sept. 1985, Duncan 194 (NBG); Aug. 1974,

Nordenstam & Lundgren 1327 (NBG). 3219 (Wuppertal): Lorraine Farm,

N of Pakhuis Pass, (-AA), July 1965, Siaens s.n. sub NBG 84814 (NBG);

Matjiesrivier, Cederberg, (-AC), Sept. 1942, Wagener 209 (NBG).

Lachenalia marginata W.F.Barker subsp. neglecta

Schltr. ex G.D.Duncan, subsp. nov. a subspecie typica inflo-

rescentia densiore elongata spicata floribus multo parviori-

bus anguste urceolatis, folioque erecto ad suberecto con-

duplicato lanceolato ad ovato-lanceolato differ!.

TYPE. — Western Cape, 3218 (Clanwilliam): Die Berg

Farm, SW of Clanwilliam, (-BB), 18-9-1966, Barker

10428 (NBG, holo.!).

Deciduous, winter-growing geophyte 110-280 mm high.

Bulb globose, 11-25 mm in diam., white with thin mem-

branous outer tunics. Leaf lanceolate to ovate-lanceolate,

erect to suberect, almost always solitary (occasionally 2),

glaucous, conduplicate, margin coriaceous, dark maroon,

undulate and crisped, upper leaf surface with depressed

longitudinal veins and irregularly scattered pale to dark

green spots, 45-200 x 18-30 mm; tightly clasping leaf

base distinctly banded or blotched with dark maroon. In-

florescence spicate, dense, many-flowered 45-185 mm

long, with several sterile flowers at tip; peduncle 40-150

mm long, erect to suberect, pale green with maroon

blotches, thickening towards base of inflorescence; bracts

white, ovate to lanceolate, 3-5 mm long. Flowers sessile,

narrowly urceolate, greenish yellow fading to dull red;

outer perianth segments oblong, 6-8 x 3-4 mm, pale

greenish yellow with minute blue speckles and conspicu-

ous dark brown gibbosities; inner perianth segments pro-

truding well beyond outer segments, obovate, 7-11 x

4-5 mm, dark greenish yellow, two upper lateral seg-

FIGURE 8. — Distribution of L. marginata subsp. marginata , •; and L.

marginata subsp. neglecta , +.

Bothalia 26,1 (1996)

FIGURE 9. — Holotype of Lachenalia marginata subsp. marginata,

Barker 6463.

ments overlapping, and 1-2 mm shorter than narrower

lower segment. Stamens declinate, included within peri-

anth; filaments white, 6 mm long. Ovary ovoid, 4 mm

long, pale green; style white, 7 mm long. Capsule ovoid,

6-8 mm long. Seed globose, with an arillode with a ter-

minal, inflated ridge of medium height. Chromosome

number : 2n = 10. Figures IF; 8 & 10.

Flowering time'. July to August.

Diagnostic characters

L. marginata subsp. neglecta differs from the typical

subspecies in its much denser, elongated, many-flowered

inflorescence of much smaller flowers and in its erect to

suberect, lanceolate to ovate-lanceolate, conduplicate leaf,

as compared to the ovate, acute leaf of subsp. marginata.

The seeds of the two subspecies have a similar medium-

sized, terminal, inflated arillode. Subsp. marginata is

found from Nieuwoudtville to as far south as the southern

Cederberg, and grows at medium to high altitude,

whereas subsp. neglecta is restricted to low-lying areas

in the Clanwilliam District.

Distribution and habitat

The first collection of this subspecies was made by

Rudolf Schlechter on 12 August, 1896, 14 km west of

Clanwilliam on the farm Zeekoe Vlei. It is one of a large

number of Lachenalia collections made by him on his

collecting trips in South Africa during the period

1891-1895, material of which was distributed to many

herbaria in South Africa, Europe and Scandinavia. Manu-

script names were given to most of these collections, some

of which have been validated in papers in the Journal of

South African Botany (Barker 1978, 1979). In the same

journal. Barker (1983) published a numerical list of

Schlechter’s Lachenalia collections made from 1891-1898,

with identifications added, to assist in bringing all his nu-

merous sets of duplicate specimens in many herbaria up

to date. Material labelled L. neglecta Schltr. was distrib-

uted to ten herbaria locally and abroad, and is here vali-

dated as a subspecies of L. marginata.

L. marginata subsp. neglecta has a restricted distribu-

tion in the Clanwilliam District of the Western Cape where

it occurs in deep sand amongst restioid fynbos at low al-

titude. Plants occur singly and have been seen growing

in association with Lachenalia mutabilis Sweet, a very

FIGURE 10. — Holotype of Lachenalia marginata subsp. neglecta.

Barker 10428.

Bothalia 26,1 (1996)

9

variable and widespread species which is common in this

area, but which flowers a few weeks later than L. margi-

nata subsp. neglecta.

Material examined

WESTERN CAPE. — 3218 (Clanwilliam): Zeekoe Vlei Farm, 14 km

W of Clanwilliam, (-BA), Aug. 1896, Schlechter 8490 (B, BM, BOL,

G, GRA, K, L, PRE, S, Z); Die Berg Farm, (-BB), Aug. 1966, Barker

10428 (NBG); 41 km on N7 road Citrusdal-Clanwilliam, (-BB), July

1986, Duncan 238 (NBG); July 1993, Duncan 365 (NBG).

ACKNOWLEDGEMENTS

This paper is dedicated to Miss W.F. Barker who died

on 27th December, 1994. 1 am very much indebted to the

staff of the Compton Herbarium, especially Dr D.A. Pat-

erson-Jones, Mr J.P. Roux and Mrs S.E. Foster for their

assistance at various stages of this study. I also thank Dr

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

diagnoses, Mrs J. Loedolff for taking the photographs of

the herbarium sheets, Mrs A. Romanowski for producing

the black and white photographs from colour slides and

Mr M. Lumley for assistance in the field.

REFERENCES

BARKER, W.F. 1978. Ten new species of Lachenalia (Liliaceae). Jour-

nal of South African Botany 44: 391 — 418.

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

Journal of South African Botany 45: 193-219.

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

Rudolf Schlechter ’s collections made on his collecting trips in

southern Africa, with identifications added. Journal of South Afri-

can Botany 49: 45-55.

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

Journal of South African Botany 49: 423-444.

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

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

African Botany 50: 535-547.

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

Hyacinthoideae) four from the Cape Province and one from southern

South West Africa/Namibia. South African Journal of Botany 53:

166-172.

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

nalia (Liliaceae-Hyacinthaceae) from the Cape Province. South

African Journal of Botany 55: 630-646.

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

bosch Botanic Gardens 1 7. National Botanical Institute, Cape Town.

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

and taxonomy. Acta Horticulturae 325: 843—845.

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

Africa 52: t. 2061.

\

Bothalia 26,1: 11-30(1996)

Saxicolous species of the genus Rinodina (lichenized Ascomycetes, Physci-

aceae) in southern Africa

M. MATZER* and H. MAYRHOFER*

Keywords: biogeography, flora, lichenized Ascomycetes, Physciaceae, Rinodina , southern Africa, taxonomy

ABSTRACT

A revision of saxicolous species of the genus Rinodina (Ach.) Gray (lichenized Ascomycetes, Physciaceae) in southern

Africa is presented. The study area covers the following countries: Angola, Namibia, Republic of South Africa, Lesotho,

Swaziland, Zimbabwe, and Mozambique. Fifteen Rinodina species are treated in detail, and three of them are new to science:

Rinodina longisperma Matzer & H.Mayrhofer, R. scabridula Matzer & H.Mayrhofer, and R. striatitunicata Matzer &

H.Mayrhofer. The general part of the study includes information on the most important characters of the species with a special

focus on ascospore types and ontogeny, and biogeographical notes are also given. A key to the species is provided. All the species

are described in detail with notes on their ascus characters, spermogonial apparatus, and chemical characteristics, and their

substrate and general distribution are indicated. The descriptions are accompanied by illustrations of the ascospores. Except for

Rinodina oxydata s.l., the distribution of all the species in southern Africa and other parts of Africa respectively, is mapped.

Several names were referred into synonymy, including Rinodina almbornii H.Mayrhofer, a synonym of R. confragosula (Nyl.

in Cromb.) Miill. Arg., and R. depressa (Vain.) Zahlbr. and R. albicans H.Mayrhofer which are synonymous to R. huillensis Vain.

A separate section encompasses nine excluded species.

INTRODUCTION

The cosmopolitan genus Rinodina (Ach.) Gray com-

prises about two hundred species (Hawksworth et al.

1983), and its species taxonomy is still not settled satis-

factorily, particularly in the southern hemisphere. This

became obvious, once again, during the preparation of

the present work, a re-assessment of saxicolous species

occurring in southern Africa. Type material of all critical

species and, to the best of our knowledge, almost all rele-

vant collections were examined. Nevertheless, the present

study should be seen as an actual inventory rather than

an ultimate revision. Given the vastness of the study area,

there are relatively few collections of saxicolous Rino-

dina species, and most of them come from the Cape

Province in the Republic of South Africa. Type speci-

mens of some species are scant or damaged, several spe-

cies are still known from a few collections only, and thus

the extent of their variation is difficult to estimate. This

is particularly problematical in cases where species which

are separated by slight morphological differences only,

have been described. Morphologically and chemically,

many of the saxicolous Rinodina species in southern Af-

rica do not exhibit ‘exciting characters’, and great simi-

larities can be observed between several species. There-

fore, determination of specimens has often proved to be

rather difficult. In the material studied, several Rinodina

species were detected which are most probably unde-

scribed. Not all of them are included in the present study,

as they are often known from a single locality only, and

their relationships to already described and similar spe-

cies could not be resolved.

However, despite the problems indicated above, many

new results are presented in this paper. Several names of

* Institute for Botany, Karl-Franzens-University Graz, Holteigasse 6,

A-8010 Graz, Austria.

MS. received: 1995-03-15.

taxa are referred into synonymy, and three new species

are introduced. The importance of minute characters such

as ascospore ornamentation and length of spermatia for

the circumscription of single species is documented. The

first data on the biogeographical affinities of saxicolous

Rinodina species occurring in southern Africa are pre-

sented.

MATERIAL AND METHODS

Standard light microscope techniques and standard

tests for thallus chemistry (tic, spot tests for colour reac-

tions) were employed. Descriptions of ascus apices are

based on observations in dilute original Lugol’s solution

(MERCK 9261) without pretreatment with KOH (K). For

the testing of the N ( = HNC>3)-reaction of the blue-green

to blackish pigment, longitudinal sections of thalli,

apothecia, and/or spermogonia respectively, were directly

mounted in this medium.

THE STUDY AREA

The main focus of this study was on species occurring

in extratropical southern Africa, mainly including Na-

mibia, the Republic of South Africa, Lesotho, and Swazi-

land. Some specimens from adjacent countries (Angola,

Zimbabwe, Mozambique) were also studied. However, as

already mentioned above, most of the specimens exam-

ined were from the Republic of South Africa, mainly from

the Cape Province, For comparative purposes it was also

necessary to study saxicolous Rinodina species described

or mentioned from other parts of Africa, mainly including

the islands of Pagalu (Annobon) and Sao Tome & Principe

in the Atlantic Ocean, and the island of Socotra in the

Indian Ocean.

12

Bothalia 26,1 (1996)

MORPHOLOGY, ANATOMY AND CHEMISTRY

A short survey is given here of the most important

characters of the species treated in detail.

Thallus and prothallus

Thalli in Rinodina species are usually crustose or crus-

tose to squamulose. The formation of blastidia is the most

important character for the separation of R. scabridula.

The presence or absence of a distinct prothallus should

not be overestimated in the circumscription of species.

Chemistry

Only a few species contain secondary lichen substances

which are detectable by thin-layer chromatography (tic).

Rinodina huillensis is the only species with gyrophoric

acid (and, in part, other secondary compounds) and is well

characterized by the C+ red reaction in the medulla of the

thallus. The thalli of R. confragosa , R. oxydata s.l., and

R. substellulata give a K+ yellow reaction. The main li-

chen compound in these taxa is atranorin.

Pigments

A blue-green to blackish pigment was often found to

be located in parts of the thallus, apothecia (proper exci-

ple, epihymenium), and/or in the spermogonia of six spe-

cies: R. confragosula , R. huillensis , R. oxydata sd., R.

striatitunicata , R. substellulata, and R. subtristis. The pig-

ment reacts N+ red, K-, and HC1+ blue after pretreatment

with K. It is most probably identical to the pigment which

was described as ‘Lecideagriin’ by Bachmann (1890), and

it occurs also in other genera of the Physciaceae, e.g. in

Buellia De Not. (Scheidegger 1993, as ‘pigment A’).

An interesting and unique character is the K+ yel-

lowish to orange reaction in the hypothecium, the epi-

hymenium (here not always), and in spermogonia (not

always) of R. reagens, which is due to the presence of a

green-yellow pigment.

Apothecia

In the genus Rinodina , a great variation in apothecial

types can be observed, and even within single species or

specimens, the formation of apothecia can be variable.

Apothecia are cryptolecanorine, lecanorine (in addition to

the thalline margin a proper margin may be developed),

and/or lecideine. Apothecia which appear to be lecideine

in surface view may contain (dead) algal cells in the ex-

ciple (e.g. in R. oxydata sd., R. subtristis, R. teichophil-

oides).

Asci

Most of the species have asci which can be described

as corresponding to the Lecanora type (compare e.g.

Honegger 1978; Rambold et al. 1994), but some variation

in the formation of ascus tips can be observed. In R. oxy-

data s.l. , the asci correspond to the Lecanora type but

show tendencies towards the Bacidia type (see e.g.

Hafellner 1984; Rambold et al. 1994). The axial body

in several species is often more or less indistinct (e.g.

in R. geesteranii, R. huillensis, R. longisperma, R. oxy-

data s.l., R. subtristis ) or even not observable (e.g. in

R. longisperma ) under the light microscope. In R. huil-

lensis, the axial body may be surrounded by a tube-like

structure, and the same phenomenon could be observed

in several other species after pretreatment with KOH.

Ascospores

The most important characters for the separation and

identification of Rinodina species are those of the as-

cospores. For terms used in the description of asco-

spores occurring in Rinodina (and other genera of the

Physciaceae) see Poelt & Mayrhofer (1979); revised

compilations of ascospore types were published by

Hafellner et al. (1979), Mayrhofer & Poelt (1979) and

Mayrhofer (1982, 1984a). With the increasing knowl-

edge on possible formation types in the ascospores of

Physciaceae, it became necessary to modify the classi-

fication systems for spores by subsequent authors. For

a better understanding, the most important changes are

briefly outlined here.

1. The presence vs. absence of a torus (i.e. a dark belt

in the region of a spore septum) is not always a reliable

character for the definition of spore types. For instance,

within several species a torus can clearly be observed

in some of the ascospores under the light microscope,

but is indistinct or even not visible in others (e.g. in

Rinodina fimbriata, R. microlepidea, R. striatitunicata,

R. substellulata); the same has been reported for other

genera and species of the Physciaceae (e.g . Amandinea

petermannii; see Matzer et al. 1994b). Scheidegger

(1993: 333) pointed out that within Buellia closely re-

lated species may differ considerably in their torus while,

on the other hand, species with only slight affinities to each

other may have a similar torus. Consequently, a distinction

of spore types which is solely based on the presence or

absence of a torus cannot be recommended. This mainly

concerns the following ascospore types: Beltraminia

type (inch Buellia type) and Physconia type (incl. Du-

byana and Sicula type); in both cases the terms which

have been introduced first into the literature should be

used (for dates of introduction see Mayrhofer 1982).

Such a procedure has been applied already by Matzer &

Mayrhofer (1993), Scheidegger (1993), and, in part, by

Rambold et al. (1994).

2. Original definitions of single spore types have been

modified or improved; the most important case in this

context is the Dirinaria type. It was introduced by

Mayrhofer (1982: 315) for ascospores lacking a torus and

with internal wall thickenings of the Physcia, Mis-

choblastia, and/or intermediate types. In subsequent

studies, the mode of spore ontogeny in this spore type

(apical internal spore wall thickenings appear before

the septum is inserted) was used more or less implicitly

as the major character for the definition of the Dirinaria

type (Giralt & Matzer 1994; Giralt et al. 1995; Rambold

et al. 1994), or this peculiar ontogeny was stressed as

being a very important character (Giralt 1994; Giralt &

Mayrhofer 1994a, b, 1995). In the present study this

concept of the Dirinaria type is adopted.

Bothalia 26,1 (1996)

13

Within the species treated in detail in the present study,

the following ascospore types were observed:

Beltraminia type (inch Buellia type): spores without in-

ternal wall thickenings, Buellia distrata.

Physconia type (inch Dubyana and Sicula type): spores

with ± pronounced septal wall thickenings, apical thick-

enings less pronounced or lacking, spore lumina at their

distal ends (i.e. near to the spore ends) rounded, R. longi-

sperma.

Physeia type: septal and apical wall thickenings well

developed, spore lumina at their distal ends concave, torus

usually developed, in young spores the septum is inserted

before wall thickenings become distinct, R. confragosa , R.

confragosula, R. scabridula , R. sp. 1.

Milvina type: similar to the Physeia type, but particularly

the apical wall thickenings are less pronounced, spore

lumina at their distal ends flattened to slightly concave, R.

confragosula , (R. geesteranii ), R. huillensis, R. scabri-

dula, R. substellulata, R. subtristis , R. sp. 1.

Mischoblastia type: septal and apical wall thickenings

strongly pronounced, R. oxydata s.l., R. reagens, R. teicho-

philoides.

Pachysporaria type: wall thickenings strongly devel-

oped around the lumina which are ± rounded, R. confrago-

sula, (R. geesteranii), R. huillensis, R. longisperma , R.

microlepidea, R. reagens, R. scabridula, R. substellulata,

R. subtristis, R. teichophiloides.

TUnicata type: spores with a markedly thickened outer

wall layer, R. striatitunicata.

Dirinaria type: when mature similar to the Physeia type

(septal and apical wall thickenings well developed, spore

lumina at their distal ends concave), but usually lacking a

torus and with a different spore ontogeny: in young spores

the septum is laid down after the formation of apical wall

thickenings, R. gennarii, (compare also R. geesteranii).

As can be seen from this compilation, in most of the

species the ascospores could not clearly be assigned to a

single spore type. Most of the spores encountered belong

to the Pachysporaria or Milvina type. In contrast to all

other species treated in detail, the insertion of the septum

in ascospores of R. geesteranii may occur before or after

internal wall thickenings become distinct.

Spermogonial apparatus

The spermogonial apparatus in all species studied in

this respect is typical of the genus Rinodina. Therefore,

we give here a general description which is not repeated

in the species accounts:

Spermogonia immersed in the thallus, with the upper parts

prominent or not, in longitudinal section ± globose, ellip-

soid, cupuliform, or flask-shaped. Spermatiophores sep-

tate, composed of spermatiogenous cells. Spermatio-

genous cells intercalar and terminal; intercalar cells ± cy-

lindrical and with a lateral projection below the upper

transverse septum; terminal cells ± flask-shaped or ob-

long-conical and with an apical projection; spermatia are

formed on the lateral and apical projections. Spermatia

bacilliform, hyaline, non-septate.

The spermatiophores correspond to type VI of Vobis

(1980, as ‘Conidiophoren’). For illustrations of sperma-

tiophores and spermatiogenous cells respectively, see

Matzer et al. (1994a) and Mayrhofer et al. (1993). The

spermogonial apparatus of R. longisperma is basically of

the same type as described above, but it differs in the way

that intercalar spermatiogenous cells could be observed

only exceptionally.

BIOGEOGRAPHICAL NOTES

According to the present, rather incomplete data,

saxicolous species of Rinodina from southern Africa

show four different biogeographical elements or affini-

ties. Particularly noteworthy is the high incidence of

endemism.

1. Endemic element: this is the dominant element and

includes 1R. confragosula (a record from New Caledonia

seems to be dubious), R. geesteranii, R. huillensis, R.

longisperma, R. microlepidea, R. scabridula, R. striati-

tunicata, R. subtristis, R. teichophiloides, and R. sp. 1.

Most of these species have been found only in the Repub-

lic of South Africa and, in part, in adjacent countries,

while R. huillensis and R. subtristis are known to extend

further north to Angola.

2. African element: R. substellulata is known to occur in

the Cape Province but also on some tropical islands adja-

cent to the African continent (Principe, Annobon, and

Socotra).

3. New Zealand-South African element: an interesting

distribution pattern is shown by R. reagens which has been

found in New Zealand and in the Cape Province in South

Africa.

4. Cosmopolitan element: this comprises R. confragosa,

R. gennarii, and R. oxydata s. 1.

Key to saxicolous Rinodina spp. in southern Africa

la Medulla of thallus C+ red (containing gyrophoric acid); ascospores of Pachysporaria type, occasionally grading

into Milvina type, small protrusions of lumina towards septum and/or spore ends may be present, (10-)12-18

(-24) x (6-)7-10(-12) fjm 5. R. huillensis

lb Medulla of thallus C-:

2a Thallus K+ yellow; (atranorin usually detectable by tic):

3a Ascospores of Physeia type, (15-)16-27 x 8-13 pm 1. R. confragosa

3b Ascospores not of Physeia type:

4a Ascospores of Pachysporaria type, occasionally grading into Milvina type, 14-20 x 7-12 ftm ... 12. R. substellulata

4b Ascospores usually of Mischoblastia type, rarely with tendencies towards Pachysporaria type, 11-25 x (7—)

8-13 /rm 8. R. oxydata s.l.

14

Bothalia 26,1 (1996)

2b Thallus K-; (lacking atranorin):

5a Ascospores of Tunicata type, internal wall thickenings corresponding to Physcia type, mature spores with striate

ornamentation, (16-)17-27 x (10-)11-17 pm 11. R. striatitunicata

5b Ascospores not of Tunicata type, mature spores without striate ornamentation:

6a Hypothecium (always), epihymenium and spermogonia (not always) in longitudinal section distinctly K+

yellowish to orange; ascospores with internal wall thickenings of Mischoblastia type, occasionally grading

into Pachysporaria type, torus absent, 22-31 x 10-19 /mi; on calcareous rocks 9. R. reagens

6b Hypothecium, epihymenium, and spermogonia K-:

7a Thallus partially blastidiate, blastidia usually situated at margins of thallus areolae; ascospores of Milvina,

Pachysporaria or Physcia type, or intermediate between these types, (12-)15-20(-24) x (6-)7-10(-13)

/<m 10. R. scabridula

7b Thallus smooth or with an irregular surface but not blastidiate:

8a In young ascospores septum is inserted after formation of internal apical wall thickenings, spores lacking

a torus in all stages of ontogeny:

9a Ascospores of Dirinaria type, in young spores septum is always inserted after formation of internal

apical wall thickenings, spores ( 11— )12— 17 x (5-)6-9 /mi; hymenium 60-80 pm tall; thallus thin,

occasionally inconspicuous, crustose, areolate or ± effuse, smooth; apothecia often numerous and

crowded 4. R. gennarii

9b Ascospores with internal wall thickenings which intervene between Pachysporaria and Milvina type,

ontogeny of spores apparently irregular; in young spores the septum is inserted before or after

internal wall thickenings become distinct, spores 14— 21(-24) x 8 — 13( — 15) /mi; hymenium 100-115

pm tall; thallus thick, crustose to slightly squamulose, areolate, large areolae and squamules with

an irregular surface 3. R. geesteranii

8b In young ascospores septum is inserted before formation of internal wall thickenings, torus in mature

spores present or absent:

10a Ascospores without torus but intense brown pigmentation in septal region may simulate presence of

a torus:

11a Ascospores at first with internal wall thickenings of Mischoblastia type, then corresponding to Pachy-

sporaria type, (16-)20-32 x 10-18 pm, in young spores septum is inserted before internal wall

thickenings become distinct, torus absent but intense brown pigmentation in septal region may

simulate presence of a torus; apothecia to 0.8 mm diam 14. R. teichophiloides

lib Ascospores with internal wall thickenings which intervene between Pachysporaria and Milvina type,

14— 21(-24) x 8— 13(— 15) /tm, ontogeny of spores apparently irregular; in young spores septum is

inserted before or after internal wall thickenings become distinct, spores always without intense

brown pigmentation in septal region which may simulate presence of a torus; apothecia to 0.5

mm diam 3. R. geesteranii

10b Ascospores usually with distinct torus (in some spores torus may be indistinct):

12a Ascospores of Physcia type, rarely grading into Milvina type, 11-17 x 6-9 /tm 15. R. sp. 1

12b Ascospores not of Physcia type or, when of Physcia type usually distinctly larger:

13a Apothecia small, to 0.3 mm diam., cryptolecanorine; ascospores small, similar to Pachysporaria

type or ± of Physconia type, occasionally with strongly pronounced apical internal wall thick-

enings, partly with protrusions of lumina to spore ends and/or septum, spores 10— 16(— 18) x

6-10(-12) /tm; spermatia comparatively long, (4— )6-9 x 1.0-1 .5 /tm 6 . R. longisperma

13b Not with this combination of characters; spermatia up to 5 /tm long:

14a Thallus whitish, yellowish, ochraceous; apothecia to 0.4 mm diam., lecanorine, rarely cryptolecanor-

ine, disc light brown to dark brown or blackish; ascospores of Pachysporaria type, (13— )15— 19

(-20) x 8-15 /tm 7. R. microlepidea

14b Not with this combination of characters; apothecia becoming larger (to 0.8 mm diam.):

15a Thallus often greyish (also grey-brown, brown, occasionally whitish due to presence of a

pruina); apothecia to 0.8 mm, usually cryptolecanorine (rarely lecanorine or lecideine), disc

usually purely black (rarely dark brown); ascospores very variable in formation of internal wall

thickenings; Pachysporaria, Milvina, or intermediate types, occasionally also of Physcia type

or spore lumina irregularly biconical in shape, spores 14-32 x 7.5-16.0 /tm ... .2 . R. confragosula

15b Thallus brown to ochraceous; apothecia to 0.7 mm diam., usually lecideine or lecanorine

(occasionally cryptolecanorine), disc dark reddish brown to black; ascospores of Pachyspo-

raria type, occasionally grading into Milvina type, (15-)16-23 x 8-16 /tm 13. R. subtristis

1. Rinodina confragosa (Ach.) Koerb. in Systema

lichenum Germaniae: 125 (1855).

For synonyms and typification see Mayrhofer & Poelt (1979) and

Mayrhofer (1984a).

Thallus thin to thick, crustose to squamulose, con-

tinuous and areolate or composed of discrete areolae

which may be ± wart-like, whitish, pale grey to ochra-

ceous; prothallus absent or present and then it may be

strongly developed, brown to black. Chemistry: thallus

K+ yellow, P ± yellowish; tic: atranorin, chloratranorin

(not always), zeorin (not always), in the material from

southern Africa an unidentified compound (rf-classes

6/6/6) was found.

Apothecia to 1.5 mm diam., lecanorine, in addition

to thalline margin a proper margin may be developed,

usually sessile, rarely adnate, disc brown to black, plane

to convex. Epihymenium 5-20 /um tall, brown. Hyme-

nium 80-110 pm tall. Hypothecium to 200 pm deep, hya-

line. Paraphyses 1-3 pm, apices 3-5 pm wide. Asci cor-

responding to Lecanora type, usually 8-spored. Ascospores

(Figure IB) of Physcia type, septum in young spores in-

serted before internal wall thickenings become distinct,

torus developed, spores finely scabrid, without septal

swellings in KOH, (15-)16-27 x 8-13 pm. Spermogo-

nia (in material from southern Africa) immersed in

thallus, ostiolar region blackish. Spermatia 4. 0-5.5 x

1.0-1. 5 pm.

Bothalia 26,1 (1996)

15

B

(d\ (o\ (d\ (Q\

loj \o/

E

(Q (Qi 6) Si

loj \Ql [g (jg

F

f0\ /o\ /o\

b b b ^

G

© A /§\ fO)

p V® ^ ^

I

FIGURE 1. — Ascospores of southern African Buellia and Rinodina species. A, Buellia distrata, lectotype (BM); B, Rinodina confragosa, Zimbabwe,

Lundi River, 27-6-1963, Kofler s.n. (LD); C, R. confragosula, isotype (BM); D, R. confragosula (PRE 2089b); E, R. geesteranii, holotype

(L); F, R. gennarii, Almborn 1249 (GZU); G, R. huillensis, Brusse 4462 (PRE); H, R. huillensis, Rambold 7389 (M); I, R. longisperma,

holotype (PRE). Scale bars: 10 pm.

Rinodina confragosa is a widespread species in Europe

including adjacent Asia (e.g. Fox & Purvis 1992; Giralt &

Barbero 1995; Mayrhofer 1984a (distribution map);

Mayrhofer & Poelt 1979; Nimis 1993; Santesson 1993), but

it is hitherto only known from two localities in southern

Africa (Zimbabwe and Western Cape; Figure 2). It is also

reported from Australia (McCarthy 1991) and North

America (e.g. Egan 1987). The record of Lecanora con-

fragosa (= Rinodina confragosa) from Sao Tome (Insula

Caprarum, H-NYL 29006) by Nylander (1887, 1889) and

Stizenberger (1890) actually refers to a taxon of the Ri-

nodina oxydata group (see under R. oxydata in the present

study. Table 1). In the area studied, R. confragosa grows

on quartzitic rocks (including quartzitic sandstone); the

specimen from Zimbabwe was associated with a Peltula

sp. and cyanobacteria indicating that this habitat more or

less regularly receives liquid water.

Among the species treated in the present study, Ri-

nodina confragosa can be recognized easily by the K+

yellow thallus reaction and by the ascospores of Phy-

scia type. As far as thallus formation is concerned, the

species shows a wide range of variation in Europe, from

thinly crustose to only slightly squamulose (these mor-

photypes are also present in southern Africa) to very

thick and then with ± wart-like to squamulose areolae;

the thallus is continuous and rimose-cracked or com-

posed of discrete areolae, and a prothallus may be ab-

sent or strongly developed. Variability can also be ob-

served in chemistry, and different chemical races appear

16

Bothalia 26,1 (1996)

FIGURE 2. — Known distribution of Rinodina confragosa in southern

Africa, □; and of R. confragosula, •. Two collection localities

of R. confragosula could not be located in detail and are therefore

not mapped: Lesotho, Mamerthes; KwaZulu-Natal, Drakensberg,

on ridge of the Sugar Loaf near The Cavern.

to exist (Hecklau et al. 1981; Mayrhofer & Leuckert

1985).

Vouchers from southern Africa: Almborn 1834 (LD); Kofler s.n. (LD).

2. Rinodina confragosula (Nyl. in Cromb.) MullArg.

in Revue Mycologique (Toulouse) 9: 79 (1887). Type: West-

ern Cape, Table Mtn, Cape of Good Hope, 9-1874, A.E.

Eaton s.n., Venus Transit Expedition (BM, lecto.!, desig-

nated by Mayrhofer: 400 (1984a); BM, iso.!); (H-NYL

28565, iso.), not seen, compare Mayrhofer (1984a: 400).

Lecanora confragosula Nyl. in Cromb.: 172 (1876b).

Rinodina almbornii H.Mayrhofer: 374 (1984a). Type: Western Cape,

Dist. Paarl, Paarl Rock near summit, 2300 ft, 26-9-1953, O. Almborn

5540 (LD, holo.; GZU, iso.!).

Thallus crustose, rimose-areolate, some areolae occa-

sionally in the form of small squamules, greyish, grey-

brown, brown, occasionally whitish due to presence of a

pruina; in longitudinal section outermost layer of the phe-

nocortex is brown or, particularly in areolae which bear

an apothecium, (blackish to) blue-green; prothallus absent

or present, black. Chemistry: no lichen substances detect-

able by tic.

Apothecia to 0.8 mm diam., cryptolecanorine, more

rarely lecanorine or becoming lecideine and then adnate,

occasionally contiguous, disc black, rarely dark brown,

occasionally whitish pruinose, the disc may be surrounded

by a small edge of white pruina, disc slightly concave,

plane, or convex. Exciple: where a proper exciple is de-

veloped it is brown and blackish to blue-green in parts or

dominantly blue -green. Epihymenium 10-30(-35) pm tall,

entirely brown to olivaceous, brown to olivaceous and

blue-green in parts, or dominantly blue-green with brown

to olivaceous spots. Hymenium 70-120(-130) pm tall. Hy-

pothecium to 250 pm deep, hyaline; adjoining parts of the

thallus to the hypothecium may be blue-green. Paraphyses

1-5 pm, apices 3-6 pm wide. Asci corresponding to Le-

canora type, with 8 or less spores. Ascospores (Figure 1C,

D) with internal wall thickenings of Pachysporaria or

Milvina type, or intermediate between these two types,

occasionally also of Physcia type or spore lumina irregu-

larly biconical in shape, septum in young spores inserted

before internal wall thickenings become distinct, torus de-

veloped, spores finely scabrid, without septal swellings in

KOH, 14-32 x 7.5-16.0 /an. Spermogonia immersed in

the thallus, with the upper parts prominent or not, often

contiguous, ostiolar region black or brown; in longitudinal

section at base and laterally hyaline, rarely pale brownish,

above brown to olivaceous and/or blackish to blue-green.

Spermatia (3-)4— 5 x 1.0-1 .5 pm.

Chemical reaction of the blue-green pigment in the

thallus, apothecia, and spermogonia: N+ red.

Rinodina confragosula grows on hard siliceous rocks

such as granite, sandstone, and quartzite. It is known with

certainty only from the Republic of South Africa and Leso-

tho where it was found to be widely distributed at altitudes

between 450 m and 3 080 m (Figure 2). The specimen cited

from Concepcion in Argentina (Muller Argoviensis 1889)

represents a Rinodina species for which an appropriate name

is not available at present. A record from New Caledonia

was given by Muller Argoviensis (1887).

R. confragosula is a quite variable species in several

respects (see description). It can be recognized best by

the usually black apothecia, the presence of a blue-green,

N+ red pigment in the phenocortex, apothecia, and sper-

mogonia, and by the absence of lichen substances detect-

able by tic. Within a single specimen, the range of spore

size is often significantly smaller than given in the de-

scription of the species above, but in other specimens

the range was found to be 18-31 x 10-16 pm ( PRE

2089b), 20.0-27.5 x 9-11 /<m ( Brusse 2547), and 22-32

x 10.5-16.0 pm ( Brusse 4575).

Other saxicolous species occurring in the study area

and showing close similarities with R. confragosula in-

clude R. longisperma, R. microlepidea, and R. subtristis.

Rinodina longisperma is distinguished from R. confrago-

sula by smaller apothecia and ascospores, and by longer

spermatia. The separation of R. confragosula from R. mi-

crolepidea and R. subtristis respectively, is discussed in

detail under both the latter species.

The differences used for the delimitation of Rinodina

almbornii from R. confragosula by Mayrhofer (1984a) are

considered to fall within the variability of R. confragosula.

Mayrhofer (1984a) pointed out that R. confragosula is

closely related to R. kozukensis (Vain.) Zahlbr. which was

described from Japan by Vainio (1921, as Melanaspicilia

kozukensis). The relationship of these two species will be

discussed in a forthcoming paper.

Rinodina argentiniana Miill.Arg. (Argentina, Concep-

cion, 1882, Lorentz, G, holo.!) differs from R. confrago-

sula by having ascospores lacking a torus and by a

different type of spore ontogeny (internal apical wall

thickenings appear before the spore septum is inserted).

The record of R. argentiniana from Lesotho (summit of

Masiti Mountain, 6300 ft, 1929-30, Hewitt, TRH) by

Mayrhofer (1984a) actually refers to R. confragosula. Ac-

cording to the material available, R. argentiniana does not

occur in the study area.

Bothalia 26,1 (1996)

17

One of the specimens cited by Mayrhofer (1984a) un-

der Rinodina depressa could not be determined with cer-

tainty, and it is treated here under R. cf. confragosula :

Western Cape, Van Rhynspass, Van Rhynsdorp, Van der

Byl 766 (W).

Vouchers: Almborn 4567 (GZU, LD), 4847 (LD), 5540 (PRE), s.n.

(GZU); Brusse 2547, 2633, 3136, 3282, 3397, 4575, 4625, 5557 (PRE);

O'Connor CH 2220 (PRE); Hean (PRE 2089b): Hewitt s.n. (TRH); Van

der Plank, CH 4568 (PRE); Schelpe 1110 (GZU); Triebel & Rambold

7896 (GZU), 7901 (M).

3. Rinodina geesteranii H. Mayrhofer in Journal

of the Hattori Botanical Laboratory 55: 412 (1984a). Type:

Western Cape, Wynberg Flats, SE of Cape Town, on gran-

ite outcrop near dusty road, 19-12-1949, R.A. Maas

Geesteranus 14669 (L, holo.!; GZU, iso.!).

Thallus thick, crustose to slightly squamulose, areolate,

brown, covered by a white pruina in parts, large areolae

and squamules with an irregular surface; thallus occasion-

ally associated with cyanobacteria; prothallus dark brown,

blackish, well developed, at margin of thallus distinctly

effigurate. Chemistry: no lichen substances detectable by

tic (Mayrhofer & Leuckert 1985).

Apothecia to 0.5 mm diam., cryptolecanorine or lecanor-

ine and then adnate, rarely with a lecideine appearance, disc

blackish, plane. Epihymenium 10-20 pm tall, brown.

Hymenium 100-115 /mi tall. Hypothecium to 120 pm deep,

usually hyaline. Paraphyses 1^4 pm, apices 4—5(-6) pm

wide. Asci corresponding to Lecanora type, axial body often

difficult to observe, asci often with 8 but also with less

spores. Ascospores (Figure IE) with internal wall thickenings

intermediate between Pachysporaria and Milvina type, on-

togeny of spores apparently irregular: septum in young

spores inserted before or after internal wall thickenings be-

come distinct, torus absent, spores finely scabrid, without

septal swellings in KOH, 14-21(-24) x 8— 13(— 15) pm. Sper-

mogonia immersed in the thallus, ostiolar region blackish.

Spermatia ± 3-4 x 1-2 pm.

Rinodina geesteranii is only known from the type lo-

cality on the Cape Peninsula where it was collected on

hard quartzitic rocks (Figure 3). It is associated with other

lichen species, most commonly with Peltula euploca

(Ach.) Poelt in Pisut, suggesting that the locality is more

or less regularly wetted by rain or dripping water (com-

pare notes on the ecology of Peltula euploca in Biidel

1987: 58). This view is also supported by the fact that

the thallus of R. geesteranii is partly associated with cy-

anobacteria; green algae containing lichens which have

connections with free-living cyanobacteria occur exclu-

sively at localities which receive liquid water (Poelt &

Mayrhofer 1988).

R. geesteranii is characterized mainly by the thickly crus-

tose to slightly squamulose thallus, and by the ascospores

lacking a torus and with internal wall thickenings interme-

diate between Pachysporaria and Milvina type. Three other

saxicolous Rinodina species with ascospores lacking a typi-

cal toms are known to occur in the study area, viz. R. gen-

narii, R. reagens, and R. teichophiloides. In R. teicho-

philoides, toms-like structures may be developed (see ac-

count of R. teichophiloides in the present study). Rinodina

gennarii is separated from R. geesteranii mainly by a thin

FIGURE 3. — Known distribution of Rinodina geesteranii, □; and of R.

gennarii in southern Africa, •.

thallus and ascospores with internal wall thickenings of

Physcia type, and R. reagens and R. teichophiloides differ

from R. geesteranii by the usually larger ascospores with

internal wall thickenings of Mischoblastia or Pachysporaria

type. In ascospore type and size close similarities can be

observed between R. geesteranii and R. argentiniana

Miill.Arg. (G, holo.!). The latter was described from Argen-

tina by Muller Argoviensis (1889). Both the species are only

known from their type collections (for R. argentiniana see

the discussion of R. confragosula in the present work). R.

argentiniana is distinguished by the thinner, exclusively

crustose thallus and larger apothecia (to 0.75 mm in di-

ameter).

4. Rinodina gennarii Bagl. in Commentario della

Societa Crittogamologica Italiana 1: 17 (1861). Type: Italy,

Liguria occidentale, alle falde del monte Faiallo dell’ Ap-

ennino sopra Voltri, Baglietto s.n. (WU, iso.), not seen.

Synonyms: see Sheard (1967, under Rinodina subexigua ), Mayrhofer

& Poelt (1979), Mayrhofer (1984a).

The following description is based only on the speci-

mens cited below.

Thallus thin, occasionally inconspicuous, cmstose, areo-

late or ± effuse, smooth, whitish, grey, ochraceous; prothallus

absent. Chemistry: no lichen substances detectable by tic.

Apothecia to 0.5 mm diam., often numerous, crowded,

lecanorine or rarely lecideine, adnate to sessile, disc dark

brown, a narrow brown border-line between disc and thalline

margin may be present, disc plane to strongly convex. Epi-

hymenium 10—15 pm tall, brown. Hymenium 60-80 //m tall.

Hypothecium to 150 /mi deep, hyaline. Paraphyses l-4(-5)

/<m, apices 3-6 //m wide. Asci corresponding to Lecanora

type, 8-spored. Ascospores (Figure IF) of Dirinaria type, sep-

tum in young spores inserted after the formation of internal

apical wall thickenings, torus absent, spores smooth or

finely scabrid, with or without septal swellings in KOH,

(11— )12— 17 x (5-)6-9 //m. Spermogonia not observed.

Rinodina gennarii is widespread in temperate regions

of the northern and southern hemisphere and is found on

18

Bothalia 26,1 (1996)

a wide variety of substrata (e.g. Fox & Purvis 1992;

Mayrhofer 1983, 1984a; Mayrhofer & Poelt 1979; Nimis

1993; Santesson 1993). The species is here recorded for

South Africa where it occurs on granitic and quartzitic

rocks from low to high altitudes (Figure 3).

This species is characterized mainly by ascospores of

Dirinaria type and the numerous, often crowded, small

apothecia. Giralt & Mayrhofer (1995) did not observe

marked differences between R. gennarii and the corticol-

ous/lignicolous R. oleae Bagl. However, we refrain here

from putting R. gennarii into the synonymy of R. oleae.

R. gennarii and R. oleae are tentatively treated as a pair

of closely related, similar species of which one is saxi-

colous and the other corticolous/lignicolous. Further ob-

servations, including detailed data on distribution and

ecology are necessary to decide whether this concept can

be maintained or not.

Rinodina gennarii and R. oleae are not the only ex-

ample for vicarious species within the genus. Another cor-

responding pair of species is formed by the saxicolous R.

beccariana Bagl. and the corticolous R. roboris (Duf. ex

Nyl.) Arnold var. roboris (Mayrhofer et al. 1993). Fur-

thermore, within R. oxydata s.l., almost all representatives

are saxicolous with the exception of R. euskadiensis A.

Crespo & M.B.Aguirre which is based on corticolous ma-

terial (Crespo & Aguirre 1984; Giralt & Matzer 1994).

Vouchers from southern Africa: Almborn 1227 , 1250, 5020 (LD),

1249 (GZU, LD); Triebel & Rambold 6714 (M, under Buellia sp.), 6732

(M, under Rinodina sp.), 8399 (M).

5. Rinodina huillensis Vain, in Catalogue of Afri-

can Plants collected by F. Welwitsch 2,2: 413 (1901).

Type: Angola, Huilla [ = Huila] (3800 ad 5500 ped.s.m.),

ad rupes juxta cataractam Ferrao prope Lopollo, 1860,

Welwitsch 50 (TUR-V 8736, holo.!).

Buellia depressa Vain.: 415 (1901). Rinodina depressa (Vain.) Zahlbr.:

510 (1931). Type: Angola, Pungo Andongo, Pedra de Cazella, ad rupes

et lapides gneissaceas, 1857, Welwitsch 64 (TUR-V 9112 , lecto.!), desig-

nated by Mayrhofer (1984a: 403); Pungo Andongo, ad rupes et lapides

dispersas mont. Praesidii, Febr. 1857, Welwitsch 61 pr. p. (BM, para.!);

Pungo Andongo, ad rupes et lapides dispersas, Febr. 1857, Welwitsch 64

pr. p. (BM, para.!); Pungo Andongo, Pedra de Cazella, ... [text illegible]

... , 1857, Welwitsch 481 pr. p. (BM, para.!).

Rinodina albicans H. Mayrhofer: 373 (1984a). Type: Western Cape,

Dist. George, 7 miles E of G., on rocks in a ravine, 14-8-1953, O. Alm-

born 2387 (LD, holo.!).

Thallus usually thin, crustose, rimose-areolate, more

rarely effuse, brown, grey-brown, grey, rarely whitish,

pale ochraceous; phenocortex of apothecia-bearing areolae

is brown, olivaceous, blackish, olivaceous to blackish pig-

ment reacts N+ red; prothallus absent or present, black.

Chemistry: medulla C+ red; tic: gyrophoric acid (often

together with lecanoric and orsellinic acid), 5-O-methyl-

hiascic acid was detected in a single specimen.

Apothecia to 0.8 mm diam., cryptolecanorine, or le-

canorine or more rarely lecideine and then adnate, occa-

sionally a proper margin is developed, disc dark reddish

brown, dark brown, black, plane to convex. Proper exciple

in its outer parts brown or olivaceous to blackish and then

N+ red. Epihymenium 10-20 tall, reddish brown,

brown, occasionally dirty olivaceous and then N+ red in

part. Hymenium 70-120 tall. Hypothecium to 150 pm

deep, hyaline. Paraphyses \-A pm , apices 3-5(-6) pm

wide. Asci corresponding to Lecanora type, axial body

easy to observe or indistinct, sometimes surrounded by a

tube-like structure, asci usually with 8 but also with less

than 8 spores. Ascospores (Figure 1G, H) of Pachysporaria

type, occasionally grading into Milvina type, small pro-

trusions of lumina towards septum and/or spore ends

sometimes present, septum in young spores inserted be-

fore internal wall thickenings become distinct, torus de-

veloped, spores smooth to finely scabrid, without septal

swellings in KOH, (10-)12-18(-24) x (6-)7-10(-12) /rm.

Spermogonia immersed in thallus, partly somewhat raised,

ostiolar region (dark) red-brown to blackish. Spermatia

3-6(-7) x 1. 0-1.5 pm.

Rinodina huillensis is a widespread species in south-

ern Africa and is so far known from Angola, the Re-

public of South Africa, and Swaziland. It occurs on

granite, quartzitic rocks and sandstone at a range of

altitudes (Figure 4).

Among the saxicolous Rinodina species known to oc-

cur in the study area, R. huillensis alone contains gyro-

phoric acid in the thallus (medulla C+ red), often

accompanied by lecanoric and orsellinic acid. It is a well-

known phenomenon in many lichens that in addition to

gyrophoric acid, at least traces of lecanoric and orsellinic

acid can be found (e.g. Schreiner & Hafellner 1992: 22).

Further saxicolous species of Rinodina containing gyro-

phoric acid occur outside southern Africa, but these are

distinguished from R. huillensis by several characters

(Matzer et al. 1994a). In a single specimen ofR. huillensis

FIGURE 4. — Known distribution of Rinodina huillensis.

Bothalia 26,1 (1996)

19

(Brusse 4497 , PRE) 5-0-methylhiascic acid was detected.

This lichen compound was also reported from Rinodina

tephraspis (Tuck.) Herre, a northern hemisphere species

which differs chemically from R. huillensis by the pres-

ence of zeorin (Mayrhofer et al. 1992).

Re-examination of the types of Rinodina albicans

and R. depressa revealed that these names are synonyms

of R. huillensis. Vainio (1901) described R. huillensis and

R. depressa (as Buellia depressa) in the same publication;

R. huillensis is selected here as the correct name as it

was mentioned first in Vainio’s paper and appropriate

generic classification was applied. The statement of

Mayrhofer & Leuckert (1985) that R. depressa does

not contain lichen substances is based on the following

specimen: Republic of South Africa, Roggeveld Moun-

tains, Brusse 3265 (PRE). We studied this specimen, and

it may represent a Rinodina species but cannot be iden-

tified with certainty as the ascospores are badly devel-

oped. From its external morphology it can be concluded

that it is not R. huillensis. One of the specimens (Van

der Byl 766, W) cited under R. depressa by Mayrhofer

(1984a) is treated as Rinodina cf. confragosula in the

present paper. Several specimens mentioned by Mayr-

hofer (1984a) under R. huillensis do not belong to this

species: one from Angola (Humpata Plateau, 6-2-1960,

Degelius, GZU) refers to R. subtristis whereas the other

one from the same country was not examined. The ma-

terial from Namibia (Haifischbucht, Fincke, W) repre-

sents Rinodina longisperma , a species newly described

in the present study. The specimen from the Northern

Province (Transvaal, Louis Trichardt, 8-10-1953, Almborn,

LD) was selected as the isotype of Rinodina scabridula

in the present work. Another specimen from Mpu-

malanga (Eastern Transvaal) was also cited: Dist. Pil-

grims Rest, 4 miles SE of P. R., on rocks near the

road, 22-10-1953, Almborn (LD). We examined two

specimens held in LD which are labelled as indicated

above but with different collection numbers: Almborn

7831 actually represents R. huillensis whereas the crus-

tose lichen species in Almborn 7836 has hyaline asco-

spores and does not belong to the Physciaceae. The

collection of R. huillensis from (Orange) Free State

(Ladybrand, Maas Geesteranus 6534, L, LD) was cited

under Rinodina microlepidea by Mayrhofer (1984a).

Vouchers: Almborn 4579 (LD, under Arthonia sp.), 7831, 7865, 8615

(LD), 6168 (PRE); Brusse 1688, 3703 (GZU), 1634 (PRE, under Rino-

dina substellulata), 1688, 1764, 2773, 2947, 3581, 3655, 4453, 4462, CH

4628 (PRE), 4497 (GZU, PRE); Van der Byl 756 (W); J. Hafellner &

A. Hafellner, 30651 (Hafellner); Maas Geesteranus 6534 (L, LD); Triebel

& Rambold 7353 (GZU, M), 7389 (M).

6. Rinodina longisperma Matzer & H.Mayrhofer

sp. nov.

Thallus saxicola, crustaceus, rimosi-areolatus, griseus

ad bmnneus, interdum pruinosus. Apothecia ad 0.3 mm

in diametro, cryptolecanorina; disci fusci ad atri, plani.

Epihymenia 5-15 pm alta, brunnea. Hymenia 50-90 pm

alta. Hypothecia ad 100 pm alta, hyalina. Paraphyses

1— 3( — 4) pm crassae, apicibus (2-)4— 5(-6) pm crassis. Asci

tholis similibus typo generis ‘Lecanora’ instructi. Asco-

sporae 10-16(-18) x 6-10(-12) pm magnae, typo Pachy-

sporaria vel Physconia similes, toro evoluto. Spermogonia

in thallo immersa. Spermatia bacilliformia, (4-)6-9 x

1.0-1 .5 pm magna.

TYPE. — Namibia, 2216 (Otjimbingwe): 10-20 km S

of Windhoek, Auasberge, Regenstein Farm, gorge behind

fort, ( — DC), on quartzite, 26-3-1984, F. Brusse 4264

(PRE, holo., with Cercidospora sp. and Dactylospora sp.;

GZU, iso.).

Thallus crustose, rimose-areolate, grey, grey-brown,

brown, occasionally whitish pruinose; prothallus occasion-

ally developed, dark brown, black. Chemistry: no lichen

substances detectable by tic.

Apothecia to 0.3 mm diam., cryptolecanorine, disc dark

reddish brown, dark brown to blackish, partly with a small

border line of white pruina, plane. Epihymenium 5-15 pm

tall, brown. Hymenium 50-90 «m tall. Hypothecium to

100 pm deep, hyaline. Paraphyses l-3(-4) pm, apices

(2-)4— 5(-6) pm wide. Asci corresponding to Lecanora

type, axial body often indistinct or not observable, asci

usually 8-spored. Ascospores (Figures II; 5A) similar to

Pachysporaria type or ± of Physconia type, occasionally

with strongly pronounced apical internal wall thickenings,

partly with protrusions of lumina to spore ends and/or

septum, septum in young spores inserted before internal

wall thickenings become distinct, torus small but well dis-

tinct, spores smooth, without septal swellings in KOH,

10— 16(— 18) x 6-10(-12) pm. Spermogonia immersed in

thallus, ostiolar region grey, brown, blackish. Spermatio-

phores septate; spermatiogenous cells usually terminal,

rarely intercalar; terminal spermatiogenous cells flask-

shaped, forming spermatia apically; intercalar spermatio-

genous cells with a lateral projection below upper

transverse septum with spermatia formed on lateral pro-

jections. Spermatia (4-)6-9 x 1.0-1 .5 pm.

Rinodina longisperma is so far known from quartzitic

rocks including sandstone from Namibia and the Republic

of South Africa (Figure 6).

Rinodina longisperma can be recognized best by a

set of characters including the small and cryptolecanor-

ine apothecia, the small ascospores similar to Pachy-

sporaria or Physconia type, and, particularly, by the

relatively long spermatia (hence the epithet ‘longis-

perma'). The bulk of Rinodina species in which sper-

matia have been observed posses spermatia with length

measurements from 3 to 6 (to 7) pm, whereas in R.

longisperma they are (4 — )6— 9 pm long. In addition, pe-

culiarities could also be observed in the spermatio-

genous apparatus of R. longisperma. Most of the

Rinodina species studied in this respect have many ter-

minal and intercalar spermatiogenous cells (e.g. see

‘morphology and anatomy’ in the general part of the

present study; Matzer et al. 1994a: 108, fig. 8). In R.

longisperma most of the spermatiogenous cells are ter-

minally arranged, whereas those with an intercalar po-

sition could be observed only exceptionally.

One of the specimens of R. longisperma from Namibia

(Haifischbucht, Fincke, W) was included in R. huillensis

by Mayrhofer (1984a).

20

Bothalia 26,1 (1996)

FIGURE 5. — Ascospores of southern African Rinodina species. A, R. longisperma, holotype (PRE); B, R. scabridula , holotype (GZU); C, D, R.

striatitunicata, holotype (GZU); D, ascospores in surface view showing their striate ornamentation. Scale bar: 10 «m.

Specimens examined :

NAMIBIA. — 2016 (Otjiwarongo): 7 km S of Otjiwarongo, (-DA),

21-3-1984, Brusse 4210 (PRE). 2017 (Waterberg): Waterberg plateau,

(-AD), 23-3-1984, Brusse 4246 (PRE). Haifischbucht, grid. ref. un-

known, Fincke s.n. (W).

NORTHERN CAPE. — 3123 (Victoria West): Dist. Victoria West,

Three Sisters, (-CC), 2-10-1953, Atmborn 5779 (LD).

7. Rinodina microlepidea MiillArg. in Flora 71:

206 (1888). Type: Mpumalanga, Eastern Transvaal, Ly-

denburg, Wilms 76 (G, holo.!).

Lecanora microlepidea (Miill.Arg.) Stizenb.: 209 (1890).

Thallus thin, crustose, rimose-areolate, continuous or

composed of discrete areolae, whitish, yellowish, ochra-

ceous; prothallus not distinct. Chemistry: no lichen sub-

stances detectable by tic.

Apothecia to 0.4 mm diam., lecanorine, rarely cryp-

tolecanorine, disc light brown to dark brown or blackish,

± plane. Epihymenium 5-20 pm tall, brown. Hymenium

± 90-100 /tm tall. Hypothecium to 180 pm deep, hyaline.

Paraphyses 1-3 pm, apices 3-4 pm wide. Asci corre-

sponding to Lecanora type, usually 8-spored. Ascospores

(Figure 7A) of Pachysporaria type, septum in young

spores probably inserted before' internal wall thickenings

Bothalia 26,1 (1996)

21

FIGURE 6. — Known distribution in southern Africa: Rinodina longi-

sperrna , •, [locality of Fincke s.n. (W) could not be located

exactly and is therefore not mapped: SWA/Namibia, Hai-

fischbucht]; R. microlepidea , ▲; R. reagens, ■; R. scabridula ,

O; and/?, striatitunicata, □.

become distinct, torus small (not always distinct), spores

smooth to finely scabrid, (13— )15— 19(— 20) x 8-15 /tm.

Spermogonia immersed in thallus, ostiolar region pale,

brown. Spermatid 4—5 x 1.0-1 .5 /um.

In its present circumscription, the species is known

only from single localities in the Republic of South Africa

and Zimbabwe where it grows on hard siliceous rocks

(Figure 6).

The status of Rinodina microlepidea is rather vague.

The description given above is based only on two rather

scant specimens and thus, not all characters (e.g. ascus

type, behaviour of ascospores in potassium hydroxide)

could be studied in full detail. Unfortunately, the type

collection is in a very poor condition, and the apothecia

in particular are very old or damaged. Therefore, no

statements can be given on the variability of R. mi-

crolepidea and only one further specimen which looks

more or less similar to the type is accepted as belonging

to that taxon. The distinction between Rinodina subtris-

tis and R. microlepidea is problematical. According to

the material available, the former species is separated by

larger apothecia (to 0.7 mm diam.) and the coloration of

the thallus: brown or ochraceous instead of whitish, yel-

lowish to ochraceous. Like R. microlepidea and R. sub-

tristis, other Rinodina species in the study area are

characterized by the lack of secondary lichen substances

and ascospores of Pachysporaria or similar types. This in-

cludes R. confragosula, R. longisperma, and R.

scabridula. Rinodina confragosula differs from R. mi-

crolepidea mainly by its larger apothecia (to 0.8 mm

diam.) and by ascospores which are rather variable in the

formation of internal wall thickenings (Pachysporaria,

Milvina, Physcia, or intermediate types or the spore lu-

mina are irregularly biconical) and exhibit a broader

size range (14-32 x 7.5-16.0 pm). Rinodina longis-

perma is separated from R. microlepidea mainly by its

often smaller ascospores, 10— 16(— 18) x 6-10(-12) pm,

and longer spermatia, 5-9 pm. Rinodina scabridula can

easily be distinguished by the formation of blastidia on

the thallus.

The following specimens were included in R. mi-

crolepidea by Mayrhofer (1984a), but for a range of rea-

sons a new settlement is proposed here: (Orange) Free

State, Ladybrand, Maas Geesteranus 6534 (L, LD) be-

longs to Rinodina huillensis. The specimen from (Orange)

Free State, Dist. Trompsburg, 2-10-1953, Almborn 5809

(LD) which is cited by Mayrhofer (1984a) with the date

‘2-10-1954’, collection number ‘ 5808 ’, and herbarium ‘L’

represents a Rinodina sp. The ascospores are usually 2-

celled but exceptionally also 3-celled spores could be ob-

served. The Rinodina species from the Western Cape,

Calitzdorp-Kruisrivier, Maas Geesteranus 6711 (L) differs

from R. microlepidea by the type of spore ontogeny in

the way that internal wall thickenings appear before the

septum is inserted. The ascospores of this specimen re-

semble those of R. geesteranii, but differences exist in the

morphology of the thallus.

Voucher: Kofler s.n. (LD).

8. Rinodina oxydata s.l.

Rinodina oxydata s.l. is distributed world-wide. The

following description is based on material from southern

Africa only. For further explanations see the discussion

given below.

FIGURE 7. — Ascospores of southern African Rinodina species. A, R.

microlepidea , holotype (G); B, R. oxydata s.l., [holotype of R.

minima ] (LD); C, R. oxydata s.l., Almborn 7104 (LD); D, R.

reagens, Hafellner 30655 (herb. Hafellner). Scale bar: 10 /(in.

22

Bothalia 26,1 (1996)

Thallus thin, crustose, areolate, continuous, yellowish,

pale ochraceous, pale greyish; prothallus absent or present,

blackish. Chemistry: thallus K+ yellow; tic: atranorin (not

found in Rinodina minima; see below).

Apothecia to 0.4 mm diam., cryptolecanorine, lecanor-

ine, or with a lecideine appearance (algal cells may be

enclosed in the exciple), when lecanorine or lecideine then

adnate, disc dark reddish brown to blackish, proper margin

black, disc usually plane. Proper exciple in its outer layer

with an olivaceous, blue-green to blackish pigment which

reacts N+ red, in addition a brown pigment (N-) may be

present. Epihymenium 10-20 pm tall, brown. Hymenium

70-100 //m tall. Hypothecium to ± 80 pm deep, hyaline.

Paraphyses 1—4 pm, apices 3-6 pm wide. Asci corre-

sponding to Lecanora type, with tendencies towards

Bacidia type, axial body may be indistinct, asci often with

8 but also with less spores. Ascospores (Figure 7B, C)

usually of Mischoblastia type, rarely with tendencies to-

wards Pachysporaria type, septum in young spores in-

serted before internal wall thickenings become distinct,

torus delicate, spores smooth or finely scabrid, with or

without slight septal swellings in KOH, 11-25 x (7-)8-13

^m. Spermogonia immersed in thallus, occasionally

slightly raised, ostiolar region dark reddish brown to

blackish. Spermatia (3-)4— 5 x 1.0— 1.5 pm.

In the study area, Rinodina oxydata s.l. is known only

from three localities (Mozambique, Transvaal, Table

Mountain) where it grows on schistic and quartzitic rocks.

The name Rinodina oxydata is commonly applied to

lichens exhibiting the following set of characters: thallus

thinly crustose or thick and nearly squamulose, occasion-

ally evanescent, containing atranorin (K+ yellow),

apothecia showing a wide variety from cryptolecanorine,

lecanorine to lecideine, ascospores (usually) of Mis-

choblastia type. Particularly, in thallus formation, in the

apothecial type, and, in part, in ascospore size, a wide

range of variability can be observed. Many binomials have

been introduced for lichens with characters mentioned

above, but none of these names can be used unequivocally

unless a thorough revision of the R. oxydata group on a

world-wide scale has been carried out (see Table 2). Such

a revision should also include R. substellulata, a species

with ascospores of Pachysporaria type occasionally grad-

ing into Milvina type, but in all other characters displaying

close similarities with R. oxydata. All characters used for

the delimitation of species (thallus formation, apothecial

type, ascospore size) are not constant and show various

transitional states, especially when many specimens are

analyzed. On the other hand, it seems to be premature to

put all these names into synonymy of R. oxydata (A.Mas-

sal.) A.Massal. As a consequence, the description given

above includes all R. oxydata- like lichens known from

southern Africa, and a survey on the ‘species’ known from

that region is given in Table 1 below.

Rambold et al. (1994) argued for a possible re-estab-

lishment of the generic name Mischoblastia A.Massal.

which may be based on Rinodina oxydata. The most im-

portant characters mentioned to be characteristic for Mis-

choblastia are: asci corresponding to Bacidia type, asco-

spores of Mischoblastia type, presence of an aeruginose,

N+ red pigment in the exciple. However, it should be

stressed here that all these characters do not justify a

separation of Mischoblastia from Rinodina (Ach.) Gray;

this is evident from the following observations: 1, asci

of R. oxydata (and of species which are considered to be

closely related to R. oxydata ) exhibit a considerable range

of variation and correspond either to Bacidia type or to

Lecanora type, or are more or less intermediate between

these two types; 2, the presence of Mischoblastia type

ascospores cannot be an argument as the formation of

pronounced internal wall thickenings is one of the most

important characters of the genus Rinodina; 3, an aerugi-

nose, N+ red pigment occurring in the exciple and/or in

the epihymenium is known not only from R. oxydata and

related species but also from other Rinodina species: e.g.

R. beccariana Bagl. (Mayrhofer et al. 1993 ),R. canarien-

sis Matzer, H.Mayrhofer & P.Clerc (Matzer et al. 1994a),

R. confragosula (Nyl. in Cromb.) Miill.Arg. (see present

study), and R. trachytica (A.Massal.) Bagl. & Car. (Mayr-

hofer et al. 1992).

Specimens examined

MOZAMBIQUE: — Sul do Save, Dist. Maputo (Lourengo Marques),

8 km E of Impamputo, on rocks near road, 18-10-1953, Almborn 7102,

holotype of Rinodina minima (LD); same locality, 18-10-1953, Almborn

7104 (LD).

NORTHERN TRANSVAAL. — On granite from the Lebombo in

Transvaal, Wilms s.n., holotype of Rinodina detecta (ZT).

WESTERN CAPE. — Cape of Good Hope, Table Mtn, September

1874, Eaton s.n., Venus Transit Expedition (BM, associated with the lec-

totype of Rinodina subtristis).

TABLE 1. — Rinodina oxydataAWe lichens known to occur in southern Africa; for further explanations see the discussion in the text

* A second specimen was cited by Mayrhofer (1984a: 440): Sao Tome & Principe: Insula Caprarum sin. Guineensis, Ilha das Cabras, 1887, Newton

(H-NYL 29006). It is in a rather bad condition and differs from the holotype by a comparatively thick thallus which is composed of more or less

discrete areolae.

Bothalia 26,1 (1996)

23

TABLE 2. — Selection of ‘species’ which have to be considered in the

course of a revision of the Rinodina oxydata group

9. Rinodina reagens Matzer & H.Mayrhofer in

Acta Botanica Fennica 150: 116 (1994). Type: New Zea-

land, Southland, Waiau River Valley, Clifden N Tuatapere,

167Q42’E, 46a02’S, limestone, 22-9-1981, H. Mayrhofer

2186 , associated with Rinodina bischoffii, (GZU, holo.!;

M, Mayrhofer, iso.!).

Exs.: Plantae Graecenses Lichenes 433, as Rinodina teichophila.

leones: Matzer & Mayrhofer (1994).

Thallus crustose to somewhat squamulose, rimose areo-

late, continuous or of discrete areolae, brown to ochraceous,

smooth but sometimes covered in part by a (?lichenized)

cyanobacterium ( Gloeocapsa sp.) resulting in a coarsely

warty appearance, occasionally associated with additional,

filiform cyanobacteria ( Scytonema sp.); prothallus indistinct.

Chemistry: no lichen substances detectable by tic.

Apothecia to 1.7 mm diam., cryptolecanorine or le-

canorine and then adnate, disc reddish brown, dark brown

to black, often with a narrow, pale reddish brown border-

line between disc and thalline margin, plane to convex,

thalline margin persistent, entire or slightly crenate. Epi-

hymenium 10-30 pm tall, brown or yellowish brown, Iv-

or K+ yellowish to orange. Hymenium 100-150 pm tall.

Hypothecium to 180 pm deep, entirely hyaline, hyaline

and green-yellow in parts, or entirely green-yellow, K+

yellowish to orange. Paraphyses 1—4 pm, apices to 6 pm

wide. Asci corresponding to Lecanora type, 8-spored or

with fewer spores. Ascospores (Figure 7D) with internal

wall thickenings of Mischoblastia type, occasionally grad-

ing into Pachysporaria type, septum in young spores in-

serted before internal wall thickenings become distinct,

toms absent, spores finely scabrid, with septal swellings

in KOH, 22-31 x 10-19 pm. Spermogonia immersed in

thallus, with a reddish brown, dark brown or blackish

ostiolar region; in longitudinal section reddish brown to

brown above, otherwise hyaline to green-yellow, K- or

K+ yellowish to orange. Spermatia (3.5-)4.0-4.5(-5.5) x

1.5-2.0 pm.

This species is hitherto known from the Western Cape

in southern Africa (Figure 6) and a few inland locations

in New Zealand (Matzer & Mayrhofer 1994). In the study

area it is confined to calcareous sandstone.

A detailed discussion of the species was provided just

recently by Matzer & Mayrhofer (1994) and thus, only a

few notes are given here. Rinodina reagens is mainly

characterized by the presence of a green-yellow pigment

in the hypothecium, epihymenium, and in the spermogo-

nia, which reacts K+ yellowish to orange. This pigment

is not known from any other Rinodina species. The large

ascospores without a torus are also a distinctive character

which separates the taxon from almost all other Rinodina

species known to occur in the study area. Similar spores

are found in Rinodina teichophiloides which is closely

related to R. reagens. These sympatric species are con-

fined to different substrates. In South Africa R. reagens

grows on calcareous sandstone, whereas R. teichophi-

loides is restricted to hard siliceous rocks.

Vouchers from southern Africa: Brusse 2874, 3852 (PRE); J. Hafell-

ner & A. Hafellner, 30654, 30655 (Hafellner).

10. Rinodina scabridula Matzer & H.Mayrhofer

sp. nov.

Thallus saxicola, crustaceus, rimosi-areolatus, brunneus

ad griseus, blastidia formans. Apothecia ad 0.45 mm in dia-

metro, lecanorina ad lecideina, immersa vel adnata; disci

brunnei ad atri, plani ad convexi. Epihymenia 5-20 pm alta,

brunnea. Hymenia 60-90 pm alta. Hypothecia ad 150 pm

alta, hyalina. Paraphyses 1-3 pm crassae, apicibus 3-5 pm

crassis. Asci tholis similibus typo generis ‘ Lecanora ’ in-

structi. Ascosporae (12-)15-20(-24) x (6-)7-10(-13) pm

magnae, typo Milvina, Pachysporaria vel Physcia similes,

toro evoluto. Spermogonia in thallo immersa. Spermatia ba-

cilliformia, ± 4—5 x 1.0-1 .5 pm magna.

TYPE. — Northern Province (Transvaal), Dist. Zout-

pansberg, Louis Trichardt, near the ‘The Punch-bowl’, ±

4500 ft, on sandstone rocks, 8-10-1953, O. Almborn 6169

(GZU, holo.); 6168 (LD, iso.).

Thallus crustose, rimose-areolate, partially blastidiate,

brown, grey-brown, or grey, occasionally associated with cy-

anobacteria; blastidia brown to dark brown, coralline when

well developed, usually situated at margin of thallus areolae,

rarely at thallus margin of apothecia; prothallus indistinct.

Chemistry: no lichen substances detectable by tic.

Apothecia to 0.45 mm diam., cryptolecanorine, or le-

canorine to lecideine and then adnate, disc dark brown to

blackish, plane to slightly convex, proper margin dark

brown to blackish. Epihymenium 5-20 /<m tall, brown.

Hymenium 60-90 pm tall. Hypothecium to 150 pm deep,

hyaline. Paraphyses 1-3 /rm, apices 3-5 //m wide. Asci

corresponding to Lecanora type, often with 8 but also with

less spores. Ascospores (Figures 5B; 8A) of Milvina,

Pachysporaria or Physcia type, or intermediate between

these types, septum in young spores inserted before inter-

nal wall thickenings become distinct, torus well devel-

oped, spores smooth to finely scabrid, without septal

swellings in KOH, (12-)15-20(-24) x (6-)7-10(-13) ^m.

Spermogonia immersed in thallus, ostiolar region black-

ish, somewhat raised. Spermatia ± 4-5 x 1.0-1. 5 pm.

24

Bothalia 26,1 (1996)

O P

o o

Rinodina scabridula is known hitherto from two localities

in the Northern Province (northern Transvaal) where it oc-

curs on sandstone and granitic rocks (Figure 6).

Rinodina scabridula is characterized mainly by the for-

mation of blastidia (for this term see Poelt 1980 and Giralt

et al. 1993a, as ‘Blastidien’) usually at the margins of

thallus areolae, and by its ascospores which are of Mil-

vina, Pachysporaria, Physcia or intermediate type. To de-

termine the species, careful observations are necessary as

blastidia are not formed by all thallus areolae and thus,

parts of the thalli are smooth. Consequently, confusion

with Rinodina subtristis (see present study) may be pos-

sible.

Other blastidiate, saxicolous Rinodina species have al-

ready been described: R. obnascens (Nyl.) Oliv. is a Euro-

pean lichen-parasitic species which is recorded growing

on species of Acarospora, Aspicilia, and Rhizocarpon (e.g.

Mayrhofer 1984a, 1987; Mayrhofer & Poelt 1979; Nimis

1993; Nimis et al. 1987). Rinodina furfurea H. Mayrhofer

& Poelt which is so far known only from the type locality

in South Tirol is separated from R. scabridula by usually

shorter ascospores lacking a torus (Mayrhofer & Poelt

1979). Rinodina blastidiata Matzer & H. Mayrhofer is a

maritime species occurring on coastal rocks in southeast-

ern Australia and in New Zealand. It is distinguished by

rather large spores (16-32 x 9-15 /<m) lacking a toms

and with internal wall thickenings of Mischoblastia type,

occasionally grading into Pachysporaria type (Matzer &

Mayrhofer 1994). For blastidiate Rinodina species grow-

ing on bark or wood see the studies of Giralt et al. (1993b,

1995) and Ropin & Mayrhofer (1995).

The specimen which has been selected as the isotype

of R. scabridula in the present work was treated under R.

huillensis by Mayrhofer (1984a).

Specimen examined

NORTHERN PROVINCE.— 2329 (Pietersburg): 20 km S of Mara

West Road near turnoff to Dendron, Farm Commissie Draai, grid. ref.

unknown, 950-1 000 m, 23-29-1-1981, Brusse 1628 (PRE).

11. Rinodina striatitunicata Matzer & H. Mayrho-

fer sp. nov.

Thallus saxicola, crustaceus, rimosi-areolatus, interdum

effusus, brunneus ad griseus. Apothecia ad 0.7 mm in dia-

metro, lecanorina ad lecideina, immersa vel adnata; disci

atri, plani ad convexi. Epihymenia 10-20 //m alta, brunnea

ad olivacea, N- vel N+ rubescens. Hymenia 90-120 pm

alta. Hypothecia ad 170 pm alta, hyalina. Paraphyses

1-3(^1) pm crassae, apicibus (2-)3-5(-6) pm crassis. Asci

tholis similibus typo generis ‘ Lecanora ’ instructs Asco-

sporae (16-)17-27 x (10— )1 1—17 pm, ad typum Tunicata

pertinentes, parietibus striatiformiter sculpturatis, toro non

vel bene evoluto.

FIGURE 8. — Ascospores of southern African Rinodina species. A, R.

scabridula, holotype (GZU); B, R. striatitunicata, holotype

(GZU), on left spore striate ornamentation is indicated; C, R.

substellulata, holotype (G); D, R. subtristis, lectotype (BM); E,

R. teichophiloides, holotype (ZT); F, R. sp. 1, Almborn 10610

(LD). Scale bars: 10 pm.

TYPE. — Western Cape, 3322 (Oudtshoorn): by the

road from Oudtshoorn to Prince Albert, near Ombinda

Karambi 7.5 km NW of the Cango Caves, (-AC),

33°23’30”, 22°08’45”E, ± 740 m. Karoo vegetation on

an E-exposed slope of a hill, 20-2-1992, J. Hafellner &

A. Hafellner 30650 (GZU, holo.).

Bothalia 26,1 (1996)

25

Thallus crustose, rimose-areolate, occasionally effuse,

dark brown, rarely grey, occasionally somewhat rusty and

rough; in longitudinal section with or without a blue-green

pigment which reacts N+ red; prothallus absent. Chemis-

try: no lichen substances detectable by tic.

Apothecia to 0.7 mm diam., cryptolecanorine, or le-

canorine to lecideine and then adnate, disc ± black, plane

to convex, proper margin black. Epihymenium 10-20 pm

tall, brown to dirty olivaceous (when dirty olivaceous N+

reddish). Hymenium 90-120 pm tall. Hypothecium to 170

urn deep, hyaline. Paraphyses 1— 3( — 4) pm, apices (2-)3-5

(-6) pm wide. Asci corresponding to Lecanora type, with 8

or less spores. Ascospores (Figures 5C, D; 8B) of Tunicata

type, internal wall thickenings corresponding to Physcia

type, septum in young spores inserted before internal wall

thickenings become distinct, torus well developed to in-

distinct, mature spores with striate ornamentation,

(16-)17-27 x (10— )11— 17 pm. Spermogonia not found.

Rinodina striatitunicata is only known from two lo-

calities in the Western Cape near the Cango Caves. It

grows on volcanic conglomerate and sandstone situated

in Karoo vegetation (Figure 6).

As indicated by the epithet, Rinodina striatitunicata is

characterized mainly by ascospores of Tunicata type with

peculiar striate ornamentation. The sculpture on the spore

surface is elongate and more or less parallel (compare

Scheidegger 1993: 335). In order to recognize the orna-

mentation in ascospores of R. striatitunicata, more or less

mature or overmature spores must be studied carefully

under the light microscope at a magnification of x 1000

or more; young ascospores are more or less smooth.

Other saxicolous Rinodina species with Tunicata type

ascospores are R. calcarea (Arnold) Arnold, R. tunicata

H.Mayrhofer & Poelt (compare Mayrhofer & Poelt 1979),

and R. filsonii H.Mayrhofer (Mayrhofer 1984b). They dif-

fer from R. striatitunicata in several respects and, as far

as the ascospores are concerned, mainly by a microrugu-

late (sculpture less than 1 pm in size, circular to elongate

and irregularly arranged) instead of a striate ornamenta-

tion.

The thallus of R. striatitunicata varies in consistency

and colour according to the rock mineralogy: a rough and

rusty morphotype was found on volcanic conglomerate

( Almborn 4320), whereas dark brown (to grey in small

parts) and compact thalli occur on sandstone ( Hafellner

30649 and 30650).

One specimen of R. striatitunicata ( Hafellner 30649)

is associated closely with small patches of another, prob-

ably undescribed Rinodina species which is distinguished

by its pale thallus and characters of the ascospores.

Specimens examined

WESTERN CAPE. — 3322 (Oudtshoom): Dist. Oudtshoom, 5 miles

N of Cango Caves, (-AC), 31-8-1953, Almborn 4320 (LD); by the road

from Oudtshoom to Prince Albert, near Ombinda Karambi 7.5 km NW

of the Cango Caves, (-AC), 33°23’30”S, 22°08'45”E, ± 740 m, 20-2-

1992, J. Hafellner & A. Hafellner 30649 (Hafellner, associated with Ri-

nodina sp.).

12. Rinodina substellulata MulLArg. in Proceed-

ings of the Royal Society of Edinburgh 11: 461 (1882).

Type: Ins. Socotra, Wadi Keschin, 650 m, 1881, Schwein-

furth s.n. (G, holo.!).

Lecanora substellulata (Miill.Arg.) Stizenb.: 209 (1890).

Rinodina quintana (Henriq. in Nyl.) Zahlbr.: 546 (1931). Lecidea

quintana Henriq. in Nyl.: 24 (1889). Lecanora quintana (Henriq. in Nyl.)

Nyl. in Hue: 55 (1891). Type: Sao Tome & Principe, Insula Principis,

1888, Quintas s.n. (H-NYL 28854, holo.!).

Rinodina quintana var. obscurior (Nyl.) Zahlbr.: 546 (1931). Lecidea

quintana var. obscurior Nyl.: 6 (1896). Type: Equatorial Guinea, Ins.

Guineensis, Annobon [ = Pagalu], 1892, F. Newton s.n. (H-NYL 10450,

holo.!).

Thallus thin, crustose, areolate or effuse, pale yellowish

or pale ochraceous; prothallus absent or present, blackish.

Chemistry: thallus K+ yellow; tic: atranorin.

Apothecia to 0.6 mm diam., cryptolecanorine, or le-

canorine or lecideine and then adnate to sessile, disc

brown, dark reddish brown to blackish, plane to strongly

convex, proper margin black. Proper exciple dark brown,

in addition a blue-green, N+ red pigment is often present.

Epihymenium 10-20 pm tall, brown, occasionally blackish

green and N+ red in parts. Hymenium 90-110 /tm tall.

Hypothecium to 200 pm deep, hyaline. Paraphyses 1-3

pm, apices 3-5(-6) pm wide. Asci corresponding to Le-

canora type, often with 8 but also with less than 8 spores.

Ascospores (Figure 8C) of Pachysporaria type, occasion-

ally grading into Milvina type, septum in young spores

inserted before internal wall thickenings become distinct,

torus often indistinct, sometimes not observable, spores

smooth to finely scabrid, without septal swellings in KOH,

14—20 x 7-12 pm. Spermogonia not found.

Rinodina substellulata is known with certainty from

some islands adjacent to the African continent (Principe

and Annobon in the Atlantic Ocean; Socotra in the Indian

Ocean), and from the Northern Province (northern Trans-

vaal) and the Western Cape in the Republic of South Af-

rica (Figure 9). It grows on hard siliceous rocks (including

quartzitic rocks, sandstone, and basalt). Further records

are given from Costa Rica (Stizenberger 1893), Indonesia

(Mayrhofer 1984a), and Australia (Mayrhofer 1984b).

R. substellulata is characterized mainly by the K+ yel-

low thallus reaction (indicating atranorin) and ascospores

of Pachysporaria type which may show tendencies to-

wards Milvina type. The species is closely related to taxa

of the Rinodina oxydata group. Rinodina oxydata and re-

lated species (see account of R. oxydata in the present

study) differ from R. substellulata mainly by having as-

cospores of Mischoblastia type. Rinodina beccariana var.

lavicola (J. Steiner) Matzer & H. Mayrhofer (Mayrhofer

etal. 1993) shares several similarities with/?, substellulata

(e.g. K+ yellow thallus reaction, lecideine apothecia, pres-

ence of a blue-green, N+ red pigment in the apothecia,

Pachysporaria type ascospores) but is separated by chemi-

cal and morphological characters: the thallus is grey to

brown and contains zeorin, the apothecia are up to 1 mm

in diameter, and the ascospores exhibit a wider size range

(14-25 x 7-14 pm). In addition, the ecology and distri-

bution of both taxa is different, with R. beccariana var.

lavicola occurring on volcanic rocks near the coast or on

26

Bothalia 26,1 (1996)

low coastal mountains in Macaronesia and in the Medi-

terranean region.

Vouchers: Almborn 3762 (LD, GZU); Brusse 1536, 1634 (PRE).

13. Rinodina subtristis (Nyl. in Cromb.) H.Mayr-

hofer in Journal of the Hattori Botanical Laboratory 55:

464 (1984a). Type: Western Cape, Cape of Good Hope,

Table Mtn, September 1874, A.E. Eaton s.n., Venus Transit

Expedition, (BM, lecto.l, associated with Rinodina oxy-

data), designated by Mayrhofer (1984a: 464); ( H-NYL

9329, iso.), not seen.

Lecidea subtristis Nyl. in Cromb.: 21 (1876a). Buellia subtristis (Nyl.

in Cromb.) Zahlbr.: 421 (1931).

Thallus thin, crustose, rimose-areolate to effuse, brown

or ochraceous; prothallus occasionally present, brown.

Chemistry: no lichen substances detectable by tic.

Apothecia to 0.7 mm diam., lecideine or lecanorine,

adnate to sessile, occasionally cryptolecanorine, disc dark

reddish brown to black, plane to convex, proper margin

black. Proper exciple dark brown to black, in addition

often with a blue-green, N+ red pigment; dead algal cells

may be present. Epihymenium 10-20 /<m tall, reddish

brown, brown, occasionally (dirty) blue-green and then

partly N+ red. Hymenium ± 80 //m tall. Hypothecium to

± 150 pm deep, hyaline. Paraphyses l-3(-4) pm, apices

(3-)4-6 pm wide. Asci corresponding to Lecanora type,

axial body occasionally indistinct, asci often with 8 but

also with less spores. Ascospores (Figure 8D) of Pachy-

sporaria type, occasionally grading into Milvina type, sep-

tum in young spores inserted before internal wall

thickenings become distinct, torus small but well distinct,

spores smooth to finely scabrid, without septal swellings

in KOH, (15-)16-23 x 8-16 pm. Spermogonia immersed

in thallus, raised, ostiolar region red-brown to blackish.

Spermatia 3-4 x 1.0-1 .5 pm.

Rinodina subtristis was originally known only from the

Cape of Good Hope (Crombie 1876a, b; as Lecidea sub-

tristis) but can now be reported also from other regions

of the Republic of South Africa and from Angola (Figure

9). The species grows on hard quartzitic rocks (including

quartzitic sandstone).

In addition to R. subtristis, there are several other Ri-

nodina species in the study area which are characterized

by the lack of secondary lichen compounds detectable by

thin-layer chromatography and by the presence of Pachy-

sporaria type ascospores with a torus, viz. R. longisperma,

R. microlepidea , R. scabridula, and R. confragosula. The

problematic separation of R. subtristis from R. mi-

crolepidea is discussed under the latter species in the pre-

sent work. Distinctive characters of R. longisperma

include the small cryptolecanorine apothecia, the small

ascospores in which the spore lumina may have small

protrusions, and the comparatively long spermatia. Rino-

dina scabridula is mainly distinguished by the formation

of blastidiate thalli. The separation of R. subtristis from

R. confragosula is based on a set of characters: grey thalli

are often found in R. confragosula but never in R. sub-

tristis; the apothecia of R. confragosula are usually purely

black and cryptolecanorine but dark reddish brown to

black and lecideine or lecanorine in R. subtristis; the as-

FIGURE 9. — Known distribution of Rinodina substellulata in Africa, •;

and R. subtristis, ■.

cospores of R. confragosula are very variable in the for-

mation of internal wall thickenings (Pachysporaria, Milv-

ina, Physcia, or intermediate types, or spore lumina

irregularly biconical in shape) and in length (14—32 ^m),

the spores of R. subtristis are usually of Pachysporaria

type and 15-23 pm long.

Three Rinodina species which were described from

West African islands share close similarities with R. sub-

tristis, viz. R. praefinita (Nyl.) Zahlbr. (Nylander 1887, as

Lecanora praefinita) and R. subanceps (Nyl.) Zahlbr.

(Nylander 1887, as Lecanora subanceps) from Sao Tome

& Principe, and R. newtonii H.Mayrhofer (Mayrhofer

1984a) from Annobon ( = Pagalu). All these species and

R. subtristis are known only from single or at best a few

specimens, and thus statements whether these taxa are

really well separated, cannot be made without some

hesitation. Linder this precondition, R. praefinita is dis-

tinguished from R. subtristis by smaller ascospores (±

13-17 x 7-10 ^m), and R. subanceps differs by the

thick, more or less squamulose thallus. A well-devel-

oped thallus and ascospores with an indistinct torus are

characteristic for R. newtonii. A detailed study of the

relationships between these taxa is beyond the scope of

the present work and was not carried out. Mayrhofer

& Leuckert (1985) recorded the presence of zeorin in

a specimen of R. subanceps from Annobon ( H-NYL

28964), but lichen compounds could not be detected by

recent tic-analyses in the holotype from the island of

Cabras (Sao Tome & Principe, H-NYL 28965) and in

another specimen from Annobon ( H-NYL 28514).

Bothalia 26,1 (1996)

27

One of the specimens now regarded as belonging to

R. subtristis was treated under R. huillensis by Mayrhofer

(1984a): Angola, Humpata Plateau, 6-2-1960, Degelius

(GZU).

Vouchers: Almborn 1720 (LD); Brusse 2719 (PRE); Degelius s.n.

(GZU); Schaefer CH 1935 (PRE).

14. Rinodina teichophiloides (Stizenb.) Zahlbr. in

Catalogus lichenum universalis 7: 557 (1931). Type: West-

ern Cape, supra saxa quartzosa schistosa ad Muizenberg

in Promontorio Bonae Spei, MacOwan 126 (ZT, holo.!).

Lecanora teichophiloides Stizenb.: 212 (1890).

Thallus thinly crustose to somewhat squamulose, oc-

casionally evanescent, effuse to rimose-areolate, light

greenish grey, grey-brown, dark brown, ochraceous, oc-

casionally associated with cyanobacteria; prothallus ab-

sent or present, brown. Chemistry: no lichen substances

detectable by tic.

Apothecia to 0.8 mm diam., often with a lecideine ap-

pearance or lecanorine, immersed to adnate, (dead) algal

cells sometimes present in the proper margin, disc black

or dark brown, plane to convex. Epihymenium 10-30 /tm

tall, dark brown. Hymenium 90-130 pm tall. Hypothecium

to 150 pm deep, hyaline, light yellow to yellowish brown.

Paraphyses 1-2 pm, apices to 3-5 pm wide; enlarged, to

± 6 pm wide ‘oil cells’ occasionally present in paraphyses.

Asci corresponding to Lecanora type, 2- to 8-spored. As-

cospores (Figure 8E) at first with internal wall thickenings

of Mischoblastia type, then corresponding to Pachyspo-

raria type, septum in young spores inserted before internal

wall thickenings become distinct, torus absent but some-

times simulated by intense brown pigmentation in the sep-

tal region, spores finely scabrid, often with septal

swellings in KOH, (16-)20-32 x 10-18 pm. Spermogo-

nia immersed in thallus, raised, ostiolar region dark red-

dish brown to blackish. Spermatia ± 4-5 x 1. 0-1.5 pm.

Rinodina teichophiloides is a characteristic species of

maritime localities in the Cape Province (Figure 10). It

grows on hard siliceous rocks such as quartzitic sandstone,

on inclined, vertical and overhanging rock faces. Occa-

FIGURE 10. — Known distribution of Rinodina teichophiloides , •; and

Rinodina sp. 1, ■.

sionally cyanobacteria were found growing at the margin

of well-developed thallus areolae.

Rinodina teichophiloides was discussed in detail re-

cently by Matzer & Mayrhofer (1994). Since then, addi-

tional material has become available and some annotations

can be given on the morphology of the species. Apothecia

with a lecideine appearance in surface view may contain

(dead) algal cells in the proper exciple. Paraphyses with

‘oil cells’ (oil paraphyses, compare Poelt & Pelleter 1984)

were observed in some specimens. The ascospores lack a

typical torus but intense pigmentation in the septal region

may simulate the presence of a torus. Rinodina teichophi-

loides can best be recognized by the often rather thin thal-

lus, by apothecia often appearing to be lecideine, the large

ascospores with internal wall thickenings of Mischoblastia

or Pachysporaria type, and by the often coloured hy-

pothecium which lacks a distinct colour change in potas-

sium hydroxide. It is closely related to R. reagens, a

species which also occurs in the Cape Province. The latter

can easily be distinguished by the presence of a K+ re-

active pigment in the hypothecium and — not always — in

the epihymenium and in the spermogonia. Both the spe-

cies also differ in substrate ecology (see discussion of R.

reagens in the present study).

Vouchers: Almborn 926, 1235, 1262, 1280, 1320 (LD), 11498 (GZU,

LD); J. Hafellner & A. Hafellner, 30656, 30658, 30659, 30661 (Hafell-

ner), 30662 (GZU); Karnefelt 8647—26 (LD); Sipman 20287 (B); Triebel

& Rambold 7708, 8156, 8186, 8190, 8204, 8227, 8230 (M), 8156 dupl.

(GZU), 8419 dupl. (GZU, M).

15. Rinodina sp. 1

Thallus crustose, effuse to areolate, continuous or com-

posed of ± discrete areolae, whitish, pale greyish, pale

brownish, areolae partly convex; prothallus absent. Chem-

istry: no lichen substances detectable by tic.

Apothecia to 0.45 mm diam., lecanorine, sessile, in ad-

dition to the thalline margin a proper margin is occasion-

ally developed, disc brown to blackish, plane or concave,

proper margin brown. Epihymenium 10-15 /an tall,

brown. Hymenium 60-70 pm tall. Hypothecium to ± 70

pm deep, hyaline. Paraphyses 1-3 pm, apices 2-5 pm

wide. Asci corresponding to Lecanora type, often with 8

but also with less than 8 spores. Ascospores (Figure 8F)

of Physcia type, rarely grading into Milvina type, septum

in young spores inserted before internal wall thickenings

become distinct, torus small but well distinct, spores

smooth to finely scabrid, without septal swellings in KOH,

11-17 x 6-9 pm. Spermogonia not found.

The species is known only from a single locality in

the Eastern Cape (Figure 10).

The above description is based on the single collection

cited below. In Mayrhofer (1984a) this specimen was in-

cluded in Rinodina interpolata (Stirt.) Sheard, but we have

some doubts whether it really belongs to that species. Ri-

nodina interpolata usually occurs in Europe on more or

less vertical or overhanging surfaces of cliffs mainly in

coastal sites but also in inland localities (e.g. Fox & Purvis

1992; Mayrhofer 1984a; Mayrhofer & Poelt 1979; San-

tesson 1993; Sheard 1973); it was also recorded from the

Antarctic region (see Lindsay 1973). A definite statement

28

Bothalia 26,1 (1996)

on the occurrence of R. interpolate! in southern Africa

only seems possible after extensive field studies and col-

lecting of more material. Only a few significant differ-

ences between Rinodina sp. 1 and R. interpolata can be

observed: the ascospores in R. interpolata are also of Phy-

scia type but, particularly when older, often grade into

Physconia type; in several specimens of R. interpolata ,

zeorin could be detected by tic (hitherto unpublished data;

Leuckert & Mayrhofer 1984). For a description of R. in-

terpolata see Fox & Purvis (1992), Mayrhofer & Poelt

(1979), and Sheard (1973).

Voucher: Almborn 10610 (LD, associated with Lichenodiplis licheni-

cola ).

EXCLUDED SPECIES

This section includes saxicolous species from southern

Africa which were referred to the genus Rinodina by vari-

ous authors, and Rinodina species which were erroneously

recorded from the study area.

1 . Buellia distrata (Nyl.) Zahlbr. in Catalogus

lichenum universalis 7: 357 (1931). Type: Western Cape,

Cape of Good Hope, Table Mtn, September 1874, E.A.

Eaton s.n., Venus Transit Expedition (BM, lecto.!) desig-

nated by Mayrhofer (1984a: 405), (BM, iso.!), ( H-NYL

9309, iso.), not seen.

Lecidea distrata Nyl. in Cromb.: 179 (1876b). Rinodina distrata

(Nyl.) C.W. Dodge: 167 (1971).

Thallus thin, crustose, areolate, light yellowish;

prothallus blackish, between the areolae well developed.

Chemistry: thallus K+ yellow, C-, P-; medulla K-, C+

orange, P-.

Apothecia to 0.3 mm diam., cryptolecanorine, disc

black, plane. Epihymenium 10-15 pm tall, brown, oli-

vaceous to dirty greenish, N+ red. Hymenium 90-110 pm

tall. Hypothecium to 100 pm deep, hyaline. Paraphyses

1-4 pm, apices 3-5 pm wide. Asci corresponding to

Bacidia type, usually with 8, rarely with less spores. As-

cospores (Figure 1A) of Beltraminia type, torus absent,

spores scabrid, ± 17-22 x 9-12 pm. Spermogonia not

observed.

Buellia distrata is known only from the type specimens

collected on Table Mountain on quartzitic rocks.

This species has ascospores which lack internal wall

thickenings, therefore it cannot be placed within Rinodina

as was done formerly (e.g. Dodge 1971; Mayrhofer

1984a). Alternative placements could be in either Aman-

dinea Choisy ex Scheid. & H. Mayrhofer or Buellia De

Not., two genera which are separated mainly by their sper-

matia: long, filiform, and curved in Amandinea and short,

bacilliform, and straight in Buellia (e.g. Matzer &

Mayrhofer 1993; Matzer et al. 1994b; Scheidegger 1993).

As spermogonia are not present in the material of Buellia

distrata now available, the best solution seems to retain

the existing name of this taxon.

2. Buellia permodica (Stizenb.) Zahlbr. in Cata-

logus lichenum universalis 7: 387 (1931). Type: Western

Cape, supra saxa arenaria prope Muizenberg in Promon-

torio Bonae Spei, MacOwan s.n. (?ZT, holo.); not seen,

citation after Stizenberger (1891: 168).

Lecidea permodica Stizenb.: 168 (1891).

This species was not validly transferred into the genus

Rinodina by Dodge (1971) as reference to the basionym

was omitted (ICBN, Art. 33.2.). Judging by the small

spores mentioned in the protologue the species probably

does not belong to Rinodina.

3. Rinodina argentiniana MiillArg. in Flora 72:

511 (1889).

See discussion of Rinodina confragosula in the present

study.

4. Rinodina atroalbida (Nyl.) C.W.Dodge in Bei-

hefte zur Nova Hedwigia 38: 165 (1971).

Lecanora sopliodes var. atroalbida Nyl.: 7 (1869). Rinodina sophodes

var. atroalbida (Nyl.) Zahlbr.: 552 (1931).

This taxon was described from Port Natal (Durban) in

the Republic of South Africa. According to Mayrhofer

(1984a) the type cannot be located.

5. Rinodina bicolor Zahlbr. in Annales de Cryp-

togamie Exotique 5: 272 (1932). Type: ?Swaziland,

Mbabana, ad lapides siliceos, V Lebzelter s.n. (W, holo.);

not seen, citation after Zahlbruckner (1932: 273).

According to Mayrhofer (1984a) this species belongs

to the genus Buellia.

6. Rinodina conspersa MiillArg. in Flora 72: 511

(1889). Type: Paraguay, Cerro Lambare prope Asuncion,

1882, Lorentz s.n. (G, lecto., iso.); not seen, citation after

Mayrhofer (1984a: 484, 485).

Lecanora conspersa (Miill.Arg.) Stizenb.: 236 (1895).

The records of this species from the Zambezi River

(Stizenberger 1895; Doidge 1950) are based on material

of Rinodina oxydata (Mayrhofer 1984a).

7. Rinodina deminutula (Stizenb.) Zahlbr. in Cata-

logus lichenum universalis 7: 508 (1931). Type: Western

Cape, supra saxa arenaria in Monte Leonis prope Cape

Town, MacOwan s.n. (ZT, holo.); not seen, citation after

Stizenberger (1890: 211).

Lecanora deminutula Stizenb.: 210 (1890).

Mayrhofer (1984a) pointed out that the scant type is in

a poor condition and a treatment of the species is not pos-

sible.

8. Rinodina microphthalma AMassa/. in Memorie

del Reale Istituto Veneto di Scienze, Lettere ed Arti 10:

61 (1861).

Bothalia 26,1 (1996)

29

This species was described from the Cape Province. Ac-

cording to the protologue it probably belongs to Buellia. The

name Rinodina microphthalma was treated as a synonym of

Lecanora microps Stizenb. by Stizenberger (1890), and of

Buellia verruculosa Mudd by Zahlbruckner (1931).

9. Rinodina procellarum (A.Massal.) H.Mayrhofer

in Beihefte zur Nova Hedwigia 79: 522 (1984b). Type:

Western Cape, Caput Bonae Spei, Wawra s.n. (VER, holo.);

not seen, citation after Mayrhofer (1984b) and Sheard (1992).

Buellia procellarum A.Massal.: 64 (1861). Hafellia procellarum

(A.Massal.) H.Mayrhofer & Sheard in Sheard: 87 (1992).

Modern taxonomic concepts clearly suggest that this

species should be included in the genus Hafellia Kalb,

H.Mayrhofer & Scheid. (Sheard 1992).

ACKNOWLEDGEMENTS

We would like to thank the directors and curators of

the herbaria who have sent type and other material on

loan: B, BM, G, GZU, H, L, LD, M, PRE, TRH, TUR,

W, ZT, and Dr J. Hafellner (Graz) for the provision of

specimens from his private herbarium. Many thanks are

also due to Dr D. Triebel and Dr G. Rambold (both Mu-

nich) who have kindly selected and provided relevant ma-

terial for the present study. We are grateful to Prof. Dr C.

Leuckert (Berlin) for his help with the identification of

5-O-methylhiascic acid in Rinodina huillensis, to Prof. Dr

J. Poelt (Graz) and Prof. Dr J.W. Sheard (Saskatoon) for

critical reading of the manuscript, to Dr M. Giralt (Bar-

celona) for valuable comments and, especially, to Dr G.

Kantvilas (Hobart) for correcting the English text. The

‘Fonds zur Forderung wissenschaftlicher Forschung (Pro-

jekte P8500-BIO and P10514-BIO)’ is acknowledged for

financial support.

REFERENCES

BACHMANN, E. 1890. Ueber nichtkrystallisierte Flechtenfarbstoffe, ein

Beitrag zur Chemie und Anatomie der Flechten. Jahrbiicher fur

wissenschaftliche Botanik 21: 1-61.

BAGLIETTO, F. 1861. Nuove specie dei Licheni. Commentario della

Societa Crittogamologica Italiana 1: 17-24.

BUDEL, B. 1987. Zur Biologie und Systematik der Flechtengattungen

Heppia und Peltula im siidlichen Afrika. Bibliotheca Liche-

nologica 23: 1-105.

CRESPO, A. & AGUIRRE, B. 1984. Rinodina euskadiensis spec, nova,

un nuevo liquen epifito. Munibe 36: 135-137.

CROMBIE, J.M. 1876a. New lichens from the Cape of Good Hope.

Journal of Botany, British and Foreign 14: 18-21.

CROMBIE, J.M. 1876b. Lichenes Capenses. An enumeration of the

lichens collected at the Cape of Good Hope by the Rev. A.E.

Eaton during the Venus-Transit Expedition in 1874. Journal of the

Linnean Society, Botany 15: 165-180.

DODGE, C.W. 1971. Some lichens of tropical Africa. V. Lecanoraceae to

Physciaceae. Beihefte zur Nova Hedwigia 38: 1-225.

DOIDGE, E.M. 1950. The South African fungi and lichens to the end of

1945. Bothalia 5: 1-1094.

EGAN, R.S. 1987. A fifth checklist of the lichen-forming, lichenicolous

and allied fungi of the continental United States and Canada.

Bryologist 90: 77-173.

FOX, B.W. & PURVIS, O.W. 1992. Rinodina (Ach.) Gray (1821). In

O.W. Purvis, B.J. Coppins, D.L. Hawksworth, P.W. James & D.M.

Moore, The lichen flora of Great Britain and Ireland: 543-552.

Natural History Museum Publications and The British Lichen

Society, London.

GIRALT, M. 1994. Key to the corticolous and lignicolous species of the

genus Rinodina present in the Iberian Peninsula and Balearic

Islands. In Cl. Roux, Homage scientifique a Georges Clauzade

80e anniversaire. Bulletin de la Societe Linneenne de Provence

45: 317-326.

GIRALT, M. & BARBERO, M. 1995. The saxicolous species of the

genus Rinodina in the Iberian Peninsula containing atranorin,

pannarin or gyrophoric acid. Mycotaxon 56: 45-80.

GIRALT, M. & MATZER, M. 1994. The corticolous species of the genus

Rinodina with biatorine or lecideine apothecia in southern Europe

and Macaronesia. Lichenologist 26: 319-332.

GIRALT, M. & MAYRHOFER, H. 1994a. Four corticolous species of the

genus Rinodina (lichenized Ascomycetes, Physciaceae) contain-

ing atranorin in southern Europe and adjacent regions. Nova

Hedwigia 59: 129-142.

GIRALT, M. & MAYRHOFER, H. 1994b. Four corticolous species of the

genus Rinodina (lichenized Ascomycetes, Physciaceae) with

polyspored asci. Herzogia 10: 29-37.

GIRALT, M. & MAYRHOFER, H. 1995. Some corticolous and lignicol-

ous species of the genus Rinodina (lichenized Ascomycetes, Phys-

ciaceae) lacking secondary lichen compounds and vegetative

propagules in southern Europe and adjacent regions. In J.-G.

Knoph, K. Schriifer & H.J.M. Sipman, Studies in lichenology

with emphasis on chemotaxonomy, geography and phytochemis-

try. Festschrift Christian Leuckert. Bibliotheca Lichenologica 57:

127-160.

GIRALT, M„ MAYRHOFER. H. & SHEARD, J.W. 1995. The corticol-

ous and lignicolous sorediate, blastidiate and isidiate species of

the genus Rinodina in southern Europe. Lichenologist 27: 3-24.

GIRALT, M., NIMIS, PL. & POELT, J. 1993a. Studien iiber einige Arten

der Flechtengattung Xanthoria mit isidiiformen vegetativen Dia-

sporen. Journal of the Hattori Botanical Laboratory 74: 271-285.

GIRALT, M„ OBERMAYER, W. & MAYRHOFER, H. 1993b. Rinodina

poeltiana spec, nova (lichenized Ascomycetes, Physciaceae), a

new corticolous blastidiate species from Austria. Herzogia 9:

709-714.

HAFELLNER, J. 1984. Studien in Richtung einer natiirlicheren Glied-

erung der Sammelfamilien Lecanoraceae und Lecideaceae. In H.

Hertel & F. Oberwinkler, Festschrift J. Poelt. Beihefte zur Nova

Hedwigia 79: 241-371.

HAFELLNER, J., MAYRHOFER, H. & POELT, J. 1979. Die Gattungen

der Flechtenfamilie Physciaceae. Herzogia 5: 39-79.

HAWKSWORTH, D.L., SUTTON, B.C. & AINSWORTH, G.C. 1983.

Ainsworth & Bisby's dictionary of the fungi (including the li-

chens), 7th edn. Commonwealth Agricultural Bureaux, Kew, Sur-

rey.

HECKLAU, C., LEUCKERT, C. & MAYRHOFER, H. 1981. Beitrage

zur Chemie der Flechtengattung Rinodina (Ach.) Gray I. Her-

zogia 5: 489-498.

HONEGGER, R. 1978. The ascus apex in lichenized fungi I. The Le-

canora-, Peltigera- and Teloschistes-types. Lichenologist 10:

47-67.

HUE, A.M. 1891. Lichenes exoticos a Professore W. Nylander descriptos

vel recognitos et in herbario musei Parisiensis pro maxime parte

asservatos in ordine systematico disposuit. Nouvelles Archives de

Museum d’Histoire Naturelle, Ser. 3, 3: 33-192.

KOERBER, G.W. 1855. Systema lichenum Germaniae. Breslau.

LEUCKERT, C. & MAYRHOFER, H. 1984. Beitrage zur Chemie der

Flechtengattung Rinodina (Ach.) Gray II. Herzogia 6: 373-385.

LINDSAY, D.C. 1973. South Georgian microlichens: I. The genera Buel-

lia and Rinodina. Bulletin, British Antarctic Survey 37: 81-89.

MASSALONGO, A. 1854. Geneacaena lichenum. Verona.

MASSALONGO, A. 1861. Lichenes Capenses quos collegit in itinere

1857-1858 Dr. Wawra. Memorie del Reale Istituto Veneto di

Scienze, Lettere edArti 10: 33-90.

MATZER, M. & MAYRHOFER, H. 1993. Generic delimitation in Phys-

ciaceae (lichenized Ascomycetes) with crustose thallus organisa-

tion including taxa with lecanorine apothecia. In A. Bellemere,

M.-A. Letrouit & J. Mouchacca, First International Workshop on

Ascomycete systematics. Programme, lectures & posters ab-

stracts and list of participants. Paris.

MATZER, M. & MAYRHOFER, H. 1994. The saxicolous Rinodina

teichophila and three closely related species from the southern

hemisphere (Physciaceae, lichenized Ascomycetes). In S. Sten-

30

Bothalia 26,1 (1996)

roos. Focus on lichen taxonomy and biogeography: a Festschrift

in honour of Teuvo Ahti. Acta Botanica Fennica 150: 109-120.

MATZER, M., MAYRHOFER, H„ SATTLER, J. & CLERC, Ph. 1994a.

Rinodina canariensis (lichenized Ascomycetes, Physciaceae), a

new species parasitic on crustose lichens in Macaronesia and the

Mediterranean region. Nordic Journal of Botany 14: 105-111.

MATZER, M„ MAYRHOFER, H. & SCHEIDEGGER, Ch. 1994b.

Notes on Amandinea petermannii comb. nov. (Physciaceae) from

Antarctica. Lichenologist 26: 39 — 46.

MAYRHOFER, H. 1982. Ascosporen und Evolution der Flechtenfamilie

Physciaceae. Journal of the Hattori Botanical Laboratory 52:

313-321.

MAYRHOFER, H. 1983. The saxicolous species of Rinodina in New

Zealand. Lichenologist 15: 267-282.

MAYRHOFER, H. 1984a. Die saxicolen Arten der Flechtengattungen

Rinodina und Rinodinella in der Alten Welt. Journal of the Hat-

tori Botanical Laboratory 55: 327-493.

MAYRHOFER, H. 1984b. The saxicolous species of Dimelaena , Rino-

dina and Rinodinella in Australia. In H. Hertel & F. Oberwinkler,

Festschrift J. Poelt. Beihefte zur Nova Hedwigia 79: 511-536.

MAYRHOFER, H. 1987. Rinodina. In PL. Nimis & J. Poelt, The lichens

and lichenicolous fungi of Sardinia (Italy). An annotated list.

Studia Geobotanica 7, suppl. 1: 208-215.

MAYRHOFER, H. & LEUCKERT, C. 1985. Beitrage zur Chemie der

Flechtengattung Rinodina (Ach.) Gray III. Herzogia 7: 117-129.

MAYRHOFER, H„ MATZER, M„ SATTLER, I. & EGEA, J.M. 1993. A

revision of the Atlantic — Mediterranean Rinodina beccariana and

related taxa (lichenized Ascomycetes, Physciaceae). Nova Hed-

wigia 57: 281-304.

MAYRHOFER, H. & POELT, J. 1979. Die saxicolen Arten der Flechten-

gattung Rinodina in Europa. Bibliotheca Lichenologica 12: 1-186.

MAYRHOFER, H„ SCHEIDEGGER, CH. & SHEARD, J.W. 1992. On

the taxonomy of five saxicolous species of the genus Rinodina

(lichenized Ascomycetes \Nordic Journal of Botany 12: 451-459.

MCCARTHY, P.M. 1991. Checklist of Australian lichens, 4th edn. Na-

tional Herbarium of Victoria, Melbourne.

MULLER ARGOVIENSIS (MULLER), J. 1882. Diagnoses lichenum

Socotrensium novorum a particibus expeditionum Prof. Bailey

Balfour et Dr. Schweinfurth lectorum. Proceedings of the Royal

Society of Edinburgh 11: 457-472.

MULLER ARGOVIENSIS (MULLER), J. 1887. Enumeration de quelques

lichens de Noumea. Revue Mycologique ( Toulouse ) 9: 77-82.

MULLER ARGOVIENSIS (MULLER), J. 1888. Lichenologische Beitrage

XXIX. Flora 71: 195-208.

MULLER ARGOVIENSIS (MULLER), J. 1889. Lichenes Argentinienses a

cl. Dr. Lorentz in Argentinia australis. Patagonica et prope Concep-

tionem lecti, additis nonnullis in Paraguay prope Assumpcion ab

eodem lectis. Flora 72: 509-512.

NIMIS, P.L. 1993. The lichens of Italy. An annotated catalogue. Museo

Regionale di Scienze Naturali, Torino, Monografia 12: 1-897.

NIMIS, P.L., MONTE, M. & TRETIACH, M. 1987. Flora e vegetazione

lichenica de aree archeologiche del Lazio. Studia Geobotanica 7:

3-161.

NYLANDER, W. 1869. Note sur les lichens de Port-Natal. Bulletin de la

Societe Linneenne de Normandie, Ser. 2, 3: 4-15.

NYLANDER, W. 1887. Lichenes nonnulli ex insula Principis. Boletim da

Sociedade Broteriana 5: 221-224.

NYLANDER, W. 1889. Lichenes insularum Guineensium (Sao Thome, do

Principe, das Cabras). Paris.

NYLANDER, W. 1896. Enumeration des lichens de l Lie Annobon. Paris.

POELT, J. 1980. Physcia opuntiella spec. nov. und die Lebensform der

sprossenden Flechten. Flora 169: 23-31.

POELT, J. & MAYRHOFER, H. 1979. Studien iiber Ascosporen-Typen der

Flechtengattung Rinodina. Beihefte zur Sydowia 8: 312-331.

POELT, J. & MAYRHOFER, H. 1988. Uber Cyanotrophie bei Flechten.

Plant Systematics and Evolution 158: 265-281.

POELT, J. & PELLETER, U. 1984. Zwergstrauchige Arten der Flechtengat-

tung Caloplaca. Plant Systematics and Evolution 148: 51-88.

RAMBOLD, G„ MAYRHOFER, H. & MATZER, M. 1994. On the ascus

types in the Physciaceae (Lecanorales). Plant Systematics and Evo-

lution 192: 31-40.

ROPIN, K. & MAYRHOFER, H. 1995. Uber einige corticole Arten der

Gattung Rinodina (Physciaceae) mit grauem Epihymenium. In E.

Farkas, R. Lucking & V. Wirth, Contributions to lichenology. In

honour of Antonin Vezda. Bibliotheca Lichenologica 58: 361-382.

SANTESSON, R. 1993. The lichens and lichenicolous fungi of Sweden and

Norway. Lund.

SCHEIDEGGER, Ch. 1993. A revision of European saxicolous species of

the genus Buellia De Not. and formerly included genera. Lichenolo-

gist 25: 315-364.

SCHREINER, E. & HAFELLNER, J. 1992. Sorediose, corticole Krusten-

flechten im Ostalpenraum. I. Die Flechtenstoffe und die gesicherte

Verbreitung der besser bekannten Arten. Bibliotheca Lichenologica

45: 1-291.

SHEARD, J.W. 1967. A revision of the lichen genus Rinodina (Ach.) Gray

in the British Isles. Lichenologist 3: 328-367.

SHEARD, J.W. 1973. Rinodina interpolata (Stirt.) Sheard, a new combina-

tion in the British and Scandinavian lichen floras. Lichenologist 5:

461-463.

SHEARD, J.W. 1992. The lichenized Ascomycete genus Hafellia in North

America. Bryologist 95: 79-87.

STIZENBERGER, E. 1890. Lichenaea Africana. Bericht iiber die Thatigkeit

der St. Gallischen Naturwissenschaftlichen Gesellschaft 1888/1889:

105-249.

STIZENBERGER, E. 1891. Lichenaea Africana. Bericht iiber die Thatigkeit

der St. Gallischen Naturwissenschaftlichen Gesellschaft 1889/1890:

133-268.

STIZENBERGER, E. 1893. Supplementa ad lichenaeam Africanam. I.

Denuo addenda et corrigenda. Bericht iiber die Thatigkeit der St.

Gallischen Naturwissenschaftlichen Gesellschaft 1891/1892: 86-96.

STIZENBERGER, E. 1895. Supplementa ad lichenaeam Africanam. II.

Addenda et corrigenda ex annis 1893-1894. Bericht iiber die

Thatigkeit der St. Gallischen Naturwissenschaftlichen Gesell-

schaft 1893/1894: 215-264.

VAINIO, E.A. 1901. Lichenes. In Catalogue of African plants collected

by F. Welwitsch 2, part 2: 396-463. London.

VAINIO, E.A. 1921. Lichenes ab A. Yasuda in Japonia collecti II. Botani-

cal Magazine (Tokyo) 35: 45-79.

VOBIS, G. 1980. Bau und Entwicklung der Flechten-Pycnidien und ihrer

Conidien. Bibliotheca Lichenologica 14: 1-141.

ZAHLBRUCKNER, A. 1931. Catalogus lichenum universalis 7. Leipzig.

ZAHLBRUCKNER, A. 1932. Lichenes in Africa lecti. Annales de Cryp-

togamie Exotique 5: 198-275.

Bothalia 26,1: 31-35 (1996)

FSA contributions 4: Agavaceae

G. F. SMITH* and M. MOSSMER*

Family Agavaceae Endl. Enchiridion botanicum:

105 (1841) nom. cons. Cronquist: 1217 (1981).

Robust, monocarpic, usually rosulate perennials arising

from short rhizome or short erect caudex. Stem commonly

with monocotyledonous type secondary growth. Leaves

usually crowded in basal rosette, leathery to succulent,

amplexicaul, persisting for many years; each vascular bun-

dle with well-developed fibrous cap at phloem pole. In-

florescence apical, tall, fast-growing, terminating in a

panicle, often massive. Flowers bisexual, actinomorphic

or somewhat zygomorphic, tubular, pedicellate, trimerous

throughout. Perianth petaloid, 3 + 3, often fleshy, united

below to form a tube. Stamens 3 + 3; anthers mostly dor-

sifixed, introrse, versatile, opening by longitudinal slits,

linear to oblong. Ovary trilocular, inferior, with septal nec-

taries, placentation axile, ovules in 2 vertical rows in each

locule; style terminal, stigma 3-lobed. Fruit a loculicidal

capsule. Seeds numerous, flattened, centrally embedded

in copious, very hard endosperm. Chromosome numbers :

2n = 60, 120, 149, 150, 180 (Bolkhovskikh et al. 1969).

When the plan for the Flora of southern Africa was

first published in 1963, it did not include the Agavaceae

as currently circumscribed. Two representatives of the ge-

nus Agave L., A. americana L. and A. sisalana Perrine,

are naturalized in South Africa, and the family as defined

here warrants inclusion in the FSA series.

The Agavaceae is a family of about 12 genera and 400

species, occurring mostly in the New World tropics and

subtropics, and India to Australia (Cordyline Comm, ex

R.Br.). Numerous genera are cultivated in southern Africa

by collectors and in amenity horticulture, but only two

species oi Agave are naturalized. The boundaries and con-

tent of this heterogeneous family is still debated; the cir-

cumscription upheld here follows Brummitt (1992), which

approximates the interpretation of Cronquist (1981).

1219000 AGAVE

Agave L., Species plantarum 1: 323 (1753); Tre-

lease: 231 (1914); Berger: 21 (1915); Gentry: 41 (1972);

Gentry: 10 (1978); Gentry: 61 (1982); Pedley & Forster:

72, 74 (1986); Couper & Cullen: 278 (1988). Type: A.

americana L.

Description as for family.

Agave is a New World genus of ± 275 species. One

introduced species, A. americana L., is naturalised around

sites of habitation, mainly in the more arid, karroid regions

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

MS. Received: 1995-04-24.

of southern Africa. A second species, A. sisalana , has

been used as a barrier plant during South Africa’s border

war and for fibre production in plantations in vari-

ous— mainly summer rainfall — savanna regions of the

subcontinent. This species has also subsequently become

naturalized in South Africa. Both species are included in

catalogues of problem plants in southern Africa (Wells et

al. 1986; Henderson 1995). Distribution maps for the spe-

cies (Figures 1 & 2) are based on those included in Hen-

derson (1995). Additional attempts to obtain a clearer

picture of naturalized distribution, uses and history of

Agave in southern Africa are being made (Smith 1995a,

b; Smith & Crouch 1996).

FIGURE 1. — Distribution map of Agave americana in southern Africa.

32

Bothalia 26,1 (1996)

The name Agave was taken from the Greek, aganos,

which means noble or admirable.

Key to species in southern Africa

Leaves lanceolate, reflexed, usually glaucous green, up to 2 m

long; marginal teeth present on mature leaves, straight or

recurved; panicle not bulbiferous; flowers 70-100 mm

long A. americcma

Leaves ensiform, rigid, usually light to dark green, up to 1.3 m

long; marginal teeth absent from mature leaves; panicle

bulbiferous after flowering; flowers 55-65 mm long . .

A. sisalana

1219000-00100 Agave americana L., Species plan-

tarum 1: 323 (1753); Berger: 151 (1915); Gentry: 278

(1982); Pedley & Forster: 74 (1986); Couper & Cullen:

280, 281 (1988). Type: ‘America calidiore’ specimen

443.1 Herb. Linnaeus [LINN, holo. non vidi; fide Gentry:

278 (1982)].

Agave complicata Trel. ex Ochot.: 100 (1913).

A. gracilispina Engelm. ex Trel.: 234 (1914).

A. melliflua Trel.: 234 (1914).

A. zonata Trel.: 234 (1914).

A. felina Trel.: 128 (1920). Type: Durango, Trelease s.n. (MO, holo.)

non vidi.

A. rasconensis Trel.: 122 (1920). Type: San Luis Potosi, Rascon, Tre-

lease 75 (MO, holo.) non vidi.

A. subzonata Trel.: 129 (1920). Type: Nuevo Leon, Monterrey, Tre-

lease s.n. (MO, holo.) non vidi.

Herbaceous, succulent multi-annual, freely suckering;

rosettes trunkless, 2^1 m in diameter, 1-2 m tall. Leaves

lanceolate to oblanceolate-spathulate, rigidly spreading to

reflexed, 1-2 x 0.3 m, light green to (more commonly)

glaucous grey-green; surfaces smooth; margins sinuous-

toothed, teeth straight or sometimes slightly recurved,

shiny blackish brown to grey, 5-10 mm long, 20-60 mm

apart; apical spine conical to subulate, 30-50 mm long,

shallowly grooved above for ± half its length, brown to

grey. Inflorescence paniculate, 4-9 m tall, with up to 35

umbellate side branches in upper half of peduncle. Flow-

ers light greenish yellow, 70-100 mm long. Perianth with

tepals subequal in length but of more or less similar shape,

spathulate, slender, outer segments cucullate at tip, 25-35

mm long. Stamens epipetalous; filaments subulate, 60-90

mm long; anthers yellow, 30-40 mm long. Ovary oblong-

ovoid, broadly trigonal, indistinctly 6-grooved, sessile,

30-45 mm long; style filiform, at length longer than fila-

ments; stigma capitate, apical, becoming well exserted,

usually considerably exceeding anthers after anthesis.

Capsule oblong, short-beaked, 40-60 x 20-30 mm. Seeds

6-9 x 5-7mm. Chromosome numbers : 2n = 60, 120, 180

(x = 30) (Granick 1944). Figure 3.

The species comprises one subspecies and seven va-

rieties (excluding the typical subspecies and variety). One

of these, Agave americana (subsp. americana) var. ameri-

cana, has become naturalised in southern Africa. The

widely cultivated A. americana var. picta which has vari-

able yellow or whitish leaf striations has not been as suc-

cessful in colonizing suitable habitats in southern Africa,

and only the typical variety is regarded as naturalized.

The form of A. americana which is naturalized in southern

Africa is morphologically and chorologically remarkably

uniform, as though from a single clone. The little variation

encountered during field observations occurs primarily in

the size of plants, depending on age, crowding, or on in-

dividual sites. A form with light glaucous green, stiff

leaves is sometimes encountered, particularly between

Montagu and Ouberg Pass in the southwestern Cape. A.

americana flowers in midsummer (late December, Janu-

ary and early February) in southern Africa.

In times of drought in some areas of the Great Karoo

the leaves of cultivated plants are cut and used as forage

for sheep. Honey produced from the nectar of A. ameri-

cana has an unpleasant taste.

A. americana occurs in Arizona and Texas in the south-

ern United States of America and in Baja California Sur,

Chihuahua, Coahuila, Durango, Hidalgo, Jalisco, Nuevo

Leon, Oaxaca, Queretaro, San Luis Potosi, Tamaulipas,

Vera Cruz and Zacatecas in Mexico (see Gentry 1982:

305, 306 for a more complete list of exsiccatae of A.

americana , including its varieties). A. americana is a com-

plex species that has been misunderstood by some authors

(Drummond 1910; Gentry 1982). The polymorphic nature

of A. americana is evidenced by the numerous infraspe-

cific segregates that have been described. It is generally

interpreted as a phylogenetically advanced species in the

section Americanae of the genus (Gentry 1982: 270).

Voucher: G. F. Smith 241 (PRE).

leones: Gentry: figs 2.8B, 12.6 & 12.7 (1982); Ullrich: figs 1-62

(1993).

Common names: English: agave, American agave,

American aloe, century plant. Afrikaans: Amerikaanse

aalwee, Amerikaanse aalwyn, blou-aalwee, gareboom,

garingboom, kaalgaarboom, makaalwyn. Sotho: lekhala.

1219000-01300 Agave sisalana Perrine in United

States of America 25th Congress, 2nd session, House of

Representatives Report No. 564 (Tropical Plants): 8, 9,

16, 47, 60, 86 (1838a); Perrine: 36, 105, 140 (1838b);

Trelease: 49 (1913); Berger: 230 (1915); Gentry: 628

(1982); Pedley & Forster: 75 (1986); Couper & Cullen:

282 (1988). Type: Ososocoautla, Chiapas, Mexico, 22

March 1957, H. S. Gentry 16434 [US, DES, neo. non

vidi; fide Gentry: 628 (1982)].

Herbaceous, succulent multi-annual, freely suckering;

rosettes with a trunk of up to 1 m, rosettes 2-3 m in

diameter, 1-2 m tall. Leaves narrowed towards base, nar-

rowly oblanceolate, rigidly spreading to slightly reflexed,

1-2 x 0.3 m, light green to (more commonly) dark grey-

green; surfaces smooth; mature leaves without marginal

teeth, young leaves with few minute teeth; apical spine

conical to subulate, 30-50 mm long, brown to grey. In-

florescence paniculate, 4—9 m tall, with up to 40 umbellate

side branches in upper half of peduncle; bulbils borne in

axils after flowering. Flowers yellow, 55-65 mm long.

Perianth with tepals equal in length, cucullate at tip,

17-18 mm long. Stamens epipetalous; filaments subulate,

60-90 mm long; anthers yellow, 30-40 mm long. Ovary

green, oblong-ovoid to globose, broadly trigonal, 6-

grooved, sessile, 30—45 mm long; style filiform, at length

Bothalia 26,1 (1996)

33

FIGURE 3. — Agave americana: A, habit, inflorescence 7m tall; B, bud, x 1; C, protandrous flower in male phase, x 1; D, developing fruit, x 1;

E, dry capsule, x 1; F, leaf margin showing recurved teeth, x 1. All drawings made from live material collected by G. F. Smith and E. J.

van Jaarsveld, deposited under G. F. Smith 241 (PRE). Artist: Jeanette Loedolff.

34

Bothalia 26,1 (1996)

FIGURE 4. — Agave sisalana: A, habil, inflorescence 7m tall; B, bud, x 1; C, protandrous flower in male phase, x 1; D, wilted flower, x 1; E, plantlet

formed on side branch of inflorescence (note small teeth on leaf margin), x 1; F, terminal portion of leaf (note entire leaf margin), x 1. All

drawings made from live material deposited under G. Condy & G. F. Smith 1 (PRE). Artist: Gillian Condy.

Bothalia 26,1 (1996)

35

longer than filaments; stigma capitate, minute, apical, be-

coming well exserted, usually considerably exceeding an-

thers after anthesis. Capsule and seeds generally lacking.

Chromosome numbers : 2n = 149, 150 (Granick 1944;

Banerjee & Sharma 1987).

A. sisalana is a sexually sterile clone, probably of hy-

brid origin. This is demonstrated by its inability to produce

seed and its pentaploid chromosome complement (Gra-

nick 1944; Gentry 1982). Although Gentry (1982: 628,

630) argues that A. sisalana might have been introduced

to various parts of the world from the Mexican state of

Chiapas (the small towns from Cintalapa to Chiapa), its

origin remains uncertain. Gentry (1982: 634) lists a

number of A. sisalana exsiccatae, many of them having

been collected from areas to which the ‘species’ has been

introduced. Agave fibre industries — based primarily on

this species — were developed in the Philippines and In-

donesia in the previous century, and more recently in East

Africa (Doughty 1937, 1938; Gentry 1982). Figure 4.

Voucher: G. Condy & G. F. Smith 1 (PRE).

leones: Doughty: fig. 2 (1937); Gentry: fig. 22.9 (1982).

Common names: English: hemp plant, sisal, sisal hemp.

Afrikaans: garingboom.

ACKNOWLEDGEMENTS

We thank Judge Selikowitz for permission to collect flow-

ering material of Agave americana for description and illus-

tration from his garden in Edinburgh Drive, Claremont, Cape

Town. We are indebted to Dr Peter Goldblatt, Missouri Bo-

tanical Garden, Ms Marinda Koekemoer, former South Af-

rican Botanical Liaison Officer at Kew, and Mr Bemd

Ullrich, Pohlheim, for providing us with useful documenta-

tion. Ms Lesley Henderson, Plant Protection Research Insti-

tute, kindly made distribution data available.

REFERENCES

BANERJEE, S. & SHARMA, A. K. 1987. Cytophotometric estimation of

nuclear DNA in different species and varieties of Agave. Cytolo-

gia 52: 85-90.

BERGER, A. 1915. DieAgaven. Beitrage zu einer Monographic. Gustav

Fischer Verlag, Jena.

BOLKHOVSKIKH, Z„ GRIF, V., MATVEJEVA, T„ ZAKHARYEVA,

O. 1969. In A. A. Federov, Chromosome numbers of flowering

plants. Academy of Sciences of the USSR, V. L. Komarov Botani-

cal Institute, Leningrad.

BRUMMITT, R. K. 1992. Vascular plant families and genera. Royal

Botanic Gardens, Kew.

COUPER, C. J. & CULLEN, J. 1988. Agave Linnaeus. In S. M Walters,

A. Brady, C. D. Brickell, J. Cullen, P. S. Green, J. Lewis, V. A.

Matthews, D. A. Webb, P. F. Yeo & J. C. M. Alexander, The

European Garden Flora, Pteridophyta, Gymnospermae, Angio-

spermae — Monocotyledons (Part 1) 1: 278-282. Cambridge Uni-

versity Press, Cambridge.

CRONQUIST, A. 1981. An integrated system of classification of flower-

ing plants. Columbia University Press, New York.

DOUGHTY, L. R. 1937. Experimental breeding of fibre agaves in East

Africa. Part I. The materials and the methods of the breeder. The

East African Agricultural Journal 3: 222-228.

DOUGHTY, L. R. 1938. Experimental breeding of fibre agaves in East

Africa. Part II. The progress of breeding at Amani. The East

African Agricultural Journal 3: 268-278.

DRUMMOND, J. R. 1910. L. Agave lurida. Kew Bulletin 1910 (No. 9):

344-349.

ENDLICHER, S. F. L. 1841. Enchiridion botanicum. Engelmann, Leipzig.

GENTRY, H. S. 1972. The Agave family in Sonora. United States Depart-

ment of Agriculture Agricultural Handbook No. 399: 1-195.

GENTRY, H. S. 1978. The Agaves of Baja California. Occasional Papers

of the California Academy of Sciences No. 130: 1-119.

GENTRY, H. S. 1982. Agaves of continental North America. The Univer-

sity of Arizona Press, Tucson, Arizona.

GRANICK, E. B. 1944. A karyosystematic study of the genus Agave.

American Journal of Botany 31: 283-289.

HENDERSON, L. 1995. Plant invaders of southern Africa. Plant Protec-

tion Research Institute, Agricultural Research Council, Pretoria.

LINNAEUS, C. 1753. Species plantarum. Salvius, Stockholm.

OCHOTERANA, I. 1913. Plantas deserticas Mexicanas. Agaves y yucas

de Durango. Memoires de la Societe “Alzate” 33: 93-113.

PEDLEY, L. & FORSTER, P. I. 1986. Agavaceae. In A. S. George, Flora

of Australia, Iridaceae-Dioscoreaceae 46: 71-88. Australian Gov-

ernment Publishing Service, Canberra.

PERRINE, H. 1838a. Tropical plants. Agave sisalana. United States of

America 25 th Congress, 2nd session — House of Representatives

Report No. 564: 8, 9, 16, 47, 60, 86.

PERRINE, H. 1838b. Agave sisalana. United States of America 25th

Congress, 2nd session — Senate Report (Unnumbered): 36, 105,

140.

SMITH, G.F. 1995a. Agave distribution maps — help needed. PlantLife

13: 6.

SMITH, G.F. 1995b. The distribution, uses and history of Agave in

southern Africa. Aloe 32: 84, 85.

SMITH, G.F. & CROUCH, N.R. in press. Agave L. (Agavaceae), the

century plants, in southern Africa. Veld & Flora 82.

TRELEASE, W. 1913. Agave in the West Indies. Memoirs of the National

Academy of Science 11: 1-55.

TRELEASE, W. 1914. Agave. In L. H. Bailey, The standard cyclopedia of

horticulture 1: 230-239. MacMillan, New York.

TRELEASE, W. 1920. 12. Amaryllidaceae. Amaryllis family. In P. C.

Standley, Trees and shrubs of Mexico (Gleicheniaceae-Betu-

laceae). Contributions from the United States National Herbar-

ium 23: 105-142.

ULLRICH, B. 1993. Agaven. Illustrationen bliihender Exemplare bis

1800. Palmengarten 21: 1-83.

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.

Bothalia 26,1: 37-40(1996)

FSA contributions 5: Buxaceae

H.F. GLEN*

Monoecious evergreen shrubs or small trees. Leaves

opposite, simple, shortly petiolate, entire; stipules 0. In-

florescences compact axillary racemes or cymes, sessile

or shortly pedicelled. Flowers radially symmetrical, uni-

sexual, small; perianth of 2 sepaloid whorls. Male flowers :

perianth lobes 4, imbricate; stamens 4—8, opposite perianth

lobes, filaments free, anthers dorsifixed near base, 2-the-

cous. Female flowers : perianth lobes 4-6, imbricate; ovary

superior, 3-locular with 2 pendulous ovules per locule,

styles 3, usually distant. Fruit a loculicidal, 3-homed cap-

sule. Seeds glossy black, triangular in section.

A mainly northern hemisphere family of four genera

and about 100 species. One genus with two species occurs

naturally within the area of the Flora of southern Aftica.

In addition species of the genera Sarcococca, Pachysan-

dra and Simmondsia (this last often regarded as constitut-

ing a separate family) are sometimes cultivated here.

4533000 BUXUS

Buxus L., Species plantarum 983 (1753); Hutch.:

375 (1915); Marloth: 142 (1925); Hutch.: 108 (1967);

R.A.Dyer : 326 (1975). Type species: B. sempervirens L.

Buxella Tiegh.: 326 (1897). No type cited.

Notobuxus Oliv.: t. 1400 (1882); Hutch.: 376 (1915); E.Phillips; 138

(1943); E.Phillips: 470 (1951). Type species: N. natalensis Oliv.

Description as for family.

la Bark longitudinally fissured, greenish brown; twigs an-

gled; leaves 12-40 x 6-18 mm, apex obtuse . .1. B. macowanii

lb Bark rough, pale brown; twigs grooved; leaves 50-100 x

20-50 mm; apex with a drip tip 2. B. natalensis

A mainly north temperate genus of some 80 species,

two of which occur naturally in our area. In addition, B.

sempervirens is occasionally cultivated here; PRE has

specimens from Tokai (Western Cape), Cedara (Kwazulu-

Natal) and Roodeplaat (Gauteng).

Friis (1989) discusses the generic delimitation of the

African species in some detail, and supports the conclu-

sion that while Notobuxus is a good subgenus, it cannot

be recognised as generically distinct from Buxus.

1. B. macowanii Oliv. in Hooker’s leones planta-

rum 16: t. 1518 (1886); Sim: 321 (1907); Hutch.: 55 (1912);

Hutch.: 375 (1915); Palmer & Pitman: 1183 (1973);

E.S.Pooley: 238 (1993). Type: Kwelegha Forest, Hutchins

* National Herbarium, Private Bag X101, Pretoria 0001.

MS received: 1994-02-08.

in MacOwan (Herbarium Nor male Austro-Africanum) 916

(K, lecto., here designated; BOL!, NBG!, isolecto.).

Buxella macowanii (Oliv.) Tiegh.: 326 (1897).

Notobuxus macowanii (Oliv.) E.Phillips: 140 (1943).

Tree 4—9 m tall, ± glabrous; stem straight, slender; bark

greenish brown, longitudinally fissured; branchlets angled.

Leaves opposite, narrowly rhombic-elliptic to ovate-ob-

long, 12-40 x 6-18 mm, apex obtuse, sometimes minutely

apiculate, base cuneate, stiffly coriaceous, dark green,

shiny; in seedlings and juvenile specimens leaves very

narrowly linear. Inflorescences ± sessile axillary raceme;

flowers very small, greenish. Male flowers clustered at

base; perianth lobes 4; stamens 4—6, filaments free; pis-

tillode minute. Female flowers apical or on separate in-

florescences; perianth lobes 4—6; ovary 3-locular; styles

3. Fruit a loculicidal, 3-homed, greenish brown capsule,

± 7 mm in diameter. Seeds glossy black, triangular in sec-

tion, ± 5.0 x 2.5 x 2.5 mm. Flowering July-October; fruit-

ing February to June. Figure 1.

Widespread from Eastern Cape to the North-West

Province and Mpumalanga (Eastern Transvaal). Figure 2.

Vouchers: Abbott 87; W.F. Barker 10902; Codd 10512; G.K. Theron

2161; H.J.T. Venter 5226.

Oliver (1886) records in the protologue that this species

is described from several gatherings made by Hutchins,

and sent to Kew by MacOwan. This implies the existence

of several syntypes, none of which is specified precisely.

Hutchinson (1912) mentions only one of these, namely

Hutchins in MacOwan 916. This implicit acceptance of

this specimen as lectotype is here made explicit.

Sterile plants of this species are similar in habit and

leaf size and arrangement to Carissa bispinosa (L.) Desf.

ex Brenan, but the latter has conspicuous branched thorns,

whereas Buxus species are unarmed. The leaves of B. sem-

pervirens are similar in size and shape to those of B. mac-

owanii, but in the exotic species the midribs and a

marginal vein are much more prominent, especially be-

neath.-

The Xhosa name garagara is recorded for this species

by Oliver (1886); the specimen Archibald 5409 (BOL)

gives the alternative spelling gala-gala. Palmer & Pitman

(1973) record, in addition to the other common names

noted in this paragraph, the names Buig-my-nie (Afri-

kaans), umGalagala and umNgquzu (Zulu). The specimen

F. Venter 2000 (PRE) records the Tswana common name

mupshane. Marloth (1925) gives the common name Cape

box, noting that the wood of Gonioma kamassi E.Mey.

was exported under the same name. He records that the

wood of Buxus macowanii is almost as suitable for mak-

ing woodcuts for printing as that of the European B. sem-

pervirens, but that logs are inclined to split while drying.

38

Bothalia 26,1 (1996)

FIGURE 1. — Buxus macowanii Oliv.: A,

branch x 0.9; B, open capsule, x

0.9; C, broad-leaved form, x 0.9;

D, cluster of one female and two

male flowers, x 2.5; E, male

flower, x 4.3; F, female flower, x

4.3. Taken from Sim (1907).

In the National Tree List (Von Breitenbach 1995) this tree

is No. 358, with standard names Kaapse buksboom and

Cape Box.

Although the first published record of this species dates

from 1886, the first written account of it known to the

present author is some 60 years older. This is a curious

handwritten account by C.H. Wehdemann (see entry under

this name in Gunn & Codd 1981) of a group of trees on

a farm near Somerset East, Eastern Cape. This account

forms part of a set dated 1 August 1826; if Wehdemann

had published his notes, this tree would have quite a dif-

ferent name today.

2. B. natalensis (Oliv.) Hutch., Genera of flowering

plants 2: 108 (1967); Palmer & Pitman 2: 1185 (1973);

E.S.Pooley 238 (1993). Syntypes: Natal, Inanda, Wood

1357 (NH, K, BOL!); Natal, Tongaat, Cooper 3465 (K).

Notobuxus natalensis Oliv.: t. 1400 (1882); Sim: 320 (1907); Hutch.:

55 (1912); Hutch.: 376 (1915).

Shrubs or small trees 1— 3(— 5) m tall, ± glabrous; main

stem straight, slender; bark rough, pale brown; young

FIGURE 2. — Distribution of Buxus macowanii Oliv.

Bothalia 26,1 (1996)

39

FIGURE 3. — Buxus natalensis (Oliv.)

Hutch.: branch with flowers and

fruit, x 0.9.

stems bright green, grooved. Leaves opposite, shortly

petiolate, elliptic, 50-120 x 20-50 mm, apex with a drip-

tip, base cuneate; midrib prominent below; thinly co-

riaceous, dark green, shiny; petiole 2-6 mm long.

Inflorescences ± sessile axillary clusters with both male

and female flowers; flowers small, white, pale grey, green

or yellow. Male flowers: perianth lobes 4; stamens 4—8,

filaments free; pistillode minute. Female flowers: perianth

lobes 4-6; ovary 3-locular; styles 3. Fruit a loculicidal,

3-horned, greenish brown capsule, ± 12 mm in diameter.

Seeds glossy black, triangular in section, ±7x3x3 mm.

Flowering August-September; fruiting September-De-

cember. Figure 3.

Endemic to KwaZulu-Natal and adjacent Transkei,

mostly coastal. Figure 4.

Vouchers: Abbott 214; A.W. Bayer 767; CJ. Ward 7153; Wells 3502;

White 10631.

Palmer & Pitman (1973) record the common names

Natal box, iGalagala and ukuXeza (Xhosa), umGalagala,

umHlulambazo, umPhicamaguma and umMgquzu (Zulu)

for this tree. In the National Tree List (Von Breitenbach

FIGURE 4. — Distribution of Buxus natalensis (Oliv.) Hutch.

40

Bothalia 26,1 (1996)

1995) this is No. 359, with standard names Natalse

Buksboom and Natal Box.

REFERENCES

DYER, R.A. 1975. Genera of South African flowering plants 1. Govern-

ment Printer, Pretoria.

FRIIS, 1. 1989. A synopsis of the Buxaceae in Africa south of the Sahara.

Kew Bulletin 44: 293-299.

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

Africa. Balkema, Cape Town.

HUTCHINSON, J. 1912. African Buxeae. Kew Bulletin 1912: 52-55.

HUTCHINSON, J 1915. Buxus and Notobuxus (Euphorbiaceae). Flora

capensis 5,2: 375-377. Lovell Reeve, London.

HUTCHINSON, J. 1967. Genera of flowering plants. Clarendon Press,

Oxford.

LINNAEUS, C. 1753. Species plantarum 1. Salvii Holmiae.

MARLOTH, H.W.R. 1925. Flora of South Africa. Darter, Cape Town.

OLIVER, D. 1882. Notobuxus natalensis. Hooker s leones plantarum 14:

78, t. 1400.

OLIVER, D. 1886. Buxus macowanii. Hooker’s leones plantarum 16: 1. 1518.

PALMER, E. & PITMAN, N. 1973. Trees of southern Africa. Balkema,

Cape Town.

PHILLIPS, E.P. 1943. Some changes in nomenclature IV. Journal of

South African Botany 9: 137-140.

PHILLIPS, E.P. 1951. The genera of South African flowering plants, edn

2. Memoirs of the Botanical Survey of South Africa No. 25.

POOLEY, E.S. 1993. The complete field guide to trees of Natal, Zululand

and Transkei. Natal Flora Publications Trust, Durban.

SIM, T.R. 1907. The forests and forest flora of the Colony of the Cape of

Good Hope. Taylor & Henderson, Aberdeen.

VAN TIEGHEM, Ph. 1897. Sur les Buxacees. Annales des sciences

naturelles, botanique, Paris, Ser. 8, 5,5: 289-338.

VON BREITENBACH, F. 1995. National list of indigenous trees, edn 3.

Dendrological Foundation, Pretoria.

Bothalia 26,1:41-52(1996)

Notes on African plants

VARIOUS AUTHORS

CELASTRACEAE

CORRECT ORTHOGRAPHY AND AUTHOR CITATION FOR ELAEODENDRON

Disagreement exists in the literature as to the correct

form and author citation of the generic name Elaeoden-

dron. Some sources refer to the taxon as Elaeodendron ,

with Jacq. f. (also abbreviated J.F. Jacq. or J. Jacq.) as

validating author (Bentham & Hooker 1862; Loesener

1942; Brummitt 1992; Greuter et al. 1993; Green 1994),

whereas others attribute the name to Jacq. or Jacq. f. ex

Jacq. (Robson 1966; Villiers 1975; Proctor 1984; Robson

et al. 1994). Kostermans (1986) argues that Elaeodendrum

Murray (1784) has priority over Elaeodendron Jacq. f.

(1787), and is therefore the correct name, a practice fol-

lowed by, for example, Loesener (1907, 1926) and Britton

& Millspaugh (1962). Jacquin f. (1787) introduced yet

another variant by referring to the genus in the genitive

case, namely Elaeodendri.

Elaeodendron was first published in 1782, without de-

scription, as a colour plate labelled Elaeodendron orien-

tate and bound in the first volume of the elder Jacquin’s

three-volumed leones plantarum rariorum (Figure 1). The

colour plates of this work were issued in fascicles of 25,

in an order unrelated to that in which they were eventually

bound. The publication date and composition of each fas-

cicle have been reviewed by Schubert (1945). Elaeoden-

dron orientale (plate no. 48) formed part of fascicle 2 and

was published in 1782. The text accompanying the plates

of this fascicle was published a few years later, possibly

in 1787 (Schubert 1945).

As source for the generic name Elaeodendron , the elder

Jacquin (in both the text and plate bound with Vol. 1 of his

leones) referred to a then still unpublished manuscript of the

younger Jacquin, intended for publication in Acta Helvetica

Physico-Mathematico-Botanico-Medica Vol. 9. This series

was apparently discontinued after Vol. 8 (Ingram 1966;

Lawrence et al. 1968; Burdet et al. 1980), a deduction sup-

ported by the fact that no reprint of any further volumes has

been found in Jacquin’s library at WU, or anywhere else

(Stafleu & Cowan 1979). Eventually, in 1787, the younger

Jacquin’s manuscript was published in the solitary issue of

a new series, Nova Acta Helvetica Physico-Mathematico-

Botanico-Medica, thus providing a description of Elaeoden-

dron, albeit with the name in the genitive case as Elaeodendri.

Although some workers (Farr et al. 1979; Stafleu & Cowan

1979; Howard 1989) proposed an earlier publication date

(1780-1784) for this new series, evidence is overwhelmingly

in favour of 1787 (Lawrence et al. 1968). Note, however,

that some authors have erroneously attributed the contribu-

tion in Nova Acta Helvetica Physico-Mathematico-Botanico-

Medica to the elder Jacquin, rather than to his son (Robson

et al. 1994).

As was the case with the elder Jacquin, Murray (1784)

had access to the younger Jacquin’s manuscript and, al-

though not intending to publish the genus, provided a brief

description of it under the name Elaeodendrum. Thus

Murray’s description of the generic name pre-dates those

of the elder and younger Jacquin by about three years.

Should Murray then not be credited as the validating

author of the generic name for the taxon under discussion,

as suggested by Kostermans (1986)?

According to Art. 42 of the Tokyo Code (Greuter et

al. 1994), Elaeodendron should, despite the lack of a de-

scription, be considered validly published just on the basis

of the colour plate in the leones plantarum rariorum, Vol.

1, t. 48 (1782). This plate meets the requirement for valid

FIGURE 1. — Copy of colour plate of Elaeodendron orientate Jacq.

published in N.J. Jacquin’s leones plantarum rariorum Vol. 1

(1782). Note name of taxon and reference to Acta Helvetica Vol.

9. Elongated leaves on juvenile branch are typical of species.

42

Bothalia 26,1 (1996)

publication in containing an analysis (defined in Art. 42.4

of the Code as a figure or group of figures, ... showing

details aiding identification,...’) and a binomial appearing

in print on it (Art. 42.3). The author citation of Elaeoden-

dron is therefore correctly referred to as Jacq. (or Jacq. f.

ex Jacq.). Recently, Brummitt & Powell (1992) proposed

J. Jacq. as abbreviation for the younger Jacquin.

The elder Jacquin ’s 1787 text accompanying his 1782

plate of E. orientate , consists of a mere list of the parts of

the illustration in Latin, but is, as in the case of the work of

Murray (1784) and Jacquin f. (1787), irrelevant in terms of

the Code. Arguments for accepting Elaeodendrum Murray

as the correct generic name are therefore not valid. Elaeoden-

drum and Elaeodendri are to be treated merely as orthog-

raphical variants and should be corrected (Art. 61). Although

the genus is often included in Cassine L. s.l, a stricter de-

limitation of the genera, including the recognition of

Elaeodendron Jacq., has recently been advocated (Archer &

Van Wyk 1993a, b).

REFERENCES

ARCHER, R.H. & VAN WYK, A.E. 1993a. Bark structure and intergen-

eric relationships of some southern African Cassinoideae (Celas-

traceae). IAWA Journal 14: 35-53.

ARCHER, R.H. & VAN WYK, A.E. 1993b. Wood structure and generic

status of some southern African Cassinoideae (Celastraceae).

IAWA Journal 14: 373-389.

BENTHAM, G. & HOOKER, J.D. 1862. Genera plantarum 1. Lovell

Reeve, London.

BRITTON, N.L. & MILLSPHAUGH, C.F. 1962. The Bahama flora.

Hafner Publishing Company, New York.

BRUMMITT, R.K. 1992. Vascular plant families and genera. Royal

Botanic Gardens, Kew.

BRUMMITT, R.K. & POWELL, C.E. 1992. Authors of plant names.

Royal Botanic Gardens, Kew.

BURDET, H.M., FELDMANN, F., VON LILIENFELD, M„ MAIUL-

LAR1, F. & MEUWLY, R. 1980. Catalogue des periodiques de la

Bibliotheque des Conservatoire etJardin botaniques de la Ville de

Geneve. Conservatoire et Jardin botaniques, Geneve.

FARR, E.R., LEUSSINK, J.A. & STAFLEU, F.A. 1979. Index nominum

genericorum, Vol. 1. Bohn, Scheltema & Holkema, Utrecht.

GREEN, P.S. 1994. Celastraceae. In A.E. Orchard, Flora of Australia 49,1:

225, 226. Australian Government Publishing Service, Canberra.

GREUTER, W„ BRUMMITT, R.K., FARR, E„ K1LIAN, P.M., & SILVA,

PC. 1993. Names in current use for extant plant families. Koeltz

Scientific Books, Konigstein.

GREUTER, W„ BARRIE, F.R., BURDET, H.M., CHALONER, W.G.,

DEMOULIN, V., HANKSWORTH, D.L., JORGENSEN, P.M.,

NICHOLSON, D.H., SILVA, P.C., TREHANE, P. & MCNEILL,

J. 1994. International code of botanical nomenclature ( Tokyo

Code). Koeltz Scientific Books, Konigstein.

HOWARD, R.A. 1989. Flora of the Lesser Antilles. Leeward and Wind-

ward Islands, Vol. 5. Arnold Arboretum, Harvard University,

Massachusetts.

INGRAM J. 1966. Notes on the cultivated Liliaceae. 4. Lachenalia.

Baileya 14: 123-132.

JACQUIN, J.F. 1787. Tria genera plantarum nova. Nova Acta Helvetica

Physico-Mathematico-Botanico-Medica 1: 34—41.

JACQUIN, N.J. 1781-1787. leones plantarum rariorum 1: t. 48 & 5.

Vindobonae [Wien].

KOSTERMANS, A.J.G.H. 1986. Notes on Asiatic Cassine L. (Celas-

traceae). Gardens' Bulletin, Singapore 39: 177-191.

LAWRENCE, G.H.M., GUNTHER BUCHHEIM, A.F., DANIELS, G.S.

& DOLEZAL, H. 1968. Botanico-Periodicum-Huntianum. Hunt

Botanical Library, Pittsburgh.

LOESENER, T. 1907. Ein neues Elaeodendrum. Botanische J ahrbiicher

40: 61.

LOESENER, T. 1926. Celastraceae. In R. Elias & T.C.E. Fries, Beitrage

zur Kenntnis der Flora des Kenia, Mt Aberdare und Mt Elgon.

VIII. Notizblatt des Botanischen Gartens und Museums zu Berlin-

Dahlem 9: 485-522.

LOESENER, T. 1942. Celastraceae. In A. Engler, H. Harms & J.

Mattfeld, Die natiirliehen Pflanzenfamilien, edn 2, 20b: 87-197.

Duncker & Humblot, Berlin.

MURRAY, J.A. 1784. Linnaeus's Systema vegetabilium, edn 14. Gottin-

gae [Gottingen],

PROCTOR, G.A. 1984. Flora of the Cayman Islands. H.M.S.O., London.

ROBSON, N.K.B. 1966. Celastraceae (inch Hippocrateaceae). In A.W.

Exell, A. Fernandes & H. Wild, Flora zambesiaca 2, 2: 355-418.

Crown Agents for Oversea Governments and Administrations,

London.

ROBSON, N.K.B. , HALLE, N„ MATHEW, B. & BLAKELOCK, R.

1994. Celastraceae. In R.M. Polhill, Flora of tropical East Africa

1-78. Balkema, Rotterdam.

SCHUBERT, B.G. 1945. Publication of Jacquin’s leones plantarum rari-

orum. Contributions from the Gray Herbarium of Harvard Uni-

versity 154: 3-23.

STAFLEU, F.A. & COWAN, R.S. 1979. Taxonomic literature, Vol. 2.

Bohn, Scheltema & Holkema, Utrecht.

VILLEERS, J.-F. 1975. Celastraceae. In A. Aubreville & J.F. Leroy, Flore du

Cameroun 19: 3-32. Museum National D'Histoire Naturelle, Paris.

R.H. ARCHER* & A.E. VAN WYK**

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

**H.G.W.J. Schweickerdt Herbarium, Department of Botany, University

of Pretoria, Pretoria 0002.

MS. received: 1995-05-26.

CROTALARIEAE

RARE OR EXTINCT SPECIES OF ARGYROLOBIUM

While revising Argyrolobium for South Africa, three

taxa which are known from single collections were rec-

ognised. Perhaps these species will join a burgeoning list

of extinctions for southern Africa. It is hoped that the

publication of this paper will alert collectors and disprove

this assertion.

1. Argyrolobium crinitum (E.Mey.) Walp. in Linnaea

13: 506 (1839); Walp.: 630 (1843); Benth.: 340 (1844); Harv.:

68 (1862). Type: Western Cape, Swaartberge, Trado, Drege

6627 (K; P!).

Chasmone crinita E.Mey.: 71 (1836).

Suffrutex up to 600 mm tall, forming clumps, branched

basally; stems annual, rufous-pilose. Leaves pilose; leaf-

lets ovate to obovate, 30-50 x 14-23 mm, apex rounded;

petiole 10-20 mm long, adaxially canaliculate; stipules

20-40 x 18-24 mm, amplexicaul, apex usually 4-lobed.

Inflorescence elongate, terminal; bracts obovate, 13 x 5

mm; bracteoles 2.00 x 0.75 mm, filiform. Calyx pilose,

deeply bilabiate; upper lobes 6 mm long, upper sinus 4.5

mm deep, lower lip 8 mm long, lobes acute, 3 mm long.

Corolla yellow; standard suborbicular, strongly reflexed,

8x8 mm, adaxial surface sparsely sericeous medially,

claw canaliculate, 4 mm long; wings oblong, 9x4 mm,

glabrous, claw 4 mm long, sculpturing in the lower basal

Bothalia 26,1 (1996)

43

FIGURE 2. — Argyrolobium crinitum,

Drege s.n. A, vegetative branch;

B, calyx; C, wing; D, keel; E,

standard; F, calyx, pedicel and

bract. Scale bars: A, 20 mm;

B-F, 2 mm.

and upper basal zones; keel cymbiform, 7x4 mm, claw

5 mm long. Stamens monadelphous, sheath split above.

Ovary narrowly oblong, 9 mm long, densely rufo-sericeous,

6-seeded; style 4 mm long. Fruit and seed unknown.

A. crinitum (Figure 2) is a distinctive species with large,

4-lobed stipules. Its distinctive morphology led Meyer (1836)

to suggest that it may represent a new genus. The closest

ally is undoubtedly A. amplexicaule (E.Mey.) Dummer

which has similar leaves, stipules and vestiture. The species

are allopatric with A. amplexicaule commonly occurring in

grasslands of the Eastern Cape and KwaZulu-Natal.

Only a single flower of A. crinitum was available for

examination and details of inflorescence structure were

therefore taken from Meyer (1836). The calyx has a trun-

cate base and the petals have well-developed claws, fea-

tures unusual for Argyrolobium. The fusion of the filaments,

the dimorphic anthers and the calyx lobing, however, support

its generic placement.

WESTERN CAPE. — 3320 (Montagu): Swaartberge, Trado, (-DC),

Drege 6627 (K. P).

2. Argyrolobium splendens ( Meisn .) Walp., Rep-

ertorium botanices systematicae 2: 845 (1843); Benth.:

348 (1844); Harv.: 76 (1862). Type: Western Cape, moun-

tainsides around Klein Rivier, Swellendam, Krauss 927

(Bf; NY!, lecto. selected here; G!, MO!, W!, isolecto.).

Chasmone splendens Meisn.: 78 (1843).

44

Bothalia 26,1 (1996)

Herb, 0.2-0. 3 m tall, erect, well branched, stems

shortly sericeous, plants forming clumps. Leaves : abaxial

surface densely sericeous, adaxially sparsely sericeous be-

coming glabrous; leaflets oblong to obovate, 20-35 x

7-14 mm, strongly revolute, apex rounded, apiculate; peti-

ole 5-12 mm long; stipules ovate, 7-9 x 4 mm, base

obliquely cordate. Inflorescence pseudo-umbellate, 1-3-

flowered, peduncle well developed, 25-60 mm long,

leaf-opposed; bracts lanceolate, 4—5 x 1. 5-2.0 mm; brac-

teoles lanceolate, 3x1 mm. Calyx shortly sericeous, up-

per lip 10-11 mm long, sinus 7-9 mm deep; lower lip

11-12 mm long, lobes 3-4 mm long, medial lobe linear.

Corolla yellow; standard orbicular, 12-14 x 9-12 mm,

adaxial surface sparsely sericeous, base obtuse, claw 3

mm long; wings oblong, 11-12 x 3 mm, with lunate-

lamellate sculpturing in upper central zone, claw 3 mm

long; keel cymbiform, 9-10 x 4 mm, upper margin

shortly ciliate, 3 mm long. Stamens monadelphous; an-

thers dimorphic. Ovary weakly arcuate, 8-9 mm long,

densely sericeous; style 4-5 mm long, basally hairy.

Fruit compressed, sericeous, 38-42 x 5 mm. Seed not

seen. Figure 3.

The obvious alliance between this species and A. lu-

nare (Meisner 1843; Harvey 1862) contradicts the use of

torulose fruits as a sectional character within Argyrolo-

bium (Bentham 1844; Harvey 1862). A. splendens is

known only from the type collection which was gathered

on the coastal fold mountains which flank the Kleinrivier

at Hermanus (Figure 4). The species is sympatric with

its ally A. lunare (L.) Druce which is distinguished by

its torulose fruits, longer petioles and non-revolute leaf

margins.

WESTERN CAPE. — 3419 (Caledon): mountainsides around Klein

Rivier, (-AD), Krauss 927 (NY, G, MO, W).

FIGURE 3. — Argyrolobiuni splen-

dens, Krauss 927. A, repro-

ductive branches; B, calyx, in-

ner surface; C, wing; D, keel;

E, standard. Scale bars: A, 20

mm; B-E, 2 mm.

Bothalia 26,1 (1996)

45

FIGURE 4. — Recorded distribution of: A. crinitum , •: A. splendens, ▲;

and A. angustissimum, ■

3. Argyrolobium angustissimum (E.Mey.) TJ.

Edwards comb. nov. Type: Western Cape, Paarlberg,

1000-2000 ft, Drege s.n. (G!, lecto. selected here; BM!,

E!, MO!, OXF!, P!, SI, W!, isolecto.).

Chasmone angustissima E.Mey.: 75 (1836).

Herb, 0.2-0.3 m tall, erect, sparsely branched, stems

weakly perennial, shortly sericeous, plants solitary, or

forming small clumps. Leaves adaxially glabrous, abaxially

shortly sericeous leaflets; linear, 25-45 x 1.0-1. 5 mm,

conduplicate, apex acute; petiole 1. 5-3.0 mm long; stipu-

les deltoid to linear, 2-4 x 0.5-1. 0 mm. Inflorescence

pseudo-umbellate, 1-6-flowered, peduncles leaf-opposed;

bracts linear, 4—5 x 0.75-1.00 mm; bracteoles linear,

3.5-5.0 x 0.75 mm. Calyx shortly sericeous, upper lip

14—15 mm long, sinus 12-14 mm deep; lower lip 15-17

mm long, lobes 4—5 mm long, medial lobe linear. Co-

rolla russet and yellow; standard obovate, 16-18 x

10-12 mm, adaxial surface sparsely sericeous, base

cuneate, claw 1-2 mm long; wings narrowly oblong to

obovate, 14.0-15.0 x 3.5 mm, with lunate-lamellate

sculpturing in upper basal zone, claw 1.0-1 .5 mm long;

keel cymbiform, 7-9 x 3.5-5. 0 mm, distally sericeous,

claw 1. 0-5.0 mm. Stamens monadelphous; anthers very

weakly dimorphic. Ovary weakly arcuate, ± 10 mm

long, densely sericeous; style 3-4 mm long, basally

hairy. Fruit not seen. Seed not seen.

FIGURE 5. — Argyrolobium acicu-

lare, Stokoe 61544: A, habit.

B, A. harveyanum, Edwards

s.n.: habit of southern form.

C-F, L: A. filiforme, Walgate

399: C, calyx; D, keel; E,

wing; F, standard; L, pistil.

G-K, A. angustissimum,

Drege 1419: G, calyx; H, keel;

I, wing; J, standard; K, pistil.

Scale bars: A, B, 20 mm; C-L,

2 mm.

46

Bothalia 26,1 (1996)

A. angustissimum is formally recognised on the basis

of its robust habit and large distinctive flowers (Figure

5). It is closely allied to A. tuberosum which has similar

leaf morphology and flower colour but differs in the

size and morphology of the flowers. The species are

allopatric, A. angustissimum is known from a single

Drege collection (Figure 4) made at Paarlberg whereas

A. tuberosum occurs sporadically in grasslands of the

Eastern Cape, KwaZulu-Natal, Mpumalanga and into

the highlands of Zimbabwe.

WESTERN CAPE. — 3318 (Cape Town): Paarlberg, (-DB), Drege

s.n. (BM, E, G, MO, OXF, P, S, W).

ACKNOWLEDGEMENTS

I wish to thank the Curators of the cited herbaria for

the loan of herbarium specimens and Kew for the provi-

sion of cibachromes of their type material. The referees

are thanked for their comments on the manuscript. The

Natal University Research Fund is gratefully acknow-

ledged for financial assistance.

REFERENCES

BENTHAM, G. 1844. Enumeration of Leguminosae, indigenous to

southern Asia, and central and southern Africa. Hooker s London

Journal of Botany 3: 338-365.

HARVEY, W.H. 1862. Leguminosae. Flora capensis 2. Hodges Smith,

Dublin.

MEISNER, C.F. 1843. Contributions towards a Flora of South Africa.

London Journal of Botany 2: 53-105.

MEYER, E.H.F. 1836. Commentariorum de plantis africae australioris 1 .

Leopoldum Voss, Leipzig.

WALPERS, G. 1839. Animadversiones criticae in Leguminosas Cap-

enses. Linnaea 13: 449-543.

WALPERS, G. 1843. Repertorium botanices systematicae 1. Friedrich

Hofmeister, Leipzig.

T.J. EDWARDS*

* Unit for Plant Growth and Development, Botany Department, Univer-

sity of Natal, P.O. Box 01, Scottsville 3209.

MS. received: 1995-05-25.

CONVOLVULACEAE

NEW RECORDS, NAME CHANGES AND A NEW COMBINATION IN SOUTHERN AFRICA

INTRODUCTION

During the preparation of the Convolvulaceae for the

Flora of southern Africa ( FSA ), Vol. 28, part 1, it became

clear that several changes and additions are necessary to

the list compiled by Welman: 588-595 (1993). As publi-

cation of this part of the FSA is not expected in the near

future, it will be useful to publish them here.

NEW RECORDS

The following species have recently been recorded as

new to southern Africa.

6979000-00200 Bonamia velutina Verde, in Kirkia

1: 27, t. Ill (1961).

Very rare perennial in the Northern Province and in the

Ingwavuma area of KwaZulu-Natal; also reported from

southeastern Botswana, Zimbabwe and Mozambique.

B. velutina can be confused with Seddera capensis (E.Mey.

ex Choisy) Hallier f. and S. suffruticosa (Schinz) Hallier

f., but can be recognized by the greyish velvety pubes-

cence on the upper side of the leaves and the golden

brown pubescence on the underside as well as on the calyx

lobes.

7003000-01450 Ipomoea consimilis Schulze-Menz in

Notizblatt des Botanischen Gartens und Museums zu Ber-

lin 14: 112 (1938).

Recorded from Tanzania (Lindi District), Mozambique

(Sofala) and Maputaland in northern KwaZulu-Natal. The

South African specimens of this species were previously

listed under I. urbaniana (Dammer) Hallier f., which does

not occur south of the Lindi District in Tanzania (Ross

1972: 296). Distinguished by its climbing, shrubby habit,

its yellow villous young parts, its large ovate leaves and

its mauve to purple flowers.

7003000-01950 Ipomoea fanshawei Verde, in Kirkia

6: 119 (1967).

Fairly rare perennial, endemic to southern Zambia and

northeastern Botswana. Distinguished by its rhomboid-

lanceolate leaves with entire to lobed margins and its 60

mm long, pale purple or mauve corollas.

7003000-02150 Ipomoea fulvicaulis (Hochst. ex

Choisy) Boiss. ex Hallier f. in Botanische Jahrbiicher 18:

128 (1893).

Perennial from Ethiopia, Kenya, Tanzania, Malawi,

Zaire, Zambia and Zimbabwe to Mozambique; also in

northern Botswana. Distinguished by its yellow-brown

pubescent stems, flowers in small heads and 25-50 mm

long purplish corollas. Only var. fulvicaulis has been re-

corded from the FSA region; var. heterocalyx (Schulze-

Menz) Verde, and var. asperifolia (Hallier f.) Verde, do

not reach southern Africa.

7003000-03350 Ipomoea ochracea (Lindi.) G.Don,

A general system of gardening and botany 4: 270 (1837).

Found throughout tropical Africa down to Ngamiland

in Botswana, where it is very rare. Only the typical var.

is known from the FSA region. Var. curtissii (House) Steam,

Bothalia 26,1 (1996)

47

a much stouter plant, comes from Uganda. The typical

variety is distinguished by being a perennial herb with

bright yellow flowers with a dark throat, arranged in

cymes.

NAME CHANGES

The following name changes have become necessary.

Turbina sensu Meeuse (1957) is sunk into Ipomoea,

sens. lat. at least as far as six of the southern African

species are concerned. The only constant difference

from Ipomoea sens, strict, is the indehiscence of the

capsules which is almost certainly attributable to an ad-

aptation associated with seed dispersal. A dehiscent

capsule, such as that of Ipomoea sens, strict., releases

several diaspores (normally 4), whereas in the African

species of Turbina , the fruit usually contains only one

or two seeds and dehiscence would therefore not add

much to the number of diaspores. If there was a ten-

dency towards fleshiness of the fruit wall and endo-

zoochory, dehiscence would not serve a useful purpose.

The southern African species formerly placed in Tur-

bina form a very heterogeneous assemblage and some

of them resemble certain species or species groups of

Ipomoea much more than the others.

7003000-02450 Ipomoea holubii Baker in Kew

Bulletin: 72 (1894).

Turbina holubii (Baker) A.Meeuse: 780 (1957).

7003000-03150 Ipomoea oblongata E.Mey. ex

Chois y in A.P. de Candolle, Prodromus systematis natu-

ralis regni vegetabilis etc.: 368 (1845).

Turbina oblongata (E.Mey. ex Choisy) A.Meeuse: 778 (1957).

/. atherstonei Baker in Baker & C.H. Wright: 53 (1904); A.Meeuse:

741 (1957).

A very common and very variable species. There are

no important morphological differences between I. oblon-

gata and I. atherstonei, there is no geographical or eco-

logical separation either.

7003000-03450 Ipomoea oenotheroides (L.f.) Raf.

ex Hallierf. in Botanische Jahrbiicher 18: 156 (1893) in syn.

Turbina oenotheroides (L.f.) A.Meeuse: 775 (1957).

7003000-04350 Ipomoea robertsiana Rendle in

Journal of Botany, British and Foreign, London 39: 18 (1901).

Turbina robertsiana (Rendle) A.Meeuse: 777 (1957).

7003000-04850 Ipomoea stenosiphon Hallier f. in

Sitzungsberichte der Akademie der Wissenschaften in

Wien 107,1: 50 (1898b).

Turbina stenosiphon (Hallier f.) A.Meeuse: 783 (1957).

7003000-04875 Ipomoea suffruticosa Burch., Trav-

els in the interior of southern Africa, Vol. 2: 226 (1824).

Turbina suffruticosa (Burch.) A.Meeuse: 776 (1957).

7003020-00100 Paralepistemon shirensis (Oliv.)

Lejoly & Lisowski, in Bulletin du Jardin botanique na-

tional de Belgique, Brussels 56: 197 (1986).

Turbina shirensis (Oliv.) A.Meeuse: 782 (1957).

This monotypic genus differs from Ipomoea as follows:

Stamens inserted on triangular, large (±2x2 mm) glan-

dular pilose scales situated near base of corolla tube.

Ovary with very short beak persistent in fruit. Style ar-

ticulated at base, caducous. Fruit indehiscent, more or less

woody.

6978000-00150 Seddera schizantha Hallier f. in

Bulletin de FHerbier Boissier 6: 532 (1898a).

Bonamia schizantha (Hallier f.) A.Meeuse: 665 (1957).

Metaporana angolensis N.E.Br.: 169 (1914).

When N.E. Brown described Metaporana, he included

two species, M. angolensis and M. densiflora (Hallier f.)

N.E.Br. These two species are not congeneric and this

confused Meeuse (1957) into accepting Metaporana for

S. schizantha, which name Brown had overlooked. The

species which Brown re-described as M. angolensis is not

quite erect but tends to climb, so that Meeuse thought it

was a Bonamia rather than something else. However, My-

int & Ward: 234 (1968) in their monograph of Bonamia

exclude B. schizantha from Bonamia and list it as a syno-

nym of S. schizantha.

6993000-00250 Convolvulus aschersonii Engl., Uber

die Hochgebirgsflora des tropischen Afrika: 349 (1892).

C. sagittanis Thunb. var. aschersonii (Engl.) Verde.: 345 (1957).

C. aschersonii can be distinguished from C. sagittatus

by its usually few-flowered inflorescences (flowers usu-

ally solitary in the latter), small flowers (8-10 mm long)

and different leaf shape (middle lobe up to 60 x 12 mm,

often crenate or sinuate, occasionally dissected; basal

lobes often toothed or multifid). These characters do not

occur together in C. sagittatus.

6993000-00600 Convolvulus capensis Burm.f,

Prodromus florae capensis: 5 (1768).

C. capensis Burm.f. var. bowieanus (Rendle) A.Meeuse: 693 (1957).

C. capensis Burm.f. var. plicatus (Desr.) Baker in Baker & C.H. Wright:

78 (1904) as to new combination only, excl. descr., synonyms and speci-

mens cited.

In view of the appreciable variation of the leaf mor-

phology on a single plant the varieties distinguished by

Meeuse (1957: 693) are presumably only growth forms.

Young shoots may have undissected leaves (before more

dissected ones have developed) and depauperate and

drought forms may have narrow leaves.

6993000-01300 Convolvulus natalensis Bernh. in

Flora 27: 829 (1844).

C. natalensis Bemh. var. transvaalensis (Schltr.) A.Meeuse: 689 (1957).

48

Bothalia 26,1 (1996)

There are intermediates between the two varieties dis-

tinguished in Meeuse (1957) and their distribution ranges

overlap. The differences in leaves and sepals do not war-

rant the retention of two varieties (which may conceivably

belong to a single population).

6993000-01500 Convolvulus ocellatus Hook.f in

Curtis’s Botanical Magazine 70: t. 4065 (1844).

C. ocellatus Hook.f. var. ornatus (Engl.) A.Meeuse: 673 (1957).

This taxon is very variable and the extreme forms have

been described as varieties; but there are many transitional

specimens, those from the drier areas tending to form

more dissected leaves and a denser tomentum.

6993000-02050 Convolvulus sagittatus Thunb.,

Prodromus plantarum capensium 1: 35 (1794).

C. sagittatus Thunb. subsp. grandiflorus (Hallier f.) A.Meeuse var.

graminifolius (Hallier f.) Baker & C.H.Wright ex A.Meeuse: 683 (1957).

C. sagittatus Thunb. subsp. grandiflorus (Hallier f.) A.Meeuse var.

grandiflorus: 683 (1957).

C. sagittatus Thunb. subsp. grandiflorus (Hallier f.) A.Meeuse var.

linearifolius (Hallier f.) Baker & C.H.Wright ex A.Meeuse: 683 (1957).

C. sagittatus Thunb. subsp. sagittatus var. hirtellus (Hallier f.)

A.Meeuse: 682 (1957).

C. sagittatus Thunb. subsp. sagittatus var. namaquensis A.Meeuse:

682 (1957).

C. sagittatus Thunb. subsp. sagittatus var. phyllosepalus (Hallier f.)

A.Meeuse: 681 (1957).

C. sagittatus Thunb. var. ulosepalus (Hallier f.) Verde.: 346 (1957).

The great range in inflorescences and leaf size and

shape has led to the description of numerous subspecies

and varieties, but examination of a larger number of speci-

mens shows that these form a continuous range rather

than discrete entities.

It is noteworthy that most of the varieties previously

described, are not geographically restricted to small areas,

but have a wide range. Without extensive field and ex-

perimental studies, it is impossible to delimit varieties in

this entangled assembly.

7003000-00800 Ipomoea bathycolpos Hallier f. in

Botanische Jahrbiicher 18: 144 (1893).

I. bathycolpos Hallier f. var. sinuatodentata Hallier f.: 53 (1899).

Specimens with larger leaves with coarse irregular teeth,

a narrow basal sinus and usually larger flowers, were de-

scribed as the var. sinuatodentata Hallier f., but this variety

is not worth maintaining, because these specimens which

seem to be restricted to the Lydenburg area (with a milder

winter climate) may simply represent a more luxuriant form.

7003000-01100 Ipomoea bolusiana Schinz in Ver-

handlungen des Botanischen Vereins der Provinz Bran-

denburg 30: 271 (1888).

I. bolusiana Schinz var. pinnatipartita Verde.: 118 (1967).

The above variety represents an extreme form of the

range of variation of the leaf morphology and does not

deserve separate rank.

7003000-02900 Ipomoea magnusiana Schinz in

Verhandlungen des Botanischen Vereins der Provinz Bran-

denburg 30: 272 (1888).

I. magnusiana Schinz var. eenii (Rendle) A.Meeuse: t. 1201 (1956a).

Two varieties were distinguished in Meeuse: (1957:

742, 743), based mainly on the size of the corolla. It now

appears that the var. eenii cannot be upheld; the dimen-

sions of the corolla overlap one another and all other fea-

tures just about agree. The large-flowered specimens are

presumably only luxuriant individuals that grew in favour-

able sites or after heavy rains.

7003000-03300 Ipomoea obscura (L.) Ker Gawl.

in Botanical Register 3: t. 239 (1817).

I. obscura (L.) Ker Gawl. var. fragilis (Choisy) A.Meeuse: t. 1222

(1956b); 747 (1957).

The plants separated as var. fragilis differ only in the

absence of a dark centre in the throat of the corolla. Such

colour variations are caused by a difference in a few genes

(or only a single one) and there is no reason to recognize

the individuals with a concolorous corolla as more than a

forma.

All the specimens from southern Africa belong to the var.

obscura with corolla concolorous or with a dark centre. The

var. sagittifolia Verde, occurs from Zimbabwe to Tanzania.

7003000-01850 Ipomoea dichroa Choisy in A.P.

de Candolle, Prodromus systematis naturalis regni vege-

tabilis etc. 9: 364 (1845); Verde.: 165 (1978).

I. arachnosperma Welw.: 588 (1859); A.Meeuse: 736 (1957).

Verdcourt (1978: 165) states: ‘Since there is no bar to

the use of the epithet dichroa in Ipomoea and Choisy pro-

vides an adequate description, the correct name for this

species would appear to be Ipomoea dichroa treating it

as a new name dating from Choisy.’

7003000-02475 Ipomoea indica (Burm.f.) Merr.,

An interpretation of Rumphius’s Herbarium amboinense:

445 (1917); Austin: 357 (1986).

Convolvulus indicus Burm.f.: 6 (1755).

I. congesta R.Br.: 425 (1810); A.Meeuse: 735 (1957).

Austin (1986: 357) gives the full synonymy of this

highly polymorphic species which had been described

several times in different genera, from specimens col-

lected in both the western and eastern hemispheres.

6995000-00050 Hewittia malabarica (L.) Suresh

in Nicolson, Suresh & Manilal, An interpretation of Van

Rheede’s Hortus malabaricus: 88 (1988).

Convolvulus malabaricus L.: 155 (1753).

Convolvulus sublobatus L.f.: 135 (1781).

Bothalia 26,1 (1996)

49

H. sublobata (L.f.) Kuntze: 441 (1891); A.Meeuse: 698 (1957).

This rather variable species is widespread in the Old

World and had been described several times from different

localities (Gonqalves 1987: 31).

7010000-00050 Stictocardia laxiflora (Baker)

Hallier f in Bulletin de l’Herbier Boissier 6: 548 (1898a).

Ipomoea woodii N.E.Br.: 101 (1894).

S. woodii (N.E.Br.) Hallier f.: 548 (1898a); A.Meeuse: 773 (1957).

5. laxiflora (Baker) Hallier f. var. woodii (N.E.Br.) Verde.: 173 (1963b).

In the past, features like the lengths of the peduncles and

the pedicels were used to distinguish S. laxiflora (Baker)

Hallier f. var. woodii (N.E.Br.) Verde. However, the descrip-

tion made by Meeuse (1957: 773) was based on a whole

series of herbarium specimens, all from a single plant grown

in the Durban Botanical Garden. The range of variation is

so broad that such distinctions cannot be made.

NEW COMBINATION

6997010-00100 Xenostegia tridentata (L.) Austin &

Staples in Brittonia 32: 533 (1980).

subsp. angustifolia (Jacq.) A.Meeuse, comb. nov.

Ipomoea angustifolia Jacq.: 367 (1789).

Merremia tridentata (L.) Hallier f. subsp. angustifolia (Jacq.) Ooststr.

var. angustifolia Verde.: 51 (1963a).

Iconotype: Jacq., leones plantarum rariorum 2: 10, t.

317 (1786-1793).

The segregation of Xenostegia from Merremia is based

on two characters, namely, the straight anthers at anthesis

(in contrast to the always at least somewhat twisted an-

thers typical of Merremia ) and the pantoporate pollen

grains rather unique in the family. The two genera also

have a different ‘look’.

EXCLUDED SPECIES

Ipomoea shupangensis Baker

Welman (1993: 594) lists this species for KwaZulu-

Natal. However, no authoritatively identified material is

known from southern Africa. I. shupangensis occurs in

tropical Africa as far south as Zimbabwe (Hwange) and

Mozambique (Gaza).

I. batatas (L.) Lam. (the ‘Sweet Potato’ from tropical

America), I. coccinea L. (annual scarlet-flowered twiner from

North America) and I. fistulosa Choisy (purple-flowered

shrub from tropical America) are also listed in Welman

(1993), but occur only as garden escapes or culture relics.

REFERENCES

AUSTIN, D.F. 1986. Nomenclature of the Ipomoea nil complex (Con-

volvulaceae). Taxon 35: 355-358.

AUSTIN, D.F. & STAPLES, G.W. 1980. Xenostegia , a new genus of

Convolvulaceae. Brittonia 32: 533-536.

BAKER, J.G. 1894. Diagnoses Africanae II. Convolvulaceae. Kew Bulle-

tin-. 67-74.

BAKER, J.G. & WRIGHT, C.H. 1904. Convolvulaceae. In W.T. Thisel-

ton-Dyer, Flora capensis, Vol. 4,2. Reeve, London.

BERNHARDI, J.J. 1844. In F. Krauss, Pflanzen des Cap und Natal-Lan-

des. Flora 27: 819-835. Regensburg.

BROWN, N.E. 1894. Decades Kewensis, Ipomoea woodii. Kew Bulletin :

101.

BROWN, N.E. 1914. Diagnoses Africanae LIX. Kew Bulletin-. 168, 169.

BROWN, R. 1810. Prodromus florae novae hollandiae, 1st edn. Richard

Taylor, London.

BURCHELL, W.J. 1824. Travels in the interior of southern Africa. Vol. 2.

Longman, London.

BURMAN, N.L. 1755. In G.E. Rumphius, Herbarium amboinense, index

universalis 7: 6. Meinard Uytwerf, Amsterdam.

BURMAN, N.L. 1768. Flora indica: cui accedit series zoophytorum indi-

corum nec non prodromus florae capensis. Amsterdam/Leiden.

DE CANDOLLE, A.P. 1845. Prodromus systematis naturalis regni vege-

tabilis 9: 323 — 465. Treuttel & Wiirtz, Paris.

DON, G. 1837. A general system of gardening and botany containing a

complete enumeration and description of all plants hitherto

known. Vol. 4. London.

ENGLER, H.G.A. 1892. Uber die Hochgebirgsflora des tropischen Af-

rika. Berlin.

GONSALVES, M.L. 1987. 117-118. Convolvulaceae-Cuscutaceae. In

E. Launert, Flora zambesiaca , Vol. 8,1. Flora Zambesiaca Manag-

ing Committee, London.

HALLIER, H. 1893. Convolvulaceae africanae. Botanische Jahrbiicher

18: 81-160.

HALLIER, H. 1898a. Beitrage zur Kenntnis der Afrikanischen Flora.

Convolvulaceae. Bulletin de VHerbier Boissier 6: 529-548.

HALLIER, H. 1898b. Sitzungsberichte derAkademie der Wissenschaften

in Wien. Vol. 107,1: 50.

HALLIER, H. 1899. Beitrage zur Kenntnis der Afrikanischen Flora.

Convolvulaceae IV. Bulletin de VHerbier Boissier 7: 41-55.

HOOKER, W.J. 1844. Convolvulus ocellatus. Purple-eyed bind-weed.

Curtis’s Botanical Magazine 70: t. 4065.

KER GAWLER, J.B. 1817. Ipomoea obscura. Hairy-stemmed East In-

dian Ipomoea. Botanical Register 3: t. 239.

KUNTZE, C.E.0. 1891. Revisio generum plantarum, Part 2. Arthur Felix,

Leipzig.

LEJOLY, J. & LISOWSKI, S. 1986. Paralepistemon , nouveau genre de

Convolvulaceae d’Afrique tropicale. Bulletin duJardin botanique

national de Belgique 56: 195-197.

LINNAEUS, C. 1753. Species plantarum exhibentes plantas rite cogni-

tas, ad genera relatas, 1st edn. Salvii, Stockholm.

LINNAEUS, C. (fil.) 1781. Supplementum plantarum. Impensis Or-

phanotrophei, Brunswick.

MEEUSE, A.D.J. 1956a. Ipomoea magnusiana var. eenii. The Flowering

Plants of Africa 31: t. 1201.

MEEUSE, A.D.J. 1956b. Ipomoea obscura var. fragilis. The Flowering

Plants of Africa 31: t. 1222.

MEEUSE, A.D.J. 1957. The South African Convolvulaceae. Bothalia 6:

641-792.

MERRILL, E.D. 1917. An interpretation of Rumphius 's Herbarium am-

boinense. Manila.

MYINT, T. & WARD, D.B. 1968. A taxonomic revision of the genus

Bonamia (Convolvulaceae). Phytologia 17: 121-239.

NICOLSON, D.H., SURESH, C.R. & MANILAL, K.S. 1988. An inter-

pretation of Van Rheede’s Hortus malabaricus. Regnum Vegeta-

bile 119.

RENDLE, A.B. 1901. Notes on African Convolvulaceae. Journal of

Botany, British and Foreign 39: 12-22.

ROSS, J.H. 1972. The flora of Natal. Memoirs of the Botanical Survey of

South Africa No. 39.

SCHINZ, H. 1888. Ipomoea magnusiana. Verhandlungen des botani-

schen Vereins der Provinz Brandenburg 30. Berlin.

SCHULZE-MENZ, G.K. 1938. Convolvulaceae in Neue und seltene

Arten aus Ostafrika leg. H.J. Schlieben XIII. Notizblatt des Botan-

ischen Gartens und Museums zu Berlin-Dahlem 14: 109-112.

THUNBERG, C.P. 1794. Prodromus plantarum capensium 1. Edman,

Uppsala.

VERDCOURT, B. 1957. Notes from the east African herbarium: V. Kew

Bulletin 2: 333-355.

VERDCOURT, B. 1961. New Convolvulaceae from the Flora zambe-

siaca area. Kirkia 1: 26-31.

50

Bothalia 26,1 (1996)

VERDCOURT. B. 1963a. Convolvulaceae. In C.E. Hubbard & E. Milne-

Redhead, Flora of tropical east Africa. Crown Agents, London.

VERDCOURT, B. 1963b. Tropical African plants: XXVII. Convolvu-

laceae. Kew Bulletin 17: 173.

VERDCOURT. B. 1967. New Convolvulaceae from the Flora zambe-

siaca area. II. Kirkia 6: 117-122.

VERDCOURT, B. 1978. Corrections and additions to the Flora of tropi-

cal east Africa: Convolvulaceae: IV. Kew Bulletin 33: 159-168.

VON JACQUIN. N.J. 1786-1793. leones plantarum rariorum 2. Vienna.

VON JACQUIN, N.J. 1789. Collectanea austriaca ad botanicum 2.

Vienna.

WELMAN, W.G. 1993. Convolvulaceae. In T.H. Arnold & B.C. de Wet,

Plants of southern Africa: names and distribution. Memoirs of the

Botanical Survey of South Africa No. 62.

WELWITSCH, F.M.J. 1859. Apontamentos phytogeographicos sobre a

flora da provincia de Angola. Imprensa nacional, Lisbon.

A.D.J. MEEUSE* AND W.G. WELMAN**

* Voorstraat 41, NL-1931 AH Egmond aan Zee, The Netherlands.

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

MS. received: 1995-08-30.

APIACEAE

A NEW SPECIES OF CENTELLA

The genus Centella is currently under revision. Field

work and a study of morphological and genetic variation

in the genus, (Schubert & Van Wyk in prep.) led to the

discovery of an up to now undescribed species, closely

related to C. macrocarpa.

Centella glauca Schubert & B.-E. van Wyk sp. nov.,

a Centella macrocarpa habitu multo robustiore, foliis

glaucis foliisque juvenilibus laminaribus non acicularibus

differt.

TYPE. — Western Cape, 3219 (Wuppertal): between

Matjies River and Dwars River on road between Ceres

and Algeria, Hugo 730 (STE holo.; PRE). See Hugo 729

(PRE, STE) which are from functionally male plants col-

lected at the same locality as the bisexual holotype of this

andromonoecious species.

Resprouting, densely virgate, robust, mostly dioecious

perennial (Figure 6), up to 700 mm tall. Leaves acicular,

somewhat succulent, (50-)80-90(-130) x 1-2 mm; dis-

tinctly glaucous, concolorous; glabrous; acicular; lamina

indistinguishable from petiole; main vein slightly sunken;

margin entire; apex acute (Figure 7D, E); coppice leaves

petiolate; petiole (30-)40-50(-100) mm long; lamina cun-

eate or narrowly elliptic, (7-) 15-25 x 2-3 mm (Figure

7A-C); base cuneate; margins sometimes inrolled, with

1-3 acute teeth on upper third of lamina; apex acute. In-

florescence mostly sparse, borne along the shoots; func-

tionally male inflorescence with 3-6 umbellules; rays

(5—) 1 0— 15(— 25 ) mm long; bracts 2, acute at apex,

glabrous; male flowers 3 on each umbellule, petals cream-

coloured, glabrous; functionally bisexual inflorescence with

single umbellule; ray (2-)15-20(-35) mm long; bracts 2,

acute at apex, 2-3 x 1 mm, shorter than fruit (Figure 7I-L).

Fruit very large, (4— )5(-7) x (3— )4(— 5) mm; widely elliptic

in commissural view, with 2 prominent, purple ribs visible

on either side of both carpels (Figure 71, J).

Judged by morphological similarities, the closest rela-

tive of C. glauca is C. macrocarpa , even though speci-

mens have mostly been identified as C. linifolia or C.

virgata in the past. Centella glauca is similar to C. macro-

carpa but differs in the much larger habit, the glaucous

leaves (hence the specific epithet) and the laminate, often

toothed, coppice leaves. The mature leaves of C. glauca

are acicular. Another distinct feature of C. glauca is that

the rays of the functionally male and female umbellules

may often be extremely long. The ribbed fruits are very

similar in shape and surface sculpturing (Figure 71, J, N)

to those of C. macrocarpa, but are usually much larger.

Centella glauca is furthermore geographically isolated

from C. macrocarpa and is restricted to the extreme north-

ern area of the Western Cape (Figure 8). It occurs on the

Witzenberg, northwards along the Grootwinterhoek

Mountains and also further north in the Cederberg near

Citrusdal and Clanwilliam.

This new species forms an interesting morphological

link between C. linifolia and the C. macrocarpa/virgata

Ott.Af.T? Schu-tve

FIGURE 6. — Holotype of Centella glauca, Hugo 730.

Bothalia 26,1 (1996)

51

FIGURE 7. — Centella glauca. A, adaxial view of tridentate coppice leaf; B, abaxial view of coppice leaf with one/two teeth; C, adaxial view of

coppice leaf with in-rolled margins; D, adaxial view of acicular mature leaf; E, abaxial view of acicular mature leaf; F, functionally male

umbellule with 3 flowers; G, male inflorescence with 3 umbellules; H, female inflorescence (infructescence) reduced to a single fruit; I,

mature fruit with bracts; J, fruit with bracts removed; K, abaxial view of bract; L, adaxial view of bract; M, habit; N, cross section of fruit.

Scale bars; A-E, G, H, J & L, M, 5 mm; F & I, 10 mm; N, 2 mm.

complex. The apparent isolation of C. linifolia from the

series Virgatae Adamson (1951) is therefore reduced. The

new species combines the glaucous leaf surface and dis-

tinct lamina of C. linifolia (at least in coppice shoots) with

the acicular leaves and virgate habit of C. macrocarpa.

The new species appears to be invariably resprouting,

whereas C. virgata and some populations of C. macro-

carpa are obligate reseeders. Further studies are in pro-

gress to examine relationships amongst these and related

species.

Material examined

WESTERN CAPE.— 3219 (Wuppertal): Heuning Vlei, (-AA),

Esterhuysen 7464 (BOL); Sneeuwkop, (-AC); Eselbank. (-AC). Taylor

FIGURE 8. — The known geographic

distribution of Centella glau-

ca, •; and Centella macro-

carpa, O.

52

Bothalia 26,1 (1996)

11357 (NBG); Cederberg, Matjies River, (-AD), Wagener 261 (NBG);

Wolfberg, (-AD), Esterhuysen 18099 (BOL); Truitjieskraal, (-AD),

Stilton & Zantovska 11496 (NBG); Zoo Ridge, (-AD), Taylor 6117

(STE); between Matjies River and Dwars River, (-CB), Hugo 729,

730 (PRE, STE); Kromme River, (-CB), Leighton 21610 (BOL);

Dasklip Pass, (-CC), Schubert & Van Wyk 101 (E, GRA, JRAU, KMG,

MO, NBG, PRE, S, WIND); mountains above Porterville, (-CC),

Barker 249 (BOL); Edwards 16149 (BOL); Esterhuysen 16116 (BOL,

NBG, PRE); Thompson 1459 (PRE, STE), 1460 (STE); Zeyher 727

(SAM). 3319 (Worcester): De Tronk, (-AA), Low 899 (STE); Visgat,

(-AA), Stokoe s.n. (SAM); Groothoek Peak, (-AA), Campbell 10362

(PRE, STE); Groot Winterhoek, (-AA), Phillips 1751 (SAM); Win-

terhoek, (-AA), Bolus 5057 (BOL); Witzenberg, (-AC), Andreae 187

(STE). Without precise locality: Marloth s.n. (PRE); Zeyher 727

(GRA, SAM), 742 (GRA).

REFERENCES

ADAMSON, R.S. 1951. Arevision of the subgenus Solandra of Centella.

Journal of South African Botany 17: 1—48.

SCHUBERT, M.T.R. & VAN WYK, B.-E. in prep. Morphological and

biochemical genetic evidence for hybridization in the genus Cen-

tella (Apiaceae), with notes on phylogenetic and taxonomic im-

plications. Bothalia.

M.T.R. SCHUBERT* and B-E. VAN WYK*

* Department of Botany, Rand Afrikaans University, P.O. Box 524,

Auckland Park 2006, Johannesburg.

MS. received: 1995-09-20.

Bothalia 26,1: 53-61 (1996)

Cytogenetic studies in some representatives of the subfamily Pooideae

(Poaceae) in South Africa. 1. The tribe Aveneae, subtribe Aveninae

J.J. SPIES*, S.K. SPIES*, S.M.C. VAN WYK*, A.F. MALAN* and E.J.L. LIEBENBERG**

Keywords: Aveneae, chromosomes, meiosis, Poaceae, polyploidy, Pooideae

ABSTRACT

This is a report on chromosome numbers for 14 species of the subtribe Aveninae, which is largely naturalized in South

Africa. This is the first chromosome number report for Helictotrichon longifolium (Nees) Schweick. (n = 14), H. longum (Stapf)

Schweick. (n = 14), H. namaquensis Schweick. (n = 14) and Lophochloa cristata (L.) Hyl. (n = 7, 21/2, 14). The subtribe has a

basic chromosome number of seven, and fewer ploidy levels occur in the naturalized species in South Africa than in the same

species in other parts of the world. All tetraploid specimens were alloploids.

INTRODUCTION

Polyploidy is one of the most important forms of

chromosomal evolution in plants (Stebbins 1971). Pre-

liminary studies on chromosome numbers of South Af-

rican grasses revealed over 80% polyploid specimens

(Moffett & Hurcombe 1949; De Wet 1954; De Wet &

Anderson 1956; Pienaar 1955; De Wet 1960; Davidse

et al. 1986; Spies & Du Plessis 1986a & b, 1987a &

b; Spies & Jonker 1987; Du Plessis & Spies 1988; Spies

& Du Plessis 1988; Spies & Voges 1988; Spies et al.

1989, 1990, 1991, 1992; Du Plessis & Spies 1992; Stry-

dom & Spies 1994; Visser & Spies 1994a-c). The pre-

sent article corroborates these findings.

The Pooideae is, for the largest part, introduced to

southern Africa. The majority of species belonging to

this subfamily is restricted to either the winter rainfall

area of South Africa or to high altitude areas. The tribe

Aveneae Dumort. comprises 57 genera and approximately

1 050 species in the world (Clayton & Renvoize 1986).

Most South African representatives (18 genera and 59

species) are naturalized (Gibbs Russell et al. 1990). Clay-

ton & Renvoize (1986) subdivided the tribe into four

subtribes, i.e. Duthieinae Potztal, Aveninae Presl, Pha-

laridinae Rchb. and Alopecurinae Dumort. The genera

representing these subtribes are listed in Table 1.

The aim of this study is to determine the chromo-

some numbers, polyploid levels and meiotic chromo-

some behaviour of the South African representatives of

the tribe Aveninae. These results will eventually be com-

pared with results of indigenous and endemic taxa to

compare the frequency of polyploidy between indige-

nous and introduced grasses.

MATERIALS AND METHODS

Cytogenetic material was collected in two different ways

for the purpose of this study. The material was either

* Department of Botany and Genetics, University of the Orange Free

State, RO. Box 339, Bloemfontein 9300.

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

MS. received: 1995-10-25.

collected and fixed in the field, or living material was

collected in the field and transplanted in the nurseries of

either the National Botanical Institute (Pretoria) or

the Department of Botany and Genetics, University of

the Orange Free State (Bloemfontein), where cytogenetic

material was collected and fixed. The material used and

their localities are listed in Table 2. Voucher specimens

are housed in the Geo Potts Herbarium, Department of

Botany and Genetics, University of the Orange Free

State, Bloemfontein (BFFU) or the National Herbarium,

Pretoria (PRE).

Young inflorescences were fixed in Camoy’s fixative

(Camoy 1886). The fixative was replaced by 70% ethanol

after 24-48 hours of fixation. Anthers were squashed in

2% aceto-carmine (Darlington & LaCour 1976). Slides

were made permanent by freezing them with liquid CCU

(Bowen 1956), followed by dehydration in ethanol and

mounting in Euparal. An Olympus Vanox-S or Nikon Mi-

crophot photomicroscope and Ilford Pan-F film (ASA 50)

were used for the photomicrographs. At least ten cells

TABLE 1. — List of subdivisions of the tribe Aveneae, indicating the

genera and number of species present in southern Africa

54

Bothalia 26,1 (1996)

TABLE 2. — Haploid chromosome numbers of representatives of the subtribe Aveninae (Poaceae, Pooideae, Aveneae) in southern Africa with the

voucher specimen numbers and their specific localities, arranged according to the system of Edwards & Leistner (1971)

Bothalia 26,1 (1996)

55

TABLE 2— Haploid chromosome numbers of representatives of the subtribe Aveninae (Poaceae, Pooideae, Aveneae) in southern Africa with the

voucher specimen numbers and their specific localities (arranged according to the system of Edwards & Leistner (1971) (continued)

per specimen were studied for each meiotic stage, ex-

cept where otherwise indicated.

Meiotic chromosome counts are given as haploid (n)

numbers to conform to the style set out by the editors of

the Index to plant chromosome numbers series, published

by the Missouri Botanical Garden.

RESULTS

All studied specimens of Aira L. were diploid (Table

2) and all meiotic cells appear normal. Seven bivalents

were present during diakinesis (Figure 1A).

Two different ploidy levels, based on seven, were pre-

sent in the Avena L. specimens studied. The species cor-

FIGURE 1. — Photomicrographs of

meiotic chromosomes in some

representatives of the genera

Aira , Avena and Corynopho-

rus. A, Aira cupaniana, Spies

4597, early metaphase I with

7n; B, Avena barbata. Spies

5287, early metaphase I with

14n; C, A. byzanthina. Spies

2481, diplotene with 21ii; D,

A. fatua, Spies 4899, diakine-

sis with 21n. E; A. fatua, Spies

4915 , anaphase I with chroma-

tid bridges; F, Corynephorus

fasciculatus, Spies 3690, diaki-

nesis with 7n. Scale bar: A & F,

35/<m: B. D & E, 30 urn: C, 28

pm.

56

Bothalia 26,1 (1996)

responded morphologically and separation of species was

sometimes doubtful. According to the classification we

received for our specimens, A. barbata Pott ex Link is

tetraploid (Figure IB), whereas A. byzanthina K.Koch

(Figure 1C) and A. fatua L. (Figure ID; 2A) are hexa-

ploid. The majority of bivalents were ring bivalents and

no multivalents were present. Up to four chromatid

bridges per cell have been observed in the three Avena

species mentioned in Spies 2481, 4808, 4899 and

4915) (Figure IE; 2). The frequent occurrence of this

phenomenon suggests that this specimen is heterozygotic

for a high number of paracentric inversions.

The only Corynephorus fasciculatus Boiss. & Reut.

specimen studied, proved to be a diploid (Figure IF), with

normal meiosis.

Haploid chromosome numbers in the genus Helic-

totrichon ranged from diploid to octaploid (Figure 3A-E).

A similar range was observed in H. turgidulum. The

FIGURE 3. — Photomicrographs of meiotic chromosomes in some representatives of Helictotrichon turgidulum. A, Spies 4721, diakinesis with 7ii,

B, Spies 4763, diakinesis with 14n; C, Spies 4678, diakinesis with 14n; D, Spies 2355, diakinesis with 14n; E, Spies 4775, diakinesis with

21n; F, Spies 4507, anaphase with 16-16 segregation of chromosomes, thus indicating the presence of B-chromosomes; G, Spies 3137, anaphase

I with a chromatid bridge and a chromosome bridge; H, Spies 3137, telophase I with a micronucleus. Scale bar: 30 p.m.

Bothalia 26,1 (1996)

57

FIGURE 4. — Photomicrographs of

meiotic chromosomes in some

representatives of the genera

Holcus and Koeleria. A, Hol-

cus lanatus, Saayman 124,

diakinesiswith 7ii; B, Koeleria

capensis, Spies 5119, diakine-

sis with 7n; C, K. capensis,

Spies 5094, diakinesis with 7ii;

D, K. capensis, Spies 5119,

metaphase I with 7n; E, K. cap-

ensis, Spies 4855, diakinesis

with 14ii; F ,K. capensis, Spies

5111, telophase II with two

micronuclei. Scale bar: A-D,

30 pm; E & F, 27 pm.

other species were all tetraploid. All the tetraploid speci-

mens were alloploids, with the observed chromosome

configurations concurring best with the expected con-

figurations for the 2:2 model of Kimber & Alonso

(1981). All specimens had x-values of 1. The 2:2

model indicates the presence of two sets of genomes,

both consisting of two genomes. The x-value can

vary from 0.5 (relative distance between the sets of

genomes equals the relative distance between the

genomes within a set) to 1 (relative distance between

sets much larger than the distance within a set). The

x-value of one, therefore, indicates that the speci-

mens are alloploid. Two specimens had B-chromo-

somes (0-4B) (Figure 3F). Chromosomes were regarded

as B-chromosomes if additional chromosomes were

observed in some cells of an individual, or if the be-

haviour of the additional chromosomes deviated from

the expected behaviour of euchromosomes. Occasion-

ally meiotic abnormalities of euchromosomes during

anaphase I have been observed. These abnormalities

included chromatid bridges (Figure 3G), anaphase

laggards and micronuclei during telophase I (Figure

3H).

Holcus lanatus was diploid (Figure 4A), with regular

chromosome behaviour. Koeleria capensis encom-

passed both diploid and tetraploid specimens (Figure

4B-E). Occasionally chromosome laggards during ana-

phase I and micronuclei during telophases I and/or II

(Figure 4F) have been observed. Lophochloa pumila

was diploid (Figure 5A), whereas L. cristata varied

from diploid to tetraploid (Figure 5B-G). One L. cristata

specimen was either triploid or it contained up to seven

B-chromosomes (Figure 5F, G; 6).

DISCUSSION

Aira is naturalized in South Africa and two species are

recognized in this country, i.e. A. caryophyllea and A. cu-

paniana (Gibbs Russell et al. 1990). Both species are dip-

loid, n = x = 7 (Table 2), with normal meiosis. This de-

viates from the somatic chromosome number of 28 usu-

ally reported for A. caryophyllea (Albers & Albers 1973;

Queiros 1974; Albers 1978; 1980; Kirschner et al. 1982).

However, a diploid specimen has been reported by

Romero Zarco (1988). This study confirms the chromo-

58

Bothalia 26,1 (1996)

FIGURE 5. — Photomicrographs of

meiotic chromosomes in some

representatives of the genus

Lophochloa. A, Lophochloa

pumila, Davidse 33272 , diaki-

nesis with 7ii; B, L. cristata,

Spies 4965, metaphase I with

7n7i; C, L. cristata, Spies

4567, diakinesis with 14n; D,

L. cristata, Spies 4965, meta-

phase I; E, L. cristata, Spies

4965, early anaphase I; F, L.

cristata. Spies 4965, late ana-

phase I with 4 laggards; G, L.

cristata. Spies 4965, late ana-

phase I with laggards. Scale

bar: A, C-G, 30 pm\ B, 21 pm.

some numbers previously described for A. cupaniana (Al-

bers & Albers 1973; Albers 1980). Although polyploidy

is frequently observed in Europe, it seems absent in the

naturalized species.

Arrhenatherum P.Beauv. is represented by one natural-

ized species in South Africa, A. elatius (L.) Presl (Gibbs

Russell et al. 1990). No specimen could be obtained for

this study. However, published results indicate that diploid

(n = 7 or 2n = 14), tetraploid (n = 14 or 2n = 28) and

hexaploid (n = 28) specimens of A. elatius have been ob-

served elsewhere (Goldblatt 1981, 1983, 1985, 1988;

Goldblatt & Johnson 1990, 1991).

Avena consists of four naturalized and a cultivated spe-

cies in South Africa (Gibbs Russell et al. 1990). All A.

barbata specimens studied, are tetraploid. This supports

the different ploidy levels, ranging from diploid to hexa-

ploid, previously described (Goldblatt 1981, 1983, 1985,

1988; Goldblatt & Johnson 1990, 1991). These results

may indicate that a single introduction, or several introduc-

tions from the same ploidy level, of A. barbata , occurred.

Avena byzantina is hexaploid, thus supporting the chro-

mosome number previously described (Morikawa 1982).

The high frequency of ring bivalents and the absence of

any multivalent suggests an alloploid origin for both spe-

cies. This alloploid origin is confirmed by the correspond-

ing values obtained during this study and when the

expected chromosome associations for the 2:2 model of

Kimber & Alonso (1981) is determined. The x-value of

1 indicates no relationship between the chromosomes of

the two different genomes.

The hexaploid chromosome number determined for A.

fatua supports the published chromosome number of this

species (Goldblatt 1981, 1983, 1985, 1988; Goldblatt &

Johnson 1990, 1991). The absence of multivalents in the

studied specimens, suggests an alloploid origin for this spe-

cies.

Bothalia 26,1 (1996)

59

The naturalized Corynophoris fasciculatus is the

only representative of this genus in South Africa (Gibbs

Russell et al. 1990). Our study confirmed the diploid

chromosome number previously described for this spe-

cies (Goldblatt 1981, 1983, 1985, 1988; Goldblatt &

Johnson 1990, 1991).

Deschampsia P.Beauv. is represented by two natural-

ized species in South Africa, D. caespitosa (L.) P.Beauv.

and D. flexuosa (L.) Trin. (Gibbs Russell et al. 1990). In

Europe D. caespitosa has haploid chromosome numbers

of 9, 12, 13 and 26 (Goldblatt 1981, 1983, 1985, 1988;

Goldblatt & Johnson 1990, 1991), whereas D. flexuosa

has haploid chromosome numbers of 7 and 14 (Goldblatt

1981, 1983, 1985, 1988; Goldblatt & Johnson 1990,

1991). It would be very interesting to study the South

African species cytogenetically to determine whether we

have both basic chromosome numbers of 7 and 13 present,

as suggested by chromosome numbers of other members

of this genus.

Helictotrichon is represented by 13 indigenous spe-

cies in South Africa (Gibbs Russell et al. 1990). Four

species were included in this study, H. longifolium,

H. longum , H. namaquensis and H. turgidulum. This

is, to the best of our knowledge, the first report of

chromosome numbers for H. longifolium, H. longum

and H. namaquensis , which are all tetraploid. Addi-

tionally to the tetraploid chromosome number de-

scribed for H. turgidulum (Hoshino & Davidse 1988),

diploid, hexaploid and octaploid specimens were stud-

ied. All tetraploid specimens (from the four species)

conform with the expected chromosome configura-

tions for the 2:2 model of Kimber & Alonso (1981).

The x-values of 1 indicate alloploid origins for all

these specimens.

Holcus is represented by one indigenous species, H.

setiger Nees, and a naturalized species, H. lanatus (Gibbs

Russell et al. 1990). Only the latter species has been stud-

ied and the species seems to be diploid in South Africa,

in contrast to the diploid and tetraploid species described

in other countries (Goldblatt 1981, 1983, 1985, 1988; Gold-

blatt & Johnson 1990, 1991).

Koeleria capensis is indigenous and the only repre-

sentative of the genus Koeleria in South Africa (Gibbs

Russell et al. 1990). This study revealed two ploidy

levels for this species, diploid and tetraploid, thus con-

firming previous results (Goldblatt 1981, 1983, 1985,

1988; Goldblatt & Johnson 1990, 1991). The tetraploid

specimens were restricted to the southern Cape, near

Bredasdorp. Their chromosome configurations con-

curred with the expected configurations for the 2:2

model with an x-value of 1. These specimens are con-

sequently alloploids.

Lophochloa is represented by two naturalized species

in South Africa, L. cristata and L. pumila (Desf.) Bor

(Gibbs Russell et al. 1990). This is, to the best of our

knowledge, the first report on chromosome numbers for

L. cristata , which has at least two different ploidy levels,

diploid and tetraploid. One specimen (Spies 4965 ) with

abnormal chromosomal behaviour was observed. This

specimen is either a triploid or it contains up to seven

B-chromosomes. The additional chromosomes do not dif-

fer morphologically from the ‘normal’ chromosomes.

However, since the number of additional chromosomes

seems to differ from one cell to another (Figure 6), we

suggest that they are B-chromosomes. The chromosome

configurations harmonized best with the expected values

obtained from the 2:2 model of Kimber & Alonso (1981).

An x-value of 1 indicated that the tetraploid specimens

FIGURE 6. — Photomicrographs of meiotic chromosomes in Lophochloa cristata. Spies 4965. A-R, various cells indicating the difficulty to determine

whether the additional chromosomes are B-chromosomes or whether they represent a third genome. Scale bar: A-R, 10 pm.

60

Bothalia 26,1 (1996)

are alloploids. This study confirms the diploid status al-

ready described for L. pumila (Goldblatt 1981, 1983, 1985,

1988; Goldblatt & Johnson 1990, 1991). However, we found

no confirmation for a basic chromosome number of 13 for

Lophochloa as often reported (Goldblatt 1981, 1983, 1985,

1988; Goldblatt & Johnson 1990, 1991).

The genus Periballia Trin. comprises a single natural-

ized species in South Africa, P. minuta (L.) Asch. &

Graebn. (Gibbs Russell et al. 1990). Reports indicate that

this species has a haploid chromosome number of 4

(Goldblatt 1981, 1983, 1985 1988; Goldblatt & Johnson

1990, 1991).

The subtribe Aveninae is largely naturalized in South

Africa. The exceptions are the genus Helictotrichon and

the species Holcus setiger and Koeleria capensis. The

subtribe has a basic chromosome number of seven, and

less ploidy levels occur in the naturalized species in

South Africa in comparison to the same species in other

parts of the world. This may be attributed to the intro-

duction of these species. The species deviating from the

basic chromosome number of the subtribe need to be

investigated thoroughly.

ACKNOWLEDGEMENTS

Dr Gerrit Davidse (Missouri Botanical Garden, St

Louis, Missouri, USA) is thanked for some material

used during this study. Financial support from the Foun-

dation for Research Development and the University of

the Orange Free State, is gratefully acknowledged.

REFERENCES

ALBERS, F. 1978. Karyologische und genomatische Veranderungen in-

nerhalb der Graser-Subtriben Aristaveninae und Airinae. Berichte

der Deutschen Botanischen Gesellschaft 91: 693-697.

ALBERS, F. 1980. Comparative karyology of the grass subtribes Aristaven-

inae and Airinae (Poaceae-Avenenae). Plant Systematics and Evo-

lution 136: 137-167.

ALBERS, F. & ALBERS, I. 1973. Diploide und tetraploide Sippen von

Aira (Poaceae) im Massif des Maures (Provence). Osterreichi-

sche Botanische Zeitschrift 122: 293-298.

BOWEN, C.C. 1956. Freezing by liquid carbon dioxide in making slides

permanent. Stain Technology 31: 87-90.

CARNOY, J.B. 1886. La cytodiere se de l’oeuf. Cellule 3: 1-92.

CLAYTON, W.D. & RENVOIZE, S.A. 1986. Genera graminum — grasses

of the world. Kew Bulletin Additional Series 13: 1-389.

DARLINGTON, C.D. & LACOUR, L.F. 1976. The handling of chromo-

somes. Allen & Unwin, London.

DAVIDSE, G„ HOSHINO, T. & SIMON, B.K. 1986. Chromosome

counts of Zimbabwean grasses and an analysis of polyploidy in

the grass flora of Zimbabwe. South African Journal of Botany 52:

521-528.

DE WET, J.M.J. 1954. Chromosome numbers of a few South African

grasses. Cytologia 19: 97-103.

DE WET, J.M.J. 1960. Chromosome numbers and some morphological

attributes of various South African grasses. American Journal of

Botany 47: 44-49.

DE WET, J.M.J. & ANDERSON, L.J. 1956. Chromosome numbers in

Transvaal grasses. Cytologia 21: 1-10.

DU PLESSIS, H. & SPIES, J.J., 1988. Chromosome studies on African

plants. 8. Bothalia 18: 119-122.

DU PLESSIS, H. & SPIES, J.J. 1992. Chromosome numbers in the genus

Pentaschistis (Poaceae, Danthonieae). Taxon 41: 706-720.

EDWARDS, D. & LEISTNER, O.A. 1971. Adegree reference system for

citing biological records in southern Africa. Mitteilungen der

Botanischen Staatssammlung, Miinchen 10: 501-509.

GIBBS RUSSELL, G.E., WATSON, M., KOEKEMOER, M„ SMOOK,

L„ BARKER, N.P., ANDERSON, H.M. & DALLW1TZ, M.J.

1990. Grasses of southern Africa. Memoirs of the Botanical Sur-

vey of South Africa No. 58.

GOLDBLATT, P. 1981. Index to plant chromosome numbers 1975-1978.

Monograms of Systematic Botany 5.

GOLDBLATT, P. 1983. Index to plant chromosome numbers 1979-1981.

Monograms of Systematic Botany 8.

GOLDBLATT, P. 1985. Index to plant chromosome numbers 1982-1983.

Monograms of Systematic Botany 13.

GOLDBLATT, P. 1988. Index to plant chromosome numbers 1984—1985.

Monograms of Systematic Botany 23.

GOLDBLATT, P. & JOHNSON, D.E. 1990. Index to plant chromosome

numbers 1986-1987. Monograms of Systematic Botany 30.

GOLDBLATT, P. & JOHNSON, D.E. 1991. Index to plant chromosome

numbers 1988-1989. Monograms of Systematic Botany 40.

HOSHINO, T. & DAVIDSE, G. 1988. Chromosome numbers of grasses

(Poaceae) from southern Africa. Annals of the Missouri Botanical

Garden 75: 866-873.

KIMBER, G. & ALONSO, L.C. 1981. The analysis of meiosis in hybrids.

III. Tetraploid hybrids. Canadian Journal of Genetics and Cytol-

ogy 23: 235-254.

KIRSCHNER, J„ STEPANEK, J. & STEPANKOVA, J., 1982. In IOPB

chromosome number reports LXXVL Taxon 31: 574-575.

MOFFETT, A.A. & HURCOMBE, R. 1949. Chromosome numbers in

South African grasses. Heredity 3: 369-373.

MOORE, R.J. 1970. Index to plant chromosome numbers. Regnum Vege-

tabile 68.

MOORE, R.J. 1971. Index to plant chromosome numbers. Regnum Vege-

tabile 77.

MOORE, R.J. 1972. Index to plant chromosome numbers. Regnum Vege-

tabile 84.

MOORE, R.J. 1974. Index to plant chromosome numbers. Regnum Vege-

table 91.

MOORE, R.J. 1977. Index to plant chromosome numbers. Regnum Vege-

tabile 96.

MORIKAWA, T. 1982. Characterization of monosomes in hexaploid

oats using double monosomies. Japanese Journal of Genetics

57: 269-279.

ORNDUFF, R. 1967. Index to plant chromosome numbers. Regnum

Vegetabile 50.

ORNDUFF, R. 1968. Index to plant chromosome numbers. Regnum

Vegetabile 55.

ORNDUFF, R- 1969. Index to plant chromosome numbers. Regnum

Vegetabile 59.

PIENAAR, R.D.V. 1955. The chromosome numbers of some indigenous

South African and introduced Gramineae. In D. Meredith, Grasses

and pastures of South Africa. Central News Agency, Johannes-

burg.

QUEIROS, M., 1974. Contribuicao para o conhecimento citotaxonomico

das Spermatophyta de Portugal. I. Gramineae. Boletim de la

Sociedad de Botanica 48: 81-98.

ROMERO ZARCO, C. 1988. Numeros chromosomaticos de plantas occi-

dentales, 472-486. Anales del jardin botanico de Madrid 45 : 273.

SPIES, J.J., DAVIDSE, G. & DU PLESSIS, H. 1992. Cytogenetic studies

in the genus Tribolium (Poaceae: Arundineae). A meric an Journal

of Botany 79: 689-700.

SPIES, J.J. & DU PLESSIS, H. 1986a. Chromosome studies on African

plants. 1. Bothalia 16: 87, 88.

SPIES, J.J. & DU PLESSIS, H. 1986b. Chromosome studies on African

plants. 2. Bothalia 16: 269, 270.

SPIES, J.J. & DU PLESSIS, H. 1987a. Chromosome studies on African

plants. 3. Bothalia 17: 131-135.

SPIES, J.J. & DU PLESSIS, H. 1987b. Chromosome studies on African

plants. 5. Bothalia 17: 257 -259.

SPIES, J.J. & DU PLESSIS, H. 1988. Chromosome studies on African

plants. 6. Bothalia 18: 111-114.

SPIES, J.J., DU PLESSIS, H„ BARKER, N.P. & VAN WYK, S.M.C.

1990. Cytogenetic studies in the genus Chaetobromus (Poaceae:

Arundineae). Genome 33: 646-658.

SPIES, J.J. & JONKER, A. 1987. Chromosome studies on African plants.

4. Bothalia 17: 135, 136.

SPIES, J.J., SAAYMAN, E.J.L., VOGES, S.P. & DAVIDSE, G. 1989.

Chromosome studies on African plants. 9. Chromosome num-

bers in Ehrharta (Poaceae: Ehrharteae). Bothalia 19: 125-132.

SPIES, J.J. & VOGES, S.P. 1988. Chromosome studies on African plants.

1. Bothalia 18: 114-119.

Bothalia 26,1 (1996)

61

SPIES, J.J., VAN DER MERWE, E„ DU PLESSIS, H. & SAAYMAN,

E.J.L. 1991. Basic chromosome numbers and polyploid levels in

some South African and Australian grasses (Poaceae). Bothalia

21: 163-170.

STEBBINS, G.L. 1971. Chromosomal evolution in higher plants. Ed-

ward Arnold, London.

STRYDOM, A. & SPIES, J.J. 1994. A cytotaxonomic study of some

representatives of the tribe Cynodonteae (Chloridoideae, Poaceae).

Bothalia 24: 92-96.

VISSER, N.C. & SPIES, J.J. 1994a. A cytotaxonomic study of the genus

Tribolium (Poaceae: Danthonieae). III. Section Tribolium. South

African Journal of Botany 60: 31-39.

VISSER, N.C. & SPIES, J.J. 1994b. A cytotaxonomic study of the genus

Tribolium (Poaceae: Danthonieae). IV. Section Uniolae. South

African Journal of Botany 60: 279 -284.

VISSER, N.C. & SPIES, J.J. 1994c. A cytotaxonomic study of the genus

Tribolium (Poaceae: Danthonieae). V. Section Acutiflorae, re-

lated genera, and conclusions. South African Journal of Botany

60: 285-292.

Bothalia 26,1: 63-67(1996)

Cytogenetic studies in some representatives of the subfamily Pooideae

(Poaceae) in South Africa. 2. The tribe Aveneae, subtribes Phalaridinae and

Alopecurinae

JJ. SPIES*, S.K. SPIES*, S.M.C. VAN WYK*, A.F. MALAN*f and E.J.L. LIEBENBERG**

Keywords: Aveneae, chromosomes, meiosis, Poaceae, polyploidy, Pooideae

ABSTRACT

This is a report on chromosome numbers for the subtribes Phalaridinae and Alopecurinae (tribe Aveneae) which are, to a

large extent, naturalized in South Africa. Chromosome numbers of 34 specimens, representing nine species and four genera, are

presented. These numbers include the first report on Agrostis avenacea Gmel. (n = 4x = 28). New ploidy levels are reported for

Phalaris aquatica L. (n = x = 1), Agrostis barbuligera Stapf var. barbuligera (n = 2x = 14 and n = 4x = 28) and A. lachnantha

Nees var. lachnantha (n = 3x = 21).

INTRODUCTION

The first paper in this series on chromosome num-

bers of representatives of the tribe Aveneae in South Af-

rica, indicated the importance of determining the ploidy

levels and basic chromosome numbers of naturalized and

endemic flora in South Africa (Spies et al. 1996). This

second paper in the series is restricted to the subtribes

Phalaridinae and Alopecurinae.

The subtribe Phalaridinae Rchb. consists of three gen-

era (Clayton & Renvoize 1986), of which two are repre-

sented in South Africa, i.e. Anthoxanthum L. and Phalaris

L. (Gibbs Russell et al. 1990). The subtribe Alopecurinae

Dumort. consists of 27 genera (Clayton & Renvoize

1986), with only six being represented in South Africa

(Gibbs Russell et al. 1990), i.e. Agrostis L., Ammophila

Host, Calamagrostis Adans., Gastridium P.Beauv., Lagu-

rus L. and Polypogon Desf. (Gibbs Russell et al. 1990).

The aim of this study was to determine the chromo-

some numbers, polyploid levels and meiotic chromosome

behaviour of the South African representatives of the tribe

Aveninae. These results will eventually be compared with

results of indigenous and endemic taxa to compare the

frequency of polyploidy between indigenous and intro-

duced grasses.

MATERIALS AND METHODS

The material was either collected and fixed in the field,

or living material was collected in the field and planted

in the nurseries of either the Department of Botany and

Genetics, University of the Orange Free State (Bloemfon-

tein), or the National Botanical Institute (Pretoria), where

the cytogenetic material was collected and fixed. The ma-

*Department of Botany and Genetics, University of the Orange Free

State, RO. Box 339, Bloemfontein 9300.

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

tCurrent address: Small Grain Centre, Private Bag X29, Bethlehem 9600.

MS. received: 1995-11-09.

terial used and the collecting localities are listed in Table

1. Voucher specimens are housed in the Geo Potts Her-

barium, Department of Botany and Genetics, University

of the Orange Free State, Bloemfontein (BFFU) or the

National Herbarium, Pretoria (PRE).

Anther were squashed in aceto-carmine and meioti-

cally analysed (Spies et al. 1996). Chromosome numbers

are presented as haploid chromosome numbers to conform

to previous papers on chromosome numbers in this journal

(Spies & Du Plessis 1986). These numbers were compared

with results published in the chromosome atlasses of Fe-

dorov (1969), Ornduff (1967-1969), Moore (1970-1972,

1974, 1977), Goldblatt (1981, 1983, 1985, 1988) and

Goldblatt & Johnson (1990, 1991, 1994). Genome homol-

ogy was determined in some tetraploid specimens accord-

ing to the models described by Kimber & Alonso (1981).

RESULTS AND DISCUSSION

The subtribe Phalaridinae Rchb. consists of three gen-

era, of which two are represented in South Africa (Clayton

& Renvoize 1986). The genus Anthoxanthum consists of

18 species, four of which are endemic to South Africa [A.

brevifolium Stapf, A. dregeanum (Nees) Stapf, A. ecklonii

(Nees ex Trin.) Stapf & A. tongo (Trin.) Stapf] and a fifth

species, A. odoratum F., which is naturalized (Gibbs

Russell et al. 1990). The genus Phalaris L. comprises 15

species, of which six are naturalized in this country [F?

angusta Nees ex Trin., P. aquatica L., P. arundinacea L.,

P canariensis F., P minor Retz. and P paradoxa L.]

(Gibbs Russell et al. 1990). No Phalaris species is indige-

nous to South Africa (Gibbs Russell et al. 1990).

Apparently only one specimen of the genus Anthoxan-

thum has sofar been cytogenetically studied and reported

in South Africa, namely of A. tongo (Trin.) Stapf for which

a haploid chromosome number of 20 + 0-5B has been

established by one of our laboratories (Spies & Voges

1988). Reports of haploid chromosome numbers of n = 5

and 10 from other parts of the world are frequent but

numbers such as n = 20, 35 or 45 are rarely reported

64

Bothalia 26,1 (1996)

TABLE 1. — Haploid chromosome numbers of representatives of the subtribes Phalaridinae, Alopecurinae (Poaceae, Pooideae, Aveneae) in south-

ern Africa, with their voucher specimen numbers and specific localities [arranged according to the system of Edwards & Leistner (1971)]

Bothalia 26,1 (1996)

65

(Goldblatt 1981, 1983, 1985, 1988; Goldblatt & Johnson

1990, 1991, 1994). In addition to these numbers based on

x = 5, three reports of n = 28 for A. borii, therefore x =

7, have been published (Goldblatt 1981, 1985, 1988). With

only three reports suggesting a basic chromosome number

of seven rather than the five indicated in 55 other reports,

we suggest that the basic chromosome number of the ge-

nus Anthoxanthum is five, as our research indicates. How-

ever, a re-investigation of this genus will help to clarify

the position of A. borii in the genus.

Three species of the genus Phalaris were included in

this study (Table 1). The only specimen of P aquatica

investigated, had both diploid (n = x = 7) and tetraploid

(n = 2x = 14) chromosome numbers (Figure 1A, B). We

attribute this apparent discrepancy to cell fusion (Spies &

Van Wyk 1995). This process involves the formation of

polynucleate microspores after cell fusion and the conse-

quent formation of polyhaploid pollen. This phenomenon

is frequently observed in grasses and was discussed by

Spies & Van Wyk (1995). Phalaris aquatica was repre-

sented by a diploid specimen (n = x = 7) in this study.

This is a lower chromosome number than the tetraploid

one previously described (Moore 1971; Goldblatt 1983,

1988). Meiosis was normal at both ploidy levels with only

bivalents observed.

One specimen of P minor was diploid (Figure 1C, D)

and the rest (10 specimens) tetraploid (Figure 1F-H). With

the exception of an anaphase I bridge in one cell of one

specimen, meiosis was normal and only bivalents were

observed in the P motor specimens. Phalaris minor speci-

mens proved to be either diploid (n = x = 7) or tetraploid

(n = 2x = 14) in this study, confirming the tetraploid level

previously described (Moore 1971; Goldblatt 1983, 1985;

Goldblatt & Johnson 1990). In a previous study by one

of our laboratories a hexaploid specimen (n = 3x = 21)

was observed (Spies & Voges 1988). In addition to these

ploidy levels octoploid specimens (n = 4x = 28) have also

been described in the literature (Moore 1977). The pres-

ence of diploid and possibly allotetraploid specimens in

the same species, should render a morphological study of

this species worthwhile.

The only P paradoxa specimen studied proved to be

a diploid (n = x = 7) with normal meiosis, thus supporting

previous reports on this species (Moore 1971, 1977; Gold-

blatt 1981, 1983). A few species of Phalaris with a basic

chromosome number of six have been reported (Fedorov

1969; Goldblatt 1981, 1983, 1985, 1988; Goldblatt &

Johnson 1994). However, in South Africa all species stud-

ied had a basic chromosome number of seven.

The subtribe Alopecurinae Dumort. consists of 27 gen-

era (Clayton & Renvoize 1986), with only six being rep-

resented in South Africa (Gibbs Russell et al. 1990).

Agrostis L. consists of 220 species worldwide (Clayton &

Renvoize 1986), eight of which are indigenous to South

Africa [A. barbuligera Stapf, A. bergiana Trin., A. con-

tinuata Stapf, A. eriantha Hack., A. lachnantha Nees, A.

polypogonoides Stapf, A. schlechteri Rendle and A. subu-

lifolia Stapf] and three species are naturalized [A.

avenacea Gmel., A. gigantea Roth and A. montevidensis

Spreng. ex Nees] (Gibbs Russell et al. 1990). Twelve

specimens, representing three species of the genus

Agrostis, were included in this study (Table 1). The three

A. avenacea specimens were octoploid (n = 4x = 28) (Fig-

ure 2A, B). This seems to be the first chromosome number

report for this species. Two A. barbuligera var. barbulig-

era specimens were tetraploid (Figure 2D) and one octo-

ploid. Both ploidy levels differ from the previously

described hexaploid (n = 3x = 21) level from one of our

laboratories (Spies & Du Plessis 1986). Four hexaploid

specimens were observed in A. lachnantha var. lachnan-

tha, as well as two octoploid specimens (Figure 2E, F).

The octoploid specimens conform with the number pre-

FIGURE 1. — Photomicrographs of meiotic chromosomes in the genus Phalaris. A, B, P. aquatica. Spies 3676, diplotene/diakinesis with 7ii and 14n

respectively; C, P. minor, Davidse 33272, diakinesis with 7n; D, P. paradoxa, Spies 5395, anaphase I with 7 chromosomes in each pole; E,

P. minor, Spies 4593, late anaphase I with a chromatid bridge; F, P. minor. Spies 4284, diakinesis with 14ii; G, P. minor, Spies 4978, metaphase

I with 14ii; H, P. minor, Davidse 33237, metaphase I with 14n. Scale bar: 10 /<m.

66

Bothalia 26,1 (1996)

FIGURE 2. — Photomicrographs of

meiotic chromosomes in the ge-

nus Agrostis. A, A. avenacea ,

Saayman 78, early metaphase I

with 28n; B, A. avenacea. Spies

3522, diakinesis with 28u; C, D,

A. barbuligera subsp. barbuli-

gera , Saayman 57; C, telophase

I cells with micronuclei; D, dia-

kinesis with 14n; E, F, A. lach-

nantha subsp. lachnantha; E,

Saayman 89, metaphase I with

28ii; F, Saayman 89, diakinesis

with 28n. Scale bar: 10 pm.

viously described by us (Spies & Du Plessis 1986). Meio-

sis was normal in almost all specimens, excepting one

telophase II cell of A. barbuligera, where micronuclei

were observed (Figure 2C).

Ammophila Host is a north temperate genus with two

species, with A. arenaria (L.) Link naturalized in South

Africa (Gibbs Russell et al. 1990). Although this species

was not represented in this study, all reports indicate that

representatives from the rest of the world are tetraploid

2n = 4x = 28 (Ornduff 1967; Moore 1970, 1972, 1977;

Goldblatt 1983; Goldblatt & Johnson 1994).

Calamagrostis Adans. is a very large genus with ± 270

species (Clayton & Renvoize 1986) with only one species

represented in South Africa, C. epigeios (L.) Roth (Gibbs

Russell et al. 1990). Chromosome number reports indicate

the presence of tetraploid to octoploid specimens for this

species (Moore 1977; Goldblatt 1981, 1983, 1985, 1988;

Goldblatt & Johnson 1990, 1994).

Gastridium phleoides (Nees & Meyen) C.E.Hubb. is

the only species of Gastridium P.Beauv. introduced to

South Africa (Gibbs Russell et al. 1990). The only chro-

mosome number report found for this species, indicates

2n = 4x = 28 (Goldblatt 1981).

The monotypic genus Lagurus L. was also introduced

to South Africa (Gibbs Russell et al. 1990). All L. ovatus

specimens studied were diploid (Figure 3) with normal

meiosis in most cells, excepting a bivalent lying away

from the metaphase plate in one cell (Figure 3D) and a

laggard in a late anaphase II cell (Figure 3E). The diploid

chromosome number observed during this study supports

previous reports (Moore 1970, 1972, 1977; Goldblatt

1981, 1983; Goldblatt & Johnson 1994).

m

B

I

FIGURE 3. — Photomicrographs of mei-

otic chromosomes in Lagurus

ovatus. A, Spies 3894, diakinesis

with 7n; B, Davidse 33570, dia-

kinesis with 7n; C, Spies 5227,

metaphase I with 1\\; D, Davidse

33570, metaphase I with one bi-

valent away from the metaphase

plate; E, Spies 3894 , telophase I

with a laggard. Scale bar: lO/rm.

Bothalia 26,1 (1996)

67

FIGURE 4. — Photomicro-

graphs of meiotic

chromosomes in the

genus Polypogon.

A, P. monspeliensis.

Spies 5199 , early

metaphase I with

14u; B, P. monspe-

liensis, Spies 3083,

early metaphase I

with 14n; C, P. viri-

dis. Spies 5201 , di-

plotene with 14n.

Scale bar: 10 pm.

The last genus of the Alopecurinae represented in this

country is Polypogon Desf. This genus comprises 18 spe-

cies worldwide and is represented by two naturalized [/I

monspeliensis (L.) Desf. and P. viridis (Gouan) Breistr.]

and two indigenous [P. griquensis (Stapf) Gibbs Russ, and

P. strictus Nees] species in South Africa (Gibbs Russell

et al. 1990). Two species of Polypogon were studied (Ta-

ble 1). Three specimens of P monspeliensis were

tetraploid (Figure 4A, B), whereas a single specimen was

found to be diploid. This confirms previous reports on

this species in which diploid to hexaploid specimens have

been described (Moore 1970-1972, 1974, 1977; Goldblatt

1981, 1983, 1985; Goldblatt & Johnson 1990, 1994). Both

P. viridis specimens studied were tetraploid (Figure 4C),

thus confirming previous reports (Goldblatt & Johnson

1990, 1994). Meiosis was normal and only bivalents were

formed in all specimens.

The two genera representing the subtribe Phalaridineae

appear to have two different basic chromosome numbers,

i.e. Anthoxanthum with five and Phalaris with both six

and seven. A thorough phylogenetic study based on mo-

lecular data of this subtribe could be very useful in de-

termining the evolutionary significance of the change in

chromosome number. A basic chromosome number of

seven is present in all the genera of the subtribe Alopecuri-

nae present in South Africa.

ACKNOWLEDGEMENTS

The University of the Orange Free State and the Foun-

dation for Research and Development are thanked for fi-

nancial assistance during this study.

REFERENCES

CLAYTON, W.D. & RENVOIZE, S.A. 1986. Genera graminum —

grasses of the world. Kew Bulletin Additional Series 13: 1-389.

EDWARDS, D. & LEISTNER, O.A. 1971. A degree reference system for

citing biological records in southern Africa. Mitteilungen der

Botanischen Staatssammlung, Miinchen 10: 501-509.

FEDOROV, A.A. 1969. Chromosome numbers of flowering plants. Acad-

emy of Sciences of the U.S.S.R., Leningrad.

GIBBS RUSSELL, G.E., WATSON, M„ KOEKEMOER, M„ SMOOK,

L„ BARKER, N.P., ANDERSON, H.M. & DALLWITZ, M.J.

1990. Grasses of southern Africa. Memoirs of the Botanical Sur-

vey of South Africa No. 58.

GOLDBLATT, P. 1981 . Index to plant chromosome numbers 1975-1978.

Monographs in Systematic Botany 5.

GOLDBLATT, P. 1983. Index to plant chromosome numbers 1979-1981.

Monographs of Systematic Botany 8.

GOLDBLATT, P. 1985. Index to plant chromosome numbers 1982, 1983.

Monographs of Systematic Botany 13.

GOLDBLATT, P. 1988. Index to plant chromosome numbers 1984, 1985.

Monographs of Systematic Botany 23.

GOLDBLATT, P. & JOHNSON, D.E. 1990. Index to plant chromosome

numbers 1986, 1987. Monographs of Systematic Botany 30.

GOLDBLATT, P. & JOHNSON, D.E. 1991. Index to plant chromosome

numbers 1988, 1989. Monographs of Systematic Botany 40.

GOLDBLATT, P. & JOHNSON, D.E. 1994. Index to plant chromosome

numbers 1990, 1991. Monographs of Systematic Botany 51.

KIMBER, G. & ALONSO, L.C. 1981. The analysis of meiosis in hybrids.

III. Tetraploid hybrids. Canadian Journal of Genetics and Cytol-

ogy 23: 235-254.

MOORE, R.J. 1970. Index to plant chromosome numbers for 1968.

Regnum Vegetabile 68.

MOORE, R.J. 1971. Index to plant chromosome numbers for 1969.

Regnum Vegetabile 77.

MOORE, R.J. 1972. Index to plant chromosome numbers for 1970.

Regnum Vegetabile 84.

MOORE, R.J. 1974. Index to plant chromosome numbers for 1972.

Regnum Vegetabile 91.

MOORE. R.J. 1977. Index to plant chromosome numbers for 1973/1974.

Regnum Vegetabile 96.

ORNDUFF, R. 1967. Index to plant chromosome numbers for 1965.

Regnum Vegetabile 50.

ORNDUFF, R. 1968. Index to plant chromosome numbers for 1966.

Regnum Vegetabile 55.

ORNDUFF, R. 1969. Index to plant chromosome numbers for 1967.

Regnum Vegetabile 59.

SPIES, J.J. & DU PLESSIS, H. 1986. Chromosome studies on African

plants. 1. Bothalia 16: 87, 88.

SPIES, J.J. & VOGES, S.P 1988. Chromosome studies on African plants.

7. Bothalia 18: 114-119.

SPIES, J.J., MOOLMAN, S.K., VAN WYK, S.M.C., MALAN, A.F. &

LIEBENBERG, E.J.L. 1996. Cytogenetic studies in some repre-

sentatives of the subfamily Pooideae (Poaceae) in South Africa. 1.

The tribe Aveneae, subtribe Aveninae. Bothalia 26: 47-55.

SPIES, J.J. & VAN WYK, S.M.C. 1995. Cell fusion: a possible mecha-

nism for the origin of polyploidy. South African Journal of Botany

61: 60-65.

Bothalia 26,1: 69-76(1996)

A tribute to Frank White (5th March 1927 to 12th September 1994)

A. ANGUS* and J.D. CHAPMAN**

Apologia

We are grateful for this opportunity to express our ap-

preciation of our mutual friend Frank White (Figure 1).

Angus worked with him early in his career from 1950

and Chapman was associated with him from the early

1960s. We all met for the first time at Chisenga resthouse

at the foot of the Mafinga Mts in Malawi in 1952. White

and Angus were on an Expedition from Oxford Univer-

sity, and Chapman, of the Nyasaland Forest Dept, was on

‘ulendo’ with his wife, returning with bulging plant

presses from a boundary survey of the Misuku Forests. It

was a momentous meeting for Chapman. He learned that

all his future collections would be named by Frank him-

self at Oxford, and it was the beginning of an association

that has been the inspiration of Chapman’s work ever

since. We have all been friends from that day, and often

enjoyed each other’s hospitality. Thus we feel qualified to

write about Frank White while we fondly remember him.

Introduction

Frank White, eminent Oxford botanist and leading

authority on African plants and vegetation died in Sep-

tember 1994 after a long struggle with emphysema. Al-

ready in 1981 he was beginning to have difficulty with

his breathing, yet despite his increasingly debilitating and

depressive illness he never let up on his work and was

productive to the end, even laying plans for the further-

ance of his unfinished projects.

Although his name is now identified with Africa, he

did, because of his Cambridge training, go on his first

expedition to Arctic Lappland, which resulted in a sig-

nificant paper in the Journal of Ecology (Coombe &

White 1951), and a radio broadcast talk about the Lapps

entitled ‘Europe’s last nomadic race’ (White 1951a).

Thereafter he turned his attentions to Africa. Towards the

end of his life, sadly too late, he had begun to widen his

interests to include tropical America and the Far East.

Career in brief

Bom in Sunderland, County Durham, and educated at

the Bede Collegiate School, he won a Scholarship to Cam-

bridge in 1945. There he obtained Firsts in Parts 1 and 2

of the Natural Science Tripos, and won the Frank Smartt

Prize in Botany. In 1948 he was appointed Demonstrator

in Forest Botany at the Imperial Forestry Institute; was

promoted to University lecturer in 1955; became Curator

of the Forest Herbarium (FHO) in 1961 and also of the

* ‘Rosebank’, Boarhills, St Andrews Fife KY16 8PR, Scotland, UK.

** ‘Braeriach’, Urlar Rd, Aberfeldy, Perthshire, PH 15 2ET, Scotland.

Fielding-Druce Herbarium (OXF) in 1971. In 1988-89 he

was awarded the E. de Wildeman Prize of the Societe

Botanique de Belgique for his work on the African Eben-

aceae, and in 1991 the degree of Sc. D. from Cambridge

University in recognition of his published work. In 1992

the Oxford University Dept of Plant Sciences conferred

on him the title of Distinguished Research Curator for his

‘outstanding research work notably on the taxonomy and

ecology of African plants’.

His work

White was one of a line of Oxford botanists interested in

Africa, beginning with J. Burtt Davy, who founded the Forest

Herbarium, and including A.C. Hoyle and J.P.M. Brenan.

Burtt Davy, incidentally, also founded the herbarium of the

Department of Agriculture of the Transvaal Province in Pre-

toria in 1903 which grew into the National Herbarium, now

part of the National Botanical Institute of South Africa.

White first became known for his works on the Ebenaceae,

FIGURE 1.— Frank White, 1927-1994.

70

Bothalia 26,1 (1996)

FIGURE 2.— White at work with P.

Baraps and J. Leonard in the

herbarium of the Jardin Bo-

tanique in Brussels, 1986.

(Photo by J. Leonard).

Chrysobalanaceae, and Meliaceae, and the forest floras of

Northern Rhodesia (Zambia) and Malawi. His work in

orthodox taxonomy, by himself and in collaboration with

others, resulted in the description of one new genus, 59

new species, and 27 new subspecies. These together with

91 new combinations and other taxonomic categories

make a total of 210 taxa covering 12 families associated

with his name. (Information supplied by S.K. Mamer cour-

tesy of Index Kewensis CD ROM). Later he became re-

nowned principally for his studies in the fields of

phytogeography, chorology, and cartography of the vegeta-

tion of the whole of Africa. To quote his AETFAT colleague,

J. Leonard (1995), ‘il aura marque la phytogeographie de

l’Afrique d’une empreinte originale et indelebile’ (Figure 2).

He was a stalwart protagonist of AETFAT (Associa-

tion pour l’Etude Taxonomique de la Flore d’Afrique

Tropicale), of which he was a founder member and

which played a big role in his life. He attended its first

plenary meeting in Brussels in 1951, and most of the

subsequent general assemblies. At the 5th Assembly in

Sept. 1963 in Italy, he became secretary of a committee

of seven (Aubreville, Barbosa, Codd, Duvigneaud,

Pichi-Sermolli, White and Wild) charged with the com-

pilation of a vegetation map to replace the more concise

earlier map of Keay (1959). The next twenty years were

devoted primarily to this work (Figure 3). Before pub-

lication the map was tested in the field by many expe-

rienced AETFAT members, but chiefly by White

himself. The finished map, with its 365-page memoir,

was a brilliant synthesis and earned great acclaim. It

was reviewed by J. Leonard (1984), and translated into

French by P. Bamps in 1986.

White derived great inspiration and help from the

AETFAT fraternity. He in turn enriched its work by his

own considerable genius. Leonard (1995) has said that

‘working on his own in relative isolation at Oxford, he

realized early that works of great breadth, like those

with which he became involved, could only be success-

fully completed with the help of numerous specialists

such as are to be found within the great family of AET-

FAT’.

White was an original thinker. Take for example his

treatment of those very variable species which defy clas-

sification because of lack of correlation of characters, a

classic example being Diospyros mespiliformis (White

1962b). He coined the term ‘ochlospecies’ to accommo-

date them, and this term has been taken up by taxonomists

around the world.

In the field of phytogeography he devoted much of

this thinking to chorology. In current usage it is the study

of the distribution of taxa and phytochoria and their his-

tories. In his memoir (White 1983c) he outlined a new

conception of African chorology. The system he used for

the Vegetation Map was concerned with the most widely

used catetory, the region (see his definition, White 1976a,

1979c), which in previous systems had been characterised

mainly by endemic families and genera. But in his opinion

the distribution of species provides a more objective clas-

sification. He always emphasised the interdependence of

taxonomy, ecology and chorology which, he maintained,

should always be studied together, and he proposed his

now famous general and chorological aphorisms which

he said all botanists should bear constantly in mind (White

1971).

He believed in studying the plants in the field, and to that

end he travelled widely and collected extensively in Mo-

rocco, Nigeria, Cameroun, Niger, Kenya, Zambia, Malawi,

South Africa and Zanzibar (Figure 4). His travel notes, con-

tained in ten documents entitled ‘ Iter africanum ’ totalling

3 000 pages of typescript (see Bibliography), and his col-

lections which run into the 14 000s, give some indication of

his work rate in the field (information supplied by A.M.

Strugnell of the Daubeny Herbarium, Oxford).

His thinking on the role of perception (or ‘intuitive

discernment’) is worth noting (White 1993b). He talks

about perception being ‘based on the capacity of the hu-

man eye and mind to detect patterns in large amounts of

visual and factual data, before they have been consciously

analysed’. ‘Perception’, he said should be based on ‘wide

experience, a critical outlook, the possession of a good

eye and a good memory’. All these qualities he had him-

Bothalia 26,1 (1996)

71

self in good measure (Figures 5 & 6). Chapman was

witness to his powers of observation in the field. ‘On

one very brief visit to the Misuku Forests (Malawi), he

found Alangium chinense, a first record for these for-

ests, an elusive tree which I had failed to discover. A

few weeks later on Mt Mulanje he spotted Olea

FIGURE 3. — White’s map of regional

phytochoria of Africa and

Madagascar (from White 1993:

237).

FIGURE 4. — White, left, at work on

his collections in camp, with

forestry officer W.D. Holmes,

Mwinilunga District, Zambia,

1952. (Photo by A. Angus).

72

Bothalia 26,1 (1996)

FIGURE 5. — Frank White, doyen of 20th century African forest bota-

nists, observing the Chinzama cedars, Widdringtonia whytei

Rendle, Mt Mulanje, Malawi, May 1981, (Photo, by J. Chapman).

europaea subsp. africana, the only record for Malawi

south of the Viphya plateau’.

One of his last accomplishments was the masterly

resume of his ideas and methods, expounded over the

years in diverse publications, which he was asked to write

by his AETFAT colleagues in Brussels (White 1993).

Leonard & Bamps (1995) note that he entitled it ‘par mod-

estie’, ‘The AETFAT Chorological Classification of Af-

rica’. This paper is a fitting finale to his writings which

were always of high quality and always bore the stamp

of scholarship.

Aside from his research. White was an excellent

teacher. He also ran the two big Oxford herbaria from

1971. Many of his students, now dispersed throughout the

world, continue to study plants inspired by his example

and using his methods (Figure 7).

The man

One might imagine from the above account of White’s

work that he must have been a very serious fellow totally

absorbed by his specimens, books and manuscripts. Far from

it. He had a lively sense of humour. He enjoyed laughter

and making laughter. A connoisseur of food and drink, he

was also an accomplished cook. A generous host, his hos-

pitality became legendary. There was always a firkin of ale

tapped ready to refresh visitors to his cottage in Taston

which he appropriately called ‘Firkins’ (Figure 8).

He was an intellectual with the common touch, equally

at home in the pub chatting with the locals as in the rooms

and corridors of the University. An eccentric, in the sense

of not conforming to the ordinary rules of behaviour, his

idiosyncracies could sometimes amuse, sometimes irritate.

For example he would, on occasion, work late into the

night and sleep well into the morning. He often ignored

public holidays. Angus remembers that on a visit to the

Mt Makulu Research Station in Zambia, on Christmas

Day he took no part in the general festivities except to

eat his Christmas dinner. Most of the day was spent col-

lecting and writing notes, and he would not be diverted

from that which he considered to be making the best use

of his time. Chapman recalls his emphatic refusal to take

a daily walk in Zomba in 1981, to prepare for the arduous

climb up Mt Mulanje. His time was too precious.

He was a renowned spinner of yams, and many of his

tales are remembered and retold by his students and col-

leagues. One of the better known anecdotes tells of him

being chased by an ostrich while riding a motor-scooter

in South Africa. Angus remembers his special brand of

‘herbarium humour’. We had numerous botanical swear

words and occasionally one might hear an oath ringing

through the herbarium — words like Hel-i-chrysum or

Sphe-r/awn-ocarpus. The reader will be able to think of

others! We invented common names for African plants,

such as ‘hedgehog with lifebelt’ for Pterocarpus angolen-

FIGURE 6. — White outside the Chinzama hut, Mt Mulanje, May 1981.

The tree is Ilex mitis (L.) Radik. (Photo, by J. Chapman).

Bothalia 26,1 (1996)

73

FIGURE 7.— White, Oxford Univer-

sity archive photograph, giv-

ing a student seminar, 1984.

sis. The tag ‘indet., mat. insuff.’ (not identified, material

insufficient), sometimes necessarily applied to specimens

sent for identification, often meant for him ‘not identifi-

able, material insufferable’.

Although a bachelor, Frank liked women, and got on

well with them. And they liked him. Chapman remembers

hearing how popular he was with the girl students at the

University of Cape Town. ‘His attraction for the ladies

had not diminished 28 years later. On one trip to the

Lichenya Forest Reserve near the foot of Mt Mulanje in

Malawi we were accompanied by a number of tea estate

and other botanically-inclined ladies. Frank obviously

FIGURE 8. — White by his cottage, ‘Firkins’, early 1970s. (Photo, by A.

Angus).

held their attention, and this was clearly not only because

of his red straw hat which they all greatly admired’. And

in his work he had much fruitful collaboration with

women, notably in the role of botanical artist; but also as

co-workers and co-authors in research. Special mention

should be made of Franqoise Dowsett-Lemaire, a Belgian

zoologist with a flair for botany. He first met her in 1981

in Malawi and with Chapman they collaborated on the

flora of Malawi until his death. Through her continuing

efforts this major work will be published. A special friend-

ship developed between them, and during her frequent

stays at his home in Taston, in the last difficult years of

his life, she was a great help to him in the mundane but

necessary affairs of life such as shopping and cooking.

She has said of him (pers. comm.) that ‘he formed me as

a taxonomist’ and she remembers him ‘above all, as won-

derful, affectionate friend’.

We who knew him well shall always remember his

laugh, his upright bearing and striding gait (he was a great

walker), and his twinkling eyes. Chapman’s wife men-

tioned his ‘twinkly eyes’ in her diary, after that first meet-

ing with him at the foot of the Mafinga Mts in 1952. That

‘twinkle’ remained with him always and was a natural

expression of his great intelligence and sense of humour.

Sir Ghillean Prance, one of his most distinguished stu-

dents has said with truth in the Funeral Oration, that ‘those

who knew Frank only for the last ten years of his life did

not know the true Frank White, because he struggled so

hard with illness that made him at times impatient, intol-

erant or outspoken. Underneath was a brave fight to con-

tinue his work in which he succeeded well’. We mourn

the passing of a botanist of great erudition and inde-

pendent mind, but more than that, a fine colleague and a

loyal friend.

REFERENCES

(see also Publications of F. White)

CRONK, Q.C.B. & SUGDEN, A.M. 1994. Frank White, an obituary. The

Independent , 18-10-1994.

KEAY, R.W.J. 1959. Vegetation map of Africa south of the Tropic of

Cancer. Oxford University Press.

74

Bothalia 26,1 (1996)

LEONARD, J, 1984. F. White: The Vegetation of Africa. A review. Taxon

33: 549-551.

LEONARD. J. 1995. Dr Frank White (1927-12 September 1994), Eng-

land. AETFAT Bulletin 42: 59, 60.

LEONARD, J. & BAMPS, P. 1995. Le Dr Frank White et le Jardin

Botanique National de Belgique. Bulletin du Jardin botanique

national de Belgique 64: 3—11.

PUBLICATIONS OF F. WHITE

Compiled by S.K. Marner*, C.F. Styles* and Q.C.B. Cronk**

BENNETT, S.T. & WHITE, F. 1990. Recent work on the chromosome

cytology of Ebenaceae and the need for continuing research.

Bulletin du Jardin botanique national de Belgique 60: 387-391.

CHAPMAN, J.D. & WHITE, F. 1970. The evergreen forests of Malawi.

Oxford Commonwealth Forestry Institute.

COOMBE, D.E. & WHITE, F. 1951. Notes on calcicolous communities

and peat formation in Norwegian Lappland. Journal of Ecology

39: 33-62.

DANIN, A., HEDGE, I.C., LEONARD, J„ WEINERT, E. & WHITE, F.

1992. Towards a new plant-geographical map of southwest Asia.

Journal of Vegetation Science 3: 566.

DE WINTER, B. & WHITE, F. 1961. New combinations in the genus

Diospyros (Ebenaceae). Bothalia 7: 457, 458.

DE WINTER, B. & WHITE, F. 1967. Ebenaceae. In H. Merxmuller,

Prodromus einer Flora von Siidwestafrika 107: 1-6.

DOWSETT-LEMAIRE, F. & WHITE, F. 1990. New and noteworthy

plants from the evergreen forests of Malawi. Bulletin du Jardin

botanique national de Belgique 60: 73-110.

HOPKINS, H.C. & WHITE, F. 1984a. The ecology and chorology of

Parkia in Africa. Bulletin du Jardin botanique national de Bel-

gique 54: 235-266.

HOPKINS, H.C. & WHITE, F. 1984b. Parkia. Distributions Plantarum

Africanarum 27: maps 893-897. Jardin botanique national de

Belgique, Meise.

LETOUZEY, R. & WHITE, F. 1969. New Diospyros (Ebenaceae) in West

and Central Africa . Adansonia, Ser. 2,9: 277-283.

LETOUZEY, R. & WHITE, F. 1970. Ebenaceae. Flore du Cameroun 11:

3-184, 205-207. Museum National d'Histoire Naturelle, Paris.

LETOUZEY, R. & WHITE, F. 1976. Chrysobalanacees nouvelles du

Cameroun et du Gabon. Adansonia. Ser. 2, 16,2: 229-243.

LETOUZEY, R. & WHITE, F. 1978a. Chrysobalanacees. Flore du

Cameroun 20: 1-138, 237-247.

LETOUZEY, R. & WHITE, F. 1978b. Chrysobalanacees. Flore du Gabon

24: 3-138, 194-201.

MOLL, E.J. & WHITE, F. 1978. The Indian Ocean Coastal Belt. In

M.J.A. Werger, Biogeography and ecology of southern Africa 31:

561-598. The Hague, Junk.

PANNELL, C.M. & WHITE, F. 1988. Patterns of speciation in Africa,

Madagascar, and the tropical Far East: regional faunas and cryptic

evolution in vertebrate-dispersed plants. Monographs in System-

atic Botany 25:639-659.

PRANCE, G.T., ROGERS, D.J. & WHITE, F. 1969. Ataximetricstudy of

an angiosperm family: generic delimitation in the Chrysobala-

naceae. New Phytologist 68: 1203-1234.

PRANCE, G.T. & WHITE, F. 1979. Resurrection of the genus Dactylade-

nia (Chrysobalanaceae). Brittonia 31: 483-487.

PRANCE, G.T. & WHITE, F. 1985. A new combination in Maranthes

(Chrysobalanaceae). Brittonia 37: 76, 77.

PRANCE, G.T. & WHITE, F. 1988. The genera of Chrysobalanaceae, a

study in practical and theoretical taxonomy and its relevance to

evolutionary biology. Philosophical Transactions of the Royal

Society of London, Ser. B, 320: 1-184.

ROBERTSON, S.A., WISE, R. & WHITE, F. 1989. Medusagyne opposi-

tifolia. Kew Magazine 5: 166-171.

STYLES, B.T. & WHITE, F. 1962. New and little known species from the

Flora zambesiaca area. XIV. Meliaceae. Boletim da Sociedade

Broteriana, Ser. 2a, 36: 71, 72.

* Department of Plant Sciences, University of Oxford, South Parks Rd,

Oxford OX1 3RB, England.

** Royal Botanic Garden, Edinburgh EH3 5LR, Scotland.

STYLES, B.T. & WHITE, F. 1989a. Three new species and a new

combination of Turraea (Meliaceae). Bulletin du Jardin bo-

tanique national de Belgique 59: 257, 258.

STYLES, B.T. & WHITE, F. 1989b. Family 124 — Meliaceae. In I. Hed-

berg & S. Edwards, Flora of Ethiopia 3: 479-489.

STYLES, B.T. & WHITE, F. 1990. Two new species of Turraea

(Meliaceae) from northeast Africa. Bulletin du Jardin botanique

national de Belgique 60: 415, 416.

STYLES, B.T. & WHITE. F. 1991. Meliaceae. Flora of tropical east

Africa. Rotterdam, Balkema.

WHITE, F. 1950. The forests of Mt Kenya. Journal of the Oxford Univer-

sity Society, Ser. 3, 5: 32-38.

-1951a. Europe’s last nomadic race. Extract from a broadcast talk on the

BBC Third Programme (1st Jan. 1951), The Listener, Jan. 11,

1951.

-1951b. Notes on five West African species of Cissus Linn. Kew Bulletin

1951: 53-62.

-1955. Distribution of the African species of Diospyros. Webbia 11:

525-540.

-1956a. Notes on the Ebenaceae. I. The genus Maba in Africa. Bulletin du

Jardin botanique de FEtat, Bruxelles 26: 237-246.

-1956b. Notes on the Ebenaceae. II. Diospyros piscatoria and its allies.

Bulletin du Jardin botanique de FEtat, Bruxelles 26: 277-307.

-1957a. Notes on the Ebenaceae. III. Diospyros monbuttensis and two

related species. Bulletin du Jardin botanique de FEtat, Bruxelles

21 (Volume Jubilaire Walter Robyns): 515-531.

-1957b. Trees, shrubs and man in Northern Rhodesia. Journal of the

Oxford University Forestry Society, Ser. 4, 5: 38-43.

-1958a. Two new combinations in Maerua Forssk. Boletim da Sociedade

Broteriana , Ser. 2a, 32: 33-35.

-1958b. The forest flora of Northern Rhodesia (description of scope and

contents of). Memorias da Sociedade Broteriana 13: 47-49.

-1959. A new Entada from tropical Africa. Boletim da Sociedade Brote-

riana, Ser. 2a, 33: 5-11.

-1961. A history of plant collecting in Northern Rhodesia. Comptes

rendus de la LVe Reunion Pleniere de FAetfat, Lisboa: 175-203.

-1962a. Forest flora of Northern Rhodesia. Oxford University Press,

London.

-1962b. Geographic variation and speciation in Africa with particular

reference to Diospyros. Systematics Association Publication No.

4: 71-103.

-1962c. Notes on the Ebenaceae. IV. Anew Diospyros from Mozambique

and Natal. Boletim da Sociedade Broteriana, Ser. 2a, 36: 97-101.

-1963a. Notes on the Ebenaceae. V. Some new names and a new species

in Diospyros. Bulletin du Jardin botanique de FEtat, Bruxelles 33:

345-367.

-1963b. Ebenaceae. In F.N. Hepper, Flora of west tropical Africa 2,2:

2-15.

-1964a. Ebenaceae. In R.W.J. Keay, C.F.A. Onochie & D.P. Stanfield,

Nigerian Trees 2: 329-344.

-1964b. Sapotaceae. In R.W.J. Keay, C.F.A. Onochie & D.P. Stanfield,

Nigerian Trees 2: 344—365.

-1965. The savanna woodlands of the Zambesian and Sudanian domains.

An ecological and phytogeographical comparison. Webbia 19:

651-681.

-1966. Melianthaceae. In A.W. Exell, A. Fernandes & A. Wild, Flora

zambesiaca 2, 3: 544—547.

-1968a. Zambia. In I. Hedberg & O. Hedberg. Conservation of vegetation

in Africa south of the Sahara. Acta Phytogeographica Suecica 54:

208-215.

-1968b. Report on the new A.E.T.F.A.T. vegetation map of Africa. In I.

Hedberg & O. Hedberg. Conservation of vegetation in Africa

south of the Sahara. Acta Phytogeographica Suecica 54: 305.

-1968c. A proposed atlas of ecologically important species in Africa. In I.

Hedberg & O. Hedberg, Conservation of vegetation in Africa south

of the Sahara. Acta Phytogeographica Suecica 54: 307-310.

-1969. Objectivity and repeatability in herbarium studies. Journal of the

Oxford University Forest Society, Ser. 6, 4: 5-8.

-1971. The taxonomic and ecological basis of chorology. Mitteilungen

der Botanisclien Staatssammlung, Miinchen 10: 91-112.

-1976a. The vegetation map of Africa: the history of a completed project.

Boissiera 24: 659-666.

-1976b. The taxonomy, ecology and chorology of African Chrysobala-

naceae (excluding Acioa). Bulletin du Jardin botanique national

de Belgique 46: 265-350.

Bothalia 26,1 (1996)

75

-1976c. Chrysobalanaceae (excluding Aciod). Distributiones Plantarum

Africanarum 9: maps 182-334. Jardin Botanique National de

Belgique, Meise.

-1977a. The underground forests of Africa: a preliminary review. Gar-

dens' Bulletin, (Singapore) 29: 57-71.

-1977b. Some new taxa in African Myrtaceae. Kirkia 10: 401^404.

-1978a. The taxonomy, ecology and chorology of African Ebenaceae. I.

The Guineo-Congolian species. Bulletin du Jardin botanique na-

tional de Belgique 48: 245-358.

-1978b. Ebenaceae. The Guineo-Congolian species of Diospyros. Dis-

tributiones Plantarum Africanarum 14: maps 440-494. Jardin du

Botanique National de Belgique, Meise.

-1978c. Flora of Panama. Part VIII, Family 155. Ebenaceae. Annals of the

Missouri Botanical Garden 65: 145-154.

-1978d. Chrysobalanaceae. In E. Launert, Flora zambesiaca 4: 33-48.

-1978e. Myrtaceae. In E. Launert, Flora zambesiaca 4: 183-212.

-1978f. The Afromontane region. In M.J.A. Werger, Biogeography and

ecology of southern Africa 31: 463-513. The Hague, Junk.

-1979a. The subdivision of Magnistipula Engl. (Chrysobalanaceae). Brit-

tonia 31: 480—482.

-1979b. The flora and vegetation of the Usambara Mountains. In W.A.

Rodgers & K.M. Homewood, The conservation of the east Usam-

bara Mountains , Tanzania: a review of biological values and land

use pressures : 8-13, 50-52, 67-79.

-1979c. The Guineo-Congolian Region and its relationships to other

phytochoria. Bulletin du Jardin botanique national de Belgique

49: 11-55.

-1980a. Notes on the Ebenaceae. VIII. The African sections of Diospyros.

Bulletin du Jardin botanique national de Belgique 50: 445-460.

-1980b. Notes on the Ebenaceae. IX. A new Diospyros from Mozam-

bique. Boletim da Sociedade Broteriana, Ser. 2, 54: 1-3.

-1981a. The history of the Afromontane archipelago and the scientific

need for its conservation. African Journal of Ecology 19: 33-54.

-1981b. Ebenaceae. The botany of the Guyana Highlands. Part XI. Mem-

oirs of the New York Botanical Garden 32: 323—329.

-1981c. Lissocarpaceae. Memoirs of the New York Botanical Garden 32:

329, 330.

-1983a. Long-distance dispersal and the origins of the Afromontane flora.

Sonderbande des Naturwissenschaftlichen Vereins in FJamburg 7:

87-116.

-1983b. Long-distance dispersal, overland migration and extinction in the

shaping of tropical African floras. Bothalia 14: 395—403.

-1983c. The vegetation of Africa: a descriptive memoir to accompany the

Unesco/AETFAT/UNSO vegetation map of Africa. Natural Re-

sources Research 20. UNESCO, Paris, [traduction francaise par P.

Bamps (1986)].

-1983d. UNESCO/AETFAT/UNSO vegetation map of Africa. Scale

1:5 000 000, 4 sheets, in colour. UNESCO, Paris.

-1983e. Ebenaceae. In E. Launert, Flora zambesiaca 7,1: 248-300.

-1983f. Ebenaceae. In B.D. Morley & H.R. Toelken, Flowering plants in

Australia : 115, 116. Rigby, Adelaide.

-1984. Afromontane elements in the flora of Sao Tome; variation and

taxonomy of some ‘nomads' and "transgressors’. Garcia de Orta,

Serie de Botanica 6, 1-2: 187-202.

-1986. The taxonomy, chorology and reproductive biology of southern

African Meliaceae and Ptaeroxylaceae. Bothalia 16: 143-168.

-1987a. Ebenaceae. Flore dAfrique Centrale ( Zaire-Rwanda-Burundi ).

Jardin Botanique National de Belgique, Meise.

-1987b. A.C. Hoyle: an appreciation. Kew Bulletin 42: 655, 656.

-1988a. The taxonomy, ecology and chorology of African Ebenaceae. II.

The non-Guineo-Congolian species Diospyros (excluding set.

Royena ). Bulletin du Jardin botanique national de Belgique 58:

325-448.

-1988b. Ebenaceae. Distributiones Plantarum Africanarum 36: maps

1191-1223. Jardin Botanique National de Belgique, Meise.

-1990. Ptaeroxylon obliquum (Ptaeroxylaceae), some other disjuncts, and

the Quaternary history of African vegetation. Bulletin du Museum

National d’Histoire Naturelle, 4 ser., 12, sect. B, Adansonia 2:

139-185.

-1991. Chapter 14: Chorology and ecology of Dombeya on the African

mainland in relation to its taxonomy. In J.H. Seyani, The genus

Dombeya (Sterculiaceae) in continental Africa: 123-161. Ar-

ranged for publication by P. Bamps & F. White. Opera Botanica

Belgica 2: 1-186.

-1993a. African Myricaceae and the history of the Afromontane flora.

Opera Botanica 121: 173-188.

-1993b. The AETFAT chorological classification of Africa: history, meth-

ods and applications. Bulletin du Jardin botanique national de

Belgique 62: 225-281.

-1993c. Twenty-two new and little-known species of Diospyros (Eben-

aceae) from New Caledonia with comments on section Maba.

Bulletin du Museum National d’Histoire Naturelle, 4 ser., 14, sect.

B, Adansonia 2: 179—222.

—1993d. Refuge theory, ice-age aridity and the history of tropical biotas:

an essay in plant geography. Fragmenta Floristica et Geobo-

tanica Supplementum 2,2: 385—409.

-1993e. Ebenaceae. Flore de la Nouvelle-Caledonie 19: 3-89.

WHITE, F. & BARNES, R.D. 1958. Generic characters in the Ebenaceae.

Oxford University Forest Society Journal, Ser. 4, 6: 31-34.

WHITE. F. & CAVENEY, A.N. 1980. Notes on the Ebenaceae. VII. Some

new names and new taxa in Africa. Bulletin du Jardin botanique

national de Belgique 50: 393—399.

WHITE, F. & GONQALVES, M.L. 1969. Melianthaceae. In A. Fernan-

des, Flora de Mozambique 52: 1-7.

WHITE, F. & LEONARD, J. 1991. Phytogeographical links between

Africa and southwest Asia. Flora et vegetatio mundi 9: 229-246.

WHITE, F. & STYLES, B.T. 1963. Meliaceae. In A.W. Exell, A. Fernan-

des & H. Wild, Flora zambesiaca 2,1: 285-319.

WHITE, F. & STYLES, B.T. 1966. Ptaeroxylaceae. In A.W. Exell, A.

Fernandes & H. Wild, Flora zambesiaca 2,2: 547-550.

WHITE, F. & STYLES, B.T. 1986. Meliaceae and Ptaeroxylaceae. In

O.A. Leistner, Flora of southern Africa 18,3: 35-61.

WHITE, F. & STYLES, B.T. 1988. Turraea obtusifolia. The Flowering

Plants of Africa 50: t. 1962.

WHITE, F, STYLES, B.T. & GONQALVES, A.E. 1979. Meliaceae. In

Flora de Mozambique 42.

WHITE, F, STYLES, B.T. & GONQALVES, M.L. 1969. Ptaeroxylaceae.

In A. Fernandes, Flora de Mozambique 53: 1—3.

WHITE, F. & VOSA, C.G. 1980. The chromosome cytology of African

Ebenaceae, with special refrence to polyploidy. Boletim da So-

ciedade Broteriana, Ser. 2, 53: 275-297 .

WHITE, F. & WERGER. M.J.A. 1978. The Guineo-Congolian transition

to southern Africa. In M.J.A. Werger, Biogeography and ecology

of southern Africa 31: 599-620. Junk, The Hague.

UNPUBLISHED SOURCES OF F. WHITE

WHITE, F. 1953. Iter africanum, 1951. 1952. Zambia and adjacent coun-

tries: botanical field notes. 320 pp. (bound manuscript).

-1960. Iter africanum. 1959, 1960. Zambia, Zimbabwe and South Africa:

botanical field notes. 695 pp. (bound manuscript and typescript).

-1963. Iter africanum. 1962, 1963. Nigeria, Cameroun, Niger: botanical

field notes. 202 pp. (bound manuscript).

-1973. Iter africanum. Kenya, Zambia, Zimbabwe, South Africa: botani-

cal field notes. 560 pp. (bound typescript).

-1974. Iter maroccanum. Botanical field notes. 105 pp. (bound type-

script).

-1976. Iter africanum. 1975, 1976. Kenya and Tanzania: botanical field

notes. 263 pp. (bound typescript).

-1978. The vegetation of Africa: a preliminary account. 729 pp. (bound

typescript).

-1979. Iter africanum. 1978, 1979. Kenya and Tanzania: botanical field

notes. 43 pp. (bound typescript).

-1981. Iter africanum. Malawi: botanical field notes. 177 pp. (bound type-

script).

-1984. Iter africanum. Malawi: botanical field notes. 49 pp. (bound type-

script).

-1985. The ecology and chorology o/Crotalaria in Africa and Madagas-

car. 127 pp. (bound typescript).

-1987. Iter africanum. Malawi: botanical field notes. 52 pp. (bound type-

script).

TUTIN, C.E.G., PARNELL, R.J. & WHITE, F. Protecting seeds from

primates: examples from the Lope Reserve, Gabon. Submitted to

Journal of Tropical Ecology.

WHITE, F„ DOWSETT-LEMAIRE, F. CHAPMAN, J.D. in prep. The

evergreen forests, forest floras and forest landscape of Africa

with special reference to Malawi, (draft, 450 pp., unbound

typescript).

76

Bothalia 26,1 (1996)

PUBLICATIONS EDITED BY F. WHITE

WHITE, F. (ed.) 1951a. The arboreal flora of Israel and Transjordan and

its ecological and phytogeographical significance, by M. Zohary.

Institute Paper No. 26. Imperial Forestry Institute, University of

Oxford.

WHITE, F. (ed.) 1951b. The grass, fern and savannah lands of Ceylon,

their nature and ecological significance, by C.H. Holmes. Insti-

tute Paper No. 28. Imperial Forestry Institute, University of

Oxford.

WHITE, F. & ANGUS, A. (eds) 1952. The vegetation of British Guiana —

a preliminary review, by D.B. Fanshawe. Institute Paper No. 29.

Imperial Forestry Institute, University of Oxford..

WHITE, F. (ed.) 1967. A taxonomic study of the Combretum collinum

Group of species. II. The subspecies of Combretum collinum , by

J.C. Okafor . Boletim da Sociedade Broteriana, Ser.2a,41: 137-150.

A Frank White Memorial Symposium on Floral Diversity in Africa and Madagascar, ar-

ranged jointly by the Royal Botanic Gardens, Kew and the Linnean Society of London, will take

place at the Department of Plant Sciences, University of Oxford on 26 and 27 September 1996.

Topics will include: chorology, vegetation structure and diversity, plant adaptation, animal interac-

tions, taxonomic complexities and application of botanical information. Sir Ghillean Prance will

introduce the meeting and speakers will include I. Friis, P. Morat, M.J.A. Werger, P. Lowry, R.M.

Polhill, J.M. Lock, F. Dowsett-Lemaire, C.E.G. Tutin, Q.C.B. Cronk, M.H.P. Jebb, D. Harris and

D.F. Cutler. Caroline Pannell will speak on Frank White’s scientific achievements.

Bothalia 26,1: 77-80(1996)

OBITUARIES

ARTHUR KOELEMAN (1915-1994)

Mr Arthur (At) Koeleman (Figure 1), succulent enthu-

siast and pioneer breeder of aloes, died on 17 July 1994

at the age of 79 from complications brought on by chronic

emphysema. ‘Oom At’, as he was affectionately known,

was born on 14 February 1915 in the district Lydenburg

in the Transvaal Province, now part of Mpumalanga. After

matriculating from the Lydenburg Secondary School in

1932, he studied at the Normaal Kollege in Pretoria and

was awarded the Transvaal Teacher’s Diploma at the end

of 1935. In 1936 Oom At started his teaching career in

Carolina, but at the end of the first term he was transferred

to Blyde and in 1937 to Linden in Johannesburg. Owing

to poor health, he was granted special leave and returned

to his parents’ home in Sabie, where he worked for a

short time at the Glen Lydenburg Mine. After regaining

his strength, Oom At returned to teaching and sub-

sequently held various teaching positions, amongst others

at the Wonderboom-Suid Primary and the Langenhoven

Secondary Schools in Pretoria.

In 1968, at the age of 53, he continued his tertiary

education and one year later obtained a B.Sc. degree from

the University of Pretoria with Botany and Genetics as

majors. A cartoon by L.N. Nel depicting Oom At ‘back

to school’, characteristically with a cigarette in one hand,

was published in the journal of the South African Succu-

lent Society, Aloe 6,1: 21, January-March 1968 (Figure

2). He then taught again for one year and eventually re-

tired from the teaching profession at the end of 1970. He

worked for the Department of Nature Conservation of the

Transvaal Provincial Administration in Pretoria (now part

of Gauteng) during 1972, at the same time registering for

a B.Sc. (Honours) degree in Botany. In 1973, the year

during which the degree was bestowed on him by the

University of Pretoria, he took up a position as Technical

Assistant (Horticulturist) in the botanical garden of the

Department of Botany at the same university. He held this

position from 1 July 1973 to 30 June 1980. Oom At

started his research for a master’s degree in Botany in

1973 under Profs P.J. Robbertse and A. Eicker, and on 6

April 1979 an M.Sc. (botany) was conferred on him for

a thesis entitled: ’n Morfologies-taksonomiese studie van

die blare van die genus Encephalartos Lehm. in Suid-

Afrika (English: A morphologic-taxonomical study of the

leaves of the genus Encephalartos Lehm. in South Africa).

In 1950 Oom At bought Plot 29, Hartebeeshoek, a

smallholding on the northern slopes of the Magaliesberg

north of Pretoria, where he stayed up to the time of his

death. Here he started to grow succulents, particularly

members of the genus Aloe. His growing interest in this

genus became a source of great pleasure and an absorbing

pastime. He systematically started to hybridize various

Aloe species and what started as a hobby, eventually de-

veloped into a full-scale research nursery. It is for this

pioneering work on the hybridization and improvement

of aloes that Oom At will probably be best remembered

(Irish 1993; Koeleman 1962, 1965). Not only was he one

of the founder members of the South African Aloe and

Succulent Society (now the Succulent Society of South

Africa) in 1962, but he was also instrumental in estab-

lishing the South African Aloe Breeders’ Association in

May 1973 (Bezuidenhout 1982; De Wet 1982a, b). Some

of the earlier volumes of the journal Aloe, official mouth-

piece of the South African Succulent Society, carried de-

scriptions of numerous new Aloe cultivars developed,

amongst others, by Oom At; the vouchers for these were

almost exclusively deposited in the National Herbarium

(PRE) in Pretoria. In fact, these were the first cultivars of

any plant group to have been registered in South Africa

(De Bruyn 1972). Oom At was also one of the first re-

searchers to establish and refine methods for multiplying

species of Aloe, Agave and Haworthia by means of tissue

culture (Groenewald, Koeleman & Wessels 1975, 1979;

Groenewald, Wessels & Koeleman 1975, 1976a, b, 1977;

Wessels, Groenewald & Koeleman 1976). Other novel

vegetative techniques, about which Oom At was very se-

cretive, were used to multiply those hybrids registered as

cultivars. In later years he also did pioneering work on

the use of tissue culture methods for the propagation of

species of the cycad genus Encephalartos (Koeleman &

Small 1982). In addition to his active role in succulent

plant studies, Oom At was for many years chairman of

the South African Aloe and Succulent Society and editor

of its journal Aloe. He was subsequently elected as one

of the honorary vice-presidents of the society, and main-

tained an active interest in the hybridization, selection and

propagation of species of Aloe up to the time of his death

in 1994.

FIGURE 1. — Arthur Koeleman (1915-1994). Photograph taken ± 1980.

78

Bothalia 26,1 (1996)

FIGURE 2. — A cartoon of Oom At Koele-

man after he resumed his graduate

studies at the age of 53. The original

caption, translated from Afrikaans,

reads ‘We hear that our chairman is

back on the school bench. All suc-

cess Oom At.’ Reproduced with the

kind permission of the editorial

board of Aloe.

In a sense Oom At was ahead of his times in that he

realized the immense horticultural potential of the South

African flora, and in particular of the genus Aloe , many

decades ago. Today most local botanists appreciate the

potential of our rich botanical heritage for, amongst others,

breeding superior cultivars for the commercial market.

But, at the time, Oom At must have been regarded as

something of an aloe crank. Sadly, he was often the victim

of unfounded and malevolent criticism (see for example

De Bruyn 1972). Not surprisingly, during his lifetime

Oom At received very little recognition for his work on

the cultivation and breeding of aloes. The only response

he received on a joint paper on the control of aloe snout

beetles (Koeleman & De Wet 1986), was a letter from a

farmer in the Karoo enquiring where such beetles could

be obtained for use as a means of eradicating aloes from

his farm (pers. comm, to G.F. Smith)!! His inability to

enthuse some of his fellow South Africans about the hor-

ticultural and other values of the genus Aloe clearly dis-

illusioned him. This is perhaps best illustrated by the fact

that he told the first author and Prof. Ben-Erik van Wyk

of the Department of Botany at the Rand Afrikaans Uni-

versity, during a visit to his nursery, Magaliesberg Na-

vorsingskwekery, that he had destroyed most of his notes

on and records of his crossing experiments in Aloe. A

great pity, indeed. Fortunately, at least some of his culti-

vars are still being grown at the Pretoria National Botani-

cal Garden and at the Lowveld National Botanical Garden

in Nelspruit, Mpumalanga, and in the private gardens of

a few succulent enthusiasts.

Oom At was a gentle, calm, soft-spoken and unassum-

ing man of great integrity. Never forward or self-seeking

in the progress with his research, he was nevertheless zeal-

ous and resolute in achieving the goals which he set him-

self and would strongly defend any views he believed

were correct. Oom At could be carried away by the splen-

dour of the Aloe hybrids he created, and new flowers were

always awaited with considerable anticipation. He was a

friendly and helpful man, always willing to share his

knowledge and inspiring enthusiasm with those who

showed interest in his work. As a trained teacher, the

youth were close to Oom At’s heart and, as can be seen

from his inspiring writings in past issues of Aloe, he al-

ways tried to instill in young people a love and apprecia-

tion for our succulent flora. On a lighter note, Oom At

was very fond of reading ‘cowboy stories’, and many an

evening he would retire with a soft cover ‘noveltjie’ and

a plate of oranges, one of his favourite fruit.

On 8 January 1944 Oom At married Maria Magdalena

Johanna Bezuidenhout (born 18 April 1917 at Kathu in

the Northern Cape). Tannie Rie, as she was familiarly

known, was Oom At’s lifelong companion. An elegant

lady with a beautiful, sparkling personality, she fully sup-

ported Oom At in his work and often assisted with the

task of cross-pollinating the Aloe plants. Tannie Rie is

commemorated in Aloe ‘Ria Koeleman’ [a hybrid between

the Madagascan A. capitata Baker (variety unknown) and

the southern African A. maculata All.], a cultivar bred by

Oom At (Koeleman 1971). Despite setbacks in health ex-

perienced by both, they greatly assisted each other during

their last years together. Tannie Rie passed away on 1

April 1993. The couple had no children.

It was Oom At’s longstanding ambition to submit his

work on the breeding of aloes for a doctoral degree at a

university. Unfortunately this ideal was not to be realized.

Arthur Koeleman will, however, long be honoured as the

man who pioneered the breeding and improvement of

Aloe. He was a man with foresight, and his life was an

inspiring model of what can be achieved through dedica-

tion, enthusiasm and hard work. He is fittingly commemo-

rated in Aloe ‘Oom At’, a handsome cultivar bred by Mr

Gawie Dednam from a cross between A. reitzii Reynolds

and A. arborescens Mill. Oom At will be remembered

with affection and respect by all those who knew him.

ACKNOWLEDGEMENTS

Mmes M. Dednam, B. Bezuidenhout, M.S. Nel and

Miss M. Nel and Mr Kotie Retief are thanked for provid-

ing information.

Bothalia 26,1 (1996)

79

REFERENCES

BEZU1DENH0UT, A.J. 1982. Latest Aloe cultivars for collectors. Farm-

ing Digest 2,17: 24, 25.

DE BRUYN, M.J. 1972. Die registrasie van aalwynkultivars. Aloe 10,2:

10-12 [English summary: p. 43],

DE WET, A. 1982a. Nuutste aalwynkultivars. Landbouweekblad 63 (No.

217) [26 Maart 1982]: 72, 73.

DE WET, A. 1982b. New cultivars: aloes for the enthusiast. Fanning

Digest 2,22: 42, 43.

GROENEWALD, E.G., KOELEMAN, A. & WESSELS, D.C.J. 1975.

Callus formation and subsequent plant regeneration from seed

tissue of Aloe pretoriensis Pole Evans. Zeitschrift fiir Pflanzen-

physiologie 75: 270-272.

GROENEWALD, E.G., KOELEMAN, A. & WESSELS. D.C.J. 1979.

The use of tissue cultures in the propagation and possible hybridi-

zation of aloes and related plants. Aloe 17,2: 37—40.

GROENEWALD, E.G., WESSELS, D.C.J. & KOELEMAN, A. 1975. 'n

Nuwe metode vir die voortplanting van Aloe pretoriensis Pole

Evans. Aloe 13,2: 48, 49.

GROENEWALD, E.G., WESSELS, D.C.J. & KOELEMAN, A. 1976a.

Embryoid formation in callus cultures of Aloe pretoriensis Pole

Evans. South African Journal of Science 72: 89, 90.

GROENEWALD, E.G., WESSELS, D.C.J. & KOELEMAN, A. 1976b.

Die gebruik van weefselkulture vir die voortplanting en moont-

like hibridisasie van aalwyne en ander naverwante plantsoorte.

Aloe 14,3: 51-58.

GROENEWALD, E.G., WESSELS, D.C.J. & KOELEMAN, A. 1977.

Callus formation and subsequent plant regeneration from seed

WERNER BAHNE GEORG JACOBSEN (1909-1995)

Werner Jacobsen, ‘Doc’ or ‘Pop’, as he was known to

colleagues and family, was born on 6 December 1909 in

Berlin, Germany (Figure 3). His schooling was completed

at the Humanistisches Gymnasium in Fiirstenwalde, Ber-

lin in 1927, where he was taught classical as well as mod-

ern languages, mathematics and science. His studies

continued at the Elniversities of Berlin, Graz, Leoben

(Austria) and Munich, wherever the most renowned pro-

fessors in their respective fields were tutoring at the time.

He majored in Botany, Geology and Chemistry in 1931.

Unable to find employment in the botanical field, which

was his first love, he chose a career in geology.

In 1932 his father, an architect, emigrated to South

Africa in search of greener pastures and settled in Cape

Town where he designed and constructed houses. Here

Werner was at home among the mountains which re-

minded him of the Alps in which he had hiked and

climbed during his student years, and he collected some

1 000 specimens of plants for his private herbarium. He

obtained employment with the Anglo-American Corpora-

tion as Assistant Geologist from 1932-1938. He was part

of the team which was instrumental in the discovery of

the Western Reefs Mine in the North-West (western Trans-

vaal). Subsequent to his marriage to Beatrice Cutts in

1936 he was seconded to the so-called Bancroft Circus

for a year, prospecting in what was then Nyasaland

(Malawi), where he was nearly killed by a wounded buf-

falo. Werner, accompanied by his family which by now

included a son, Jens, returned to Germany in 1939 to write

up his doctoral thesis on the geology of the Ventersdorp

System entitled ‘Tektonik und Lithogenese der Venters-

tissue of an Agave species (Agavaceae). Zeitschrift fiir Pflanzen-

physiologie 81: 369-373.

IRISH, G. 1993. Bitter beauty. Signature 26,7: 38-42.

KOELEMAN, A. 1962. Die veredeling van die aalwyn. Lantern 12,2:

82-91 [English summary: p. 91].

KOELEMAN, A. 1965. The selection and breeding of aloes. Cactus and

Succulent Journal (New South Wales) 5,2: 30-32; 5,3: 47, 48.

KOELEMAN, A. 1971. Die registrasie van aalwynkultivars. Aloe 9,3:

4-12.

KOELEMAN, A. & DE WET, A. 1986. Snuitkewers: aalwyn gevaar kan

gekeer word. Landbouweekblad 68 (No. 436) [4 Julie 1986]: 34,

35.

KOELEMAN, A. & SMALL, J.G.C. 1982. A note on callus formation by

stem and root tissue of some Encephalartos species. South Afri-

can Journal of Botany 1: 165, 166.

WESSELS, D.C.J., GROENEWALD. E.G. & KOELEMAN, A. 1976.

Callus formation and subsequent shoot and root development

from leaf tissue of Haworthia planifolia var. cf. var. setulifera v.

Poelln. Zeitschrift fiir Pflanzenphysiologie 78: 141-145.

G.F. SMITH* and A.E. VAN WYK**

* Plant Systematics Research Subdirectorate, National Botanical Insti-

tute, Private Bag X101, Pretoria 0001.

** H.G.W.J. Schweickerdt Herbarium, Department of Botany, University

of Pretoria, Pretoria 0002.

dorp Formation im Klerksdorp — Wolmaransstad Distrikt

des siidwestlichen Transvaal und im nordwestlichen

Oranje-Freistaat’, obtaining his degree with distinction.

FIGURE 3. — Werner Bahne Georg Jacobsen (1909-1995).

80

Bothalia 26,1 (1996)

Unfortunately his timing placed him in the path of the

oncoming conflict of the Second World War and he was

drafted into service spending most of the war years work-

ing for the Reichsamt fin Bodenforschung in Norway, in

the Balkans and in Russia. After the retreat of German

forces from Russia he surrendered to the American forces

on crossing the Elbe River into what became known as

West Germany. As a prisoner of war his linguistic ability

led to his appointment as interpreter to the Allied Forces.

In 1949 he returned to South Africa to rejoin his family

which now included a second son, Niels. They had man-

aged to return to South Africa the year before. Here he

worked as a consultant geologist until 1955, when he

joined the Messina Transvaal Development Corporation

and moved to Mangula (now Mhangura) in Southern Rho-

desia (Zimbabwe). He was resident geologist on the local

mine for the next 15 years. During this time he was able

to indulge his first love, studying the plants of copper-rich

anomalies. This activity resulted in three papers on the

subject (Jacobsen 1968a, b, 1970). At the same time he

made a collection of the flora of the area which resulted

in another publication (Jacobsen 1973).

In 1970 he moved back to South Africa, working from

the headquarters of the Messina Corporation in Johannes-

burg, visiting prospective claims in Namibia and Natal as

well as Zimbabwe until his retirement at the age of 68.

Throughout this time he collected ferns wherever he went,

particularly during holidays when he could visit specific

fern-rich areas, which resulted in a paper on the subject

(Jacobsen 1978). His retirement meant that he was now

free to fulfil his ideal of working on plants, and on ferns

in particular. He attempted to obtain a post at the then

Botanical Research Institute but, much to his disappoint-

ment, he was unsuccessful due to a lack of funds. He now

concentrated his efforts on writing a book on the southern

African ferns, of which he had collected many specimens

during his working years. At the same time he also had

the collection of the National Herbarium at his disposal.

This work resulted in the Ferns and fern allies of southern

Africa (1983) which was the first treatise on this subject

since T.R. Sim’s (1914), Ferns of South Africa. This was

followed by several publications on the taxonomy and dis-

tribution of ferns (Jacobsen 1991; Jacobsen & Jacobsen

1985, 1986, 1988a, b, 1989).

During his later years, his health was gradually eroding

his ability to move about. This resulted in ‘Pop’ spending

most of his time working at his desk on geographical and

other aspects, including the growth of human populations,

shipwrecks around the South African coast, mythological

and clerical figures and a host of other topics. His know-

eldge of Greek and Latin made it possible for him to

compile a 600 page manuscript (unpublished) on the deri-

vations of plant names of the flora of Namibia. His general

and scientific knowledge was exceptional. He was always

the perfect gentleman, a patient teacher to his sons and

to all who worked with him, in many instances encour-

aging his staff to further their studies. He was loved and

respected by all his colleagues. During his retirement he

also presented a series of lectures on ferns to the Wit-

watersrand University Botany honours class, and he wrote

articles on ferns in the newsletters of the SWA Scientific

Society.

Although specialising in ferns in his later years, ‘Pop’

collected about 4 500 plant specimens, mostly from Zim-

babwe and South Africa but also from Mozambique,

which are deposited in the National Herbarium, Zim-

babwe, the National Herbarium, Pretoria and the Senck-

enberg Herbarium, Germany. The 1 000 specimens

collected in the Cape in 1932 were destroyed by fire in

Germany during the war.

‘Pop’ died in his sleep in a Pretoria hospital after a

short illness on the morning of the 24th October 1995.

BOTANICAL PUBLICATIONS BY W.B.G. JACOBSEN

JACOBSEN, W.B.G. 1968a. The influence of the copper content of the

soil on trees and shrubs of Molly South Hill, Mangula. Kirkia 6:

63-84.

JACOBSEN, W.G.B. 1968b. The influence of the copper content of the

soil on the vegetation at Silverside North, Mangula area. Kirkia 6:

259-277.

JACOBSEN, W.B.G. 1970. Further notes on the vegetation of copper

bearing soils at Silverside. Kirkia 7: 285-290.

JACOBSEN, W.B.G. 1973. A checklist and discussion of the flora of a

portion of the Lomagundi District, Rhodesia. Kirkia 9: 139-207.

JACOBSEN, W.B.G. 1978. Some problems of South African Pterido-

phyta. Journal of South African Botany 44: 157-185.

JACOBSEN, W.B.G. 1983. The ferns and fern allies of southern Africa.

Butterworths, Durban, Pretoria.

JACOBSEN, W.B.G. 1991. Phytogeographical analysis of the pterido-

phytes of some randomly selected areas of the world. Thaizia 1:

69-94.

JACOBSEN, W.B.G. & JACOBSEN, N.H.G. 1985. A new species of

C heilanthes (Pteridophyta, Adiantaceae) from the eastern Trans-

vaal. South African Journal of Botany 51: 145-148.

JACOBSEN, W.B.G. & JACOBSEN, N.H.G. 1986. Cheilanthes del-

toidea Kunze in the Waterberg, Transvaal (Adiantaceae). Bothalia

16:41,42.

JACOBSEN, W.B.G. & JACOBSEN, N.H.G. 1988a. The Cheilanthes

hirta complex and allied species (Adiantaceae/Pteridaceae) in

southern Africa. Bothalia 18: 57-77.

JACOBSEN, W.B.G. & JACOBSEN, N.H.G. 1988b. Doryopteris pilosa ,

a new record for South Africa (Adiantaceae/Pteridaceae). Botha-

lia 18: 90-92.

JACOBSEN, W.B.G. & JACOBSEN, N.H.G. 1989. Comparison of the

pteridophyte floras of southern and eastern Africa, with special

reference to high altitude species. Bulletin du Jardin botanique

national de Belgique 59: 261-317.

N.H.G. JACOBSEN* and J.B.E. JACOBSEN

* P.O. Box 49193, Hercules, Pretoria 0030.

Bothalia 26,1: 81 (1996)

Book Review

AUSTRALIAN VEGETATION, 2nd edn, edited by R.H. Groves. 1994.

Cambridge University Press, The Pitt Building, Trumpington St, Cam-

bridge CB2 1RP. Pp XVIII + 562. Price: hard cover, £50; paperback,

£17.95.

The second edition of this well-known publication is, like its prede-

cessor, designed to be ‘a reasonably concise book on Australian vegeta-

tion’ to quote its editor, R.H. Groves. It is just that, but also incorporates

new research data published during the 12 years that have elapsed since

the appearance of the 1st edition.

This book is not a descriptive account of all Australian vegetation

types at community level. For such data one should consult Noel Beadle’s

The vegetation of Australia (Cambridge 1981). Australian vegetation on

the other hand deals more with the dynamics, ecology and biology of

the principal vegetation types found in Australia.

There are some 18 chapters each prepared by a specialist in their

field. After an introduction dealing with phytogeography in the Australian

region. Quaternary vegetation history and a short overview on alien spe-

cies, the bulk of the book is devoted to accounts of the major vegetation

types followed by chapters on extreme habitats such as coastal dunes,

salt marshes and mangroves, inland wetlands and alpine habitats. Some

of these chapters are specifically devoted to those peculiarly Australian

features, e.g. Acacia woodlands, shrublands and open forests, and Euca-

lyptus forests and shrublands. Each chapter concludes with an ample list

of relevant references. R.L. Specht rounds off the book with an excellent

chapter on biodiversity and conservation.

This is an elegantly produced high quality publication in a robust

binding, amply illustrated with diagrams and clear halftone plates. As a

general overview of the subject it would be hard to beat. Regrettably the

hard covered version at £50 sterling (nearly R300) is probably beyond

the means of impoverished South African acacdemics. A paperback edi-

tion at £17.95 is apparently only available in Australia.

J.P. ROURKE

BOTHALIA

Volume 26,1 May 1996

CONTENTS

1. Four new species and one new subspecies of Lachenalia (Hyacinthaceae) from arid areas of South

Africa. G.D. DtJNCAN 1

2. Saxicolous species of the genus Rinodina (lichenized Ascomycetes, Physciaceae) in southern

Africa. M. MATZER and H. MAYRHOFER 11

3. FSA contributions 4: Agavaceae. G.F. SMITH and M. MOSSMER 31

4. FSA contributions 5: Buxaceae. H.F. GLEN 37

5. Notes on African plants:

Apiaceae. A new species of Centella. M.T.R. SCHUBERT and B.-E. VAN WYK 50

Celastraceae. Correct orthography and authbr citation for Elaeodendron. R.H. ARCHER and

A.E. VAN WYK 41

Convolvulaceae. New records, name changes and a new combination in southern Africa. A.D.J.

MEEUSE and W.G. WELMAN 46

Crotalarieae. Rare or extinct species of Argyrolobium. T.J. EDWARDS 42

6. Cytogenetic studies in some representatives of the subfamily Pooideae (Poaceae) in South Africa. 1.

The tribe Aveneae, subtribe Aveninae. J.J. SPIES, S.K. SPIES, S.M.C. VAN WYK, A.F. MALAN

and E.J.L. LIEBENBERG 53

7. Cytogenetic studies in some representatives of the subfamily Pooideae (Poaceae) in South Africa. 2.

The tribe Aveneae, subtribes Phalaridinae and Alopecurinae. J.J. SPIES', S.K. SPIES, S.M.C. VAN

WYK, A.F. MALAN and E.J.L. LIEBENBERG 63

8. A tribute to Frank White (5th March 1927 to 12th September 1994). A. ANGUS and J.D. CHAPMAN 69

9. Obituaries: Arthur Koeleman (1915-1994). G.F. SMITH and A.E. VAN WYK 77

Werner Bahne Georg Jacobsen (1909-1995). N.H.G. JACOBSEN and J.B.E. JACOBSEN 79

10. Book review 81

Abstracted, indexed or listed in • AETFAT Index • AGRICOLA • BIOSIS: Biological Abstracts/RRM • CAB: Herbage Abstracts , Field Crop

Abstracts • CABS: Current Advances in Plant Science • ISI: Current Contents, Scisearch, Research Alert • Kew Record of Taxonomic Literature • Taxon:

Reviews and notices.

ISSN 0006 8241

© Published by and obtainable from: National Botanical Institute, Private Bag X101, Pretoria 0001, South Africa. Typesetting and page layout:

S.S. Brink (NBI). Reproduction and printing: Aurora Printers, P.O. Box 422, Pretoria 0001. Tel. (012) 327-5073.