Bothalia

’N TYDSKRIF VIR PLANTKUNDIGE NAVORSING

A JOURNAL OF BOTANICAL RESEARCH

Vol. 25,1

May/Mei 1995

TECHNICAL PUBLICATIONS OF THE NATIONAL BOTANICAL INSTITUTE, PRETORIA

TEGNIESE PUBLIKASIES VAN DIE NASIONALE BOTANIESE INSTITUUT, PRETORIA

Obtainable from the National Botanical Institute, Private Bag X101,

Pretoria 0001. Republic of South Africa. A catalogue of all available

publications will be issued on request.

BOTHALIA

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

Minister of Agriculture of the Union of South Africa. This house journal

of the National Botanical Institute, Pretoria, is devoted to the furtherance

of botanical science. The main fields covered are taxonomy, ecology,

anatomy and cytology. Two parts of the journal and an index to contents,

authors and subjects are published annually.

A booklet of the contents to Vols I -20 is available.

STRELITZIA

A series of occasional publications on southern African flora, replacing

Memoirs of the Botanical Survey of South Africa and Annals of Kirsten-

hosch Botanic Gardens. Published: 1 & 2.

MEMOIRS OF THE BOTANICAL SURVEY OF SOI

The memoirs are individual treatises usually of an ecological nature, but

sometimes dealing with taxonomy or economic botany. Published: Nos

1-63 (some out of print). Discontinued after No. 63.

Verkrygbaar van die Nasionale Botaniese Instituut, Privaatsak X101,

Pretoria 0001, Republiek van Suid-Afrika. ’n Katalogus van alle beskik-

bare publikasies kan aangevra word.

Bothalia is vernoem ter ere van Generaal Louis Botha, eerste Eerste Minister

en Minister van Landbou van die Unie van Suid-Afrika. Hierdie lyfblad van

die Nasionale Botaniese Instituut, Pretoria, is gewy aan die bevordering van

die wetenskap van plantkunde. Die hoofgebiede wat gedek word, is tak-

sonomie, ekologie, anatomie en sitologie. Twee dele van die tydskrif en 'n

indeks van die inhoud, outeurs en onderwerpe verskyn jaarliks.

'n Inhoudsopgawe tot Vols 1-20 is beskikbaar.

’n Reeks ongereelde publikasies oor die flora van suidelike Afrika.

Vervang Memoirs van die Botaniese Opname van Suid-Afrika en Annals

of Kirstenbosch Botanic Gardens. Gepubliseer: 1 & 2.

H AFRICA

’n Reeks van losstaande omvattende verhandelings oor vemaamlik eko-

logiese, maar soms ook taksonomiese of plantekonomiese onderwerpe.

Gepubliseer: Nos 1-63 (sommige uit druk). Gestaak na No. 63.

ANNALS OF KIRSTENBOSCH BOTANIC GARDENS

A series devoted to the publication of monographs and major works

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

published as Supplementary volumes to the Journal of South African

Botany). Discontinued after No. 19.

FLOWERING PLANTS OF AFRICA (FPA)

This serial presents colour plates of African plants with accompanying

text. The plates are prepared mainly by the artists at the National

Botanical Institute. Many well known botanical artists have contributed

to the series, such as Cythna Letty (over 700 plates), Kathleen Lansdell,

Stella Gower, Betty Connell, Peter Bally and Fay Anderson. The Editor

is pleased to receive living plants of general interest orof economic value

for illustration.

From Vol. 55, twenty plates will be published at irregular intervals.

An index to Vols 1-49 is available.

FLORA OF SOUTHERN AFRICA (FSA)

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

Swaziland, Namibia and Botswana. The FSA contains descriptions of

families, genera, species, infraspecific taxa, keys to genera and species,

synonymy, literature and 1 imited specimen citations, as well as taxonomic

and ecological notes.

Contributions to the FSA will also appear in Bothalia.

PALAEOFLORA OF SOUTHERN AFRICA

A palaeoflora on a pattern comparable to that of the Flora of

southern Africa. Much of the information is presented in the form

of tables and photographic plates depicting fossil populations. Now

available:

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

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

gepubliseer as Supplementary volumes van die Journal of South African

Botany). Gestaak na No. 19.

Hierdie reeks bied kleurplate van Afrikaanse plante met bygaande teks.

Die skilderye word meestal deur die kunstenaars van die Nasionale

Botaniese Instituut voorberei. Talle bekende botaniese kunstenaars het

tot die reeks bygedra, soos Cythna Letty (meer as 700 plate), Kathleen

Lansdell, Stella Gower, Betty Connell. Peter Bally en Fay Anderson. Die

Redakteur verwelkom lewende plante van algemene belang of ekono-

miese waarde vir afbeelding.

Vanaf Vol. 55 sal 20 plate op ’n keer met ongereelde tussenposes

gepubliseer word..

'n Indeks tot Vols 1-49 is beskikbaar.

’n Taksonomiese verhandeling oor the flora van die Republiek van Suid-

Afrika, Lesotho, Swaziland, Namibie en Botswana. Die FSA bevat be-

skry wings van families, genusse, spesies, infraspesifieke taksons, sleutels

tot genusse en spesies, sinonimie, literatuur, verwysings na enkele ek-

semplare, asook beknopte taksonomiese en ekologiese aantekeninge.

Bydraes tot die FSA sal ook in Bothalia verskyn.

'n Paleoflora met ’n uitleg vergelykbaar met die van die Flora van

suidelike Afrika. Baie van die inligting word aangebied in die vorm van

tabelle en fotografiese plate waarop fossiele populasies afgebeeld word.

Reeds beskikbaar:

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

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

Prodromus of South African Megafloras. Devonian to Lower Cretaceous, by/deur J.M. & H.M. Anderson. Ob-

tainable from/Bcskikbaar van: A. A. Balkema Marketing, Box/Posbus 317, Claremont 7735, RSA.

=5

MARY GUNN LIBRARY

NATIONAL BOTANICAL INSTITUTE

PRIVATE BAG X101

REPUBLIC OF SOUTH AFRICA

BOTHALIA

'N TYDSKRIF VIR PLANTKUNDIGE NAVORSING

A JOURNAL OF BOTANICAL RESEARCH

Volume 25,1

Scientific EditorAVetenskaplike Redakteur: O.A. Leistner

Technical Editor/Tegniese Redakteur: B.A. Momberg

NATIONAL

NASIONALE BOTANIESE

INSTITUUT

Private Bag X101 PRETORIA 0001

1995 -05- 15 I

Privaatsak X 101 PRETORIA 0001

NATIONAL BOTANICAL

INSTITUTE

T A N I C A L

INSTITUTE

2 Cussonia Avenue, Brummeria, Pretoria

Private Bag X101, Pretoria 0001

ISSN 0006 8241

May/Mei 1995

Editorial Board/Redaksieraad

D.F. Cutler

B.J. Huntley

P.H. Raven

J.P. Rourke

M.J. Werger

Royal Botanic Gardens, Kew, UK

National Botanical Institute, Cape Town, RSA

Missouri Botanical Garden, St Louis, USA

Compton Herbarium, NBI, Cape Town, RSA

University of Utrecht, Utrecht, Netherlands

CONTENTS— INHOUD

Volume 25,1

1. Systematic studies in the genus Mohria (Pteridophyta: Anemiaceae). VI. Taxonomic review. J.P. ROUX 1

2. Studies in the Marchantiales (Hepaticae) from southern Africa. 8. The genus Plagiochasma (Aytoniaceae:

Aytonioideae) and six local taxa. S.M. PEROLD 13

3. FSA contributions 2: Asphodelaceae/Aloaceae. 1029010 Chortolirion. G.F. SMITH 31

4. FSA contributions 3: Asphodelaceae/Aloaceae, 1028010 Poellnitzia. G.F. SMITH 35

5. Notes on the typification of some species of Aloe (Asphodelaceae/Aloaceae). H.F. GLEN and G.F. SMITH 37

6. Solanum (Solanaceae) in Uganda. Z.R. BUKENYA and J.F. CARASCO 43

7. A new species and a change in status in Ophioglossum (Ophioglossaceae: Pteridophyta) in Africa.

J.E. BURROWS and T.J. EDWARDS 61

8. The generic delimitation of Lachnaea and Cryptadenia (Thymelaeaceae). J.B.P. BEYERS and J.J.A.

VAN DER WALT 65

9. Studies in the Ericoideae (Ericaceae). XVI. Six new species of Erica from the Western Cape, South Africa.

E.G.H. OLIVER and I.M. OLIVER 87

10. Notes on African plants:

Asphodelaceae/Aloaceae. Typification of Aloe species described by B.H. Groenewald. H.F. GLEN,

G.F. SMITH and D.S. HARDY 97

Asteraceae. A new species of Trichogyne from Namaqualand. J.B.P. BEYERS 107

Geraniaceae. Lectotypification of Pelargonium multibracteatum. P. VORSTER and lb. FRIIS . . 102

Rosaceae. A new species of Cliffortia from the Swartberg. A.C. FELLINGHAM 104

Verbenaceae. A new species in the genus Clerodendrum. P.J.J. HERMAN 100

11. Preliminary ethnobotanical studies of the Rwenzori Mountain forest area in Bundibugyo District. Uganda.

H. ORYEM-ORIGA, E.K.Z. KAKUDIDI, A.B. KATENDE and Z.R. BUKENYA Ill

12. Vegetation structure and small-scale pattern in Miombo Woodland. Marondera, Zimbabwe. B.M. CAMP-

BELL, R.N. CUNLIFFE and J. GAMBIZA 121

13. Book reviews 127

14. In memoriam: Jan Kornas, Ellaphie Ward-Hilhorst, Frank White 131

15. Valuable books missing 132

Digitized by the Internet Archive

in 2016

https://archive.org/details/bothaliavolume2525unse

Bothalia 25,1 : 1-12(1995)

Systematic studies in the genus Mohria (Pteridophyta: Anemiaceae). VL

Taxonomic review

J.P. ROUX*

Keywords: Anemiaceae. Mohria , taxonomy

ABSTRACT

A taxonomic review of the genus Mohria Sw. is presented, including diagnostic features, distribution and variation. A key

to the taxa in the genus is provided and each species described.

UITTREKSEL

n Taksonomiese oorsig van die genus Mohria Sw., waaronder kenmerkende eienskappe, verspreiding en variasie word

aangebied. 'n Sleutel tot die taksons in die genus word verskaf en elke spesie word beskryf.

INTRODUCTION

The genus Mohria Sw. is largely an African one,

occurring from the Western Cape in South Africa, along

the eastern mountain ranges as far north as Kenya. Al-

though the genus has a largely eastern distribution, two

species, M. lepigera (Baker) Baker and M. vestita Baker,

also occur on the Bihe Plateau in southern Angola. Moh-

ria, furthermore, occurs on Madagascar and Reunion Is-

land.

Mohria was first described by Plukenet (1700) as Filicula

geranii arvensis folio et facie aethiopica. It was not until

1771 that Linnaeus in his Mantissa plantarum altera de-

scribed Polypodium cajfrorum from a Konig collection.

Konig, a pupil of Linnaeus, was on his way to Tranquebar

(India) but spent a short period (April 1-28, 1768) at the

Cape of Good Hope, where lie collected on Lion’s Head,

Table Mountain, Devil’s Peak, the flats towards Constantia

and along the sea shores (Gunn & Codd 1981).

The younger Linnaeus, however, in his Supplementum

plantarum (1781), transferred Polypodium cajfrorum L. to

the genus Adiantum L. P. cajfrorum has since been trans-

ferred to genera such as Lonchitis L. by Bemhardi (1801)

and Colina by Greene (1893). Before describing the genus

Mohria in 1806, Olaf Swartz initially placed what is today

known as Mohria cajfrorum in the genus Osmunda L.

(Swartz 1801). At the time of publication of the genus

Mohria , M. thurifraga was the only species known.

The genus Mohria honours Daniel Matthias Mohr,

German botanist and later professor of philosophy at the

University of Kiel (Stafleu & Cowan 1981).

A review of the genus has never been undertaken, with

the result that all the collections, till recently, have either

been placed in M. cajfrorum or M. lepigera. The review

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

Claremont 7735.

MS. received: 1993-12-01.

presented here is the result of studies on the morphology

and anatomy of the rhizome and frond (Roux et al. 1992),

vestiture morphology (Roux 1992a), sporangium and

spore morphology (Roux 1992b) and karyology (Roux

1994).

TAXONOMIC REVIEW

Mohria Sw., Synopsis filicum: 6, 159 (1806). Type:

M. thurifraga Sw., nom. superfl. [= Polypodium cajfrorum

L.; now M. cajfrorum (L.) Desv.J.

Terrestrial or lithophytic. Rhizome dictyostelic, pros-

trate or creeping, irregularly branched, scaled, often

stoloniferous, stolons amphiphloic siphonosteles. Fronds

polystichous, erect or spreading, closely spaced or

crowded, homomorphic to dimorphic; vernation circinate

or non-circinate. Stipe terete, firm, variously set with

naviculate trichomes, hairs and/or scales, fertile ones

usually longer than sterile ones. Lamina pinnate to 3-

pinnatifid, herbaceous to coriaceous. Rachis terete or

adaxially shallowly sulcate, set with indumentum

similar to that on stipe. Pinnae proximally short-stalked

and widely spaced, distally sessile and overlapping,

adaxially with clavate and naviculate trichomes, often

also with hairs and/or small scales, abaxially variously

set with clavate and naviculate trichomes, hairs and

scales. Pinnules opposite to alternate, widely spaced or

overlapping, venation free, ending near margin in lobe

or teeth apices. Stomata anomo-, copolo-, desmo-,

euperi-, eupolo-, para- and/or tripolocytic, amphistomic,

hypostomic in M. nudiuscula. Sporangia exindusiate,

usually borne on distal pinnae, borne singly near vein

endings on short, massive stalks, globose; annulus ter-

minal, 8-22-celled; stomium well defined. Spores

tetrahedral, trilete, radially symmetrical, with ridges

and grooves parallel to the equatorial plane, variously

sculptured, 70-120 pm in diameter. Chromosome

number. 2n = 76, 152.

Bothalia 25,1 (1995)

KEY TO SPECIES

la Pinnae adaxially sparsely to densely hirsute:

2a Lamina pinnate, oblanceolate to linear-attenuate; abaxially hirsute or with a few narrow to broad-ovate scales along rachis;

proximal pinnae not reduced in size to base, closely spaced; rhizome stoloniferous 2. M. marginalis

2b Lamina 2-pinnatifid to 2-pinnate, narrowly elliptic; abaxially sparsely to densely set with lanceolate to broad-ovate, pale to

dark brown, entire scales; proximal pinnae widely spaced, decurrent, rhizome not stoloniferous 1. M. lepigera

lb Pinnae adaxially glabrous or sparsely set with short or long hairs largely along secondary rachises and veins:

3a Vernation circinate; stipe and lamina scales spreading; exine ridges fossulate 4. M. saxarilis

3b Vernation non-circinate; scales appressed or twisted but never spreading; exine ridges rounded:

4a Fertile frond stipe significantly longer than sterile frond stipe; adaxial surface of pinnae with hairs only; hairs with sinuate

transverse walls 3. M. caffronim

4b Fertile and sterile frond stipes of near equal length; adaxial surface of pinnae glabrous or with hairs and/or small scales,

especially along secondary rachises; hair cells sinuous-walled or ossiform:

5a Scales on abaxial surface of lamina narrowly linear to narrowly lanceolate, twisted; lamina usually >200 mm long, erect:

6a Lamina herbaceous; pinnae adaxially sparsely hairy, especially along secondary rachises and veins; pinnule margins

strongly dentate 6. M vestita

6b Lamina herbaceous to coriaceous; pinnae adaxially and abaxially glabrous or nearly so; pinnule margins shallowly

dentate to crenate 7. M. nudiuscula

5b Scales on abaxial surface of lamina lanceolate to ovate, appressed; lamina usually < 200 mm long, erect to spreading

5. M. rigida

1. Mohria lepigera ( Baker > Baker in Annals of

Botany 5: 498 (1891).

Notochlaena lepigera Baker: 53 (1884). Type: Mount Dzomba, Zam-

besi-land (Malawi, Zomba Mountain), Kirk s.n. (K, lecto.!, here desig-

nated).

Mohria caffronim (L.) Desv. var. multisquamosa Bonap.: 85 (1917).

M. lepigera (Baker) Baker var. madagascarica Tardieu: 10 ( 1952). Type:

Madagascar, Mont Lntangabalala, pres Ihosy, I 000 m, Perrier de la

Bdthie 7849 (P, holo. !).

Terrestrial or lithophytic. Rhizome prostrate, short, up

to 4 mm in diameter, densely set with roots, stipe bases

and scales; scales 0. 8-5.0 x 0.2-0.9 mm, scarious, cas-

taneous, adnate, linear to narrowly lanceolate, entire.

Fronds few, erect, crowded; vernation non-circinate. Stipe

terete, stramineous, up to 57.0 x 1.2 mm, set with hairs

and scales; hairs up to 2.4 mm long, sinuous-walled;

scales 0.9-3. 8 x 0.3-14 mm, stramineous, adnate, cordate

or cordate-imbricate, broadly ovate to lanceolate, entire.

Lamina narrowly elliptic, up to 402 mm long, 2-pinnatifid

to 2-pinnate, membranous to firmly herbaceous, with 12-

39 pairs of pinnae. Rachis firm, proximally terete, distally

adaxially shallowly sulcate, abaxially sparsely to densely

set with stramineous scales similar in structure to those

on abaxial lamina surface, reducing in size to frond apex,

adaxially with naviculate trichomes and gland-tipped,

straight- walled hairs similar to those on adaxial lamina

surface. Pinnae 9. 0-3.5 x 6-14 mm, opposite to alternate,

proximally widely spaced and decurrent, crowded near

apex, ovate to linear, abaxially sparsely to densely set with

hairs and scales; hairs 0.6-2.2 mm long, sinuous-walled;

scales 0.2-1. 3 x 1. 3-3.6 mm, lanceolate to broadly ovate,

adaxially with naviculate trichomes and sparsely to den-

sely hirsute, with hairs 0.4— 2.6 mm long. Pinnules or seg-

ments opposite, 4—8 pairs per pinna, widely spaced to

overlapping, ovate to circular, margins shallowly crenu-

late or lobed, terminating in 1 or 2 acute, falcate or obtuse

teeth. Spores stramineous, broadly ridged, ridges closely

spaced, colliculate. Chromosome number unknown. Fig-

ure 1 A-H.

Diagnostic featu res

Mohria lepigera is characterized by its generally long

and narrow frond outline, the short stipe in relation to the

lamina length (x= 1:10; n = 30), the often greatly reduced

and widely spaced proximal pinnae and the large, densely

set stramineous to cream-coloured sinuous-walled scales

which often cover the entire abaxial pinna surface. Clavate

trichomes are 30.5-(42.52)-48.8 pm long and naviculate

trichomes are 50.0— ( 1 1 3.44)— 147.5 pm long.

Distribution and habitat

Mohria lepigera occurs in Tanzania, Burundi, Zaire,

Malawi, Mozambique, Zimbabwe and Madagascar (Fig-

ure 2). M. lepigera occurs from 1 200-2 400 m in habitats

ranging from rock crevices or boulder bases and riverine

fringes to moist open ground. Komas (1979) dealt with

the ecology of the taxon in Zambia.

Variation

Mohria lepigera shows a fair amount of plasticity in

the degree to which the lamina is divided, lamina size,

texture and indumentum composition and density. Collec-

tions recorded from moist shaded conditions at higher

elevations have relatively short, less dissected (2-pin-

natifid) membranous fronds, whereas those from xeric

environments have relatively long, 2-pinnate, firmly her-

baceous fronds which are borne in an erect manner. The

densely scaled abaxial surface which is typical of M.

lepigera in some instances only bears filiform scales and

hairs.

Vouchers: Brass 16484 (PRE, SRGH); Drummond & Hemsley 2557

(PRE); Pichi Sermolli 7042 (BOL. SRGH); Polhill & Paulo 1634 (K);

Taylor 3279 (NBG).

2. Mohria marginalis (Savigny) J.P Rota in South

African Journal of Botany 56: 401 (1990b).

Osmunda marginalis Savigny: 655 (1798). Type: Bourbon (Reunion).

Commerson s.n. (P, lecto.!, designated by Roux 1990b).

O. thurifraga Comm, ex Savigny: 655 (1798), nom. nud.

O. thurifraga Comm, ex Bory: 348 (1804), nom. nud.

O. crenata Desv.: 307 (1811). Type: habitat in insula Bourboniae

(Reunion), sine coll. (P, lecto.!, here designated).

Mohria hirsuta J.P. Roux: 435 (1984). Type: Qwa Qwa, Tsheseng,

near cave on road to The Sentinel, Roux 907 (NBG, holo.!; K, PRE,

iso.!).

Bothalia 25,1 (1995)

3

Terrestrial or lithophytic. Rhizome prostrate, short, up

to 3 mm in diameter, irregularly branched, stoloniferous,

sparsely set with scales; scales 0.8-3.3 x 0.2-0.5 mm,

stramineous, scarious, cordate, linear to narrowly lanceo-

late, entire. Fronds erect, crowded; vernation non-cir-

cinate. Stipe terete, firm, stramineous, up to 36 x 1 mm,

sparsely set with naviculate trichomes, hairs and scales;

hairs up to 2.7 mm long, sinuous-walled; scales 1. 2-2.8

x 0.3-0. 9 mm, cordate, narrowly lanceolate, narrowly

ovate to linear-acuminate, entire or shallowly and uneven-

FIGURE 1. — Frond scales in Mohria. A-H. M. lepigera. Burrows 1686 (NBG). A-C, stipe; D, E, rachis; F, adaxial lamina surface; G, H, abaxial

lamina surface. I-Q, M. marginalis, Roux 907 (NBG). I-M, stipe; N, O, rachis; P, adaxial lamina surface; Q, abaxial lamina surface. Scale

bars: A. C, D-G, I, K, M-Q, 1 mm; B. H. J, L, 0.2 mm.

4

Bothalia 25,1 (1995)

10 20 30 40 50 60

FIGURE 2. — Distribution of Mohria lepigera , O; M. marginalis, ■; and

M. nudiuscula , □.

ly crenulate. Lamina 33-131 mm long, oblanceolate to

linear-attenuate, pinnate, firmly herbaceous, with 8-21

pairs of pinnae. Rachis terete, firm, distally somewhat

flexuose, adaxially often shallowly sulcate, sparsely set

with naviculate trichomes, hairs and filiform scales, which

are absent at apex. Pinnae 4.4-14.7 x 5.5-8 mm, opposite

to alternate, proximally widely spaced, crowded and over-

lapping distally, proximally short-stalked and circular to

reniform, distally sessile and ovate to lanceolate, adaxially

sparsely to densely hirsute, with straight-walled, often

eglandular hairs, abaxially with sinuous-walled hairs up

to 1 mm long and filiform scales; scales 1. 6-2.6 x 0. 1-0.2

mm, segments opposite, 2-5 pairs per pinna, usually over-

lapping slightly, rotund to ovate-obtuse, less deeply dis-

sected towards lamina base and apex, shallowly crenulate

to bluntly dentate. Spores stramineous, ridges widely

spaced, ridges and grooves granulate, up to 82 pm in

diameter. Chromosome number. 2n = 76. Figure 1I-Q.

Bory has always been considered as the author of Os-

munda thurifraga (Schelpe 1970, 1977; Schelpe & An-

thony 1986) but the name was published without a

description. Bory (1804: 348) merely noted 'Dans les en-

virons je trouvai une belle fougere, dont les feuilles

froissees repandaient l’odeur de fencens: je l’ai vue

depuis dans l’herbier de M. de Jussieu, elle lui venait de

Commerson, qui l’avait appelee Osmunda thurifraga .’ It

is therefore evident that he did not describe, nor had any

intention to describe, the species, as in other parts of this

work he either provided a short description or gave a ref-

erence to a previous work where he thought a taxon to

be new. The name is therefore a nomen nudum.

Savigny (1798) also published Commerson’s manu-

script name Osmunda thurifraga as a nomen nudum , citing

it as one of the elements used in his description of O.

marginalis, the other being a Sonnerat collection from

Reunion. The Commerson specimen in the herbarium of

the Museum National d’Histoire Naturelle, Paris (P) was

designated as the lectotype of O. marginalis Savigny

(Roux 1990b).

Desvaux (1811), in his description of Mohria crenata,

gave the locality of the species as 'Habitat in insula

Bourboniae’ (Reunion), with no collector given. The

specimen designated here as lectotype of M. crenata is a

sheet which formerly formed part of Desvaux’s herbarium

and is currently housed in the herbarium of the Museum

National d’Histoire Naturelle, Paris (P).

Diagnostic features

Mohria marginalis is characterised by the relatively

long and narrow, less dissected (2-pinnatifid) fronds, the

usually eglandular, short-celled, straight-walled hairs on

the frond, as well as the filiform scales and sinuous-walled

hairs on the stipe, rachis and abaxial surface of the lamina.

Clavate trichomes are 37.5— (48.75)— 57.5 pm long and

naviculate trichomes are 57.0— ( 104.3)— 145.8 pm long.

Distribution and habitat

Mohria marginalis occurs from the southern Drakens-

berg in the northeastern Cape along the mountain ranges

into central Africa, as far north as Burundi. It also occurs

on Reunion (Figure 2). In the Drakensberg the taxon oc-

curs in seasonally moist, often exposed situations at the

rim of vegetation pockets commonly found in shallow

depressions in the Clarens Sandstone formation, at boulder

bases or among grass tussocks at higher elevations. Fur-

ther north it has been recorded from rock crevices. The

plants usually form large masses and may often grow

alongside xerophytic ferns such as Cheilanthes eckloniana

(Kunze) Mett., Ophioglossum lancifolium C. Presl and M.

rigida J.P. Roux.

Variation

Slight variations occur in frond length, the number of

segments per pinna and in the indumentum density on the

stipe and rachis. Narrow hair-like scales often replace

some of the hairs on the abaxial lamina surface.

Vouchers: De Mame s.n. (NBG); Leighton 3224 (BOL); Reekmans

8789 (PRE); Roux 907, 1524 (NBG).

3. Mohria caffrorum (L.) Desv. in Memoirs de la

Societe Linneenne de Paris 6, 2: 198 (1827). Type: habitat

ad Cap. b. Spei., Konig s.n. [LINN 1251/67, lecto.!, desig-

nated by Schelpe & Anthony (1986)].

Polypodium caffrorum L.: 307 (1771); Adiantum caffrorum (L.) L.f.:

447 (1781); Lonchitis caffrorum (L.) Bemh.: 124 (1801); Colina caf-

frorum (L.) Greene: 247 (1893); Mohria thurifraga Sw.: 159, 385 (1806)

nom. superfl.

Osmunda thurifera Sw.: 105 (1801). Type: C.B.S. Thunberg s.n.

[UPS, lecto.!, designated by Schelpe & Anthony (1986)].

Cheilanthes fuscata Blume: 136 (1828). Type: Cape (L. holo.!).

Mohria thurifraga Sw. var. achilleifolia (as achilliaefolia) E.J. Lowe:

104, t. 62B (1862). Typa; t. 62B (icono.!).

Lithophytic or terrestrial. Rhizome prostrate, short and

stout or slender and creeping, 2-6 mm in diameter, ir-

regularly branched, stoloniferous, clothed with roots and

scales; scales 1.6-2. 8 x 0.3-0.8 mm, castaneous, adnate,

lanceolate to ovate, acute to obtuse, entire. Fronds erect,

approximate to crowded, fertile ones usually longer than

sterile ones; vernation non-circinate. Stipe terete, firm,

castaneous throughout or proximally castaneous and dis-

Bothalia 25, 1 (1995)

5

tally stramineous, 17-162 mm long, up to 1.7 mm in

diameter, fertile one significantly longer than sterile one,

densely set with naviculate trichomes, uniseriate hairs and

scales, becoming glabrous later; scales 0.5-3. 8 x 0.1 -1.5

mm, of two types, variable, stramineous — larger scales

linear, narrowly elliptic or lanceolate or broadly ovate,

entire — smaller scales sessile or cordate, irregularly

shaped or with fimbriate base. Sterile lamina 31-235 mm

long, elliptic to oblanceolate, fertile lamina 75-270 mm

long, narrowly elliptic to narrowly oblanceolate, 2-pin-

natifid to 3-pinnate, with 8-26 pairs of pinnae. Rachis

proximally terete or shallowly sulcate, narrowly winged

and distally often flexuose, densely set with indumentum

similar to that on abaxial pinna surface. Pinnae 9-30 x

6.5-12.6 mm, proximally widely spaced, distally crowded

and overlapping, ovate to oblong, proximally short-

stalked, becoming sessile distally, adaxially with clavate

and naviculate trichomes and few short or long sinuous-

walled hairs, 0.4-1. 4 mm long, largely on secondary

rachis; secondary rachis narrowly winged, abaxially with

clavate and naviculate trichomes, hairs and scales; hairs

and scales sparsely to densely set, pale brown to whitish;

scales 0.2-3.5 x 0.1-1. 3 mm, cordate to cordate-imbricate,

narrowly linear to lanceolate, ovate or circular at base,

terminating abruptly in a long and narrow entire apex,

proximally with short and long, straight or curved mar-

ginal outgrowths. Pinnules opposite to alternate, 4—9 pairs

per pinna, widely spaced or overlapping, segments lobed,

each terminating in 1 or 2 cuneate or falcate teeth. Spores

stramineous, ridges and grooves glebulate, pustulate, fine-

ly verrucous or reticulate-verrucous. Chromosome num-

ber: 2n = 76. Figure 3A-0.

Mohria thurifraga Sw. (1806) is a superfluous name

for Polypodium caffrorum L. (1771). However, the con-

cept of this species as construed by Swartz ( 1806) includes

two different elements, namely a Cape taxon and a

Reunion taxon. The latter was described as Osmunda mar-

ginalis Savigny based on collections by Commerson and

Sonnerat from Reunion and is here upheld as a distinct

species.

The description and illustration published by Lowe

(1860) is that of M. vestita (No. 6). Here he mentions a

dwarf form of this species which was known as M. acliil-

leifolia. This he described as M. thurifraga var. achil-

leifolia in 1862. A Sim collection annotated by him as M.

thurifraga var. achilleifolia from Zwaartberg near Pieter-

maritzburg, Natal, is housed in Pretoria (PRE) but this

proved to be a variant of M. vestita. However, the illustra-

tion (t. 62B ) provided by Lowe which is based on a Sim

collection, judging by the size and laminar segmentation,

appears to be a collection of M. caffrorum. The collection

has obviously not been preserved and I therefore designate

plate 62B as iconotype of M. thurifraga var. achilleifolia.

Diagnostic features

Mohria caffrorum is characterized by the branched,

often widely creeping stoloniferous rhizome, the marked

degree of dimorphism in the fertile frond stipe which is

strikingly longer than the sterile frond stipe, indumentum

structure and distribution, and spore ornamentation.

Clavate trichomes are 55.0— (59. 1 5)— 67.5 pm long and

naviculate trichomes are 147.5 — ( 1 79.8)— 222.5 pm long.

Distribution and habitat

Mohria caffrorum is found in the Northern, Western

and Eastern Cape where it occurs in a wide range of

ecological and climatic conditions (Figure 4). It grows at

elevations ranging from almost sea level at Cape Point to

± 1 000 m in the Cedarberg. Plants are generally found

in small clonal clusters at boulder bases, in rock crevices

or in light shade of shrubs, or in large stands in seasonally

moist exposed situations. Fires, which are prevalent in

most of the vegetation types in which the species occurs,

appear to have no ill effect on the plants but rather stimu-

late them.

Variation

Scale density and scale outline or structure on the

abaxial lamina surface show the most striking degree of

variation. When scale variation throughout the distribution

is viewed, a gradation in scale outline is detected from

the west, becoming more complex to the east. Scale varia-

tion is especially apparent in the Gifberg/Vanrhynsdorp

ecotype where the abaxial surface of the mature fronds

of some plants is densely covered by large pale brown to

whitish scales with sinuate-walled cells (Figure 3J-0).

M. caffrorum shows the highest degree of frond dimor-

phism. The difference between stipe length of the sterile

and fertile fronds is the most significant. A relationship

exists between frond and stipe length in the sterile frond

(r = 0.724, P <0.001) and the fertile frond (r = 0.530,

P < 0.001). Fertile fronds proved to have a larger number

of pinnae and pinnules than sterile fronds.

Vouchers: Barker 9790 (NBG); Dahlstrand 792 (GRA); Goldblatt

3667 (PRE); Perry 42 (NBG); Taylor 11833 (PRE. STE).

4. Mohria saxatilis J.P Roux in Journal of South

African Botany 56: 399 (1990a). Type: Cape Province,

3219 (Wuppertal): Wuppertal, west-facing cliff near ex-

perimental forest plot on road to Heuningvlei, (-AA),

Rowe 2012 (NBG, holo.!; K, PRE, iso.!).

Lithophytic. Rhizome prostrate, short, up to 5 mm in

diameter, densely clothed with roots, closely spaced per-

sistent stipe bases and scales; scales 1. 0-3.7 x 0.3-0.4

mm, castaneous, adnate or cuneate, linear to narrowly lan-

ceolate, entire. Fronds few, erect, caespitose, fertile ones

slightly longer than sterile ones; vernation circinate. Stipe

terete, firm, stramineous, 22.0-61.5 mm long, up to 1.2

mm in diameter, fertile one slightly longer than sterile

one, densely set with scales; scales, 1 .0-3.2 x 0.5-1. 1 mm,

stramineous, spreading, cuneate to cordate, narrowly lan-

ceolate to ovate, shallowly and irregularly crenulate.

Lamina elliptic to oblanceolate, 67-117 mm long, 2-pin-

natifid, with 8-15 pairs of pinnae. Rachis proximally

terete, distally narrowly winged and shallowly sulcate,

adaxially with naviculate trichomes and sinuous-walled

hairs, abaxially densely set with naviculate trichomes and

scales similar in structure to those on stipe. Pinnae 11-24

x 8-10 mm, opposite to alternate, oblong to ovate, obtuse,

with 4—6 lobe pairs per pinna, each lobe terminating in a

single tooth or paired acute teeth, abaxially densely set

with naviculate trichomes and scales; scales 0.9-3.4 x

0.3-0.9 mm, stramineous, spreading, cuneate to cordate,

narrowly lanceolate to ovate, entire or shallowly and un-

6

Bothalia 25,1 (1995)

FIGURE 3. — Frond scales in Mohria. A-I, M. caffrorum, Roux 2014 (NBG). A-C, stipe; D-F. rachis; G, abaxial lamina surface; H, I, adaxial lamina

surface. .1-0, M. caffrorum, Roux 2015 (NBG). J, stipe; K, rachis; L, M, abaxial lamina surface; N, O, adaxial lamina surface. Scale bars: A,

C, D, F, G-L, N, O, I mm; B, E, M, 0.2 mm.

Bothalia 25.1 (1995)

7

FIGURE 4. — Distribution of Mohria caffrorum, ■: M. saxatilis, 0;and

M. rigicla , A.

evenly crenulate, adaxially with naviculate trichomes and

sparsely set with straight or sinuous-walled hairs along

broadly winged secondary rachis, 0.9-3. 3 mm long.

Spores stramineous, ridges broad, fossulate, sparsely and

irregularly verrucous, 75.4—92.8 pm in diameter. Chromo-

some number: n = 76. Figure 5A-K.

Diagnostic features

Mohria saxatilis is distinguished by the short prostrate

rhizome, short stipe, the crowded fronds, the large spread-

ing scales on the stipe and on the abaxial lamina surfaces,

and the broadly winged costa. Diagnostic micromor-

phological characters are the larger clavate trichomes

which are 6 1 . 1— (68. 1 1 )— 76.4 pm long (Roux 1992a), and

epidermal cells and the occurrence of copolocytic and

tripolocytic stomata not found in any of the other taxa

(Roux et al. 1992). Naviculate trichomes are 97.7-

( 1 39.05)— 1 77. 19 pm long.

Distribution and habitat

Mohria saxatilis is confined to the Table Mountain

Sandstone formation and occurs from the Cedarberg to

the Langeberg in the Worcester District. The species

grows at elevations ranging from ± 1 000 to 1 500 m

(Figure 4).

Vouchers: Bean 259 (in part, A & B only) (BOL); Compton 17993

(NBG); Roux 2000 (NBG); Roux 2003 (in part) (NBG): Roux 2012 (K,

NBG, PRE).

5. Mohria rigida J.P. Rota in South African Jour-

nal of Botany 56: 268 (1990a). Type: Qwa Qwa, at

Clarens Sandstone cave on road to The Sentinel, Roux

1910 (NBG, holo.!).

Lithophytic or terrestrial. Rhizome prostrate, short, up

to 5 mm in diameter, densely clothed with roots, persistent

stipe bases and scales; scales 3.6-5 .6 x 0.3-0.6 mm, cas-

taneous, adnate, narrowly lanceolate, entire. Fronds erect

or spreading, caespitose, fertile ones slightly longer than

sterile ones, vernation non-circinate. Stipe terete, firm,

proximally castaneous, distally stramineous, 16-88 mm

long, up to 1 .3 mm in diameter, fertile one slightly longer

than sterile one, with sparse indumentum, becoming

glabrous later; scales 0.5-3.4 x 0.3-0. 8 mm, stramineous,

variable, truncate, cordate or cordate-imbricate, narrowly

lanceolate, ovate, broadly ovate or transversely broadly

elliptic, terminating abruptly in long filiform, often

uniseriate hair-like apex, entire or shallowly and irregular-

ly crenulate or with a few long and/or short outgrowths

at base. Lamina 43-197 mm long, elliptic to narrowly

elliptic, 2-pinnate, with 7-21 pairs of pinnae. Rachis terete

and rigid or distally shallowly sulcate, sparsely to densely

set with naviculate trichomes and scales, adaxial scales

similar in structure to those on abaxial surface. Pinnae

10-35 x 4.5-9.5 mm, opposite to approximate, proximally

often alternate, distally alternate and closely spaced, often

overlapping, mostly short-stalked throughout, narrowly

ovate, adaxially glabrous, with naviculate trichomes or

with few long ossiform-celled hairs, 0.9-2.7 mm long, or

with small scales (similar to those on rachis); secondary

rachis shallowly sulcate, distally winged, curves apically,

abaxially sparsely to densely set with naviculate trichomes

and scales; scales 0.5-3. 1 x 0.4— 0.9 mm, stramineous, cor-

date to cordate-imbricate, narrowly linear, triangular, lan-

ceolate or ovate, crenulate or fimbriate. Pinnules opposite

to alternate, 4—8 pairs per pinna, not or slightly overlap-

ping, segments up to 3 pairs per pinnule, lobed, each ter-

minating in 1 or 2 acute teeth. Spores stramineous, ridges

widely spaced, ridges and grooves, granulate.

Chromosome number. 2n = 152. Figure 5L-T.

Diagnostic features

Mohria rigida is characterised by abaxially appressed,

often broadly ovate, cordate-imbricate, often sinuous-

walled scales, the crowded, caespitose frond arrangement,

the rigid unwinged rachis, the often proximally unwinged

secondary rachis, and sporangia generally borne on the

entire frond or only the proximal pinnae may be sterile.

In M. rigida the stipe/lamina ratio averages 1:2.45 (n =

45) whereas in M. vestita, with which this species may

be confused, it averages 1:3.6 (n = 61). Clavate trichomes

are 35.0— (45.76)— 61 . 1 pm long and naviculate trichomes

are 77.5— ( 1 1 2.86)— 1 64.97 pm long.

Distribution and habitat

Mohria rigida is restricted to elevations of 1 800-

2 438 m along the Drakensberg, the adjacent mountainous

Lesotho and the Transvaal Highveld and escarpment (Fig-

ure 4). In the Drakensberg the plants usually occur in-

dividually or in small groups consisting of a few plants

in the alpine and subalpine vegetation belts as defined by

Killick (1963). Almost without exception they occur in

seasonally moist, often exposed, rock crevices.

Variation

Variation occurs in frond and pinna length and in the

density, size and outline of the scales on the abaxial and

adaxial lamina surfaces. In Mohria rigida the stipe/lamina

ratio ranges from 1: 0.84 to 1: 7.0 (x = 2.45; n = 45).

Vouchers: Roberts 3611 (PRE); Roux 1013. 1075. 1904 (NBG);

Schweickerdt s.n. (PRE).

6. Mohria vestita Baker in Transactions of the Lin-

nean Society, Botany 2; 328 (1887). Type: Tanzania.

Bothalia 25,1 (1995)

Kilimanjaro, ad rupium fissuras, 6 000 ft. Johnston s.n.

(K, holo.!).

M. caffrorum (L.) Desv. var. vestila (Baker) F. Ballard: 561 (1954).

M. thurifraga sensu E.J. Lowe: 197, t. 70 (1860).

Lithophytic or terrestrial. Rhizome prostrate, short, up

to 8 mm in diameter, irregularly branched, clothed with

roots and scales; scales 1. 7-6.3 x 0.5- 1.7 mm, castaneous

to feixugineous, adnate, lanceolate to ovate, entire. Fronds

erect, crowded, fertile ones generally longer than sterile

A. B. C. E. F. G. H, J, K, L. M, 0, P, R

FIGURE 5.— Frond scales in Mohria. A-K, M. saxalilis , Roux 2002 (NBG). A-D, stipe; E, F, rachis; G-I, abaxial lamina surface; J, K, adaxial lamina

surface. L-S, M. rigida , Roux 1910 (NBG). L, stipe; M, N. rachis; O-Q, abaxial lamina surface; R-T, adaxial lamina surface. Scale bars:

A-C, E-H, J-M, O, P, R-T, 1 mm; D, I, N, Q, 0.2 mm.

Bothalia 25,1 (1995)

9

ones; vernation non-circinate. Stipe terete, firm, proximal-

ly castaneous, distally stramineous, 20-132 mm long, up

to 4 mm in diameter, with naviculate trichomes and scales

becoming glabrous later; scales 0.5-6.8 x 0. 1-1.5 mm,

stramineous to castaneous, variable, cordate or cordate-

imbricate, filiform or narrowly linear, shortly attenuate,

hastate, or broad at base terminating abruptly in a long

filiform apex, entire or with few short or long outgrowths

at base curving apically. Lamina 53.5-380.0 mm long,

herbaceous, sterile, narrowly elliptic, fertile, 7SM-64 mm

long, narrowly oblanceolate, 2-pinnatifid to 2-pinnate,

with 10-29 pairs of pinnae. Rachis proximally terete, dis-

tally shallowly sulcate and winged, sparsely to densely

set with scales similar to those on abaxial lamina surface.

Pinnae 8^44 x 6.5-19.0 mm, opposite to alternate, widely

spaced proximally, distally crowded, often overlapping,

ovate, proximally short-stalked, distally sessile; secondary

rachis straight, winged throughout, shallowly sulcate

adaxially, with clavate and naviculate trichomes and

sinuous-walled and ossiform-celled hairs, 0.4— 2.2 mm

long, abaxially sparsely to densely set with scales; scales

0.7-3.2 x 0. 1-0.7 mm, castaneous, variable, filiform or

adnate, linear and shortly attenuate, or cordate to cordate -

imbricate, hastate to narrowly lanceolate, entire, irregular-

ly crenulate or with short or long outgrowths at base,

curving apically. Pinnules opposite to alternate, 4—9 pairs

per pinna, generally not overlapping, segments up to 5

pairs, lobed, terminating in a single or paired, acute, often

obtuse and falcate tooth. Spores stramineous, widely

spaced ridges and grooves puncticulate. Chromosome

number. 2n = 152. Figure 6A-L.

Diagnostic features

Mohria vestita is characterized by the short stipe/

lamina ratio, the reduction in size of the pinnae towards

the lamina base, ossiform-celled hairs which adaxially are

confined to the secondary rachis and veins, and the rela-

tively small, generally hastate and sparsely set twisted

scales which are also confined to the secondary rachis and

veins on the abaxial surface. Clavate trichomes are 30.0-

(5 1 .3 1 )— 63.4 pm long and naviculate trichomes are 85.0-

(138.32)- 196.54 pm long.

Distribution and habitat

The distribution of Mohria vestita ranges from the

southern Cape along the central mountain ranges as far

north as Kenya. It also occurs on the higher lying areas

of Angola and in Madagascar (Figure 7). M. vestita

grows in habitats ranging from coastal subtropical con-

ditions to elevations exceeding 3 000 m along the Natal

Drakensberg. On Mount Kilimanjaro, M. vestita has

been recorded at elevations ranging from 1 200-1 400

m and at Humpata in Angola it occurs at 2 220 m. Fires

are common throughout the distribution area of M. ves-

tita, but appear to have no effect on the subterranean

rhizome.

Variation

Variation in frond length can be ascribed to the wide

distribution and diverse environmental conditions

throughout the range. In hostile environments the fronds

may be merely 109 mm long but at lower, more

favourable conditions the fronds may measure up to 690

mm. Irrespective of variation in frond length, a relation-

ship exists between stipe and frond length in the sterile

(r = 0.683, P <0.001) and fertile (r = 0.711, P <0.001)

fronds. Variation in the density and distribution of in-

dumentum is most apparent. Hairs on the adaxial and

abaxial lamina surfaces appear to increase as the distribu-

tion extends to the north. Slight variation in the cell struc-

ture may also be evident.

Vouchers: Daubenberger s.n. (PRE); Faden. Evans & Kabuye

70/344A (BOL); Janse 106 (GRA, PRE); Nicholas 497 (SRGH); Roux

2233 (NBG).

7. Mohria nudiuscula J.P Roux in South African

Journal of Botany 56: 266 (1990a). Type: Natal, Weenen

Division, damp streambanks in the Mnweni area,

Drakensberg, Esterhuysen 14523 (NBG, holo.!; BOL,

PRE, iso.!).

M. caffrorum (L.) Desv. va r.ferruginea J.E. & S.M. Burrows in J.E.

Burrows: 168 (1989). Type: Natal, (2929) Underberg: Drakensberg

Mountains, Injasuti, below Women Grinding Corn, (-AB), Burrows 360

(BOL, holo.!; K, PRE, iso.).

Terrestrial. Rhizome prostrate, short, up to 10 mm in

diameter, densely clothed with roots and scales; scales

4.7-6.9 x 0.3-0.5 mm, castaneous to ferrugineous, adnate,

narrowly lanceolate, entire. Fronds few, caespitose, spread-

ing or erect, fertile ones slightly longer than sterile ones;

vernation non-circinate. Stipe terete, firm, proximally cas-

taneous, distally stramineous, 20-220 mm long, up to 3

mm in diameter, sparsely to densely set with naviculate

trichomes and scales, becoming glabrous later; scales 0.5-

7.5 x 0.1 -1.2 mm, stramineous to castaneous, narrowly

linear, narrowly lanceolate, triangular or hastate, terminat-

ing in a long filiform apex, entire, usually lignified at

point of attachment. Lamina 53-635 mm long, often

coriaceous, narrowly elliptic, pinnate to 2-pinnate, with

7-34 pairs of pinnae. Rachis shallowly sulcate for most

of its length, densely set with naviculate trichomes and

scales, similar but smaller than those along stipe. Pinnae

10-80 x 6-15 mm, proximally opposite to alternate, al-

ternate and closely spaced distally, often overlapping,

ovate to narrowly ovate, adaxially glabrous or with few

naviculate trichomes and hairs, up to 1.8 mm long, often

also with irregularly shaped filiform scales, up to 2.0 x

0.2 mm, along secondary rachis, abaxially with clavate

and naviculate trichomes, and few or numerous scales;

scales stramineous to castaneous, 0.4-3. 2 x 0. 1-0.8 mm,

attenuate or hastate, filiform or narrowly lanceolate;

secondary rachis straight, winged throughout, shallowly

sulcate adaxially, sparsely to densely set with scales

similar to those along rachis. Pinnules opposite to alter-

nate, 3-12 pairs per pinna, basal pair often smaller than

the next, distant to crowded and overlapping, lobed. seg-

ments up to 3 pairs per pinnule, margins slightly revolute,

shallowly crenulate or obtusely toothed. Spores strami-

neous, broad ridges and deep narrow grooves sparsely to

densely set with verrucae or spherical deposits, 70-90

pm in diameter. Chromosome number. 2n = 152 (Figure

6M-W).

Mohria nudiuscula was recently described by Burrows &

Burrows in Burrows (1989) as M. caffrorum var. ferruginea.

Unaware of this study, 1 described the same species as M.

nudiuscula (Roux 1990a). However, when my manuscript

10

Bothalia 25,1 (1995)

was in press, M. cajfrorum (L.) Desv. var. ferruginea J.E.

Burrows & S.M. Burrows was published. Although the

latter name antedates M. nudiuscula 1 found it to be a

distinct taxon and believe it not to be related to M. caf-

frorum. In accordance with Article 11.2 of the Interna-

tional Code of Botanical Nomenclature ( 1994) the name

M. nudiuscula is adopted.

Diagnostic features

Diagnostic features in Mohria nudiuscula are the thick,

coriaceous texture of the lamina, the often less dissected

lamina, the shallowly crenulate or obtuse teeth and the

almost glabrous adaxial lamina surface (hence the specific

epithet). In most cases, however, a few long hairs, or small

FIGURE 6.— Frond scales in Mohria. A-L. M. vestita , Roux 2219 (NBG). A-D, stipe; E, G, rachis; H-J, abaxial lamina surface; K, L, adaxial lamina

surface. M-W, M. nudiuscula , Esterhuysen 14523 (NBG). M— P, stipe; Q-S, rachis; T, U, abaxial lamina surface; V, W, adaxial lamina surface.

Scale bars: A, C, E, G, H. O, Q, S, 0.5 mm; D, J-N, T-W, 1 mm; B, F, I, P, R, 0.2 mm.

Bothalia 25.1 (1995)

Vouchers: Brass 17286 (PRE, SRGH); Brummitt & Synge 65

(SRGH); Last s.n. (K); Nicholson 97 (BOL); Roux 786 & 790 (NBG).

FIGURE 7.-Distribution of Mohria vestita.

scales, may occur adaxially along the secondary rachis;

abaxially long filiform scales also occur along the veins

on the stipe and rachis. Diagnostic micromorphological

characters are the absence of stomata from the adaxial

lamina surface, the presence of pericytic and paracytic

stomata and the cuticular ridges occurring adaxially along

the major veins (Roux et al. 1992). Clavate trichomes are

37.0-(50.5)-62.5 pm long and naviculate trichomes are

1 20.8— f 1 56.4)— 1 90.2 pm long.

Distribution and habitat

Mohria nudiuscula is largely confined to higher eleva-

tions with the distribution ranging from the Amatola

Mountains in the Eastern Cape, along the KwaZulu-Natal

Drakensberg, where this species has been recorded at

elevations up to 2 400 m, and along the escarpment be-

tween the northeastern Orange Free State and KwaZulu-

Natal. It is also widespread throughout the higher parts

of Lesotho. Further north it occurs along the mountain

ranges between Zimbabwe and Mozambique at elevations

ranging up to 2 545 m, the Nyika Plateau in Malawi and

the adjacent Rungwe Mountain range in Tanzania. In Zim-

babwe, M. nudiuscula also occurs on serpentine-derived

soils along the Great Dyke and as far west as the Victoria

Falls (Figure 2). The habitat is subject to frequent bums

which is evident from the fire scars borne by some col-

lections.

Variation

When material throughout the distribution is viewed,

an increase in frond length and pinna number becomes

apparent towards the north. Fronds of plants occurring

along the higher reaches of the Drakensberg are generally

caespitose, firmly coriaceous and less dissected (2-pin-

natifid) with the pinnae and pinnules crowded and over-

lapping. To the north, however, and especially along the

mountains in eastern Zimbabwe, the fronds become erect

and rigid. They are also more dissected (2-pinnate) with

the pinnae and pinnules often distally spaced. The lamina

texture along the northern limits of the distribution area

also appears to be less coriaceous.

ACKNOWLEDGEMENTS

I wish to express my sincerest appreciation to directors

and keepers of the BOL, GRA, K, P, PRE, SRGH and

STE herbaria for making their collections available to me.

REFERENCES

BAKER, J.G. 1884. New plants from Zambesi Country. Journal of Botany

22: 53.

BAKER, J.G. 1887. Enumeration of plants collected by Mr H.H. Johnston

on the Kilimanjaro Expedition, 1884. Transactions of the Linnean

Society of London, Botany 2: 327-355.

BAKER, J.G. 1891. A summary of the new ferns which have been

discovered or described since 1874. Annals of Botany 5: 497, 498.

BALLARD, F. 1954. Ferns and fern allies: miscellaneous notes. Kew

Bulletin 9: 559-561.

BERNHARDI, J.J. 1801. Tentamen alterum filices in genera redigendi.

Journal of Botany (Schrader) 1800,2: 124.

BLUME, C.L. 1828. Enumeratio plantarum Javae. Leiden.

BONAPARTE, R. 1917. Herbier de M. Henri de la Bathie. Notes Pterido-

logiques 4: 85.

BORY DE SAINT VINCENT, J.B.G.G.M. 1804. Voyage dans les quatre

principales lies des mers d'Afrique I: 348. Paris.

BURROWS, J.E. 1989. New taxa, combinations and records of Pteridophyta

from southern and central Africa. Bothalia 19: 167-174.

DESVAUX, N.A. 1811. Observations sur quelques nouveaux genres de

fougeres et sur plusieures especes nouvelles de la meme famille. Der

Gesellschaft naturforschender Freunde zu Berlin Magazin fur die

neuesten Entdeckungen in der gesammten Naturkunde 5: 307.

DESVAUX, N.A. 1827. Prodrome de famille des fougeres. Memories de

la Societe Lineenne de Paris 6, 2: 198.

GREENE, E.L. 1893. Corrections in nomenclature. IV. Erythea I: 247.

GREUTER, W.. BARRIE, F.R.. BURDET, H.M., CHALONER, W.G.,

DEMOULIN, V., HAWKSWORTH, D.L., JORGENSEN, P.M.,

NICOLSON, D.H.. SILVA, PC., TREHANE. P. & MCNEILL, J.

(eds) 1994. International code of botanical nomenclature, adopted

by the Fifteenth International Botanical Congress, Yokohama,

August-September 1993. Regnum Vegetabile 131.

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

Africa. Balkema, Cape Town.

KILLICK, D.J.B. 1963. An account of the plant ecology of the Cathedral

Peak area of the Natal Drakensberg. Memoirs of the Botanical

Survey of South Africa No. 34.

KORNAS, J. 1979. Distribution and ecology of the pteridophytes in

Zambia. Krakow.

LINNAEUS, C. 1771. Mantissa plantarum altera. Stockholm.

LINNAEUS, C. fil. 1781. Supplementum plantarum. Bmnsvigae.

LOWE, EJ. 1860. Natural history’ of British and exotic ferns, Vol. 8.

Groombridge, London.

LOWE, E.J. 1 862. A natural history of new and rare ferns. Groombridge,

London.

PLUKENET, L. 1700. Almagesti botanici mantissa. London.

ROUX, J.P. 1984. Mohria hirsuta, a new species from the Drakensberg.

Journal of South African Botany 50: 435 — 44 1 .

ROUX, J.P. 1990a. Description of two new species of Mohria (Schizae-

aceae: Pteridophyta) from South Africa. South African Journal of

Botany 56: 266-270.

ROUX J.P. 1990b. A new species and combination in Mohria (Schizae-

aceae: Pteridophyta). South African Journal of Botany 56: 399-

402.

ROUX, J.P. 1992a. Systematic studies in the genus Mohria (Pteridophyta:

Anemiaceae). II. Comparative vestiture morphology and phylo-

geny. South African Journal of Botany 58: 215-219.

ROUX, J.P. 1992b. Systematic studies in the genus Mohria (Pteridophyta:

Anemiaceae). III. Comparative sporangium and spore morphol-

ogy. Bothalia 22: 199-204.

ROUX, J.P. 1994. Systematic studies in the genus Mohria (Pteridophyta:

Anemiaceae). V. Karyology. Bothalia 24: 97-99.

ROUX, J.P, VAN DER WALT, J.J.A. & VAN DER MERWE. R.B. 1992.

Systematic studies in the genus Mohria (Pteridophyta: Anemi-

aceae). I. Comparative morphology and anatomy of the rhizome

and frond. South African Journal of Botany 58: 83-89.

SAVIGNY, M.J.C.L. 1798. In Encyclopedic methodique, Botanique Vol.

4. Paris.

12

Bothalia 25,1 (1995)

SCHELPE, E.A.C.L.E. 1970. Pteridophyta. In A.W. Exell & E. Launert,

Flora zambesiaca. Crown Agents, London.

SCHELPE. E.A.C.L.E. 1977. Pteridophyta. In R.B. Fernandes, E. Launert

& E.J. Mendes, Conspectus florae angolensis. Junta de Investi-

gates Cientificas do Ultramar, Lisboa.

SCHELPE, E.A.C.L.E. & ANTHONY, N.C. 1986. Pteridophyta. In O.A.

Leistner, Flora of southern Africa. Department of Agriculture and

Water Supply, Pretoria.

STAFLEU, F.A. & COWAN, R.S. 1981. Taxonomic literature, edn 2. Vol.

3: Lh-O. Bohn, Scheltema & Holkema, Utrecht.

SWARTZ, O. 1801. Genera et species filicum. Journal of Botany (Schrader)

1800,2:4-109.

SWARTZ, O. 1806. Synopsis filicum. Kiliae.

TARDIEU-BLOT, M.-L. 1952. Schizeacees. In H. Humbert, Flore de

Madagascar et des Comores. Museum National d'Histoire Naturelle,

Paris.

Bolhalia 25,1 : 13-29 (1995)

Studies in the Marchantiales (Hepaticae) from southern Africa. 8. The genus

Plagiochasnia (Aytoniaceae: Aytonioideae) and six local taxa

S.M. PEROLD*

Keywords: Hepaticae, Marchantiales, Plagiochasnia appendiculatum, P. beccarianum, P. eximium, P. microcephalum var, microcephalum,

P. rupestre var. rupestre, P. rupestre var. volkii, southern Africa

ABSTRACT

A taxonomic account is given of the genus Plagiochasnia Lehm. & Lindenb. and its two subgenera. Micropylum Bischl.

and Plagiochasnia. The first subgenus is represented in southern Africa by P. rupestre var. rupestre (J.R. & G. Forst.) Steph. and

P. rupestre var. volkii Bischl.; the second by P. appendiculatum Lehm. & Lindenb. (newly recorded for the region), P.

beccarianum Steph., P. eximium (Schiffn.) Steph. and P. microcephalum (Steph.) Steph. var. microcephalum. Descriptions and

illustrations of these taxa together with distribution maps and a key to the subgenera and species are provided.

U1TTREKSEL

’n Taksonomiese verslag oor die genus Plagiochasnia Lehm. & Lindenb. en sy twee subgenusse Micropylum Bischl. en

Plagiochasnia word gegee. Eersgenoemde subgenus word in Suider-Afrika deur P. rupestre var. rupestre (J.R. & G. Forst.)

Steph. en P rupestre var. volkii Bischl. verteenwoordig en laasgenoemde deur P. appendiculatum Lehm. & Lindenb. (nuut

vermeld vir die streek), P. beccarianum Steph., P. eximium (Schiffn.) Steph. en P. microcephalum (Steph.) Steph. var.

microcephalum. Beskrywings en illustrasies van hierdie taksons sowel as verspreidingskaarte en 'n sleutel tot die subgenusse en

spesies word verskaf.

Plagiochasma Lehm. & Lindenb. in Lehm., Novarum et

minus cognitarum stirpium pugillis quaitus: 1 3 ( 1832); Nees:

33, 40 (1838); Gottsche et al.: 511 (1846); Steph.; 775

(1898); A. Evans: 262 (1915); Sim: 16 (1926); K. Mull.:

331 (1951-1958); Hassel de Menendez: 83 (1963); S.W. Ar-

nell: 65 (1963); Bischl.: 71 (1977); R.M. Schust.: 264(1992).

Type species: P. cordatum Lehm. & Lindenb.

Aytonia J.R. & G. Lorst.: 147 (1776); Lindb.: 291 (1868);

Schiffn.: 30 (1893). Type species: A. rupestris.

Aitonia J.R. & G. Lorst.: 46, 73 (1787), orth. var. [not

of Thunb.: 166 (1776)].

Rupinia L. f.: 69 (1781) nom. illegit.

Ruppinia L. f.: 204 (1783) orth. var. Type species: R.

rupestris.

Antrocephalus Lehm.: 682 (1838). Type species: A.

nepalensis.

Teldea Mont, ex Webb & Berthel.: 59 (1840) nom.

illegit. Type species: T. elastica.

Thalloid, smallish to medium-sized to large, Hat to slight-

ly concave, not grooved, firm and somewhat leathery,

glaucous to purplish and then dull and waxy to velvety, or

else bright green to yellowish green, dorsally hardly percep-

tibly areolate to distinctly so, in crowded, extensive mats, on

calcrete soil, or in rocky crevices, on ledges, under over-

hangs, at foot of large boulders, at seepages or along stream

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

MS. received: 1994-07-04.

banks. Branches lingulate and simple to pseudodicho-

tomously or variously furcate, with lateral or apical innova-

tions from keel, sometimes articulated; thickened over

midrib, thinning toward attenuate, narrowly purple, scalloped

margins, apically notched, with scale appendages recurved

over edge. Dorsal epidermis mostly lacking chloroplasts,

thick- or thin-walled with trigones, roughened or smooth and

with or without waxy, granular deposit externally. Air pores

simple, sometimes ± stellate, minute and very inconspicuous

or larger and slightly raised, encircled by 1 ring or by 2(3)

concentric rings of (4)5-8 cells in each, radial walls of cells

often forming continuous lines which may be thickened,

pores leading below into small, compact, empty air chambers

in several irregular layers, bounding walls chlorophyllose,

some scattered cells nearly filled with a single oil body, also

present in storage tissue, where cells are closely packed;

rhizoids ventral, some smooth, others pegged. Scales purple-

red to violet, in 2 forwardly directed ventral rows, large,

extending beyond thallus margins or not, basal portion broad-

ly ovate, apically with 1 or 2(3) appendages, variable in

shape, linear-lanceolate or ovate to orbicular, sometimes con-

stricted or folded at base, containing a few scattered oil cells;

scale margin entire or toothed and with papillae in subgenus

Plagiochasma , but without in subgenus Micropylum.

Monoicous, autoicous or paroicous. Androecia with an-

theridia sunken in tumid, sessile, crescentic to broadly U-

or V-shaped dorsal cushions, base encircled by short

paleae. Arche goniophores dorsal, single to several acrope-

tally arranged along length of leading branch, usually with

tuft of slender paleae around base and eventually mostly

at apex of very short to long unfurrowed stalk, bearing

carpocephalum with ( 1— )2 — ^ 4(— 5) capsules, each on a short

seta, capsule wall unistratose, cells lacking thickening

bands, dehiscing by an irregularly decaying lid and

14

Bothalia 25,1 ( 1995)

covered by ± spherical, bilabiate involucre with vertically

slightly overlapping, somewhat swollen lips, top of car-

pocephalum slightly raised to nearly flat or somewhat

depressed, 3- or 4-sided, with compound air pores. Spores

70-105 pm, yellow to brown, triangular-globular, winged,

both faces coarsely reticulate, proximal face with distinct

triradiate mark. Elaters 175-300 pm long, 7.5-20.0 pm

wide in middle and tapering to ends, bi- or tri-spiral.

The genus Plagiochasma is classified in the family

Aytoniaceae Cavers, and in the subfamily Aytonioideae,

whereas the other four genera of the family, namely

Asterella , Cry pt omit rium, Mannia and Reboulia are placed

in subfamily Reboulioideae Grolle (Grolle 1972).

Plagiochasma was previously said to include ± 34 species

worldwide (Evans 1915), but the number of taxa has been

considerably reduced by Bischler (1977, 1978, 1979).

Most are confined to the rather warmer, dry regions of

the world, although P. rupestre var. rupestre is nearly

cosmopolitan. The other taxa that also occur in southern

Africa, are more narrowly restricted in their distribution.

Two subgenera, namely Micropylum and Plagiochasma

have been instituted by Bischler (1977), based mainly on

the compactness and colour of the thalli, the structure of

the dorsal air pores and on the shape, size and margins

of the scale appendages.

Key to subgenera and species of Plagiochasma

la Thallus glaucous to greyish green, dull and velvety, dorsally granular, with numerous air pores, tiny and obscure, bounded

by a single ring of 4-6 cells but never by an inner hyaline ring of collapsed cells; ventral scales with 1-2(3) narrow

appendages, hardly or not constricted at base, with margins not differentiated and lacking teeth and/or papillae (subgenus

Micropylum Bischl.):

2a Ventral scales reddish pink to purple, rarely more than 2000 pm long, including pink, purple or hyaline appendage(s) ( 1

or 2), narrowly to broadly triangular, up to 900 pm long, apically shortly pointed, with 1 or 2 (rarely 3) cells in

series, thin-walled and quadrate or rectangular; spores up to 92.5 pm in diameter la. P. rupestre var. rupestre

2b Ventral scales dark red or purple, large, up to 3000 pm long, including hyaline appendage(s) (1-3), narrowly triangular, up

to 1500 pm long, apically with a long point, with 3-5 elongated cells in series, thick-walled; spores more than 92.5

pm in diameter lb. P. rupestre var. volkii

lb Thallus green to yellow-green, shiny and dorsally smooth with fewer air pores, large and quite conspicuous, raised and

bounded by an inner hyaline ring of collapsed cells and then by 2 or 3 concentric rings of 5-8 cells in each; ventral

scales with 1-3 appendages, wide or narrow, often constricted or folded at base with margins differentiated into

smaller cells, teeth and/or papillae (subgenus Plagiochasma):

3a Scales mostly with a single appendage, sometimes with two, round or oval or broadly triangular, widest across middle,

constricted or folded at base; along margins 1 or 2 rows of regular, smaller cells or alternating with them cells of

usual size, lacking teeth; elaters with evenly thickened spirals:

4a Thallus robust; carpocephalum on short, thick stalk; scale appendage single, rarely double, rounded, large, hyaline above,

red or pink below, markedly constricted at base; margins smooth, with small cells here and there alternating with

somewhat larger ones 2. P. appendiculatum

4b Thallus medium-sized; small carpocephalum on thin, short or long stalk; scale appendage single or double, oval or broadly

triangular, pink or violet, slightly constricted or folded or horizontally pleated at base; margins with 1 or 2 rows of

smaller, regular cells, occasionally with papillae 3. P. microcephalum var. microcephalum

3b Scales with 1, 2 (or 3) appendages, narrowly triangular, slightly constricted to folded at base or evenly tapered; sometimes

toothed along margins; elaters with spirals interrupted or evenly thickened:

5a Thalli robust but carpocephala relatively small; dorsal air pores of thalli surrounded by hyaline ring and 3 concentric rings

of cells; scale appendages red, not acuminate, slightly constricted or folded at base; elaters with spirals often interrupted

4. P. eximium

5b Thalli less robust and carpocephala of usual size; air pores surrounded by hyaline ring and 2 concentric rings of cells;

scale appendages purple, acuminate, lanceolate, not constricted at base; elaters with spirals evenly thickened . . .

5. P. beccarianum

Subgenus Micropylum Bischl.

Micropylum is characterized by very compact, velvety

thalli, the dorsal surface covered with a water-repellent,

granular deposit and the numerous air pores tiny and in-

conspicuous; the ventral scales have acuminate ap-

pendages with undifferentiated margins, lacking teeth and

papillae. Plagiochasma rupestre var. rupestre and P.

rupestre var. volkii are placed here.

1. Plagiochasma rupestre (J.R. & G. Forst.) Steph.

in Bulletin de FHerbier Boissier, Ser. I, Vol. 6.10: 783

(1898); K. Mull.: 332 (1952); A. Evans: 277 (1915); Sim:

18 (1926); Hassel de Menendez: 84 (1963); S.W. Arnell:

67 (1963); Bischl.: 64 (1978); Volk: 237 (1979); Bischl.

& Sergio: 173 (1984); R.M. Schust.: 292 (1992). Type:

Madeira, Funchal, Quinta do Bom Sucesso, Sergio &

Nobrega 3873 [LISU, neo.! selected by Bischl. & Sergio

(1984)]; PC, BM, G, MADJ, iso.).

P. abyssinicum Gola: 62 (1914). Type: Ethiopia, Erythraea, in regione

Hamasen prope Asmara n. 132, 24. IV. 1909, leg. Chiovenda.

P. algericum Steph.: 780 (1898). Type: Algeria, leg. Trabut.

P. capense Sim: 17 (1926). Type: S Africa. Herschel, Cape Province,

5000 ft, leg. Hepburn.

P. dschallanum Steph.: 778 (1898). Type: Tanzania. Kilimandscharo,

ad lacum Dschalla, leg. Holst.

P. muricatum Steph.: 310 (1901a). nom. illegit.

P. tenue Steph.: 779 (1898). Types: S Africa, Transvaal, leg. MacLea,

Wilms ; Tanzania, Usambara, leg. Holst: Angola, leg. Welwitsch.

The above list of synonyms includes only African

plants and was taken from Bischler (1978). Schuster

(1992) gives an extensive list of ± all synonyms estab-

lished largely on the basis of detailed studies by Evans

(1915) and Bischler (1978).

Bothalia 25,1 (1995)

15

Thallus medium-sized to quite large, nearly Hat to

somewhat concave with the sides slightly raised, oblong

to lingulate (Figure 1A), compact, glaucous to greyish

green, dull, surface ± water-repellent, narrow purple edge

along margins, pores very inconspicuous, almost imper-

ceptible as minute dots, subdorsal air chamber walls hard-

ly visible, when wet; thallus margins incurved or clasped

together, exposing deep purple to dull black, transversely

striate underside of wings, toward apex sometimes

covered with a dull bloom and medianly always by scales,

when dry; in crowded, gregarious patches, simple or once,

rarely twice furcate or with apical or latero-ventral in-

novations and then articulated or jointed. Branches 8-25

x (3-)4— 6 mm, 375-650 pm thick over midrib, laterally

thinning out into attenuate wings (Figure IF); apex

notched, with appendages of 8-10 deep pinkish purple or

purple, sometimes partly decolorate scales recurved over

edge; margins acute, thin, scalloped and undulate; flanks

sloping obliquely, purple; ventral face medianly keeled,

green, with row of purple-red or purple scales on either

side (Figure IE).

Dorsal epidermal cells unistratose, hyaline, polygonal,

5-7-sided, 25^-7 x 20-37 pm. thin-walled but thickened

at comers, in transverse section 25-37 pm thick, exter-

nally covered with granules, occasionally containing an

oil body, along margins cells rectangular (Figure II), ±

27 x 15 pm, air pores numerous, 88-250 pm distant from

each other, not raised (Figure 1H), simple, tiny, mostly

appearing plugged, but lacking small inner hyaline ring

of collapsed cells, surrounded in one series by 3-5 small,

bluntly triangular cells (Figure 1G), 10-20 pm long, 17-

25 pm across widest part, partly overlying dorsal cells,

radial walls often thickened. Assimilation tissue 175-220

pm thick, ± 1/3 the thickness of thallus, air chambers

empty, in several layers, vertical medianly, sloping obli-

quely toward the sides, 37— 55(— 125) pm wide, cells in

bounding walls rounded, rather irregular in size, but

averaging 25 x 22-25 pm, some with oil bodies, these

also present in storage tissue which occupies remaining

% thickness of thallus, cells angular, 25-32 pm wide;

rhizoids mostly pegged, 20-30 pm wide, occasionally

smooth, 17.5 pm wide. Scales reddish pink or purple, oc-

casionally with a faint bloom, appendages and base some-

times hyaline, arranged in 2 forwardly directed ventral

rows, one on either side of midrib, rather asymmetric,

obtusely triangular with flatly arched base, gradually nar-

rowed above, without constriction into 1 (Figure IK) or

2(3) (Figure 1J) acuminate to ovate-lanceolate ap-

pendages, margins entire, total length (including ( 600—)

900-1450 pm long appendage) 1200-3000 pm, width at

base (600—) 900- 1250 pm, cells in body of scale 4—6-

sided, 30-62(-75) x 17-25 pm, becoming smaller toward

base and rectangular or not along margin, 30-40 x 20

pm, sometimes bulging outwards, where cells join, at apex

ending in l^f uniseriate cells, scattered throughout scale

several cells that had contained oil bodies, 20-37 x 25

pm.

Monoicous. Androecia in sessile cushions, round or

crescentic, up to ± 2 x 3 mm, medianly on leading branch

or on apical or latero-ventral innovations with immersed

antheridia that open above via raised, 37.5 pm high, con-

ical papillae (Figure ID), at base encircled by tapering,

purple paleae (Figure IN), up to 600 pm long, 180 pm

wide at base, cells 4- or 5-sided, ± 50 x 30-37 pm, apical

cell ± 32 x 20 pm. Archegoniophores in acropetal se-

quence medianly along main branch, or paired on pseu-

dodichotomously furcate branches, enclosed by tall tuft of

tapering, purple-red paleae, up to 1500 pm long, 220 pm

wide at base, most of them later carried to tip of stalk

(Figure 1M), cells in body of paleae rectangular, 37-53

x 20-25 pm, toward apex elongating and narrowing, up

to 62 x 17 pm, 3 or 4 serially arranged. Caipocephala

2-3 mm wide, on top initially slightly raised (Figure IB),

later becoming depressed, air pores compound, with 1-

3(— 5) lobes, involucral lips vertical and swollen, slightly

overlapping, each enclosing a single oval capsule, 1500

x 1000 pm, on a short seta and with decaying lid, wall

unistratose, cells 25.0-37.5 x 22.5-32.5 pm, polygonal,

thin-walled with comers thickened, lacking thickening

bands (Figure 1L); stalk at maturity usually remaining

very short, only rarely up to 6 mm long, lacking rhizoidal

furrow, 640 x 430 pm in transverse section (Figure 1C),

cortical cells thicker-walled externally, rounded, 17.5 x

15.0 pm, medullary cells thin-walled, angular, ± 25 pm

wide. Spores 80-105 pm in diameter, triangular-globular,

polar, light brown to yellow brown, translucent, wing up

to 12 pm wide, margin undulate, crenulate; ornamentation

similar on both faces: distal face (Figure 2 A, F) with ± 4

areolae across, ± 20 pm wide, areolar walls wide and

studded with granules (Figure 2B, G); proximal face with

narrow triradiate mark, clearly elevated and distinct (Fig-

ure 2C), each of 3 facets with 6-9 areolae, up to 25 pm

wide, walls raised and wide, studded with granules (Figure

2D, H). Elaters light brown to yellow brown, 175-235

pm long, 12.5 pm wide in middle, tapering toward ends,

± 5 pm wide, bispiral (Figure 2E, I). Chromosome num-

ber: n = 9, 18 (Bischler 1978).

Two varieties are recognized: P. rupestre var. rupestre

and P. rupestre var. volkii. They are distinguished by the

longer, acuminate, hyaline appendages of the ventral

scales and slightly larger spore dimensions of the latter.

la. P. rupestre var. rupestre

Obliquely lunate ventral scales rarely longer than 2000

pm and 1050 pm across base; 1 or 2 narrowly to broadly

triangular appendages up to 900 pm long, 1.7-2. 7 times

longer than wide, and apically acuminate with uniseriate

1 or 2(3) tenninal cells, quadrate or rectangular in shape

and thin-walled; cells in curved margin of body of scale

rectangular, sometimes bulging slightly outward where 2

cells join; male paleae around basal part of androecium

550-580 pm long, 1 10-180 pm wide at base, tapering to

a narrow tip, with 1-3 cells in series, ± 32 x 20 pm (Fig-

ure IN); female paleae 1030-1375 pm long, 150-220 pm

wide at base, tapering to a narrow tip, with up to 4 cells

in series, 30-40 x 17 pm (Figure 1M); spores 80.0-92.5

pm in diameter (Figure 2A, D).

The typical variety is subcosmopolitan and widely dis-

tributed. especially in xerothermic regions. Frey &

Kiirschner (1988) regard it as a xerothermic Pangaean

taxon. In southern Africa it is quite common and frequent-

ly collected in rocky crevices, moist ledges, under

boulders, or at seepages, on calcareous substrates or on

soil overlying cave sandstone or dolomite. It sometimes

grows together with Targionia hypophylla , Athalamia

16

Bothalia 25.1 (1905)

FIGURE I. — A-N. Plagiochasma rupestre var. rupestre : A, E, F, I, thallus: A, dorsal face, with archegoniophore and carpocephalum; E, ventral

face; F, t.s.; I, margin. B, carpocephalum, side view; C, l.s. of stalk; D, androecium at apex of latero-ventral branch; G, air pore and dorsal

cells from above; H, t.s. of air pore, dorsal cells and air chambers. J, K, ventral scale: J, with 2 appendages; K, with 1 appendage. L, cells in

capsule wall; M, female paleae; N, male palea. O-R, Plagiochasma rupestre var. volkii. O, P, ventral scale: O, with I appendage; P, with 2

appendages. Q, male palea; R, female palea. A, B, D, E, L, N, HeilgendorffCH 1361 1 ; C, Manning CH 13590\ F, H, Anderson 1230\ G, I-K,

M, S.M. Perold 305ti\ O-R, Mogg 37590. Scale bars: A, B, D, E, 2 mm; F, 1 mm; C, J, K, M-R, 250 pm; G-I, L, 50 pm.

Bothalia 25,1 (1995)

17

FIGURE 2. — SEM micrographs of spores and elater. A-E, Plagiochasma mpestre var. rupestre: A, distal face; B, much enlarged view of some

areolae and walls on distal face; C, side view; D, proximal face; E, part of elater much enlarged. F-I, P. rupestre var. volkii : F, distal face; G,

much enlarged view of some areolae and walls on distal face; H, proximal face; I, part of elater much enlarged. A, G.W. Sim CH 1145: B,

neotype, Sergio & Nobrega 3873: C, Volk 00639: D, Pole Evans 458: E, Cooper 962: F-I, Mogg 37590. A, x 452; B, G, x 1 150; C, x 464;

D, x 440; E, I, x 936; F. H. x 402.

spathysii and Riccia spp. It is known from Namibia, the

Northwest, Northern and Eastern Transvaal, Gauteng

(PWVi. Oranee Free State. Lesotho. KwaZulu-Natal, as

FIGURE 3. — Distribution of Plagiochasma rupestre var. rupestre, •;

and P. rupestre var. volkii, □, in southern Africa.

well as from Northern. Western and Eastern Cape (Figure

3). Its range in Africa extends northward into Zimbabwe

from where it was also reported by Best (1990) and by

Bischler (1978), who reported it from Kenya, Tanzania,

Uganda. Ethiopia, Djibouti, Sudan, Chad. Morocco, Al-

geria and Angola, as well as from the islands of Madeira,

Azores, Ascension, Cape Verde, St Helena and Reunion

(Bischler 1990).

Plagiochasma rupestre var. rupestre is easily identified

by its dull, velvety and glaucous thalli with simple, very

inconspicuous pores and by its reddish pink or purple

scales, with ovate-lanceolate or acuminate appendages, the

margins of which are entire.

lb. P. rupestre var. volkii Bischl. Type: Namibia,

Neudamm bei Windhoek, Volk 948 (JE).

Purple ventral scales larger (up to 3000 x 1350 pm)

and more conspicuous than those of typical variety, espe-

cially 2 or 3 hyaline appendages, which are narrowly tri-

angular, 1370-1450 pm long and 4 or 5 times longer than

18

Bothalia 25,1 (1995)

FIGURE4.— Plagiochasmaappendicululum. A-D, thallus: A, dorsal lace with archegoniophore and earpocephalum; B, dorsal face with androecium;

C, ventral face; D, transverse section. E, transverse section of air pore, dorsal cells and air chambers; F, air pore and surrounding cells from

above; G, margin of thallus; H, I, scales; J, scale appendage; K, transverse section of stalk; L, male paleae; M, female paleae. A, C, F, G, K,

M, Bottomley CM 268\ B, D, E, H-J, L, S.M. Perold 854. sValc bars: A-C, 2 mm; D, 1 mm; E, F, G, 50 pm; H-M, 250 pm.

Bothalia 25,1 (1995)

19

wide (Figure lO, P). Apex rather fragile, 3-5 elongated

cells in series, walls thickened; cells at rounded margin

of body of scale irregular in shape and size, cross walls

often oblique; male paleae ± 600 pm long, 150 pm wide

at base, tapering to a narrow tip where up to 4 cells, 50

x 15 pm, are in series (Figure IQ); female paleae up to

1500 pm long, 100-150 pm wide at base, tapering to a

narrow tip with 3 or 4 cells, 37.5^42.5 x 12.5-17.5 pm,

in series (Figure 1R); spores (Figure 2F-H) 92.5-105.0

pm in diameter, slightly larger but very similar in ap-

pearance to those of typical variety.

This variety is quite rarely collected in southern Africa,

but fairly frequently both varieties grow together. In the

present investigation specimens of P. rupestre var. volkii

from Namibia, the Northwest, Gauteng (PWV), Northern

Transvaal, as well as KwaZulu-Natal have been examined

(Figure 3). Bischler (1978) had also studied plants from

the Western Cape, Orange Free State, Lesotho and Zim-

babwe, so that it occurs throughout most of southern

Africa. Schuster (1992) states that P. rupestre var. volkii

also occurs in Peru and Argentina and it is thus not en-

demic to southern Africa.

In P. rupestre var. volkii the thallus is generally some-

what narrower than in the typical variety, but otherwise

it is very similar in colour, appearance and composition.

The very long, decolorate scale appendages are con-

spicuous, however, and make it easily separable.

Subgenus Plagiochasma

Most species are assigned to subgenus Plagiochasma ,

which is characterized by less compact green or yellow-

green thalli, dorsally with quite large, raised air pores,

surrounded by a hyaline ring and 2 or 3 concentric rings

of (5)6-8 cells in each, radial walls generally forming

continuous radiating lines that can be somewhat thick-

ened; air chamber walls faintly visible from above and

scale and appendage margins differentiated with smaller

cells, teeth or papillae.

2. Plagiochasma appendiculatum Lehm. & Lin-

denb. in Lehmann, Novarum et minus cognitarum stir-

pium pugillus quartus: 14 (1832); Gottsche et ah: 517

(1844-1847); Steph.: 782 (1898); Kashyap: 318 (1914);

Bischl.: 228 (1978). Type: Nepal, Punjab, Dehra Doon,

Wallich.

P. appendiculatum Lehm. & Lindenb. var. erythraeum Gola: 62

(1914). Type: Ethiopia, Eritrea, Hamasen, sul Monte Bizen nella valle

Nabaret a Mai Electi, Ragazzi 253.

P. fischerianum (Steph.) Steph.: 786 (1898) (P. fischeri). Type: Kenya,

Ligaijo, Fischer 692 [as Aitonia fischeriana Steph.: 301 (1895)].

For a complete synonymy see Bischler (1978).

Thallus large, robust, flat with sides sometimes curved

slightly downwards or upwards, broadly Ungulate (Figure

4A, B), bright green, shiny, with narrow purple edge along

margins, pores visible, small and slightly raised, when

wet; thallus margins incurved or indexed, exposing shiny,

reddish purple to deep purple transversely striate and

wrinkled underside of wings, not covered by scales, when

dry; in crowded, gregarious patches, simple or once.

sometimes twice furcate, rarely jointed with apical or

latero- ventral innovations. Branches 10-20 x 5-8 mm,

750-925 pm thick over midrib, laterally thinning out into

attenuate wings (Figure 4D); apex notched, with large or-

bicular reddish or partly hyaline scale appendages

recurved over edge in 2 layers; margins acute, thin, scal-

loped and slightly undulate; flanks sloping very obliquely,

reddish or purple; ventral face medianly keeled, green,

with row of deep red or purple red scales on either side

(Figure 4C). Dorsal epidermal cells unistratose, hyaline,

± rectangular to polygonal, 22.5^42.5 x 15.0-27.5 pm,

walls thin but thickened at corners, in transverse section

30.0-37.5 pm thick, smooth externally, along margins 2

or more rows of cells, rectangular, up to 25 x 12 pm or

shorter than broad, ± 15 x 22 pm (Figure 4G); air pores

not so numerous, 125-225 pm distant from each other,

slightly raised (Figure 4E), simple, 7.5-10.0 pm wide, sur-

rounded by an innermost ring, ± 2.5 pm wide, of tiny

collapsed cells and then by 2 (occasionally partly by 3)

concentric rings of larger cells, 5 or 6 inner ones transver-

sely oval or round, 10-15 x 15-20 pm (Figure 4F) partly

overlying outer row of 5 or 6 bluntly triangular cells, ±

22.5 x 30.0 pm across widest part, radial walls not thick-

ened. Assimilation tissue 375-500 pm thick, ± Vi thick-

ness of thallus, air chambers empty, in several layers,

upper ones ± 25 pm wide, lower down wider, ± 62.5 pm

wide, cells in bounding walls 30-45(-50) x 22-37 pm,

some with a brown oil body, 25-30 pm wide; storage

tissue occupying ventral Vl of thickness of thallus, cells

angular, up to 45 pm wide, with spaces between them,

some smaller cells also with oil bodies; rhizoids either

smooth, 12.5-25.0 pm wide, or pegged, 12.5 pm wide.

Scales deep red, appendages mostly decolorate and some-

times base as well, arranged in 2 forwardly directed

ventral rows, one on either side of midrib, asymmetric,

obtusely triangular with flatly arched base, gradually nar-

rowed above, deeply constricted and folded where joined

with large, mostly single, orbicular appendage (Figure 4H,

I), the latter up to 750 pm long, 550 pm across widest

part in middle, 300-375 pm wide at base, at margin 1 or

2 rows of small rectangular cells 10.0-17.5 x 7.5-12.5

pm, alternating with somewhat larger cells, in centre of

appendage toward base, cells large, rectangular, ± 75.0 x

37.5 pm, surrounded by several rows of irregularly shaped

cells, variable in size (Figure 4J); body of scale up to

1250 pm long, 1 100 pm across base, cells rectangular or

5-sided, ± 70 x 25 pm, 6 or 8 smaller, scattered ones with

remains of oil bodies, ± 27.5 x 20.0 pm; at margins cells

small, ± 25.0 x 12.5 pm, walls thin, curved, occasionally

with long, projecting papillae.

Monoicous, but male and female receptacles often on

separate plants. Androecia in sessile cushions, oval, horse-

shoe- or heart-shaped, 1. 5-2.5 x 2.0 mm, on leading

branch medianly, near apex, proximally partly surrounded

by shallow curved groove in thallus, base encircled by

blunt, hyaline or partly purple paleae, 550-580 x 130-180

pm (Figure 4L), cells rectangular or 5-sided, ± 57.5 x

20.0 pm, toward apex smaller, quadrate, ± 25 x 25 pm,

near to base margins with some projecting papillae, 25.0

x 12.5 pm. Archegoniophores single or several in acrope-

tal sequence medianly along main branch, initially

enclosed by arching hyaline paleae, ± 850 x 120 pm (Fig-

ure 4M), lower cells mostly rectangular. 40-45 x 22 pm,

toward apex smaller, ± 20 x 15 pm and at margin 15 x

20

Bothalia 25,1 (1995)

FIGURE 5. — Plagiochasma appendiculatum. SEM micrographs of spores and elater, A-F. A, distal face; B, much enlarged view of some areolae,

walls, wing and pore on distal face; C, side view, distal face above; D, proximal face; E, side view, proximal face above; F, part of elater

much enlarged. A, F, Hook 8204: B-E, S.M. Perold & M. Koekemoer 3135. A, x 5 1 1 ; B. x 959; C, x 577; D, E. x 534; F, x 940.

25 pm, lacking papillae. Carpocephala 2x2 mm when

4 lobes present, raised on stalk. 1.25-2.00 mm long, ±

750 pm in diameter (Figure 4K), in transverse section 1

or 2 rows of cortical cells, 17.5-27.5 x 17.5-30 pm,

medullary cells angular, up to 37.5 pm wide, thin-walled.

Spores 75-85 pm in diameter, triangular-globular, polar,

light brown, translucent, wing ± 10 pm wide, porate at

corners, margin undulate, finely crenulate; ornamentation

similar on both faces: distal face (Figure 5A-C) with 5

areolae across, ±22 pm wide, walls finely granular;

proximal face (Figure 5D, E) with very narrow triradiate

mark, each of 3 facets with 6 or 7 areolae, walls wide

and studded with granules. Elaters pale brown, 187-240

pm long, 10 pm wide in middle, tapering toward ends, 5

pm wide, bi- or tri-spiral (Figure 5F). Chromosome num-

ber: n = 9 (Bischler 1978).

After P. rupestre, P. appendiculatum was one of the

earliest species in the genus to be described. It is generally

a large plant and is easily recognized by its mostly single,

very rarely double, large, orbicular scale appendages.

Bischler (1978) examined a large number of specimens

and found quite considerable variation in the size of the

thallus and spores as well as in the thickness of the radiat-

ing cell walls at the air pores. In the present study only

three southern African specimens were available for study;

a fourth one is from Zimbabwe and was collected at Es-

sexvale by Borle (also reported by Bischler), but the exact

locality could not be traced. Three other specimens seen,

were from elsewhere. Plagiochasma appendiculatum has

not previously been reported from southern Africa, al-

though it had been collected by Bottomley in 1930 at

Pelindaba, the specimen being held at PRE. Unfortunately

it was misidentified as a species of Asterella and was not

sent on loan to Bischler, when she was revising the genus.

Recently I have collected specimens of it twice (the

second time with M. Koekemoer) in Bakker’s Pass, east

of Thabazimbi in southwestern Northern Transvaal (Fig-

ure 6), where it grew between boulders in a shady, damp

gulley, together with Fissidens sp. and Philonotis sp. Ar-

nell (1963) refers to P. fischeri, the synonym of P. appen-

diculatum., in his key to the species of Plagiochasma , but

there is no description or illustration of the species.

Plagiochasma appendiculatum is chiefly an Asiatic

species and has been reported from Afghanistan, Burma,

Celebes, China, Taiwan [Formosa], India, Kashmir, Nepal,

Pakistan, Philippines, Sikkim and Vietnam by Bischler

(1978). She has also reported it from Yemen and Socotra

as well as from Ethiopia, Kenya and Zimbabwe, where it

is rare. Bizot et at. (1985) reported it as new from Tan-

zania and regard it as a palaeotropical species. Frey &

F1GURE6. — Distribution ol /J. appendiculatum, O; P. beccarianum , LI;

P. eximium , •; and P. microcephamn var. microcephalum , ▲.

in southern Africa.

Bothalia 25,1 (1995)

21

Kiirschner (1988) also reported it from the Arabian Penin-

sula and from Socotra; they assume that it is of xerother-

mic Pangaean origin.

3. Plagiochasma microcephalum (Steph.) Steph. in

Bulletin de l'Herbier Boissier, Ser. 1, Vol. 6.10: 781

(1898); Bischl.: 237 (1978); Volk: 240 (1979).

P. dinteri Steph.: 762 (1901b). Type: Namibia (= Deutsch-Siidwest-

Afrika). Hereroland: Kransfontein. leg. Dinter.

Aitonia microcephala Steph.: 301 (1895). Type: Tanzania, Usambara,

leg. Holst 362 (G. holo. ! ).

Thallus smallish to medium-sized, flat to slightly con-

cave medianly. with the sides somewhat raised or gently

arching downwards, ligulate to oblong (Figure 7A), bright

green, along margins red-brown to purple; finely and ir-

regularly areolate, with pores faintly visible, when wet;

thallus margins tightly inflexed, exposing shiny, deep

purple or dark red transversely striate underside of wings,

not covered by scales except partly toward apex, when

dry; in crowded, gregarious patches, simple to several

times pseudodichotomously furcate, occasionally with

apical or latero-ventral innovations. Branches 6-12 x 3-

4(— 6) mm, 450-850 pm thick over midrib, laterally thin-

ning out into attenuate wings (Figure 7E); apex notched

with broadly ovate, purple or partly decolorate appendages

recurved over edge; margins acute, thin, somewhat

transversely pleated or crinkled, scalloped; Hanks sloping

obliquely, deep purple or dark red, ventral face medianly

keeled, green, partly obscured by 2 rows of purple scales,

one on either side (Figure 7B).

Dorsal epidermal cells unistratose, hyaline, 5- or 6-

sided or polygonal, 22.5-35.0(-40.0) x 17.5-25.0 pm,

rarely containing an oil body, walls thin, except at comers,

where thickened and rounded with trigones, in transverse

section 25.0— 32.5(— 37.5) pm thick, smooth externally,

along margins generally 2 or 3(4) rows of ± quadrate or

transversely rectangular cells, 12.5-20.0 x 12.5-25.0 pm

(Figure 7H); air pores varying from not so numerous to

numerous, distances between them ( 5 0— ) 1 00— 185 pm,

slightly raised (Figure 7F), simple, round or oval. 12.5 x

7.5- 10.0 pm, surrounded by an innermost ring, ± 2.5 pm

wide, of small collapsed cells and then by 2 concentric

rings of 5-7(8) cells in each (Figure 7G), inner cells

transversely oval, 12.5-17.5 x 15.0-20.0 pm, partly over-

lying next row of bluntly triangular cells, 12.5-22.5 pm

long, 25.0-32.5 pm across widest part, narrowing to base

of ± 12.5 pm wide, radial walls sometimes thickened (Fig-

ure 7G). Assimilation tissue 200-400 pm thick, ± Vl thick-

ness of thallus, air chambers empty, in several layers,

upper ones smaller, 30-50 pm wide, lower down up to

85 pm wide, cells in bounding walls 20.0-37.5 x 27.5-

35.0 pm, some with a spherical oil body almost filling

cell; storage tissue occupying ventral Vl of thickness of

thallus, cells angular, closely packed, 35.0-52.5 x 35.0-

42.0 pm, a few scattered ones also containing an oil body;

rhizoids either smooth, 17.5-25.0 pm wide, or pegged.

12.5- 17.5 pm wide. Scales and appendages purple or red-

dish or partly decolorate, arranged in 2 forwardly directed

ventral rows, one on either side of midrib, asymmetric,

obliquely triangular with base hardly arched, narrowed

above into 1 (Figure 7J) or sometimes 2 (Figure 71) broad-

ly ovate or triangular appendages (Figure 7K. L), equal

in size or not, 350-600 pm long, 350-450 pm across mid-

dle, constricted or folded or transversely pleated at base

where joined to rest of scale, cells toward centre quadrate

or rectangular and fairly regular, 25.0-40.0 x 22.5-37.5

pm, apically small and blunt, 10 x 10 pm, 1 or 2 rows

along margins ± 17.5 x 20.0 pm, some transversely rec-

tangular, ± 15 x 35 pm, thinner-walled, occasionally with

an outwardly projecting papilla; body of scale up to 675

pm long, 800-1500 pm across base, cells 4—6-sided, 37.5-

70.0 x 25.0-30.0 pm, ± 10 with remains of oil body, 42.5

x 32.5 pm, cells smaller and thinner-walled toward mar-

gins, sometimes curved or bent, a few with projecting

papilla.

Monoicous. Androecia in small sessile cushions,

rounded or V-shaped. ± 1 mm wide, medianly along

length of thallus and alternating with archegoniophores,

occasionally paired at pseudodichotomy of branches, an-

theridia opening above via prominent papillae, basally

cushions encircled by shallow groove in thallus and sur-

rounded by inconspicuous purple paleae that taper slightly

toward apex (Figure 7M), ± 400 x 200 pm. cells 32.5-

37.5 x 15.0-20.0 pm, several mucilage papillae projecting

along margins and from apex. Archegoniophores acrope-

tally arranged along length of thallus and alternating with

androecia, enclosed by arching, hyaline or purple paleae,

± 500 x 150 pm, tapering toward apex (Figure 7N), cells

rectangular, 35.0-45.0 x 17.5-22.5 pm, sometimes with

a few mucilage papillae at margins. Carpocephala small,

1.0 x 2.0 mm, when 2 lobes present (Figure 1C), raised

on stalk, up to 15 mm long and variously twisted, striate

and dark red, ±310 pm in diameter, in transverse section

(Figure 7D) cortical cells in 1 row, thick-walled externally

and rounded, 12.5-17.5 x 12.5-15.0 pm, medullary cells

thin-walled, angular, 12.5-17.5 pm wide. Spores 70-75

pm in diameter, triangular-globular, polar, light brown,

translucent, wing ± 10 pm wide, pore at comers occa-

sionally present, margins undulate, finely crenulate, or-

namentation similar on both faces: distal face (Figure 8A,

B) with 4 areolae across, 15-20 pm wide, walls narrow,

only ±2.5 pm wide, almost smooth to granular; proximal

face with distinct triradiate mark, arms thin, 5. 0-7. 5 pm

high, 4 areolae on each of 3 facets, walls almost smooth

(Figure 8C-E) to granular. Elaters light brown, 212-300

pm long, 7.5 pm wide in middle, tapering toward ends,

± 5.0 pm wide, bi-or trispiral (Figure 8F). Chromosome

number: n = 9 (Bischler 1978).

Bischler (1978) recognized two varieties of P. micro-

cephalum, namely var. microcephalum and var. tunesicum

Bischl. The latter appears to be restricted to Tunisia,

whereas she reported the former from the following

African countries: Ethiopia, LIganda, Angola, Namibia and

South Africa. It is also known from Madagascar, Yemen

and southwest India. In the present investigation, a few

more specimens from Northern Transvaal, Gauteng

(PWV), the Northwest and Northern Cape have been iden-

tified (Figure 6). It can be distinguished by the green

colour of the fresh thallus and by the generally large,

broadly ovate or triangular scale appendages which have

one or two rows of smaller cells along the margin.

Bischler (1978) reported that plants growing in dry

habitats have scales with a single, broadly ovate ap-

pendage which is constricted at the base, whereas those

from more humid sites usually have one or two triangular

22

Bothalia 25,1 (1995)

FIGURE 7. — Plagiochasma microcephalum var. mierocephalum. A, dorsal face of thallus with young receptacles; B, ventral face of thallus; C,

carpocephalum on stalk; D, transverse section of stalk; E, transverse section of thallus; F, transverse section of air pore, dorsal cells and air

chambers; G, air pore and dorsal cells from above; H, margin of thallus; I, scale with 2 appendages; J, scale with single appendage; K, L,

scale appendages; M, male paleae; N, female paleae. A, B, F, G, Candy 47; C, D, Holst 362; E, S.M. Perold 2976 ; H-K, M, N, Bottomley CH

175; L, Burn Davy 15176. Scale bars: A-C, E, I mm; D, 100 pm; F-H, 50 pm; I-N, 250 pm.

Bothalia 25,1 (1995)

23

FIGURE 8. — Plagiochasma microcephalum var. microcephalum. SEM micrographs of spores and elater. A, distal face; B. much enlarged view of

some areolae and walls on distal face; C, E, side view; D, proximal face; F, part of elater much enlarged. A-F, Holst 362. A, x 600; B, x 1357;

C, x 679; D, x 585; E, x 647; F, x 940.

scale appendages horizontally folded at the base and hard-

ly constricted. My observations appear to confirm this.

This species is equally well adapted to dry or to damp

habitats. It grows in dry river beds between limestone

boulders or on granite hills or on damp stream banks, but

has not been frequently collected. Plants with mature

female receptacles are rare. The carpocephala are small

(hence the specific epithet) and are raised on a slender,

short or long, dark red stalk.

Frey & Kiirschner (1988) regard it as a palaeotropical

taxon with a tropical African distribution. Sim (1926)

stated that P. dinteri (placed in synonymy under P.

microcephalum by Bischler ( 1978)] was not known to him

and he quoted a few particulars from Stephani (1901b).

Arnelfs (1963) description of P. dinteri contains several

inexactitudes as already observed by Bischler (1978): thal-

lus dimensions ‘15 cm long and 16 mm wide; peduncle

apical, spores 40-50 pm in diameter, without a wing or

with a rudimentary one, occasionally with a wing up to

4 pm wide; surface irregularly areolatef One of the

specimens seen by him [Essendale (sic)] is the Borle col-

lection from Essexdale and belongs to P. appendiculatum.

The collection from Hennops River, of which he illustrates

the spores, has not been traced. The Holst specimen (the

type of P. microcephalum) was seen by Amell and under

P. tenue he states that it belongs to another species, which

is certainly correct, since P. tenue is a synonym of P.

rupestre , but he failed to notice the similarity between P.

microcephalum and P. dinteri.

4. Plagiochasma eximium (Schiffn.) Steph. in Bul-

letin de l'Herbier Boissier, Ser. 1, Vol. 6.10: 781 (1898);

Bischl.: 248 (1978).

P. schimperi Steph.: 788 (1898). Type: Ethiopia (Abyssinia). In monte

Semen, leg. Schimper.

Aitonia eximia Schiffn. in Steph.: 300 (1895). Type: Kamerun, Buea,

an den Hohlen eine Stunde ostiich von Manus-Quelle, 2 500 m, 4-II-

1891, leg. Preuss 731 (FI, lecto. !).

Grimaldia abyssinica Gola: 63 (1914). Type: Abyssinia, leg. Chioven-

da 2993 (FI).

Thallus large, robust, ± Hat to sometimes medianly

concave with sides raised or slightly incurved, broadly

ovate (Figure 9B) to lingulate (Figure 9C), yellow green,

along margins narrowly to widely dark red; finely and

irregularly areolate with pores slightly raised, when wet;

thallus margins tightly inflexed or incurved, exposing

shiny, dark red, somewhat wrinkled and transversely

striate underside of wings, ventrally covered in wine red

or deep pink scales, when dry; in crowded, gregarious

patches, simple or once pseudodichotomously furcate

(Figure 9A), sometimes with one or more apical innova-

tions and articulated (Figure 9B). Branches 16.5-22.0 x

5.0-8.5 mm, ± 750 pm thick over midrib, laterally thin-

ning out into attenuate wings (Figure 9E); apex notched

with lanceolate, reddish pink or dark red scale appendages

recurved over edge; margins acute, thin, scalloped and

undulate; flanks sloping obliquely, dark red, ventral face

medianly keeled, green, with a row of deep red scales on

either side (Figure 9C).

Dorsal epidermal cells unistratose, hyaline, oblong,

ovate or polygonal, walls with trigones at comers, 25-40

x 20-35 pm, in transverse section 30.0-37.5 pm thick,

smooth externally, along margins 2(3) rows of cells (Fig-

ure 9H), rectangular, 25-35 x 10-20 pm, or shorter than

broad, ± 17.5 x 35.0 pm; air pores not numerous, distan-

ces between them variable, 230-440 pm, slightly raised

(Figure 9F), simple, round or oval, 12.5-27.5 x 15.0-27.5

pm, surrounded by an innermost ring, ± 7.5 pm wide, of

24

Bothalia 25.1 (1995)

FIGURE 9 - Plagiochasma eximium. A, lhallus willi 2 carpocephala; B, thallus with androecium; C, ventral face of thallus; D, transverse section of

stalk; E, transverse section of thallus; F, transverse section of air pore, dorsal cells and air chambers; G, air pore and dorsal cells from above;

H. margin of lhallus; I, scale with single appendage; I, scale with 2 appendages; K, scale appendage; L, M, male paleae; N, female paleae.

A-C, L. S.M. Perolcl 249S\ D-I, N, Sim CH 1163; .1, K, H. Anderson CH 4498; M, Sim CH 1200. Scale bars: A-C, 2 mm; D, 100 pm; E, 1

mm; F-H, 50 pm; I-N, 250 pm.

Bothalia 25,1 (1995)

25

small collapsed cells and then by 3 concentric rings of 7

or 8 larger cells in each (Figure 9G), inner cells transver-

sely oval, ± 12.5 x 17.5 pm, partly overlying next row

of transversely oblong cells, ± 12.5-15.0 x 25.0-32.5 pm,

outermost ring of cells ± 17.5 x 40.0-45.0 pm, radial

walls sometimes thickened. Assimilation tissue 375-475

pm thick, ± 1/2 or more than 1/2 thickness of thallus, air

chambers empty, in several layers, upper ones ± 35 pm

wide, lower down ±110 pm wide, cells in bounding walls

22.5-27.5 x 25.0-27.5 pm, gradually enlarging to 42.5 x

32.5 pm, some with spherical dark brown oil body, ± 32.5

pm in diameter, almost filling cell, these also present in

storage tissue, which occupies ventral part of thallus, cells

angular, up to 37.5 pm wide; rhizoids some smooth. 1 2.5—

17.5 pm wide, others pegged, 10.0-17.5 pm wide. Scales

and appendages deep pink to wine red, arranged in 2 for-

wardly directed ventral rows, one on either side of midrib,

asymmetric, obtusely triangular with arched base and nar-

rowed above into 1 (Figure 91) or 2 (Figure 9J) and some-

times 3 tapering appendages, 650-900 pm long, slightly

constricted or pleated at base (Figure 9K), 250—350(^400 )

pm wide, cells long-rectangular or hexagonal, 37.5-55.0

x 20.0-25.0 pm, apex not acuminate but often unicellular,

20.0-30.0 x 20.0-27.5 pm; body of scale ± 950 pm long,

1650 pm across base, cells 4—6-sided. 37.5-62.5 x 27.5-

35.0 pm, ± 14 with remains of oil body, scattered and

smaller, 25.0 x 22.5 pm, at margins cells narrowly rec-

tangular, ± 22.5 x 1 0.0 pm, occasionally with a projecting

papilla, 15 x 15 pm.

Monoicous, but male and female receptacles some-

times on separate plants. Androecia in sessile cushions,

kidney- or sausage-shaped, 1 mm wide, medianly at base

of smaller articulated, apical innovation (Figure 9B) of

leading branch, proximally partly surrounded by shallow

curved groove in thallus, encircled by apically tapered.

dark red paleae, 250-500 x (90—) 1 00— 1 20 pm, cells rec-

tangular, 40-55 x 17-20 pm, smaller toward apex, 32.5

x 15.0 pm, and along upper part of margin 25.0-37.5 x

12.5-15.0 pm, sometimes with a projecting papilla sup-

ported on a narrow base and also above at apex. Ar-

che goniophores single or occasionally paired when

branches bifurcate (Figure 9A), medianly near apex of

branch, enclosed by arching, hyaline paleae, ± 550 x 80

pm, tapering toward apex with single cell. ± 22.5 x 12.5

pm, lower down cells larger, rectangular, up to 42.5 x

15.0 pm, margins sometimes with a papilla. Carpocephala

3.25 x 3.25 mm, when 4 lobes present, raised on stalk.

3 — 6(— 9) mm long, striate and dark red, in transverse sec-

tion (Figure 9D) ± 850 x 750 pm, cortical cells in 1 row,

± 20.0 x 17.5 pm, thick-walled externally, medullary cells

thin-walled, angular, up to 37.5 x 30.0 pm. Spores 82.5-

90.0 pm in diameter, triangular-globular, polar, golden

brown, translucent, wing up to 12.5 pm wide, porate at

comers, margins undulate, finely crenulate; ornamentation

similar on both faces: distal face (Figure 10A-C) with 4

or 5 areolae across, 22.5-32.5 pm wide, walls wide and

finely granular; proximal face (Figure 10D, E) with dis-

tinct triradiate mark, arms ± 5 pm high, mostly 4 shallow

areolae on each of 3 facets, walls sparingly sprinkled with

granules. Elaters light brown, 175-225 pm long, 15-20

pm wide in middle, tapering toward ends, 5.0-7. 5 pm

wide, laxly bispiral or spirals interrupted (Figure 10F).

Chromosome number: n = 9 (Bischler 1978).

Plagiochasma eximium has previously been reported

from southern Africa by Bischler (1978), who had ex-

amined a specimen, Sim CH 1163 , from Mont-aux-Sour-

ces which had earlier been identified as P. rupestre. Two

other specimens, Sim CH 1200 (Pietermaritzburg) and Sim

CH 1186 (Victoria Falls), have now been identified as P.

eximium, as well as some later collections from the

FIGURE 1 0. — Plagiochasma eximium. SEM micrographs of spores and elater. A, B, distal face; C, much enlarged view of margin, some areolae and

walls on distal face; D. proximal face; E, side view of proximal face; F, elater. A, C, D, T.R. Sim CH 1 163: B. E, F. T.R. Sim CH 1186. A, x

470; B, x 482; C, x 925; D, x 459; E, x 501; F, x 308.

26

Bothalia 25,1 (1995)

FIGURE 1 I . — Plagiochasmabeccarianum. A, thallus with mature carpocephalum; B, thallus with archegoniophore and androecium; C, ventral face

of thallus; D, transverse section of stalk; E, transverse section of thallus; F, transverse section of air pore, dorsal cells and air chambers; G,

air pore and surrounding cells from above; H, margin of thallus; I, scale with 2 appendages; J, scale with 3 appendages; K, scale appendage;

L, male paleae; M, female paleae. A, B, D-K, M, Volk 00950\ C, S.M. Perold 2992 ; L, S.M. Perold 2995. Scale bars; A-C, 2 mm; D, 100

pm; E, I mm; F-H, 50 pm; I-M, 250 pm.

Bothalia 25,1 (1995)

27

Drakensberg by H. Anderson, E. Esterhuysen, S.M. Perold

and O.H. Volk (Figure 6). Amell (1963) gave no descrip-

tion or illustration of P. eximium , merely listing it in his

key to the species of Plagiochasma. This species is

widespread in Africa and has been reported by Bischler

(1978) from Sierra Leone, Guinea, Cameroons, Zaire,

Djibouti, Ethiopia, Uganda. Kenya, Tanzania, Malawi and

the following islands: Cape Verde, Reunion and Madagas-

car (Bischler 1990). Its distribution extends to the Arabian

Peninsula: Saudi Arabia, Yemen, Oman and to Socotra.

Frey & Kiirschner (1988) consider P. eximium to be a

palaeotropical taxon with tropical African distribution.

Many of the collections are from high altitudes, such

as the Drakensberg in southern Africa, where the plants

grow in shady kloofs on muddy rock faces or on soil

covering rocks or under boulders. As far as could be as-

certained, the specimen, Sim CH 1186 , from Victoria Falls

seems to be the first record of this species from Zim-

babwe.

Plagiochasma eximium can be distinguished by its

robust, yellow-green thalli with large, deep pink to wine-

red ventral scales narrowed above into ( 1 )2 or 3 tapering

appendages which are only a little constricted or pleated

at the base. Ventrally the flanks are wrinkled and deep

red, not dark purple. Elaters from the few spore-bearing

plants studied are bispiral, but Bischler (1978) also found

them to be 3- or 4-spiral and generally distinct along only

a small part of their length.

Bischler (1978) states that herbarium specimens of P.

eximium can look quite different from each other. Those

from damp, shady places (e.g. a cave) have thin, olive-

green thalli and decolorate margins, large epidermal cells

with thin walls, and smaller, pale scales. Others have

thicker, yellowish green thalli with pigmented margins,

slightly smaller epidermal cells with well-developed

trigones and large, dark red scales. Bischler (1978) con-

siders them to be ecological variants of the same species.

Other characters that are quite variable are the width of

the thalli, the thickness of the radial walls at the air pores,

the presence or absence of teeth at the base of the scale

appendages, the spore ornamentation and the presence or

absence of spiral thickenings in the elaters.

5. Plagiochasma beccarianum Steph. in Bulletin

de l'Herbier Boissier, Ser. 1, Vol. 6.10: 781 (1898);

Bischl.: 257 (1978); Volk: 240 (1979). Type: Abyssinia

(Bogos), Keren, in Monte Deban, inter 4500' & 55002

1870, Beccari (FI, holo.; 009583G, iso.!).

Thallus medium-sized to rather large, flat to very

slightly concave medianly and gently arching downward

toward margins, ligulate to lingulate or broadly ovate (Fig-

ure 11 A, B), bright green, shiny, along narrow edge of

margins purple; finely and irregularly areolate, pores dis-

tinctly visible, raised, when wet; thallus margins frequent-

ly tightly inflexed or sometimes incurved, exposing shiny,

ink-black or very dark purple, transversely striate under-

side of wings, partly covered by purple scales, when dry;

in crowded, gregarious patches, once or twice pseudo-

dichotomously furcate, sometimes with apical innovations

and then articulated. Branches 8.0-20.0 x 3.5-5.0(-6.0)

mm. 600-850 pm thick over midrib, laterally thinning out

into attenuate wings (Figure 1 1 E); apex notched, with

several purple ovate-lanceolate scale appendages recurved

over edge; margins acute, thin, sparingly scalloped and

slightly undulate; flanks sloping obliquely, black or

purple, ventral face medianly keeled and green, partly

obscured by 2 rows of purple scales, one on either side

(Figure 11C).

Dorsal epidermal cells unistratose, hyaline, rectangular

to polygonal, 25.0-37.5 x 20.0-27.5 pm, walls thin, ex-

cept at corners where thickened and rounded with

trigones, in transverse section 30-35 pm thick, smooth

externally, along margins (Figure 1 1 H) 2 or 3 rows of

short rectangular cells, ± 25.0 x 22.5 pm; air pores not

so numerous, distances between them quite variable. 1 37—

287 pm, slightly raised (Figure 1 1 F), simple, round or

oval, 15.0-30.0 x 17.5-25.0 pm, surrounded by an inner-

most hyaline ring, 5 pm wide, of collapsed cells, and then

by 2 (or partly by 3) concentric rings of 6-8 cells in each,

inner cells ± rounded or transversely oval, 10.0-15.0 x

15.0- 27.5 pm, partly overlying next row of bluntly trian-

gular cells, 12.5-17.5 x 25.0-35.0 pm, outermost row of

cells (if present), 20-25 x 35 pm, radial walls not thick-

ened (Figure 11G). Assimilation tissue 300-400 pm thick,

± 1/2 thickness of thallus, air chambers in several layers,

30.0- 57.5 pm wide, cells in bounding walls 22.5-32.5 x

22.5-27.5 pm. occasionally with dark brown oil body al-

most filling cell, 37.5 x 35.0 pm; storage tissue occupying

ventral part of thallus, cells transversely oval to rounded

or angular, 30.0-37.5 pm wide, walls mostly faintly red-

dish and thickened at corners, intercellular spaces here

and there, some cells containing an oil body; rhizoids

either smooth, 12.5-25.0 pm wide, or pegged, 10.0-12.5

pm wide. Scales and appendages deep purple to violet,

in 2 forwardly directed ventral rows, one on either side

of midrib, ± asymmetric, with flatly arched base, gradually

tapered above into 2 (Figure 111) or 3 (Figure 1 1 J) lan-

ceolate appendages up to 1000 pm long, not constricted

at base (Figure 1 1 K), ± 400 pm wide, cells mostly rec-

tangular to 6-sided, 62.5-100.0 x 27.5 -30.0 pm, at apex

often a single conical cell, 22.5 x 20.0 pm, at margins

lower down a few prominent teeth, ± 50 x 20 pm. some-

times basally supported on 1 or 2(3) cells in series; body

of scale ± 850 pm long, 1750 pm across base, cells 4-

or 6-sided, 50-75 x 25-30 pm, at margin narrower, ±

17.5 pm wide and thinner-walled, also present a few

(± 10) scattered smaller cells, which had contained an oil

body.

Monoicous, but receptacles often on separate plants.

Androecia in single sessile cushions, kidney-shaped or

rounded, up to 2 mm wide, medianly and generally just

proximal to apical innovation (Figure 1 IB), basally sur-

rounded by purple paleae, 300—100 x 160-200 pm, api-

cally pointed or blunt (Figure 1 1 L) , cells 32.5-37.5 x

15.0- 20.0 pm, occasionally containing an oil body and

some marginal ones with a papilla. Archegoniophores

generally single, medianly near apex of main branch (Fig-

ure 1 1 A) or on apical innovation, sometimes 2 in acropetal

sequence, basally surrounded by dark paleae with pointed

apex (Figure 11M), up to 500 x 80 pm, cells rectangular,

± 45.0 x 22.5 pm, sometimes a few papillae at marginal

cells. Carpocephala up to 3.5 x 3.5 mm, when 4 lobes

present (Figure 1 1A), raised on stalk, 1.2-3. 5 mm long,

in transverse section (Figure 1 ID), 825 x 675 pm, cortical

28

Bothalia 25.1 ( 1995)

cells in 1 row, thick-walled externally, 15-20 x 15-20

|inr, medullary cells thin-walled, rounded or polygonal

and then angled, up to 37.5 x 25.0 pm. Spores 80-90 pm

in diameter, triangular-globular, polar, light brown, trans-

lucent, wing 10 pm wide, porate at corners, margin finely

crenulate; ornamentation similar on both faces: distal face

(Figure 12A-C) with 3 or 4 areolae across, 20-25 pm

wide, walls ± 7.5 pm wide and finely granular; proximal

face (Figure 12D, E) with distinct triradiate mark, ridge

±7.5 pm high, each of 3 facets with 4 areolae, walls wide

and sprinkled with granules. Elaters (Figure 12F) light

brown. 195-258 pm long, 12.5 pm wide in middle, taper-

ing toward ends, 7.5 pm wide, occasionally branched,

bi spiral except at tips. Chromosome number, n = 9

(Bischler 1978).

Plagiochasma beccarianum is regarded as quite a

heterogeneous species, but so closely resembling P. ex-

imium that the two species are often difficult to distinguish

when sterile (Bischler 1978). In the few southern African

specimens of P. beccarianum available for study, it ap-

pears that the fresh thalli are a clear green dorsally, not

yellow-green, and the underside of the wings as well as

the scales are deep purple and not dark red as in P. ex-

imium. The two or three scale appendages of P. be-

ccarianum are ± acuminate or narrowly triangular and not

constricted at the base; their margins are irregularly

toothed.

This species is rare and lias been reported from rela-

tively few places in Africa; except for Namibia (Volk

00950), it is mostly found along the eastern part of the

continent, namely Ethiopia, Tanzania and Zambia

(Bischler 1978). It has also been reported from (he

Arabian Peninsula (Frey & Kiirschner 1988) and from

Socotra (Bischler 1978; Frey & Kiirschner 1988). Some

specimens from the Northern and Eastern Transvaal have

recently been collected (Figure 6), where they grew in a

shady kloof under boulders or in a rocky crevice in Blyde

River Canyon respectively. Frey & Kiirschner (1988)

regard it as a palaeotropical taxon with a tropical African

distribution.

Bischler (1978) states that P. beccarianum belongs to

a complex represented by several species in Asia and

America. She thinks that this group probably diversified

more rapidly on these two continents than in Africa and

that the heterogeneity of P beccarianum could be due to

a slower African evolution of the complex which has not

as yet achieved the separation of distinct taxonomic units.

SPECIMENS EXAMINED

(held at PRE, unless otherwise indicated)

H. Anderson 1230 , 1245, 1254 (la); CH 13477, CH 13512 (lb); CH

13588 (la); CH 4498-CH 4500 (4). T. Anderson 13 (la).

Badenhorst CH 4360 ( la). Bean & Oliver 2354 ( la). Bester 15 ( la). Bews

CH 1132, CH 1152, CH 1232 (la). Bode CH 1340 (2). Botha 136=145,

CH 11 70, CH 13240 (la). Bottomley CH 155, CH 160, CH 173 (lb); CH

175 (3); CH 1 98 (la); CH 268 (2); CH 269 pp. ( lb); CH 269, CH 3661

(la). Brueckner 225 (3) BOL. Brusse4123, 4124, 4126^1130, 4261 (la).

Burrows 2363, 2520 (la). Burtt Davy 15176 (3).

Condy 45 (la); 47 (3); 82, 83 (lb); 84 (3); CH 13629, CH 13631 (la).

Cooper 962 ( 1 a).

Dieterlen 793C (la). Doidge CH 169, CH 3601 (la). Du Preez 2107,

2108 (la).

Edwards CH 1153 ( la). Esterhuysen 26166A (la); 26166 (4) BOL. Eyles

937, 1181 (la).

Germishuizen 5393, 5385 ( la). Giess, Volk & Bleissner 6768 ( la, lb, 3).

Giffen 2 (CH 1228) ( la). Glen 1250 (5); 1644 (lb); 1722, 2241, 2846,

3171 ( la). Gotze 16 (la). Graham CH 1174 (la).

FIGURE 12. — Plagiochasma beccarianum. SEM micrographs of spores and elater. A, B, distal face; C, much enlarged view of margin, pore, some

areolae and walls; D, proximal face; E, side view; F, part of elater much enlarged. A, Glen 1250\ B-F, Volk 00950. A, x 437; B, E, x 472; C,

x 936; D, x 460; F, x 1014.

Bothalia 25,1 (1995)

29

Hansen 3241 ( 1 a). Hedberg 2150 BOL (4). Heilgendorff CH 13611 ( 1 a).

Hepburn 3 (= CH 1155) (la). Herman 338 (la). Heymann, Cloete &

Burgoyne 67 (lb); 68 (3). Hilton 54706 (la) BOL.

Jacot Guillarmod CH 3675, CH 3791, CH 4187 (la). Jacot Guillarmod,

Getliffe & Mzamane 114, 287 (la). Jensen 442 (la).

Koekemoer 104, 296 (la); 637 (lb); 842, 912, 1024a (la). Kreiner ( =

Volk 89-31) (lb).

Leighton 3284 (la) BOL. Liebenberg 7593 (la). Lyle 7085 (la).

Magill 6372, 6398. 6413. 6418. 6431, 6480 (la). Manning CH 13590

(la). Mogg 144 (la); CH 3720, 37590 ( lb). Moore CH 55 (la). Morley

358 (la). Moss CH 1256 (la). Mott 860 ( la).

Oliver 1449 BOL. 1464, 10133 (la). Ortendahl 696A (la).

Pearson 9849 (la). S.M. Perold 55 ( la); 176-178 (lb); 224 (3); 225 (lb,

3); 229 (3); 260, 301, 852 (la); 854 (2); 947, 948, 1292, 1297, 1477, 1504.

1514, 1876,2167, 2168 (la); 2498 (4); 2508 (la); 2586 (lb); 2799, 2803

(la) ; 2974 (lb); 2976 (3); 2984 (la); 2955 (lb); 2992, 2995 (5); 2997,

2998 (lb); 3002, 3058. 3062 (la). Perold & Koekemoer 2944, 2949,

2950, 2972 (la); 3135 (2). Perold, Koekemoer & Smook 3020 (la).

Pieterse 100 (la). Playford CH 1264 (la). Pole Evans CH 11, CH 14

(lb) ; CH 457, CH 1154 (la). Potts CH 1168, CH 1173, CH 1230, CH

1258 (la).

Retief 1543, 1675a (la). Retief & Germishuizen 170, 408 ( la). Retief &

Shearing 1228 (la).

Saaiman 309 (la). Schelpe s.n. ( BOL54717 ), 2019 (la); 4781 (lb); 4782.

4852 BOL, 5283, 5290 BOL (la); 5587 BOL (lb); 5823, 5834, 6360 ( 1 a).

Scott 21 (la). G.W. Sim 8109 ( la). T.R. Sim CH 1157 (la); CH 1163 (4);

CH 1166, CH 1180-CH 1185 (la); CH 1186 (4); CH 1189, CH 1199(\ a);

CH 1200 (4); CH 1201, CH 1209, CH 1225 (la). Smook 4425, 4486,

5176a, 7929, 7952, 8240a, 8639, 8745 (la).

Tidmarsh CH 3175 (la, lb).

University of Durban-Westville 2152 (la).

Vahrmeijer CH 13145 (la). Van der Bijl 498 (la). Van der Westhuizen &

Deetlefs 1 ( la) BOL. Van Rooy 690, 734, 779, 791, 1166, 1347, 1467,

2025, 2366, 2379. 2408, 2423, 2525, 2595, 2621. 2657, 2679, 2726, 2771,

2777, 2782, 3160, 3197, 3212, 3293. 3542 (la). CM. Van Wyk2679, 3190

(la). Venter 8612 (la). Viljoen CH 4524 (la). Vlok 2662 (la). Volk 212,

00482, 00554, 00639, 00684, 00909 (la); 00948 (lb); 00950 BOL (5);

01000. 01348 (la); 5020, 6105, 6657, 6854 ( lb); 6900, 11361, 11363,

11406, 12728, 81/063, 81/093 { la); 81/183 (3); 81/194, 81/274, 81/287

(la); 84/630, 84/650 (la, 4); 84/724 (la). Vorster 655 (la).

V. Wager 100 ( la). Welman CH 1141 (la).

Young CH 1164 (la).

ACKNOWLEDGEMENTS

I wish to sincerely thank Dr H. Bischler for refereeing

this article. Prof. O.H. Volk of Wurzburg University and

the curators of BOL and LISU for the loan of specimens;

also my colleagues at NB1 for collecting specimens, the

artist. Ms G. Condy. the typist, Mrs J. Mulvenna, and the

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

tributions.

REFERENCES

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

Science Council, Stockholm.

BEST. E.B. 1990. The Bryophyta of Zimbabwe — an annotated checklist.

Kirkia 13; 293-318.

BISCHLER. H. 1977. Plagiochasma Lehm. & Lindenb. I. Le genre et ses

subdivisions. Revue bryologique et lichenologique 43: 67-109.

BISCHLER, H. 1978. Plagiochasma Lehm. & Lindenb. II. Les taxa

europeens et africains. Revue bryologique el lichenologique 44:

223-300.

BISCHLER, H. 1979. Plagiochasma Lehm. & Lindenb. III. Les taxa

d’ Asie et d’Oceanie. Journal of the Hattori Botanical Laboratory

45: 25-79.

BISCHLER, H. 1990. Quelques Marchantiales de L'ile de la Reunion.

Cryptogamie, Btyologie et Lichenologie 11: 169-171.

BISCHLER, H. & SERGIO. C. 1984. Un neotype pour Plagiochasma

nipestre (J.R. & G. Forst.) Steph. Cryptogamie, Btyologie et

Lichenologie 5: 173, 174.

BISCHOFF, W. 1835. Bemerkungen liber die Lebermoose, vorziiglich

aus den Gruppen der Marchantieen und Riccieen. Nova Acta

Academiae Caesareae Leopoldino-Carolinae Germanicae Naturae

Curiosorum 17: 911-1088.

BIZOT. M„ POCS, T. & SHARP, A.J. 1985. Results of a bryogeographi-

cal expedition to East Africa in 1968, III. The Bryologist 88:

135-142.

EVANS, A.W. 1915. The genus Plagiochasma and its North American

species. Bulletin of the Torrey Botanical Club 42: 259-308.

FORSTER, J.R. & G. 1776. Characteres generum plantarum. White,

Cadell & Elmsly, London.

FORSTER, G. 1787. Plantae Atlanticae. Commentationes Societas

Regiae Scientiarum, Gottingen 9: 46-74.

FREY, W. & KURSCHNER. H. 1988. Bryophytes of the Arabian Penin-

sula and Socotra. Studies in Arabian bryophytes 12. Nova Hed-

wigia 46: 37-120.

GOLA, G. 1914. Epatiche dell’ Abissinia. Annali di Botanica (Rome) 13:

59-75.

GOTTSCHE, C M., LINDENBERG, J.B.G. & NEES AB ESENBECK,

C.G. 1 844—1847. Synopsis Hepaticarum. Hamburg, Meissner.

GROLLE, R. 1972. Die Namen der Familien und Unterfamilien der

Lebermoose (Hepaticopsida). Journal of Bryology 7: 201-236.

HASSEL DE MENENDEZ, G.G. 1963. Estudio de las Anthocerotales y

Marchantiales de la Argentina. Opera Lilloana 7: 1-297.

KASHYAP, S.R. 1914. Morphological and biological notes on new and

little known West Himalayan liverworts. II. New Phytologist 13:

308-323.

LEHMANN, J.G.C. 1832. Novarum et minus cognitarum stirpium pugil-

lus quartus. Meissner, Hamburg.

LEHMANN, J.G.C. 1838. Antrocephalus, eine neue Gattung der Leber-

moose aus der Gruppe der Marchantieen. Nova Acta Academiae

Caesareae Leopoldino-Carolinae Germanicae Naturae Curio-

sorum 18: 679-684.

LINDBERG, S.O. 1868. Musci novi scandinavici. Notiser Sdllskap pro

Fauna et Flora Fennica Forhandlingar 9: 255-299.

LINNAEUS, C. fil. 1781. Supplementum plantarum systematis vegetabi-

lium editionis decimae tertiae. Brunswiga.

LINNAEUS, C. fil. 1 783. Acta Med. Suecica 1 : 204.

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

Kryptogamen-Flora 6:41 6-47 1 .

NEES AB ESENBECK, C.G. 1838. Naturgeschichte der europaischen

Lebermoose 4: 1—540.

SCHIFFNER, V. 1893. Hepaticae. Naturliche Pflanzenfamilien 1: 3-141.

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

America. Vol. 6. Chicago.

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

Royal Society of South Africa 15: 1-475.

STEPH ANI, F. 1895. Hepaticae africanae. Botanische Jahrbiicherfiir Sys-

tematic Pflanzengeschichte und Pflanzengeographie 20: 299-301.

STEPHANI, F. 1898. Species hepaticarum: Plagiochasma. Bulletin de

I’Herbier Boissier, Ser. 1, Vol. 6,10: 775-790.

STEPHANI, F. 1901a. Hepatics. In Hieron, Catalogue of the African

plants collected by Dr F. Welwitsch in 1853-61, 2: 310-321.

STEPHANI, F. 1901b. Hepaticae. In H. Schinz, Beitrage zur Kenntnis der

Afrikanischen Flora (Neue Folge). XHI. Bulletin de I’Herbier

Boissier , Ser. 2, Vol. 1,8: 757-788.

THUNBERG, C.P. (‘1776’) 1780. In Physiographic Sdllskap Handlingar

1,3: 166.

TREVISAN DE SAINT-LEON. V. 1877. Schema di una nuova clas-

sificazione delle Epatiche. Memorie del Reale lstituto Lombardo

di Scienze e Lettere, Ser. 3,4: 383-451.

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

tiales) aus Siidwesafrika (Namibia). 1 . Mitteilungen der Botani-

schen Staatssammlung, Munchen 15: 223-242.

WEBB. P.B. & BERTHELOT, S. 1840. Plantes cellulaires. Histoire

Naturelle des lies Canaries 3,2. Bethune. Paris.

-

m

Bothalia 25,1: 31-33 (1995)

FSA contributions 2: Asphodelaceae/Aloaceae, 1029010 Chortolirion

G.F. SMITH*

Chortolirion A. Berger in Das Pflanzenreich 4, 38, III,

II (Heft 33): 72 (1908); E. Phillips: 149 (1926), 187

(1951); Oberm.: 119 (1973). Type: Haworthia angolensis

Baker, i.e. Chortolirion angolense (Baker) A. Berger.

Herbaceous perennial; acaulescent, with leaves origi-

nating from a short, simple (rarely once branched),

cylindrical, subterranean butt ± 9 mm in diameter; roots

few, fusiform, fleshy, up to 8 mm in diameter; bulb with

few, loosely packed, membranous scales covering the

inner, slightly fleshy leaf bases, ovoid-oblong, 30-40 mm

long, ± 20 mm in diameter. Leaves rosulate, slender, grass-

like, flaccid, erect, deciduous, ± 10, light green to

glaucous green, usually once or twice twisted, upper 5-10

mm of leaves often dry, ± 150 mm long, diameter ± 2

mm; upper surface canaliculate, immaculate or with very

few white spots near base; lower surface convex, copious-

ly white-spotted near base, the spots often slightly tuber-

culate-subspinulescent; margins armed with soft, white,

decurved teeth, ± 0.5 mm long, larger low down, smaller

upwards, 1-2 mm distant throughout. Inflorescence ± 360

mm tall; peduncle simple, diameter ± 2 mm, ± 200 mm

long; sterile bracts membranous, ovate, abruptly long

acuminate, erect, clasping the peduncle, keeled with 1-3

reddish brown vein(s), ± 8 mm long; raceme ± 150 mm

long, ± 14 spirally arranged flowers and buds, 3 open

simultaneously; floral bracts membranous, mucronate,

keeled, clasping the pedicels, longer than the pedicels, 5

mm long; pedicels erect, persistent, brownish green, 1-2

mm long, diameter 1 mm. Flowers erect, zygomorphic,

greenish, brownish or pinkish white with greenish keels

to the segments, base obtuse; perianth funnel-shaped, tube

straight, constricted to 3 mm above, 14 mm long, ± 2 mm

across; segments greenish white with darker green veins,

not free to the base, closely adhering for two thirds of the

length, limb bilabiate; upper-outer segments strongly

recurved, retuse, spoon-shaped at tips; upper-inner seg-

ment slightly recurved, obtuse, spoon-shaped at tip, lower-

outer segments recurved, lower-inner segment strongly

recurved, tips flared; bud narrow, straight, decurved and

pinkish at tip; stamens 6, of ± equal length, inserted within

the perianth tube, attached below ovary, ± 7 mm long;

filaments white, thinner towards apex; anthers yellow,

dorsifixed, dehiscing longitudinally and introrsely; ovary

green, sessile, 3 mm long, diameter 2 mm; style white,

straight, capitate, 4 mm long. Fruit light green, capsule

trilocular, cylindric, apically acute, dehiscing loculicidally,

chartaceous when dry, ± 15 mm long, 5-6 mm in

diameter. Seed dark brown to black, angled, shortly

winged, ± 3 mm long. Chromosome number. 2n = 14.

Figure 1.

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

MS. received: 1994-03-24.

Monotypic. In southern Africa it occurs in Namibia,

Botswana, Lesotho and in all the provinces of the

Republic of South Africa except the Western Cape. For

KwaZulu-Natal the genus is known from a single acces-

sion only [grid reference unknown: Zululand, Anon. s.n.

(K)!] which is not shown in Figure 2. It also occurs in

Angola and Zimbabwe. Chortolirion is found from near

sea level up to altitudes of more than 2 000 m; the general

habitat of Chortolirion is the climatically severe inland

area above the Great Escarpment. The genus is adapted

to sparse or dense grasslands in which a wide variety of

graminoids and forbs dominate. These grasslands are

usually subject to natural or deliberate seasonal burning.

Chortolirion is morphologically quite distinct from

Haworthia , especially with regard to the presence of an

underground bulbous rootstock. Furthermore, it is the only

haworthioid taxon of which the leaves are deciduous and

die back to ground level after fires or frost.

The name Chortolirion means ‘heath lily’ and refers

to the fact that plants of the genus usually occur in

grassland and, especially when not in Bower, can easily

be mistaken for small tufts of grass.

Chortolirion angolense (Baker) A. Berger in Das

Pflanzenreich 4, 38, III, II (Heft 33): 73 (1908). Type:

Angola, Huilla, regio subtemperata, in dumetis arenosis,

Welwitsch 3756 (BM, holo.!, PRE. photo.!).

Description as for the genus.

Haworthia angolensis Baker: 263 (1878); Baker: 210 (1880): Baker:

469 (1898); Oberm.: 119 (1973). Type: as above.

H. tenuifolia Engl: 2 (1888); Baker: 355 (1896). Chortolirion

tenuifolium (Engl.) A. Berger: 73 (1908). Type: Betschuanaland. Man-

jering pr. Kuruman, in arenosis alt. 1 200 m, Marloth 1049 (B, holo.!).

H. stenophylla Baker: sub t. 1974 (1891); Baker: 355 (1896). Chor-

tolirion stenophyllum (Baker) A. Berger: 72 (1908); Dyer: t. 932 (1944).

Type: Transvaal, grassy mountain slopes of the Saddleback range near

Barberton, Galpin 858 (K, holo.!).

H. saundersiae Baker: sub t. 1974 (1891). nom. nud.

H. subspicata Baker: 998 (1904). Chortolirion subspicatwn (Baker)

A. Berger: 74 (1908). Type: Transvaalkolonie, Modderfontein, Conrath

645 (Z, holo.!).

Chortolirion bergerianum Dinter: 24 ( 1914). Type: Deutsch-Sudwest-

Afrika (Namibia), the Farm Voigtland, 20 km to the east of Windhoek,

K. Dinter, Neue und wenig bekannte Pflanzen Deutsch-Siidwest-Affikas

t. 12 (1914) (holo.!, icono.!).

leones: Dyer: t. 932 (1944); Fabian & Germishuizen: t. 13a (1982).

Vouchers: Dinter 4295 (B); Hanekom 1843 (PRE); Smith 8 (PRU);

Smith 12 (PUC); Ubbink 318 (PUC).

32

Bothalia 25, 1 (1995)

FIGURE 1. — Chortolirion angolense: A.

habit, x 1 ; B, inflorescence, x I ; C,

longitudinal section of flower (one

stamen removed), x 3; D, flower

face showing reflexed segments, x

2; E, fruit, acuminate capsule, x

1.5. All drawings, except E, were

made from live material collected

by Craib, deposited under Smith

234 (PRE). Fruit drawn from

Leeman s.n. (PRE 34956). Artist:

Gillian Condy.

Bothalia 25,1 (1995)

33

FIGURE 2. — Distribution of Chortolirion angolense in southern Africa.

REFERENCES

BAKER, J.G. 1878. Report on the Liliaceae, Iridaceae, Hypoxidaceae,

and Haemodoraceae of Welwitsch’s Angolan herbarium. Transac-

tions of the Linnean Society of London, Botany , Ser. 2, 1 : 245-

273.

BAKER, J.G. 1880. A synopsis of Aloineae and Yuccoideae. Journal of

the Linnean Society, Botany 18: 148-241.

BAKER. J.G. 1891. Haworthia stenophylla. Baker. Liliaceae. Tribe

Aloineae. Hooker’s leones Plantarum 20, Ser. 3, 10: t. 1974.

BAKER, J.G. 1896. Liliaceae, Haworthia. Flora capensis 6,2: 332-355.

Reeve, London.

BAKER. J.G. 1898. Liliaceae, Haworthia. Flora of tropical Africa 7,3:

467. Reeve, London.

BAKER, J.G. 1904. Liliaceae. Beitrage zur Kenntnis der Afrikanischen

Hora, Neue Folge, XVI. Bulletin de L'Herbier Boissier, Ser. 2, 4:

996-1002.

BERGER, A. 1908. Liliaceae-Asphodeloideae-Aloineae. 3. Chor-

tolirion. Das Pflanzenreich 4, 38, III. II (Heft 33): 72-74. Wil-

helm Engelmann, Leipzig.

DINTER, M.K. 1914. Neue und wenig bekannte Pflanzen Deutsch-

Siidwest-Afrikas unter besonderer Beriicksichtigung der Suc-

culenten. Im Selbstverlag, Okahandja.

DYER, R.A. 1944. Chortolirion stenophyllum. Flowering plants of South

Africa 24: t. 932.

ENGLER, A. 1888. Plantae Marlothianae. Ein Beitrag zur Kenntnis der

Flora Siidafrikas. I. Monokotyledonae und Dikotyledonae ar-

chichlamydae. Botanische Jahrbiicher fur Systematik, Pflanzen-

geschichte und Pflanzengeographie 10: 1-50, t. 1.

FABIAN, A. & G ERM I S HU1ZEN , G. 1982. Transvaal wild flowers.

MacMillan, Johannesburg.

OBERMEYER. A.A. 1973. Aloe, Chamaealoe, Haworthia, Astroloba.

Poellnitzia and Chortolirion (Liliaceae). Bothalia 11: 119.

PHILLIPS. E.P. 1926. The genera of South African flowering plants.

Memoirs of the Botanical Survey of South Africa No. 10.

PHILLIPS. E.P. 1951. The genera of South African flowering plants, 2nd

edn. Memoirs of the Botanical Survey of South Africa No. 25.

Bothalia 25,1: 35-36 (1995)

FSA contributions 3: Asphodelaceae/Aloaceae, 1028010 Poellnitzia

G.F. SMITH*

Poellnitzia Uitewaal in Succulenta 22: 61 (1940);

D. R. Hunt: sub t. 804 (1981); G. F. Sm.: 75 (1994b); G.

F. Sm. & B-E. van Wyk: t. 2119 (1995). Type: Apicra

rubriflorci L. Bolus, i.e. Poellnitzia rubriflora (L. Bolus)

Uitewaal.

Herbaceous perennial; caulescent, offsetting at and

near the base, stems erect when young, ascending when

old, up to 250 mm long, ± 10 mom in diameter; roots

terete, 2-3mm in diameter. Leaves thick, hard, usually

four-ranked in spirally arranged rows, squarrose-imbri-

cate, ovate-squamiform, apex triquetrous, pungent-

acuminate, dark green to glaucous green, coated with a

waxy layer, 20^-0 mm long, ± 20 mm broad near base,

up to 5 mm thick; upper surface concave or flat, immacu-

late or with very few lighter green spots near base; lower

surface convex, keeled towards tip, usually with few

lighter green spots, spots sometimes confluent to form lon-

gitudinal lines; margins and keel apex minutely scabrid.

Inflorescence up to 500 mm long; peduncle simple, erect,

slender, ± 300 mm long, ± 3 mm in diameter; sterile

bracts membranous, lanceolate-acuminate, keeled with

one reddish brown vein, ± 7 mm long; raceme borne

horizontally, ± 200 mm long, ± 28 spirally arranged

flowers and buds, 3-8 accessible to pollinators simul-

taneously; floral bracts membranous, lanceolate, keeled,

clasping pedicels, shorter than pedicels, 3-5 mm long;

pedicels erect, persistent, brown, 6 — 8 mm long, ± 1 mm

in diameter. Flowers borne secundly erect, narrowly

tubular, elongate, slightly constricted above ovary and

below segment lobes, upper one-third slightly decurved,

tube red to orange, upper one-tenth of perigone members

dark green, connivent. reduplicate-valvate, tips of seg-

ments scarcely separated, ± 20 mm long; 3 mm in

diameter; segments red to orange with brownish midveins,

not free to base, fused, closely adhering where free, api-

cally spoon-shaped, outer segments slightly larger than

inner, margins apically minutely crenulate -erase, yellow;

stamens 6, subequal, inserted within perianth tube, at-

tached below ovary, ± 18 mm long; filaments light green;

anthers yellow, dorsifixed, dehiscing longitudinally; ovary

green, sessile, 6-7 mm long, 3 mm in diameter; style

white, straight, capitate, ± equalling anthers, ± 12 mm

long. Fruit green, capsule trilocular, cylindric, apically

retuse, dehiscing loculicidally, chartaceous when dry, ± 16

mm long, 3-4- mm in diameter. Seed dark brown to black,

angled, shortly winged, ± 4 mm long. Chromosome num-

ber: 2n - 14. Figure 1.

Monotypic. Poellnitzia is endemic to the Robertson

Karoo (districts of Robertson, Bonnievale and McGregor)

which is one of the drier, predominantly winter rainfall

areas bordering the Fynbos Biome. The vegetation in the

* National Botanical Institute, Private Bag XI 01. Pretoria 0001.

MS. received: 1994-03-24.

habitat of Poellnitzia can be broadly classified as Karroid

Shrublands and the entire range of distribution of the

genus falls within the Succulent Karoo Biome. Poellnitzia

is found ± 150-250 m above sea level. Figure 2.

On vegetative morphological grounds Poellnitzia

shows affinities with some representatives of Aloe ,

Astroloba and Haworthia. However, the flower morphol-

ogy of Poellnitzia is unique in the family Aloaceae in that

the dark green, free portion of the segment lobes is con-

nivent and reduplicate-valvate with the very tips of the

segments scarcely separated.

The genus name Poellnitzia honours Dr Joseph Karl

Leopoldt Arndt von Poellnitz (4 May 1896-15 February

1945). He was a German agriculturist and botanist with

FIGURE 1. — Poellnitzia rubriflora : A, habit, x 0.7; B, terminal portion

of raceme, x 0.7; C, lateral view of flower, x 1 .4; D, longitudinal

section of flower (three stamens removed), x 1 .4; E, fruit,

trilocular capsule, x 1. All drawings, except E, were made from

live material of Smith 184 (PUC). Fruit drawn from C.J. Burgers

78 (PRE). Artist; Gillian Condy.

36

Bothalia 25,1 (1995)

a general interest in succulent plants, particularly the fami-

ly Aloaceae. Both von Poellnitz and his only daughter

died tragically in bombing raids during World War II.

Poellnitzia rubriflora (L. Bolus ) Uitewaal in Suc-

culenta 22: 61 (1940); D. R. Hunt: sub t. 804 (1981); G.

F. Sm.: 75 (1994b); G. F. Sm. & B-E. van Wyk: t. 2119

(1995). Type: Western Cape: southwestern region; Swel-

lendam Div., Bonnie Vale, Smith s.n. ( National Botanic

Gardens , No. 2/17) (Herbarium Bolusianum 45213) (BOL,

holo.!).

Apicra rubriflora L. Bolus: 13 (1920); Uitewaal: 28 (1939).

Astroloba rubriflora (L. Bolus) E. Lamb: 230 (1955) nom. illegit.

Haworthia rubriflora (L. Bolus) Parr: 196 (1971). Aloe rubriflora (L.

Bolus) G. D. Rowley: 2 (1981).

Apicra jacobseniana Poelln.: 95 (1939). Poellnitzia rubriflora var.

jacobseniana (Poelln.) Uitewaal in Jacobsen & Rowley: 80 (1955).

Haworthia rubriflora var. jacobseniana (Poelln.) Parr: 89 (1972)

comb, inval. Type: "Kapland: Worcester, leg. Mrs Florence Morris; Typ

(= Triebner 34) im Botanischen Garten Kiel kultiviert,' no specimen

preserved [unnumbered photograph in Kakteenkunde (1939): 95

lecto. !, icono.f

leones: Hunt: t. 804 (1981); Court: 140 (1981); Smith: 17 (1994a).

Vouchers: Acocks 14098 (PRE): Burgers 78 (PRE); Smith 9 (PRU);

Smith 174. 177 (PUC).

Description as for the genus.

REFERENCES

BOLUS, H.M.L. 1920. Novitates africanae: Apicra rubriflora L. Bolus

(Liliaceae-Aloineae). Annals of the Bolus Herbarium 3: 1-14; t. 2d.

COURT, D.G. 1981. Succulent flora of southern A frica. Balkema,

Cape Town.

HUNT. D.R. 1981. Poellnitzia rubriflora Liliaceae. Curtis's Botanical

Magazine 183: t. 804.

JACOBSEN. H. & ROWLEY, G.D. 1955. Some name changes in

succulent plants. Part II. National Cactus & Succulent Journal

10: 80, 81.

LAMB, E. 1955. The illustrated reference on cacti and other suc-

culents, Vol. 1. Blandford Press, Poole.

PARR. C.A.E. 1971. Revision of the genus Astroloba. Part II. Bulletin of

the African Succulent Plant Society 6: 195-197.

PARR, C.A.E. 1972. Revision of the genus Astroloba. Part IV. Bulletin of

the African Succulent Plant Society 7: 89.

ROWLEY, G.D. 1981. Re-name that succulent. Cactus & Succulent

Journal of Great Britain 43: 2.

SMITH, G.F. 1994a. Conservation — is there a solution? Aloe 31: 17-20.

SMITH, G.F. 1994b. Taxonomic history of Poellnitzia Uitewaal, a unispecific

genus of Alooideae (Asphodelaceae). Haseltonia 2: 74-78.

SMITH. G.F. & VAN WYK, B-E. 1995. Poellnitzia rubriflora. Flowering

Plants of A frica 54: t. 2119.

UITEWAAL, A.J.A. 1939. Het geslacht Apicra (Willd.) Haw. c.m. (Ver-

volg). Succulenta 21: 25-29.

UITEWAAL, A.J.A. 1940. Een nieuw geslacht der Aloineae. Succulenta

22: 61-64.

VON POELLNITZ, J.K.L.A. 1939. Apicra jacobseniana v. P. Kak-

teenkunde (1939): 95.

Bothalia 25,1: 37-42(1995)

Notes on the typification of some species of Aloe (Asphodelaceae/Aloaceae)

H.F. GLEN and G.F. SMITH

ABSTRACT

Lecto- or neotypes are designated for Aloe thompsoniae Groenew., A. micrqkantha Haw., A. schmidtiana Regel (a synonym

of A. cooperi Baker), A. longistyla Baker, A. aristata Haw., A. longiaristata Schult. & Schult. f., (a synonym of the preceding

name), A. humilis (L.) Mill. var. incurva Haw., A. glauca Mill., A. muricata Schult. (a synonym of the preceding name), A.

saponaria ( Aiton) Haw. var. latifolia Haw. (a new synonym of A. metadata All.) and A. pluridens Haw. The confusing citation

of the type of A. thorncroftii Pole Evans is clarified.

UITTREKSEL

Lekto- of neotipes word aangewys vir Aloe thompsoniae Groenew., A. micracantha Haw., A. schmidtiana Regel (’n

sinoniem van A. cooperi Baker), A. longistyla Baker, A. aristata Haw., A. longiaristata Schult. & Schult. f. (’n sinoniem van die

voorafgaande naam), A. humilis (L.) Mill. var. incurva Haw., A. glauca Mill., A. muricata Schult. (’n sinoniem van die

voorafgaande naam), A. saponaria (Aiton) Haw. var. latifolia Haw. (’n nuwe sinoniem van A. maculcita All.) en A. pluridens

Haw. Die verwarrende sitering van die tipe-eksemplaar van A. thorncroftii Pole Evans word opgeklaar.

INTRODUCTION

In the course of studies directed towards both a revision

of the southern African species of Aloe and the data cap-

ture for PRECIS-FLORA, it was found that the typifica-

tion of several well known names was ambiguous. Details

necessary to remove the ambiguity in three cases were

recently published in some detail by Smith (1990a, b) and

Glen & Hardy (1991). but these are only three of the more

involved cases. In a significant number of instances, all

that is required is the choice of a lectotype or neotype —

which hardly merits a separate paper, or even a separate

short note, for each name. The purpose of this paper, then,

is to gather up all these problems into a single worthwhile

publication.

Typification of extant yet untypified names is not

obligatory (Art. 9.9 of the International Code of Botanical

Nomenclature , but see Art. 10.2). In a taxonomically dif-

ficult genus such as Aloe , however, typification is often

necessary to establish the application of the plethora of

available names. Furthermore, it fixes the concepts of the

present authors regarding selected names in Aloe that

should be reduced to synonymy.

The names in question are considered in a taxonomic

order approximating that used by Berger (1908) and

Reynolds (1950). They are dealt with in groups roughly

following those employed by these authors. In all cases,

lectotypes and neotypes have been chosen bearing in mind

two criteria. In order of importance, these are: 1, preser-

vation of current usage, as far as possible establishing the

intent of the original author, and 2, accessibility of

material.

National Botanical Institute, Private Bag X101 , Pretoria 0001 .

MS. received: 1994-03-24.

NOTES ON TYPIFICATION

Aloe thompsoniae Groenew. in Tydskrif vir Wetenskap

en Runs 14: 64 (1936) sphalm. thompsoni. Lectotype: Trans-

vaal, Haenertsburg, Thompson s.n. in PRE 274 (PRE. lecto.!,

here designated).

Although not cited by Groenewald ( 1936), who did not

designate a type, the specimen designated here (Figure 1)

is the only one in PRE that was collected by the person

commemorated in the name. Dr (Mrs) Sheila Clifford

Thompson of Haenertsburg, Petersburg District (Reynolds

1946; Prinsloo 1972) and is therefore relevant type

material. In this context it is also noteworthy that Dr F.

Z. van der Merwe, an avid collector of aloes in the mid-

1900’s, passed most of his newly described species of

Aloe on to Mr B. H. Groenewald who described them and

deposited the specimens in PRE (Gunn & Codd 1981).

In general it is therefore not worth searching other her-

baria for material suitable as lectotypes of the names of

species described by Groenewald.

Aloe micracantha Haw. (microcantha auett.) in Sup-

plemental! plantarum succulentarum: 105 (1819). Neo-

type: Burchell 4482 (K!. here designated).

Haworth (1819) cites no type for this species, but states

that he saw a small plant collected in the wild and planted

in the Royal Botanic Gardens, Kew. Sims (1821) states

‘For this very rare species, which is supposed not to exist

in any of our other collections except in that of Kew,

where it was introduced about two years since from the

Cape of Good Hope, we are indebted to Thomas Kitchin,

Esq., of Norwich, in whose garden, rich in succulent

plants, it flowered in July last.’ Therefore one is ultimately

forced to rely on Sims’s skill and good faith in having

compared the Kew and Norwich plants (neither of which

is accessible any more). This problem of speculative

typification based on circumstantial evidence can for-

tunately be circumvented due to the existence in Kew of

38

Bothalia 25,1 (1995)

\

' A

FIGURE 1. — Thompson s.n. (PRE 274), the lectotype specimen of Aloe

thompsoniae.

two Burchell specimens of A. micracantha (Figure 2).

They are both mounted on a single sheet, and are clearly

identifiable with locality material nowadays referred to A.

micracantha (Smith 1993). Selection of one of these

specimens gives the name an accessible, readily identifi-

able type that stabilizes current usage. In addition, this

specimen is the element that comes closest historically to

being a live plant with which Haworth was familiar. Selec-

tion of either the Botanical Magazine plate of the species

(Sims 1821), or an excellent colour plate kept at Kew of

A. micracantha (Figure 3) as neotype for the name was

considered, because, especially in the case of succulent

plants, a drawing is often more diagnostic than a specimen

and therefore more useful for fixing the application of a

name (Smith 1990a). The Kew plate has often been at-

tributed to Franz Bauer, but is by an unknown artist (G.

LI. Lucas pers. comm.). In the case of A. micracantha ,

the Burchell specimens are of such exceptional quality

that one of them is the preferred choice (Figure 2). Of

the two specimens available we chose Burchell 4482 (a

whole plant) rather than Burchell 4564 (an inflorescence

and two separate leaves), because the former gives a better

idea of the appearance of the plant.

Aloe schmidtiana Regel in Gartenflora t. 970 (1879).

Holotype: Gartenflora 97: t. 970.

It is unlikely that a herbarium specimen was prepared

of the leaf and flower material referred to in the

protologue. A very attractive illustration depicting A.

cooperi Baker, however, accompanies the protologue.

Since this plate is the one illustration used by the author

(Art. 9.3 of the International Code of Botanical Nomen-

clature) it is here treated as the holotype.

Aloe longistyla Baker in Journal of the Linnean

Society, Botany 18: 158 (1880). Lectotype: Cape, Graaff-

Reinet, Bolus 689 (K, lecto.!; PRE, photo!, here desig-

nated).

Baker (1880) cites two specimens in the protologue of

this species. The specimen Bolus 689 is chosen as lec-

totype rather than Drege 8640 because of its more detailed

locality data (Figure 4).

Aloe aristata Haw. in Philosophical Magazine 66:

280 (1825). Neotype: Cape, near Steynsburg, Reynolds

1024 (PRE!, here designated).

Aloe longiaristata Schult. & Schult. f. in Systema vegetabilium 7,1:

684 (1829). Neotype: Salm-Dyck. Monographia generum Aloes et

Mesembryanthemi, Fasc. 2, fig. 21 [Sect. 15, fig. 7 (1837), here desig-

nated].

FIGURE 2. — Both Burchell specimens, 4482 and 4564, are mounted on

the same sheet. The neotype of Aloe micracantha, Burchell 4482,

is the complete one in the middle (arrowed).

Bothalia 25,1 (1995)

39

This is another case where a neotype is chosen for a

name considered to be a synonym of another — the

varieties of A. humilis intergrade so thoroughly as not to

be worth upholding. Haworth (1804) has here, as is often

the case, supplied no hint of the material he saw. Ker

Gawler (1805) tells us that his plant came from a nursery

in Kensington, which must surely have been known to

Haworth, who at that time lived in Chelsea in the same

part of the then southwestern outskirts of London. Again,

the earliest available element is chosen, to stabilize current

usage.

Aloe glauca Mill, in The Gardeners' Dictionary No.

16 (1768). Neotype: Cape, hills south of Wyke, Reynolds

1967 (PRE!, here designated).

FIGURE 3. — Reduced photograph of an unpublished painting of Aloe

micracantha by an unknown artist, probably not Franz Bauer as

claimed by Reynolds (1950) (G. LI. Lucas pers. comm.).

Reproduced with permission of the Controller, Her Majesty’s

Stationery Office, and The Trustees, Royal Botanic Gardens,

Kew.

As usual. Haworth gives no clue as to what material

he had available when drawing up his description of A.

aristata. In the absence of any contemporary or near-con-

temporary illustrations or specimens (no published refer-

ence was made to the name for 70 years after the

protologue), a good modem specimen, Reynolds 1024 , is

chosen as neotype (Figure 5). One of its synonyms, A.

longiaristata , is another name that is not in general use

in the current taxonomy of the genus. However, it shows

clearly how in the previous century names were based on

etiolated plants grown ex situ thousands of miles away

from their natural habitats (Salm-Dyck 1837). The plate

chosen here as neotype of the name is the only obvious

one that can serve the purpose. J. A. & J. H. Schultes

(1829) inexplicably included the validly published name,

A. aristata , in the synonymy of A. longiaristata.

Aloe humilis (L.) Mill. var. incurva Haw. in Trans-

actions of the Linnean Society 7: 15 (1804). Neotype

(icono.): Curtis’s Botanical Magazine 21: t. 828 (1805),

here designated.

Miller's protologue (1768) of this species gives a ref-

erence to a plate in Commelin (1703), which Wijnands

(1983) shows clearly to represent a completely different

species. Miller gives no other indication of a type for this

name. As Wijnands (1983) points out, one is faced in this

case with a choice of two options, both of which have

undesirable features, namely: 1, drawing the conclusion

from the evidence so ably marshalled by Wijnands ( 1983)

that Miller suffered a series of errors at the crucial point

here, and stabilize current usage by conserving the name

Aloe glauca Mill, with a completely new type, rejecting

all references to his confused citation of Commelin il-

lustrations; or 2, reject this well known name as a nomen

confusion in favour of the next-oldest, securely identified

name, A. rhodacantha DC. It seems to us that the first

course is the less harmful, and so neotypes are proposed

here for both A. glauca Mill. (Figure 6) and A. muricata

FIGURE 4. — Bolus 689. lectotype of Aloe longistyla.

40

Bothalia 25,1 ( 1995)

FIGURE 5. — Reynolds 1024, neotype of Aloe aristata.

Schult., the basionym of one of its infraspecific taxa (Fig-

ure 7; see below).

Aloe glauca Mill. var. muricata (Schult.) Baker in

Journal of the Linnean Society 18: 161 (1880). Neotype:

Cape, Piketberg Division, De Hoek, Reynolds 4749 (PRE!;

SAM! isoneo.)

A. muricata Schult. in Observationes botanicae: 70 (1809).

No type is cited for A. muricata Schult. and, as in the

previous case, neotypification stabilizes current usage of

the name.

Aloe saponaria (Aiton) Haw. var. latifolia Haw. in

Transactions of the Linnean Society 7: 1 8 ( 1804). Neotype

(icono.): Ker Gawler in Curtis’s Botanical Magazine 34:

t. 1346 (1811), here designated.

It is proposed here that the name should be included

in the synonymy of Aloe maculata All. Ker Gawler (1811)

states ‘We have to thank Mr Haworth for the present

specimen.’ Although Haworth (1804), as usual, gives no

clue as to the material he described under var. latifolia, it

would be taking caution altogether too far to assume that

the plant figured was not at least part of the material of

this taxon known to Haworth at the time he coined the

Aloe striata Haw. in Transactions of the Linnean

Society of London 7: 18 (1804). Neotype: Cape, near

Grahamstown, Bottomley s.n. in PRE 27 (PRE!), here

designated.

Haworth (1804) mentions, in a cryptic note, a Masson

plant we have had no success in tracing. One must there-

fore assume that this specimen, if it ever existed, has long

since disappeared and a neotype may therefore be chosen.

This is done here in such a way as to stabilize current

usage. The specimen chosen is mounted on two sheets;

to obviate future confusion we designate the sheet shown

in Figure 8 as iectoneotype.'

Aloe thorncroftii Pole Evans

In the introduction to a paper in which he describes

several species of Aloe that are now well known, Pole

Evans (1917) states that ‘the accompanying descriptions

have been made from plants growing in the rockeries in

the grounds of the Botanical Laboratories of the Union

of South Africa’; that is, the lower western part of the

Union Buildings garden in Pretoria. He gives a collector

and locality (sometimes more than one) for each species,

but never cites a specimen number. This makes locating

the intended type specimen somewhat problematical in

some cases.

name.

FIGURE 6. — Reynolds 1967. neotype of Aloe glauca.

Bothalia 25,1 (1995)

41

FIGURE 7. — Reynolds 4749, neotype of Aloe muricata.

The protologue of Aloe thomcroftii forms part of Pole

Evans’s paper, and in this case the only indication of a

type is the following paragraph: ‘All the plants (7) which

so far have been flowered in Pretoria have borne an un-

branched inflorescence, but Mr George Thomcroft, who

first sent me specimens of this Aloe and after whom it

has been named, writes me that the inflorescence is

branched in the case of some plants.’

One deduces that the type specimen of this species, if

one exists, would be marked as having been collected by

Thomcroft before 1917, and possibly as having been

grown in the Union Buildings grounds. As there are no

other specimens from the Union Buildings gardens known

in any herbarium except PRE, and as Pole Evans was

Chief of the then Division of Botany (later renamed the

BRI and finally the NBI), one would search only in PRE

for such a specimen. Such a search was undertaken in the

course of studies on the southern African species of Aloe

by the senior author. Only one specimen fitting these re-

quirements was found (Figure 9). The label is typical of

the Pole Evans period, and matches those of many

specimens giving their locality only as ‘Union Buildings

gardens.’ The specimen was collected by Thomcroft in

1914 (long enough before the date of publication to allow

quite a small plant to grow to flowering size), and is num-

bered 247 in an unknown series. Neither Pole Evans’s

collecting register nor the PRE accessions register has a

species of Aloe as No. 247, therefore one wonders if the

number is from the accessions register of the Vredehuis

Garden (predecessor to the Pretoria National Botanical

Garden). No register of this date from this garden has

survived. It seems that this is the type specimen of Aloe

thomcroftii and its correct citation is: TYPE: Transvaal,

Barberton, 1914, G. Thomcroft s.n. in PRE 247 (PRE,

holo.!).

Aloe pluridens Haw. in The Philosophical Maga-

zine 64: 299 (1824). Neotype: Cape, Ettrick Hills near

Carlisle Bridge, Reynolds 1425 (PRE!, here designated).

As usual, Haworth gives no clue as to what material

he had available when drawing up his description of this

species. Just as in the case of A. aristata , this name was

ignored in print for 70 years after the publication of the

protologue. In the absence of any contemporary or near-

contemporary illustrations or specimens, a good modem

specimen is chosen as neotype (Figure 10).

ACKNOWLEDGEMENTS

Permission granted by the Director of the Royal

Botanic Gardens, Kew, to use copyright material in this

article, is gratefully acknowledged.

iyl ... .!.) 0 '<

FE3 . ... -•

I Kit RE 8. — Bottomley s.n. ( PRE 27), neotype of Aloe striata.

42

Bothalia 25.1 ( 1995)

FIGURE 9. — Thomcroft s.n. ( PRE 247), holotype specimen of Aloe

thomcroftii.

REFERENCES

BAKER, J.G. 1880. A synopsis of Aloineae and Yuccoideae. Journal of

the Linnean Society, Botany 18: 148-241.

BERGER, A. 1908. Liliaceae-Asphodeloideae-Aloineae. In A. Engler,

Das Pflanzenreich 4, 38, III, II (Heft 33). Engelmann, Leipzig.

COMMELIN, C. 1703. Praeludia botanica. Haringh, Leiden.

GLEN, H.F. & HARDY, D.S. 1991. The type specimen of Aloe sout-

pansbergensis (Liliaceae/Asphodelaceae). Botluilia 21: 151, 152.

GROENEWALD, B.H. 1936. ’n Nuwe aalwyn van die Wolkberg,

Transvaal. Tydskrif vir Wetenskap en Kuns 14: 64—66.

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

Africa. Balkema, Cape Town.

HAWORTH, A. H. 1804. A new arrangement of the genus Aloe. Transac-

tions of the Linnean Society of London 7 : 1-28.

HAWORTH, A.H. 1819. Supplementum plantarum succulentarurn. Hard-

ing, London.

HAWORTH, A.H. 1824. Decas secunda novarum plantarum succulen-

tarum. Philosophical Magazine 64: 298-302.

HAWORTH, A.H. 1825. Decas quinta novarum plantarum succulen-

tarum. Philosophical Magazine 66: 279-283.

KER GAWLER, J.B. 1805. Aloe humilis var. incurva. Curtis's Botanical

Magazine 2 1 : t. 828.

KER GAWLER, J.B. 1811. Aloe saponaria var. latifolia. Curtis 's Botani-

cal Magazine 34: t. 1436.

MILLER, R 1768. The Gardeners' Dictionary, edn 8. Rivington, London.

FIGURE 10. — Reynolds 1425, neotype of Aloe pluridens.

POLE EVANS, I.B. 1917. Descriptions of some new aloes from the

Transvaal, part II. Transactions of the Royal Society of South

Africa 5: 703-712.

PRINSLOO, H.D.A. 1972. A rare aloe. Veld & Flora 2: 16, 17.

REGEL, E.A. VON, 1879. B. Aloe Schmidtiana Rgl. Liliaceae. Gar-

tenflora 97: t. 970.

REYNOLDS, G.W. 1946. Aloe thompsonii. The Flowering Plants of

Africa 25: t.980.

REYNOLDS, G.W. 1950. Aloes of South Africa. Aloes of South Africa

Book Fund, Johannesburg.

SALM-DYCK, J.M.F.A.H.I. 1837. Monographia generum Aloes et

Mesembryantliemi, Aloe longiaristata. Fasc. 2, fig. 21 (Sect. 15,

fig. 7). Diisseldorf.

SCHULTES, J. A. 1809. Observationes botanicae. Innsbruck.

SCHULTES, J.A. & SCHULTES, J.H. 1829. Classis VI, Hexandria Mono-

gynia Genera, 1417. Aloe. In J.J. Roemer & J.A. Schultes, Sys-

tema vegetabilium 7,1: 684. Sumptibus J.G. Cottae, Stuttgartiae.

SIMS, J. 1821. Aloe microcantha. Curtis’s Botanical Magazine 48: t.

2272.

SMITH, G.F. 1990a. Neotypification of Aloe bowiea (Asphodelaceae:

Alooideae). South African Journal of Botany 56: 4 1 5—4 1 8.

SMITH, G.F. 1990b. Neotypification of Bowiea myriacantha, basionym

of Aloe myriacantha (Asphodelaceae: Alooideae). Botanical Bul-

letin of Academia Sinica (new series) 31: 315-320.

SMITH, G.F. 1993. Notes on the taxonomic and conservation status of

Aloe microcantha (Asphodelaceae: Alooideae). Haseltonia 1:

55-60.

WIJNANDS, D.O. 1983. The Botany of the Commelins. Balkema, Rotter-

dam.

Botha] ia 25,1: 43-59(1995)

Solarium (Solanaceae) in Uganda

Z.R. BUKENYA* and J.F. CARASCO**

Keywords: food crops, indigenous taxa, key, medicinal plants, ornamentals. Solatium, Solanaceae, Uganda, weeds

ABSTRACT

Of the 41 species, subspecies and cultivar groups in the genus Solatium L. (Solanaceae) that occur in Uganda, about 30 are

indigenous. In Uganda several members of the genus are utilised as food crops while others are put to medicinal and ornamental

use. Some members are notorious weeds. A key to the species and descriptions of all Solatium species occurring in Uganda are

provided.

UITTREKSEL

Van die 41 spesies, subspesies en kultivargroepe in die genus Solatium L. (Solanaceae) wat in Uganda voorkom, is sowat 30

inheems. Verskeie lede van die genus word as voedselgewasse benut, terwyl ander vir geneeskundige en omamentele gebruike

aangewend word. Sommige lede is welbekend as onkruide. 'n Sleutel tot die spesies en beskrywings van al die Solatium- spesies

wat in Uganda voorkom word voorsien.

CONTENTS

Introduction 44

Materials and methods 45

Key to species 45

Solatium L 46

A. Subgenus Solatium 46

Section Solatium 46

1. S. nigrum L 46

la. S. nigrum subsp. nigrum 47

2. S. americanum Mill 47

3. S. scabrum Mill 47

4. S. sarrachoides Sendtn 48

5. S. villosum Mill 48

6. S. grossedentatum A. Rich 48

7. S. florulentum Bitter 48

8. S. tarderemotum Bitter 48

Section Afrosolanum Bitter 49

9. S. terminate Forssk 49

9a. S. terminate Forssk. subsp. termiiiale Heine 49

9b. S. termiiiale subsp. sanaganum (Bitter) Heine 49

9c. S. termiiiale subsp. inconstans (C.H. Wright)

Heine 49

10. S. welwitschii C.H. Wright 49

1 1 . S. nakurense C.H. Wright 49

Section Benderianum Bitter 50

12. S. benderianum Schimp. ex Dammer 50

13. S. runsoriense C.H. Wright 50

B. Subgenus Brevantherum (Seithe) D'Arcy 50

Section Brevantherum Seithe 50

14. S. mauritianum Scop 50

* Department of Botany, Makerere University, P.O. Box 7062. Kampala,

Uganda.

** Department of Biochemistry, Makerere University, P.O. Box 7062,

Kampala, Uganda.

MS. received: 1993-06-29.

C. Subgenus Leptostemonum (Dunal) Bitter 50

Section Acantliophora Dunal 51

15. 5. mammosum F 51

16. S. aculeatissimum Jacq 51

Section Aculeigerum Seithe 51

17. S. wendlandii Hook 51

Section Melongena Dunal 51

18. S. melongena L 51

19. S. aculeastrum Dunal 52

20. S. incanum F 52

21. S. macrocarpon F 52

22. S. wrightii Benth 53

Section Monodolichopus Bitter 53

23. S. coagulans Forssk 53

Section Oliganthes (Dunal) Bitter 54

24. S. anguivi Fam 54

25. S. aethiopicum F 54

25a. S. aethiopicum Gilo group 54

25b. S. aethiopicum Shum group 55

26. S. albicaule Kotschy ex Dunal 55

27. S. cyaneo-putpureum De Wild 55

28. S. hastifolium Hochst. ex Dunal 55

29. S. taitense Vatke 55

30. S. usatnbarense Bitter ex Dammer 56

Section Tor\>a Nees 56

31. S. giganteum Jacq 56

32. 5. kagehense group 56

33. S. renschii Vatke 56

D. Subgenus Potatoe (G. Don) D'Arcy 57

Section Petota Dumort 57

34. S. tuberosum F 57

Section Jasmino solatium Bitter ex Seithe 57

35. S. seaforthianum Andrews 57

Conclusions 57

Acknowledgements 58

References 58

44

Bothalia 25,1 (1995)

INTRODUCTION

The genus Solatium L. belongs to the family Sola-

naceae, and contains about 2 000 species of which about

35 occur in Uganda. Several species are important food

crops, yielding edible fruits and leaves whereas others are

ornamentals or weeds (Heine 1963).

The genus also contains plants of medicinal value. For

example the fruits of S. anguivi Lam. contain alkaloids

used in the treatment of a number of diseases including

chronic respiratory diseases (Bector et al. 1971). Walters

(1965) observed that Solarium alkaloids have antifungal

effects. Thus it is possible that some of these alkaloids

could be used as antibiotics. Beaman-Mbaya & Muham-

med (1976) reported that alkaloids from fruits of S. in-

canum L. are used in treatment of cutaneous mycotic

infections and other pathological conditions in Kenya. In

Uganda, the soup from green fruits of S. anguivi Lam. is

popular especially among women, for it is believed to

cure hypertension (Sengendo 1982).

The genus is widely distributed throughout the world

with major species representation in America, Australia

and Africa. The genus was first studied by Dillenius

(1732) and later by Linnaeus (1753). Since 1753 the genus

has been reclassified innumerable times and a multitude

of varieties, subspecies and species have been named,

especially in the section Solanum. For example, Dunal

(1813) in his monograph of the genus, described 60

species belonging to section Solanum. Bitter (1912, 1913,

1917, 1919, 1921, 1922, 1923) was the second worker to

attempt to monograph the genus; he is criticized for being

‘more of a splitter than Dunal, and he described more

than 60 new Solanum species from the Americas alone'

(Edmonds 1977). He recognised 20 sections for the genus,

and revised Solanum in Africa utilizing mainly collections

from German expeditions. He erected a partial classifica-

tion of Solanum.

The validity of some of Bitter’s varieties has been

questioned because they were based on minor variations

which are of very limited taxonomic value. However, his

work is the most detailed treatment so far available on

African Solanum.

D’Arcy (1972) provided a modem classification of the

genus Solanum into subgenera, sections and series and his

classification is widely accepted today. It is also followed

here.

Although the above major works and others attempted

to streamline the taxonomy of Solanum , the genus is

taxonomically difficult, due to various factors. These in-

clude the difficulty of associating the names of Solanum

used by earlier taxonomists with plants of today due to

early descriptions being brief, often vague and frequently

lacking in characters now considered to be diagnostic.

Another problem is that some of the early names, for ex-

ample many of the names of Linnaeus and those before

him, are difficult to typify (Hepper 1979).

Another problem is the occurrence of polyploid series

within the section Solanum (Edmonds 1977), such as

tetraploids and hexaploids occurring within the S. nigrum

complex. These may provide a barrier to hybridization

FIGURE I. — Geographical divisions,

U 1 , U2, U3 and U4, of the Flora

of tropical East Africa and the

main towns of Uganda.

Bothalia 25,1 (1995)

45

between morphologically similar plants leading to

cytoraces which are difficult to differentiate using classical

methods.

There is also considerable phenotypic plasticity within

species and hybridization between closely related species.

Hybridization followed by inbreeding may result in for-

mation of new populations different from either parent.

This is particularly true for the cultivated species, as for

example found in sections Melongena and Oliganthes. A

large number of ‘microspecies’ or ‘semispecies' (Grant

1971) occur in section Solanum and it is difficult to decide

which of these deserve taxonomic recognition.

In recent years, there has been an explosion of data

from various taxonomic research projects aimed at at-

tempting to solve some of the above problems and im-

proving the knowledge of the genus. The majority of these

studies has been made outside Africa. Jaeger (1985)

recognized 80 species of Solanum in Africa. To a large

extent we accept the taxonomic treatment of the taxa he

discussed. The information in this work, especially on

typification and synonymy, was very useful in the present

study. No comprehensive taxonomic study had been made

on the genus in Uganda. Lind & Tallantire (1975)

provided short descriptions of only three Ugandan species

of Solanum, i.e. S. terminate Forssk., S. incanum L. and

5. nigrum L. Sengendo ( 1982) produced some data on the

two species S. anguivi and S. aethiopicum L., while

Bukenya (1991) gave a comparative account of a few

Solanum fruit and leaf vegetables.

To date the Solanaceae has not yet been treated for the

Flora of tropical East Africa. Hence, to identify a species

of Solanum in Uganda one has to use other regional floras,

particularly the Flora of tropical West Africa, edn 2 (Heine

1963). However in this Flora, not all the species of

Solanum occurring in Uganda are included and the

descriptions of species are brief.

It was necessary to carry out a comprehensive study

of these species so that critical descriptions and a key to

the species could be provided.

MATERIALS AND METHODS

The species descriptions were largely based on the

study of herbarium material in the Botany Department,

Makerere University; the Royal Botanic Gardens, Kew;

the Forest Herbarium, Oxford, and the Institute of Sys-

tematic Botany, Munich.

Figure 1 shows the map of Uganda divided into the

four geographical divisions of the Flora of tropical East

Afiica. Table 1 gives the infrageneric classification of

Ugandan Solanum species based on D’Arcy (1972).

Key to species

la Plants usually armed:

2a Hairs stellate:

4a Mature leaves ± glabrous; inflorescence corymbose, cyme subfasciculate or paniculate:

5a Leaves ± 280 x 120 mm; inflorescence paniculate 31. S. giganteum

5b Leaves < 200 x 80 mm; inflorescence cymose or corymbose:

6a Inflorescence cymose; flowers often subumbellate or branched, peduncle 10-20-flowered; leaves up to 150 x 70 mm

32. S. kagehense group

6b Inflorescence corymbose, 20-50-flowered; leaves 50-80 x 30-40 mm 33. S. renschii

4b Mature leaves mostly hairy; flowers solitary or inflorescence few-flowered, racemose, lateral:

7a Fruit dry, usually completely enclosed by heavily armed accrescent calyx, seeds shiny black 23. S. coagulans

7b Fruit not dry, seeds yellow-brown:

8a Corolla 17^45 mm long; fruit 20-130 x 30-100 mm:

9a Fruit red when mature; prickles absent or present on young leaves 25a. S. aethiopicum Gilo group

9b Fruit yellow when mature:

10a Plants 4-10 m high:

11a Leaves about 150 x 100 mm, upper surface glabrous 19. S. aculeastrum

lib Leaves up to 300 x 240 mm, both surfaces hairy 22. S. wrighrii

10b Plants < 4 m high:

12a Petiole < 30 mm long; prickles present or absent 21. S. marcocarpon

12b Petiole 40-70 mm long; prickles present or absent:

13a Prickles almost always absent; fruit 60-130 x 30-100 mm 18. S. melongena

13b Prickles always present; fruit 27 x 30 mm 20. 5. incanum

8b Corolla 10-15 mm long; fruit 8 x 10 mm:

14a Leaves glabrous; corolla 5-7 mm long; fruit ± 15 mm diam 25b. S. aethiopicum Shum group

14b Leaves hairy; corolla > 7 mm long; fruit < 15 mm diam.:

15a Plants clambering or scandent:

16a Inflorescence racemose, 3-10-flowered 27. 5. cyaneo-purpurewn

16b Inflorescence not racemose:

17a Inflorescence umbelliform, 2-6(-10)-flowered; leaves ovate 28. S. hastifolium

17b Inflorescence (1) 2^t-flowered; leaves lanceolate 29. S. taitense

15b Plants not clambering or scandent:

18a Leaves rather small, 20-70 x 10 mm; fruits yellow when ripe 26. S. albicaule

18b Leaves 100-200 x 50-150 mm; fruits red when ripe:

19a Plants generally non-prickly, occasionally a few prickles present; flowers solitary or 5-15 in a raceme -like

cyme 24. 5. anguivi

19b Plants heavily armed with prickles; inflorescence 8- to many-flowered, rachis simple or branched, racemose

30. S. usambarense

46

Bothalia 25,1 (1995)

2b Hairs simple, sometimes glandular or absent, occasionally with stellate hairs interspersed:

20a Plants climbing; leaves compound; inflorescence many-flowered (50+ ), inflorescence axis very long, ± 300 mm

17. S. wendlandii

20b Plants not climbing; leaves simple; inflorescence few-flowered (< 10), a short unbranched raceme:

21a Corolla violet or blue, ± 12 mm long; fruit with terminal nipple or mammilla 15. S. mammosum

21b Corolla white, ± 17 mm wide; fruit with no terminal nipple or mammilla 16. 5. aculeatissimum

lb Plants mostly unarmed:

3a Hairs stellate; fruits pubescent 14. 5. mauritianum

3b Hairs simple, sometimes branched or glandular, or absent; fruits not pubescent:

22a Leaves compound:

23a Tuberous stolons present; plant not climbing; inflorescence with simple hairs; fruits ± 7 mm diam 34. S. tuberosum

23b Tuberous stolons absent; plant climbing; inflorescence glabrous; fruits ± 10 mm diam 35. 5. seaforthianwn

22b Leaves not compound:

24a Plants herbaceous < 2 m high:

25a Fruiting calyces strongly adherent-accrescent, enclosing at least lower half of berries 4. 5. sarrachoides

25b Fruiting calyces only slightly accrescent, reflexed or adhering to base of berries:

26a Berries longer than broad, yellow, orange or red 5. S. villosum

26b Berries globose or broader than long, black, purple or green:

27a Inflorescence branched, up to 20-flowered 7. S. florulentum

27b Inflorescence not branched, less than 20-flowered:

28a Inflorescence racemose 8. 5. tarderemotum

28b Inflorescence an umbellate cyme:

29a Leaves glabrous, up to 250 x 200 mm 3. S. scabrum

29b Leaves not glabrous, less than 150 x 100 mm:

30a Fruiting pedicel erect; seeds 1 x 1 mm 2. S. americanum

30b Fruiting pedicel recurved:

31a Corolla about 10 mm diam la. S. nigrum subsp. nigrum

31b Corolla about 15 mm diam 6. S. grossedentatum

24b Plants shrubs 2-7 m high:

32a Inflorescence a lax cymose panicle:

33a Leaves hairy on both surfaces 13. S. runsoriense

33b Leaves more or less glabrous 12. 5. benderianum

32b Inflorescence spiciform, umbellate or subracemose:

34a Erect subshrub; stem verrucose; leaves 20-70 x 10-40 mm; inflorescence umbellate, 5-10(-20)-flowered

11.5. nakurense

34b Climbing shrubs; stem not verrucose; leaves 80-170 x 40-80 mm; inflorescence various, (2-7)-50-flowered:

35a Inflorescence terminal, spiciform, up to 200 mm long 10. 5. welwitschii

35b Inflorescence not spiciform:

36a Inflorescence subracemose, leaf-opposed or in leaf axil, 2-7-flowered 9c. 5. terminate subsp. inconstans

36b Inflorescence not as above, many-flowered:

37a Peduncles with terminal umbel, sometimes a few lateral ones as well; inflorescence sometimes with more

than 50 flowers 9a. 5. terminate subsp. terminate

37b Peduncle terminal or subterminal, strongly branched, giving inflorescence a paniculate appearance

9b. 5. terminate subsp. sanaganum

Solanum L., Species plantamm: 184-188 (1753);

L.: 85 (1754); D’Arcy: 85 (1973).

The genus is recognized by the 5-partite calyx and by

long, often connivent anthers dehiscing by terminal pores

and with short filaments. Other characters useful for

recognition are the frequently rotate, 5-lobed corolla, the

fruit being a berry with flattened seeds; the often stellate-

pubescent prickles; and the often extra-axillary, usually

cymose inflorescence.

A. Subgenus Solanum D’Arcy in Annals of Mis-

souri Botanical Garden 60: 733 (1973). Type species: S.

nigrum L.

Leaves subentire or shallowly lobed, often mem-

branaceous; indumentum simple, hairs rarely branching.

Prickles absent. Flowers mostly small. Corolla mostly deep-

ly lobed. Filaments often pubescent; anthers short, dehiscing

introrsely by large, often oblique terminal pores and some-

times ultimately by longitudinal slits in the upper portion.

Ovary glabrous. Fruit rather small, 7-15 mm diam.

The subgenus is represented in Uganda by three sec-

tions: Solanum , Afrosolanum and Benderianum.

Section Solanum Seithe in Botanische Jahrbiicher 81:

261-336 (1962).

1. Solanum nigrum L., Species plantarum: 186

(1753); Heine: 335 (1963); E.M. Lind & Tallantire: 130

(1975); Hepper: 12 (1976). Type: LINN 248.18 (IDC

microf. 1775 138: III.4). All the above authors have taken

S. nigrum in a broad sense.

Herb, about 1 m high. Leaves ovate-elliptic, 60-100 x

40-70 mm; lamina glabrous when mature, margin entire

or toothed, repand or with 2 or 3 pairs of short lobes;

petiole 10-30 mm long. Inflorescence 6-8-flowered;

peduncle ± 10 mm long; pedicels ± 5 mm long. Corolla

3^4- mm long. Fruit usually broadly ovoid, 5-8 mm diam.,

shiny black when ripe; fruiting pedicel ± 10 mm long,

recurved. Seeds ±1x2 mm.

The name S. nigrum has been used in Uganda in a broad

sense until the work of Edmonds (Gray 1968; Edmonds

1971, 1972) and Jaeger (1985). Edmonds, at Oxford Univer-

sity, identified some of the specimens from the Botany

Department herbarium, Makerere. Studies of this complex

were carried out with specimens from Uganda, kept at

Kew Herbarium and at Makerere Herbarium. Those

studies have led to the recognition of eight Ugandan taxa

Bothalia 25,1 (1995)

47

TABLE 1. — Infrageneric classification of Ugandan Solatium spp.

based on D'Arcy (1972)

from the S. nigrum complex. These are S. nigrum subsp.

nigrum, S. americanum Mill., S. scabrum Mill., S. sar-

rachoides Sendtn., S. villosum Mill., S. grossedentatum A.

Rich., S. florulentum Bitter and 5. tarderemotum Bitter.

la. S. nigrum L. subsp. nigrum; Edmonds: 141—

178 (1977).

Herb about 1 m high, with abundant mostly appressed

simple hairs when young, subglabrescent when mature.

Stem robust. Leaves usually bearing simple hairs on both

surfaces and margin; margin repand or crenate with teeth

± 3 mm long; petiole 20-35 mm long. Inflorescence

lateral, lax cymes, 6-8-flowered. Corolla ± 4 mm long.

Fruit ± 7 mm diam.; fruit stalk decurved, deep purple to

shiny black when ripe. Seeds ±1x2 mm. 2n = 72 (Ed-

monds 1977). Pollen diam. 25.1-28 pm.

This subspecies and S. nigrum as a whole constitute a

Eurasian taxon. It has, however, spread to all other con-

tinents. The leaves are eaten in Uganda. Fruits are, how-

ever, reported to be poisonous (FAO 1988).

Vouchers: Ul: Karamoja. fl. & fir. Jan. 1958, Beadle 8. U3: Busoga

(0° 15', 33° 54') fl. Sept/ 1973. Wandera 10. U4: 0.5 km E of Port Bell

Pier, fl. & fr. Jan. 1969, Lye 1155.

2. Solanum americanum Mill, in Gardener’s dic-

tionary edn 8: (1768); D’Arcy; 735 (1973); Edmonds:

141-178 (1977). Type: cultivated Chelsea Physic Garden,

introduced from Virginia, North America, Miller s.n.

(BM).

The names S. nodiflorum Jacq. and 5. nigrum L. have

been widely and incorrectly used for this species. A

detailed list of synonyms has been given by Edmonds

(1972, 1979a).

Herb about 1 m high. Stem with simple hairs when

young, glabrous when mature. Leaves 50—1 30 x 30-60

mm; mature lamina glabrous, margin repand or with 2-3

pairs of lobes, ± 5 mm long; petiole up to 50 mm long.

Inflorescence lateral, umbellate cymes 2-10-flowered:

peduncle up to 30 mm long. Corolla ± 3 mm long, up to

10 mm wide. Fruit globose, up to 9 x 10 mm, shiny black

when ripe; fruit stalk 10-13 mm long, erect. Seeds 1 x 1

mm. 2n = 24 (Edmonds 1977). Pollen diam. 17.7-19.2

pm.

S. americanum is a morphologically variable species.

Edmonds (1977) divided it into two varieties, i.e. var.

americanum and var. patulum (L.) Edmonds. The former

is pilose and the latter glabrescent.

S. americanum is an introduced weed. Its leaves are

collected and eaten. It occurs in all the four geographical

regions of Uganda. It is also widely distributed throughout

Africa and around the world. It is of South American

origin, probably recently introduced into Africa. It is

believed to have donated at least 2 genomes to the

hexaploid S. nigrum L. (Edmonds 1979b).

Vouchers: Ul: Maraca Kijomoro W. Nile, fl. & fr. May 1971. Batia

19. U2: Rwenzori Mtn, fl. Aug. 1938, Purseglove 308. U3: Nakaloke

Mbale, fl. Aug. 1991, Bukenya 136. U4: M. 12 Kla-Ebb Rd, fl. & fr.

May 1932, Eggeling 414.

3. Solanum scabrum Mill, in Gardener's dictionary

edn 8: (1768); Edmonds: 141-178 (1977). Type: cul-

tivated Chelsea Physic Garden, origin North America,

Miller s.n. (BM) (fide Henderson 1974).

The synonymy has been discussed by Henderson

(1974) and Edmonds (1979a).

Bushy subshrub 1.0-1. 5 m high. Stem with simple hairs

when young, glabrous when mature. Leaves up to 250 x

200 mm; lamina glabrous; petiole up to 70 mm long. In-

florescence lateral, umbellate to racemose cymes, usually

6-23-flowered, often more; peduncle 20-25 mm long;

pedicel + 5 mm long. Corolla ± 7 x 15 mm. Anthers

purplish brown. Fndt broadly ovoid, 13 x 17 mm, dull

purple-black when ripe; fruit stalk erect or recurved 30-40

mm long. Seeds 1 x 1 mm. 2n = 72 (Edmonds 1977).

S. scabrum is a rare species in Uganda and is recorded

only from the extreme southwestern part. Its leaves are

eaten. In other parts of Africa, the West Indies and In-

donesia, it is cultivated for its leaves and young shoots

(FAO 1988). The fruits are eaten in Europe, where it is

known as ‘garden huckleberry’.

48

Bothalia 25,1 (1995)

Its origin is unknown (Edmonds 1979b). Some authors,

however, describe it as a native of Guinea. Heine (1960)

has pointed out that S. scabrum , ‘does not occur wild in

Guinea, but is cultivated there as a pot-herb and for medi-

cal purposes; nor apparently is it a native of any part of

Africa’. Morphologically it is distinct and genetically iso-

lated from other hexaploid solanums (Edmonds 1979b).

Vouchers: U2: Kigezi D.F.I., fl. & fr. Aug. 1972, Goode 3/72\ Bugan-

giri Ruzhumbura Kigezi, fl. & fr. Feb. 1949, Purseglove 2712.

4. Solanum sarrachoides Sendtn. in Martius, Flora

braziliensis 10: 18 (1846). Types: Brazil, Sellow s.n. (Bf;

P. lecto.) (fide Edmonds 1972, 1986). The typification of

this species is discussed by Edmonds (1986).

The synonymy is discussed by Edmonds (1972, 1986).

Erect and spreading herb up to 750 mm high with

dense glandular hairs. Leaves ovate, sinuate dentate, 30-

70 mm x 15-40 mm. Cymes simple, shortly racemose,

3-8-flowered. Flowers up to 10 mm diam. Fruiting

pedicels reflexed; fruiting calyces strongly adherent, ac-

crescent, enclosing at least the lower half of the fruit. Fruit

green to brownish purple when mature, less than 9 mm

diam. 2n - 24 (Edmonds 1972).

S. sarrachoides is a relatively rare species in Uganda.

It is not edible. It is originally from South America but

now established across tropical Africa and naturalized in

Europe.

Edmonds (1979b) suggested that this species might be

a diploid progenitor of the tetraploid S. villosum Mill, and

therefore may have played a role in the origin of S. nigrum

L. It is morphogenetically completely isolated from all

other diploid solanums (Edmonds 1977).

Vouchers: U2: nr footbridge, Nyabitaba Rwenzori, fl. Jan. 1969, Lye

1365\ Kigezi D.F.I., fl. & fr. Aug. 1972, Goode 5/72. U3: N.E. Elgon,

fl. & fr. Nov. 1951, Tweedie 1068.

5. Solanum villosum Mill, in Gardener’s dictionary

edn 8: (1768). Type: cultivated Chelsea Physic Garden,

introduced from Barbados. Miller s.n. (BM). [Lectotype

fide Edmonds 1979a; neotype fide Henderson (1977)].

Erect herb, sometimes woody, much branched, up to

0.9 m high, glabrescent to villose, with glandular or eglan-

dular hairs. Leaves ovate, entire or dentate 30-100 x 15-

40 mm. Inflorescence simple, 3-9-flowered; flowers

10-17 mm diam. Fruiting pedicels usually deflexed. Fruit

longer than broad, 6-10 mm diam., yellow, orange or red.

2n = 48 (Edmonds 1977).

Edmonds (1977, 1984) subdivided S. villosum into two

subspecies: subsp. villosum and subsp. miniatum (Bernh.

ex Willd.) Edmonds. The former has dense, mostly patent

glandular hairs whereas the latter has few to many ap-

pressed eglandular hairs. S. villosum is believed to have

hybridized with S. americanum and given rise to a sterile

triploid which through chromosome doubling gave rise to

S. nigrum (Edmonds 1979b). The origin of S. villosum is

not clear. Stebbins (1950) speculated that S. americanum

might have played a role in its origin. Edmonds (1979b)

suggested the possibility of S. sarrachoides being the

second parent of S. villosum.

S. villosum is native to Europe. It has become estab-

lished in Africa. In Uganda it occurs in three of the four

geographical regions. Its leaves are eaten as spinach.

Vouchers: Ul: Moroto Mtn. fl. & fr. Jan. 1959, Wilson 633 ; Kidepo

N.P. Dodoth, fr. Jan. 1973, Synnot 1404. U2: Ruizi R., fl. & fr. Apr.

1951, Jarrett 454. U3: Bujagali Falls, Jinja, fl. & fr. July 1952, Lind 82.

6. Solanum grossedentatum A. Rich., Tentamen

florae abyssinicae 2: 101 (1850). Type: Ethiopia, ‘crescit

in provincia Tchelikote', A. Petit s.n. (P).

Semiprocumbent herb up to 700 mm high, villose with

rust-coloured hairs. Leaves ovate, dentate to incised, 30-

70 x 20-40 mm. Cymes simple, subumbellate, 3-4-

flowered, pedicels reflexed. Corolla about 15 mm diam.

Fruiting pedicels reflexed. Ripe fruit black.

S. grossedentatum seems to be native to Africa. Its

ploidy level is unknown.

In Uganda it is fairly widely distributed, occurring in

three of the geographical regions. It is a crop weed. The

leaves are used as a vegetable and the fruits eaten by

children.

Vouchers: Ul: Paida W. Nile, fl. & fr. Aug. 1953, Chancellor 185.

U2: Muhokya Rwenzori, fl. Dec. 1925, Maitland 1290. U3: Budadiri

Bugishu, fl. & fr. Jan. 1932, Chandler 405.

7. Solanum florulentum Bitter in Feddes Reper-

torium 10: 544 (1912). Type: Tanzania, Albers 189 (Bf,

EA?).

Scrambling herb, up to 1 .5 m high, sparsely pubescent

with simple hairs. Leaves ovate to lanceolate up to 100 x

60 mm, petiole up to 20 mm long. Inflorescence branched

once, racemose, 8-20-flowered; inflorescence stalk ± 17

mm long; pedicels ± 8 mm long. Corolla 6-10 mm wide,

lobes ± 3 mm long. Fruit stalk reflexed, ± 10 mm long.

Fruit globose, 6x6 mm, purple to black when ripe.

Ploidy level unknown.

S. florulentum is fairly common in geographical

regions U2 and U4 of Uganda and it seems to be native

to East Africa. Its leaves are eaten. It has been confused

with S. nodiflorum Jacq. and S. nigrum L. sensu lato.

Vouchers: U2: Kigezi D.F.I., fl. & fr. Aug. 1972, Goode 2/72. U4:

Kituza, 35 ml SE of Kla, fl. & ff. June 1957, Griffiths 47.

8. Solanum tarderemotum Bitter in Feddes Reper-

torium 10: 547 (1912). Type: Tanzania Winkler 3856

(WRSL?).

Erect or scrambling herb, up to 2.5 m high, glabrous

or sparsely pubescent with simple hairs. Leaves ovate to

lanceolate, entire or sinuate-dentate, 70-180 x 35-70 mm.

Inflorescence simple, racemose, 7-12-flowered. Flowers

7-9 mm diam. Fruiting pedicels reflexed. Fruit greenish

yellow or purple when ripe, 4-6 mm diam. Ploidy level

unknown.

Bothalia 25, 1 (1995)

49

S. tarderemotum is closely related to S. florulentum hut

differentiated from it by its simple inflorescence. It also

seems to be native to East Africa. In Uganda it is repre-

sented in all the geographical regions. Its leaves are eaten.

Vouchers: Ul: Terego W. Nile, fl. & fr. Apr. 1938, Hazel 486. U2:

Kigezi D.F.I., fl. & fr. Aug. 1972, Goode 1/72. U3: Budadiri Bugishu,

fl. & fr. Jan. 1932, Chandler 458. U4: Nakyesanja nr Kawanda, fl. & fr.

Mar. 1972, Damulira 35.

Section Afrosolanum Bitter in Botanische Jahr-

biicher 54: 44CM87 (1917); D'Arcy: 266, 274 (1972).

Type species: S. tenninale Forssk.

This is a rather difficult section, complicated by species

plasticity and the existence of numerous specific and in-

fraspecific names. Heine (1960) adopted a broad view of

the species. He considered all material in this section from

the FWTA (Heine 1963) area to be S. tenninale. This was

split into three subspecies: inconstans, sanaganum and

welwitschii. Jaeger (1985) recommended three species for

this section: S. nakurense, S. tenninale and S. welwitschii.

The subspecies of S. tenninale are not very clear cut.

9. Solanum terminale Forssk., Flora aegyptiaco-

arabica: 45 (1775); Bitter: 301 (1922); Heine: 247 (1960).

Type: Yemen, Mokhaja, Forsskdl s.n. (C, IDC microf.

2200 102: II.3-6).

9a. Solanum terminale Forssk. subsp. terminale

Heine in Kew Bulletin 14: 247 (1960).

The synonymy is discussed by Heine (1960, 1963) and

Jaeger (1985).

Liana about 4 m tall. Leaves up to 110 x 65 mm.

petiole up to 80 mm long. Inflorescence very many-

flowered, often > 50 flowers; pedicels to ± 10 mm long;

peduncles with terminal umbel, occasionally also with a

few lateral umbels; lateral umbels subsessile or oc-

casionally with the lowest on short branches. Corolla ± 8

mm long, whitish purple or bluish purple. Fruit globose,

± 8 mm diam., red when ripe.

Subsp. tenninale is quite widely distributed in Uganda

and eastern Africa from Ethiopia to South Africa.

Vouchers: U2: Kigezi, fl. Feb. 1956, M.C. 909\ Kalinzu Forest, Igara,

fr. June 1970, Katende 352. U3: Bunya, Busoga, fl. Nov. 1937. Webb

62. U4: 2 km E of Bujuko-Mubende Rd. fl. Feb. 1969, Lye 1950.

9b. Solanum terminale subsp. sanaganum ( Bitter )

Heine in Kew Bulletin 14: 248 (1960). Type: Cameroon,

nr Deng Deng, Mildbraed 8619 (K!).

For synonymy see Heine (1960, 1963) and Jaeger

(1985).

Climber about 6 m high. Leaves about 90 x 40 mm;

petiole about 40 mm long. Inflorescence many(-50)-

flowered; peduncles terminal or subtenninal, strongly

branched giving the inflorescence a paniculate ap-

pearance; pedicels ± 10 mm long. Corolla ± 8 mm long.

whitish purple. Fruit globose or slightly longer than wide,

8x6 mm.

Subsp. sanaganum is quite close to subsp. terminale

in both vegetative and floral characters. The main dif-

ference between them is that the former has a paniculate,

the latter an umbellate type of inflorescence. In Uganda

it is less common than subsp. terminale but it is widely

distributed in upland forests of tropical Africa.

Vouchers: U2: Echuya F.R. Kigezi, fl. Apr. 1970, Katende 221. U4:

Entebbe, fl. & fr.. Brown 14\ Mabira Forest, fl. Feb. 1972, Katende 1526.

9c. Solanum terminale subsp. inconstans (C.H.

Wright) Heine in Kew Bulletin 14: 247 (1960); Heine:

331 (1963); Gbile: 118 (1979). Syntypes: Fernando Po,

Mann 62 (K); Cameroons, Kalbreyer 172 (K).

For synonymy see Heine (1960).

Slender woody climber. Stem flexuose with long inter-

nodes. Leaves up to 100 x 50 mm; petiole up to 18 mm

long. Inflorescence subracemose, leaf opposed or in leaf

axil; few (2-7)-flowered; peduncle up to 35 mm long;

pedicel ± 14 mm long. Corolla 8-10 mm long. Fruit spindle-

shaped, ± 25 x 10 mm; fruit stalk ± 28 mm long.

Subsp. inconstans is rare, found in disturbed forest. In

Uganda it has been collected from Mabira Forest.

Vouchers: U4: Mabira Forest, fl. Sept. 1938, Loveridge 75.

10. Solanum welwitschii C.H. Wright in Kew Bul-

letin 1894: 126 (1894). Syntypes: Angola, Welwitsch 6081,

6098 (K!).

A list of synonyms is provided by Jaeger (1985).

Slender woody climber. Leaves elliptic, up to 170 x

80 mm; petiole up to 120 mm long. Inflorescence ter-

minal, spiciform, up to 200 mm long; flowers > 50, bluish

purple, in cymules, subsessile on axis. Corolla 8-10 mm

long. Fruit globose, to 10 mm diam., red when ripe; fruit

stalk to 6 mm long.

S. welwitschii is found in the forests of western and

central Uganda. It also occurs in secondary forests of

western tropical Africa.

Vouchers: U2: Siba Forest Kinkizi, fl. & fr. May 1951, Dawkins 751 .

U4: Mabira Forest, fl. Sept. 1933, Bransnett s.n.

11. Solanum nakurense C.H. Wright in Kew Bul-

letin: 275 (1897). Type: Kenya, Nakuru, Scott Elliott 6800

(K!).

For synonymy see Jaeger (1985).

Erect subshrub to 2 m high. Stem verrucose. Leaves

variable, 20-70 x 10-40 mm, ovate, with scattered simple

hairs on upper surface; lower surface sparse to densely

hairy. Inflorescence umbellate; peduncle rarely branched,

5-10(-20)-flowered. Flowers white, orange to violet;

corolla 6-10 mm long, to 15 mm diam.; pedicel to 12

50

Bothalia 25,1 (1995)

mm long, strongly reflexed at anthesis. Fruit red, globose,

8 mm diam.

S. nakurense is morphologically similar to S. terminate.

It differs from the latter by being erect, and having smaller

leaves which are more hairy than in S. terminate. Its in-

florescence is also generally simpler than in S. terminate.

S. nakurense is relatively rare in Uganda. It generally

prefers the upland woodlands and open habitats.

Vouchers: U2: Kanaba gap Kabale-Kisoro, fl. Oct. 1960, Miller 448.

U3: Namasindwa, Mt Elgon, fl. May 1924, Snowden 889/a-, nr Kapkwata

Forest St. Sebei, fl. Jan. 1969, Lye 1571.

Section Benderianum Bitter in Botanische Jahrbiicher

54: 487-489 (1917). Type species: S. benderianum Schimp.

ex Dammer.

1 2. Solanum benderianum Schimp. ex Dammer in

Botanische Jahrbiicher 38: 184 (1906). Syntypes: Ethiopia,

near Gaffat, Schimper 1227 (1863) (E); Uganda, Rwenzori

Mts, Scott Elliott 7733 (K! ).

Climbing shrub. Leaves lanceolate, ± glabrous. In-

florescence terminal, a lax cymose panicle with > 50

flowers. Corolla violet, ± 20 mm diam.

The more or less glabrous S. benderianum is closely

related to the hairy S. runsoriense. It is very rare in Ugan-

da, only recorded from the Rwenzori Mts. It is common

in Ethiopia, growing at an altitude of 2 500-3 600 m. S.

macrothyrsum Dammer from the Comoro Islands is

probably synonymous with S. benderianum (Jaeger 1985).

Voucher: U2: Rwenzori, fl. & fr. 1916, Fyffe 20.

13. Solanum runsoriense C.H. Wright in Johnston,

Uganda Protectorate 1: 362 (1902). Type: Uganda, Rwen-

zori Mts, Doggett s.n. (K!).

Climbing shrub to 4 m high, with dense mealy pubes-

cence of much branched hairs. Leaves lanceolate upper

surface with light cover of mostly simple hairs; lower sur-

face with a heavy cover of branched hairs. Inflorescence

terminal, a lax cymose panicle with ± 50 or more flowers.

Corolla light blue to purple, ± 20 mm diam.; filaments

sometimes 3 mm long, anthers slightly longer, dehiscing

by terminal pores and longitudinal slits running down-

wards from the pore.

S. runsoriense is a montane forest species occurring in

Uganda on Rwenzori Mtn in the west and Mt Elgon in

the east at or above the bamboo zone. It also occurs in

Kenya at an altitude of 2 500-3 000 m.

Jaeger (1985) suggested that S. longipedicellatum De

Wild., S. dewildemanianum Robyns and S. keniense

Standi, are probably later synonyms of S. runsoriense.

Vouchers: U2: Mubuku Valley, Rwenzori, fl. July 1938, Eggeling

3792. U3: above bamboo zone Elgon, II. Apr. 1930, Liebenberg 1637.

B. Subgenus Brevantherum (Seithe) D'Arcv in An-

nals of Missouri Botanical Garden 59: 267-274 (1972);

D’Arcy: 713 (1983). Type species: S. erianthum D. Don.

Plants unarmed. Hairs branched or stellate. Leaves

mostly entire or nearly so. Anthers stout, opening by large

terminal pores and sometimes ultimately by longitudinal

slits. Ovary glabrous to tomentose. Fruit rather small,

7-20 mm diam. The subgenus is represented in Uganda

by one section: Brevantherum.

Section Brevantherum Seithe in Botanische Jahrbiicher

81: 297 (1962). Type species: S. erianthum D. Don.

14. Solanum mauritianum Scop., Deliciae florae

et faunae insubricae 3: 16 t. 8 (1788). Type: Scop.: t. 8

(1788) (fide Roe 1972).

For synonymy see Heine (1963).

Shrub or small tree 4-7 m high, unarmed, young stem

densely covered with sessile stellate hairs, stem with axil-

lary subactive buds bearing small leaves. Leaves elliptic-

ovate, entire, up to 250 x 100 mm, apices acute, bases

attenuate; both surfaces of leaves densely covered with

stellate hairs; petiole to 70 mm long. Inflorescence ter-

minal, paniculate, many(> 50)-flowered. Flowers purple;

pedicels ± 5 mm long. Corolla 15 mm diam., lobes 5 mm

long. Anthers differing slightly in length, i.e. 1 short (±

1.8 mm), 2 or 3 medium (2 mm) and 2 longest (2.4 mm).

Fruit spherical, pubescent, green when young, yellow

when mature, 12 x 11 mm; stalk ± 5 mm long. Seeds 2

x 2 mm. 2n = 24 (D’Arcy 1974).

S. mauritianum is a widespread weed in Uganda and

other parts of tropical Africa. The closely related species

S. erianthum D. Don. and S. umbellatum Mill., established

elsewhere in Africa (Bukenya & Hall 1988), have not yet

been recorded for Uganda.

S. mauritianum, S. erianthum and S. umbellatum are

native to the Americas. Their spread to Africa has been

associated with the 16th century Spanish and Portuguese

trade routes (Roe 1979). These species are colonizers of

open ground: forest openings, stream borders and areas

of human disturbance such as roadsides.

Reproduction in these plants is not only by seed, but

apparently more commonly by adventitious roots from

shallow roots to form large colonies. They are self-com-

patible, another characteristic of successful weeds. The

colourful berries and frequently isolated plants suggest

bird dispersal of seed (Roe 1979).

S. mauritianum has been widely known in the past as

S. auriculatum Aiton but the publication of S. mauri-

tianum antedates this name by one year.

Voucher: U4: Nakiyaga Masaka, fl. & fr. Sept. 1989. Bukenya 49.

C. Subgenus Leptostemonum (Dunal) Bitter in

Botanische Jahrbiicher 55: 69-89 (1919); D’Arcy: 684

(1973); Whalen: 179-282 (1984). Lectotype: S. mam-

mosum L. (fide D’Arcy 1972).

Bothalia 25,1 (1995)

51

Indumentum often stellate, prickles usually present.

Anthers mostly slender, tapering to the tip and opening

by small terminal pores or, if stout, narrowing abruptly to

a small tip and also often opening by longitudinal slits

near base, dehiscing introrsely or extrorsely by outward

bending of the tips. Ovary glabrous. Fruit often large (7-

) 1 0— 80(— 90— 1 30 mm).

This subgenus is the largest in Uganda, represented by

six sections: Leptostemonum ( Acanthophora ), Aculeigerum,

Melongena, Monodolichopus, Oliganthes and Torva.

Section Acanthophora Dunal, Histoire Naturelle,

Medicale et Economique des Solatium : 131, 132 (1813);

D'Arcy: 909 (1973). Type species: S. mammosum L. (fide

D'Arcy 1972).

15. Solanum mammosum L., Species plantarum:

187 (1753); D’Arcy; 712 (1973); Nee: 576 (1979). Fec-

totype: Pluk.: t. 226, fig. 1 (1696). (Typotype: Herb.

Sloane Vol. 98 fol. 59 (BM). For reasons for type selection

see Jaeger ( 1985).

A list of synonyms is given by Whalen (1984).

Shrub about 1.5 m high. Stems densely covered with

simple hairs ± 2 mm long, and slightly decurved prickles

that are 5 mm long with base 2 mm broad. Leaves ±110

x 90 mm, lobed or doubly lobed to about Vl width of

leaf; lobes triangular; prickles on midrib straight, ± 17

mm long, base ± 1 mm broad, on primary lateral veins

3-8 mm long, hairs on upper surface mainly simple, on

lower surface stellate mixed with simple hairs; petiole up

to 70 mm long with simple hairs and straight prickles ±

13 mm long, base 1 mm broad. Inflorescence 3-4-

llowered; pedicel ± 7 mm long. Corolla violet or blue, ±

12 mm long. Fruit up to 50 mm wide, bearing a terminal

nipple or mammilla, 5 mammillae or protuberances often

present at base. Seeds brown. 2n = 22, 24 (Heiser 1971).

S. mammosum is an introduced ornamental which is

rarely found in Uganda, being native to the Caribbean

region of central America where it is found in disturbed

habitats and where it is cultivated for its curious fruits

which are used as a medicine and as a cockroach poison

(Duke 1970).

Vouchers: U3: Serere, Teso, fr. Dec. 1931, Chandler 203. U4:

Makerere Univ., fl. & fr. Feb. 1992, Bukenya 143 (seed from Nakasero

nr Kampala).

16. Solanum aculeatissimum Jacq., leones plan-

tarum rariorum 1: 5, t. 41 (1781); C.H. Wright: 228

(1906); Bitter: 148 (1923); Dalziel: 432 (1937); Heine:

535 (1963); Gbile: 115 (1979). Type: cultivated plant at

Vienna, Jacquin s.n. (W).

For synonymy see Whalen (1984).

Shrub ± 1.5 m high, densely pubescent with simple

hairs 0. 1-1.0 mm long, decurved prickles up to 12 mm

long with base 0.5 mm broad. Leaves ± 180 x 160 mm,

lobed or doubly lobed to Vi width of leaf, with 3-4 pairs

of major triangular lobes; middle lobe up to 55 mm long;

simple pilose hairs on both surfaces or leaves, rare stellate

hairs on underside; prickles on midrib and petiole slender,

± 15 mm long, base 0.8 mm broad, purple near base,

upper part yellow; petiole 4-80 mm long. Inflorescence

3-6-flowered. Corolla white, ± 17 x 30 mm, peduncle

0.5 mm long; pedicel ± 1 1 mm long. Fruit globose, ± 35

mm diam. Seeds 3x4 mm, brownish.

S. aculeatissimum generally grows in forest clearings.

It is widely distributed throughout Africa. It also occurs

in southeastern Brazil (Whalen 1984). It is likely to have

been introduced to Africa several hundred years ago

(Jaeger 1985).

Vouchers: Ul: Kidepo N.P., fl. & fr. Sept. 1972, Synnott 1269. U2:

Kizimba Bundibugyo, fl. Sept. 1991, Bukenya 146. U4: Entebbe Region,

Mar. 1921, Kew Herb, sheet No. 681.

Section Aculeigerum Seithe in Botanische Jahr-

biicher 81: 291, 292 (1962). Type species: S. wendlandii

Hook.

17. Solanum wendlandii Hook, in Curtis's Botani-

cal Magazine 113: t. 6914 (1887). Type: cultivated at Kew

from Costa Rican seed (K! ).

Woody climber, armed with small recurved prickles on

stem and midribs of leaves. Leaves up to 150 x 120 mm.

broadly elliptic, compound; at base, leaflets separate,

towards apex lamina only deeply divided. Inflorescence

many (50+)-flowered. Inflorescence axis very long, ± 300

mm. Flowers purple; pedicel 20 mm long. Calyx 3 mm

long. Corolla 50 mm wide. Stamens of different filament

lengths: 5 mm (1), 3 mm (2) and 2 mm (2); anther tips

purple, lower part yellow. 2n = 24 (Federov 1969; Whalen

1984).

S. wendlandii is widely cultivated in the tropics as an

ornamental for its showy flowers. It flowers regularly but

the senior author has never seen it fruiting. It is probably

a native of Costa Rica (Whalen 1984).

Vouchers: U4: Ml 2 Gayaza Rd. fl. Aug. 1990, Bukenya 53\ Buganda

Road RS„ fl. Apr. 1990. Bukenya 44\ Masaka City, fl. May 1952, Lye

6840.

Section Melongena Dunal, Histoire Naturelle. Medi-

cinale et Economique des Solanum : 208-218 (1813);

D'Arcy: 698 (1972). Type species: S. melongena F. (fide

D'Arcy 1972).

18. Solanum melongena L., Species plantarum:

186 (1753); Wright: 242 (1906); Bitter: 292 (1923);

Heine: 322 (1963); D'Arcy: 704 (1973); Hepper: 122

(1976); Khan: 630 (1979). Type: lectotypified by D'Arcy

(1974) with LINN 248.28 (IDC microf. 139: ff.2).

A list of synonyms is given by Heine (1963) and

Whalen (1984).

Shrub + 1.5 m high. Stem with stellate hairs of 8-10

unequal arms. Leaves ± 150 x 100 mm, bearing on both

surfaces short, stalked, stellate hairs, margin with 2-3

pairs of lobes up to 30 mm long; petiole about 40 mm

52

Bothalia 25. 1 (1995)

long. Flowers usually solitary or inflorescence 2-3-

flowered; pedicel ± 25 mm long. Corolla 20-22 mm long,

purple, petal tips apiculate, incurved. Calyx about 10 mm

long, normally not prickly but prickles 2-3 mm long may

occur. Fruit ovoid or globose, 60-130 x 30-100 mm,

green, with white patches, white or purple when young,

orange-yellow to brownish when ripe; fruit stalk 20-80

mm long, decurved. Seeds 3x4 mm. 2n = 24, but

tetraploid forms occur (Whalen 1984).

S. melongena is an important commercial fruit/

vegetable in Uganda. The species is believed to have

originated in Asia (Indo-Burma) and is now cultivated on

all continents for its edible fruits. S. melongena is believed

to have evolved from S. incanum sensu lato which

migrated into Asia from northeastern Africa and the Mid-

dle East (Lester & Hasan 1991).

Vouchers: U2: Kyembogo D.F.I., fl. & fr. Oct. 1990, Bukenya 108.

U3: Kyabirwa, Budondo-Jinja, fl. Aug. 1991, Bukenya 130. U4:

Bukanaga, Mityana, fl. & fr. Aug. 1990, Bukenya 71.

19. Solanum aculeastrum Dunal in A.P. de Can-

dolle, Prodromus systematis naturalis regni vegetabilis

13,1; 366 (1852). Type: South Africa, Cape Province,

Morley, Drege (Vb, as S. sodomeum ) (G-DC, IDC microf.

2091: III.4).

For synonymy see Heine (1963), Whalen (1984) and

Jaeger (1985).

Shrub to small tree up to 7 m high. Leaves with deep

lobes about 30 mm long, upper surface glabrous, lower

with a heavy cushion of stellate hairs; petiole short, about

20 mm long. Inflorescence few- to 10-flowered, only 1

or 2 flowers functionally female. Flowers white, ± 25 mm

diam., lobes + 10 mm long; pedicel ± 15 mm long. An-

thers ± of equal length. Fruit warted, yellow when ripe,

40 x 42 mm; stalk slightly decurved, about 30 mm long.

Seeds 4x3 mm. 2n = 24 (Whalen 1984).

S. aculeastrum is a native African species extending

eastwards from Imatong Mts in Sudan to the Cape in

South Africa and westwards from Cameroon highlands. It

is rather variable. Jaeger (1985) divided it into 4 sub-

species (subsp. I ; subsp. 2; subsp. aculeastrum and subsp.

thomsonii). It has many traditional uses (Bukenya 1993).

Vouchers: U2: Mushungyero, Bufumbira, 0. & fr. 1970, Katende 153.

U3: Buwenda-Busoga, fl. July 1945, Wakabi 41; Iganga Town, fl. & fr.

Aug. 1991, Bukenya 135. U4: Wattuba nr Kiboga. fl. & fr. Aug. 1970,

Katende 498.

20. Solanum incanum L., Species plantarum: 188

(1753); C.H. Wright: 238 (1906); Bitter: 200 (1923); Dal-

ziel: 433 (1937); Heine: 332 (1963); Gbile: 118 (1979).

All the above authors have taken S. incanum in a broad

sense.

The typification of S. incanum L. is discussed in a

paper by Hepper & Jaeger (1985), who selected a neotype

(Herb. J. Burser vol. 9 No. 20 UPS IDC microf. series

No. 1064 51: 1.4).

An extensive list of synonyms is provided by Heine

(1963), Whalen (1984) and Jaeger (1985).

Shrub 1. 0-3.0 m high; stem with stellate floccose hairs

with 8-10 more or less equal arms; prickles ± 5 mm long

with base ± 2 mm broad. Leaves ± 240 x 120 mm; margin

sinuate or with 3 pairs of short lobes; middle lobe up to

20 mm long; both surfaces with stellate hairs; prickles on

midrib ± 3 mm long with base ± 0.5 mm broad; petiole

up to 70 mm long. Inflorescence up to 10-flowered; 1-5

flowers functionally female; pedicel ± 10 mm long. Corol-

la violet, ± 30 x 27 mm. Calyx, especially on the lower-

most flower, very prickly, prickles ± 2 mm long. Fruits

globose, 30 x ± 27 mm, green with light green patches

when young, yellow when ripe.

S. incanum sensu lato is extremely common in Uganda,

occurring in all the geographical regions and different

habitats. It is polymorphic, and its intraspecific variation

requires a thorough study, especially experimental work.

Jaeger (1985) treated S. incanum in Africa as a species

aggregate and divided it into five groups. These include

S. incanum group which occurs in the dry country scrub

of NE Africa and the middle East; S. lichtensteinii group

which is found in wooded grasslands of southern Africa;

S. panduriforme group which occurs in eastern southern

Africa; S. campylacanthum group, a common shrub of

disturbed ground in the grasslands of central. East and

southern Africa and S. cerasiferum group which occurs in

NE Africa extending to northern Nigeria. Lester & Hasan

(1991) divided S. incanum sensu lato into four groups:

Group A, including S. lojeru Dunal, S. campylacanthum

Dunal, S. delagoense Dunal and many other species

recognised by Bitter (1923); Group B containing S. pan-

duriforme Dunal only and has the narrowest leaves of all

the species; Group C being S. incanum L. sensu stricto

(Hepper & Jaeger 1985); and Group D containing S. lich-

tensteinii Willd. and allied taxa.

The Ugandan material of S. incanum sensu lato pos-

sibly belongs to Jaeger's S. panduriforme group (Lester

& Hasan’s Group B and Jaeger’s S. campylacanthum

group (Lester & Hasan’s Group A).

Vouchers: Ul: Amolotar Lango, fl. Sept. 1946, Dawkins 218. U2: 5

km NE of Nyarushanje, fl. & fr. Dec. 1968, Lye 1082. U3: Serere, tl.

Sept. 1970, Edam 2. U4: Mulago, fl. Sept. 1967. Mweheire 20; Kasa

Forest nr L. Wamala, IT Nov. 1949, Dawkins 454.

21. Solanum macrocarpon L., Mantissa altera: 205

(1771); C.H. Wright: 214 (1906); Bitter: 195 (1923);

Burkill: 333 (1925); Dalziel: 343 (1937); Heine: 234

(1963). Lectotype: Sheet 248.11 in LINN (IDC microf.

138: II. 3). The selection of this lectotype is discussed by

Jaeger ( 1985).

The synonymy is discussed by Burkill (1925), Heine

(1963), Whalen (1984), Jaeger (1985) and Bukenya

(1993).

Subshrub or shrub, 0.5-1. 5 m high. Stem terete,

glabrous or with stellate hairs; not prickly or with prickles

± 6 mm long. Leaves 150-460 x 80-300 mm, entire or

with short lobes, ± 10 mm long to deeply doubly lobed

Bothalia 25,1 (1995)

53

with major lobes up to 80 mm long; young leaves bear

on upper surface simple, or stellate hairs, either singly or

in combination; lower surface with stalked stellate or more

or less sessile stellate hairs; mature leaves glabrous or with

simple hairs and stellate hairs; prickles present or absent

on leaves, when present, principally on midrib and lateral

veins; petiole very short to 70 mm long. Inflorescence

lateral, racemose, 3-12-flowered. Flowers : lowermost

flower or flowers hermaphrodite, larger than the rest and

functionally female, distal flowers with short styles, func-

tionally male. Normally 1— 2(— 5) hermaphrodite plus 1-4

functionally male flowers present. Corolla infundibuli-

form-rotate or campanulate, 20-35 mm long, white, light

purplish or blue. Calyx not prickly or with prickles ± 10

mm long; fruiting calyx often accrescent, 15-50 mm long.

Ovary glabrous or with short-stalked or sessile glandular

hairs. Fruit depressed globose, 20-60 x 30-100 mm,

green, ivory or purplish white with dark stripes; when

ripe, yellow to brownish; stalk erect or decurved, 10-40

mm long. Seeds 3.0-4.5 x 2.0-3.6 mm. 2n = 24.

The S. macrocarpon complex is extremely variable. In

this treatment the complex is taken as a combination of

S. macrocarpon L. and S. dasyphyllum Schum. & Thonn.

This is because Bukenya (1993) obtained fully fertile FI

and F2 hybrids (pollen stainability was 80-100%) be-

tween what was previously known as S. macrocarpon and

S. dasyphyllum. All accessions of these taxa and of hybrids

between them showed 12 bivalents at metaphase- 1 of

meiosis and regular disjunction indicating that they belong

to the same biological species. Bukenya (1993) split the

complex in Uganda into four major groups: two are cul-

tivars, one is a semi-wild and the fourth is the wild group

(formerly called 5. dasyphyllum).

Seme (1983), Bista (1983) and Jaeger (1985) were also

of the opinion that the separation of S. macrocarpon and

S. dasyphyllum is no longer justifiable. The S. macrocar-

pon complex is closely related to S. sessilistellatum Bitter

which is endemic to Kenya, but distantly related to the S.

incanum group (Whalen 1984).

The S. macrocarpon complex is native to Africa from

where cultivars were introduced to other parts of the

world. In Uganda, the complex, especially the wild group,

is widely distributed.

S. macrocarpon (cultivated group)

Vouchers: U3: Nkoma Bujoloto, fl. Aug. 1991, Bukenya 138. U4:

Nabingo, fl. & fr. Sept. 1990, Bukenya 80.

S. macrocarpon (semi-wild group)

Vouchers: U3: Budondo Jinja, fl. Aug. 1991, Bukenya 131. U4:

Nabingo, fl. & fr. Oct. 1990. Bukenya 83.

S. macrocarpon (wild group)

Vouchers: Ul: Napak Karamoja, fl. & fr. July 1948, Philip 203. U2:

Mile 8 Kingumya Bwamba, fl. & fr. Sept. 1991, Bukenya 148. U4:

Kivuuvu-Mityana, fl. & fr. Aug. 1990, Bukenya 61. 62.

22. Solanum wrightii Benth. in Flora hongkongen-

sis: 243 (1861); Bitter: 180 (1923); Heine: 248 (1960);

Irvine: 733 (1961); Heine: 335 (1963); Gbile: 1 19 (1979).

Type: Hong Kong, C. Wright 489 ( K !).

For synonyms see Heine (1960, 1963) and Jaeger

(1985).

Small tree up to 10 m high. Stem bearing prickles, and

stellate hairs on setae ± 1 mm long. Leaves ± 300 x 240

mm, with 2-3 pairs of prominent lobes up to 90 mm long,

upper surface bearing simple hairs ± 1 mm long, lower

surface with stellate hairs with 5-8 unequal arms on setae

± 0.5 mm long; prickles on midrib 10-20 mm long, base

4 mm broad; petiole ± 100 mm long; prickles on petiole

up to 20 mm long with base 6 mm broad. Inflorescence

> 20-flowered. Corolla blue to violet, turning white with

age, up to 45 x 80 mm. Calyx ± 20 mm long; pedicel ±

25 mm long. Anthers of different length (2 ± 20 mm long,

2 medium, 1 short). Fruit globose, green when young with

light green patches, yellow when ripe, 50 x 55 mm; fruit-

ing calyx about 20 mm long.

S. wrightii is an introduced decorative tree which is a

native of Bolivia. It has been introduced to other tropical

areas of the world.

Vouchers: U2: Kyembogo D.F.I., fl. & fr. Oct. 1990, Bukenya 107.

U3: Jinja town, fl. Aug. 1991, Bukenya 125. U4: Mityana S.S., fl. & fr.

Aug. 1990, Bukenya 69. 70.

Section Monodolichopus Bitter in Feddes Reper-

torium Beih. 16: 297-307 (1923). Type species: S.

coagulans Forssk. [ fide Seilhe (1962) as S. dubium

Fresen.j.

23. Solanum coagulans Forssk. Flora aegyptiaco-

arabica 108 and 47. Cent. 1155 (1775). Type: Yemen,

Forsskdl s.n. (C).

Perennial herb, often with several stems up to 0.7 m

high. Stems with white stellate hairs, usually covered with

dense straight yellow prickles. Leaves very variable, up

to 100 x 50 mm or more, ovate-lanceolate, prickly and

hairy. Inflorescence racemose, with 6-10 flowers. Corolla

blue-violet, 10-15 mm diam. Stamens unequal, one fila-

ment slightly longer than rest. Fruit yellow, globose, 10

mm diam., dry, usually completely enclosed by heavily

armed accrescent calyx. Seeds shiny black.

Jaeger (1985) took a broad view of the species to in-

clude S. dubium Fresen., S. thruppi C.H. Wright, S.

depression Bitter and S. ellenbeckii Dammer. On the other

hand Whalen (1984) used S. thruppi for S. dubium and

commented that: ‘S. dubium was published by Fresenius

in 1834 but was pre-dated by S. dubium Dunal (1813) an

unrelated species. A still earlier name is S. coagulans

Forssk. (1775), but that epithet has been persistently

misapplied and probably should be rejected’. This con-

troversy needs to be resolved.

S. coagulans is not common in Uganda. It has been

collected from Ankole and Karamoja. These are pastoral

areas and the species is noted to withstand overgrazing

pressures due to its heavy armature and creeping nature.

On the African continent, it occurs from Egypt to Tan-

54

Bothalia 25,1 f 1995)

zania. It has no clear relatives among the Old World

solanums (Whalen 1984).

Vouchers: Ul: Kangole, Karamoja, fl. & fr. May 1940, Thomas 3482.

U2: Mbarara, 11 & fr. 1925, Maitland 1377.

Section Oliganthes (Dunal) Bitter in Feddes Reper-

toriumBeih. 16: 1 (1923); D’Arcy: 272 (1972). Lectotype:

S. indicum auct. non L. (= S. anguivi Lam.), fide D’Arcy

(1972).

24. Solanum anguivi Lam. in Tableau Encyclo-

pedique et Methodique: 23 (1794); Hepper: 287-292

(1978). Type: Madagascar, Commerson s.n. Holotype:

MPU, syntype P, fide Hepper (1978). According to

D'Arcy & Rakotozafy (1994), Hepper (1978) has chosen

a lectotype, and not indicated the holotype, because a

specimen exists in P-JU which Lamarck is sure to have

examined.

For synonymy see Whalen ( 1984) and Bukenya & Hall

(1988).

Shrub 1 .5-3.0 m high. Stem bearing small, sessile stel-

late hairs with 4-8 ± equal arms; sometimes prickly.

Leaves elliptic-ovate, 100-200 x 50-100 mm, sinuate to

distinctly lobed, 2-4 pairs of lobes, lobes 20-30 mm long,

apex acute to obtuse, base oblique, occasionally truncate

or subcordate; both surfaces of leaves have ± sessile stel-

late hairs with 6-10 ± equal arms; petiole 20-60 mm long,

with dense stellate hairs. Inflorescence a raceme-like

cyme, 5 to > 15-flowered, occasionally flowers solitary,

extra-axillary, flowers mostly hermaphrodite, occasionally

distal flowers with short styles, functionally male;

peduncle 0-3-6 mm long; pedicel 4—15 mm long, bearing

stellate hairs. Corolla ± 10 x 10 mm, white, occasionally

with light purple veins on outer surface; with stellate hairs

outside, ± glabrous on inner surface. Fruit 7-18 x 8-12

mm, mostly globose, smooth, green or white when young,

red when ripe; stalk 8-15 mm long, usually erect, oc-

casionally horizontal or decurved. Seeds 1 .5-2. 1 x 1 .9-2.9

mm.

S. anguivi is a rather polymorphic species. It exhibits

tremendous variation in features such as prickliness,

pubescence and inflorescence. This variation is possibly

partly due to domestication and partly to selection. There

has been a shift from prickly, many-flowered and small-

fruited types to prickless, less-flowered and large-fruited

types (Bukenya 1980).

Bitter (1923) recognised more than 10 subspecies and

several varieties for this species. Jaeger ( 1985) recognised

five subspecies of S. anguivi. A revision of Bitter’s in-

fraspecific classification is necessary, following ex-

perimental work. Many of Bitter’s infraspecific names are

likely to be reduced to synonyms. No infraspecific com-

binations of Bitter’s subspecific names have yet been pub-

lished for S. anguivi (Jaeger 1985).

Lester & Niakan (1986) demonstrated that S. anguivi

is most likely the wild progenitor of the red or orange-

fruited garden eggplants (S. aethiopicum) commonly cul-

tivated in Africa.

In Uganda S. anguivi is a minor crop, grown for its

fruits. It also grows as a weed, possibly dispersed by birds.

S. anguivi is widely distributed on the African continent

and its neighbouring islands, e.g. Madagascar.

Vouchers: Ul: Napak. Karamoja, fl. & fr. June 1950, Eggeling 5956.

U2: Nyaruzinga Bushenyi, fl. & fr. Oct. 1990, Bukenya 104 ; Kalinzu

Forest, fr. Mar. 1970, Synnott 415. U3: Budondo Jinja, fr. Aug. 1991,

Bukenya 126. U4: Bukenaga Mityana, IF & fr. Aug. 1990, Bukenya 67.

25. Solanum aethiopicum L., Amoenitates aca-

demicae: 307 (1759); Dalziel: 432 (1937); Heine: 332

(1963); Gbile: 115 (1979). Lectotype: ‘Habitat in

Aethiopia’, Burser vol. 9, No. 17 (IDC microf. 1064, 51:

2) selected by Hepper & Jaeger (1985).

Lester & Niakan’s (1986) biosystematic studies

showed that all the African taxa of Solanum section

Oliganthes series Aethiopica recognised by Bitter (1923):

S. gilo Raddi (including S. olivare Paill. & Bois); S. zuc-

cagnianum (= S. aethiopicum L. sensu Bitter); S. aethio-

picum L. and S. aethiopicum var. aculeatum (- S. integri-

folium auct. non Poir.), comprise a single species. They

gave these four taxa non-Linnaean names under S.

aethiopicum L. These are Gilo group, Shum group,

Kuirtba group and Aculeatum group, respectively.

The Gilo group is cultivated in Africa for its fruits;

Aculeatum group is grown in several European gardens,

not African; Shum group is a leafy vegetable in tropical

Africa, and Kumba group is cultivated for its large fruits

and leaves, especially around the Niger River.

The Gilo and Shum groups occur in Uganda. Lester

& Niakan’s system (1986) is followed.

25a. Solanum aethiopicum Gilo group

For synonymy see Heine (1963), Whalen (1984) and

Jaeger (1985).

Shrub 1-2 m high; stem bearing tiny sessile stellate

hairs of 4—5 ± equal arms. Leaves elliptic-ovate, 100-240

x 70-180 mm, margin sinuate-repand to distinctly lobed

with 2-4 pairs of lobes up to 50 mm long, apex acute,

base oblique; both surfaces but especially the lower, with

± sessile stellate hairs with 7-9 unequal amis; petiole 15-

18 mm long; prickles may occasionally be present on

young leaves. Flowers usually solitary or two next to each

other, rarely 3-5 or more arising from a common stalk,

extra-axillary and hermaphrodite. Corolla ± 15 x 20-30

mm, white, occasionally light purple; petal tips apiculate,

incurved; inner and outer surface of corolla with stellate

hairs; pedicel ± 15 mm long. Fruit ellipsoid, globose or

oval, 13-70 x 15-50 mm, smooth or with shallow lon-

gitudinal grooves, usually solitary, occasionally two or

more together, green, white or purple when young, red

when mature; fruit stalk 16-40 mm long, mostly decurved.

Seeds 3x4 mm.

The Gilo group is by far the most widely grown cul-

tivar group of Solanum species throughout southern Ugan-

da. Its fruits are used in soup or stew preparation. It is

preferred to S. melongena because it has softer flesh than

S. melongena. Within the Gilo group there is considerable

Bothalia 25,1 (1995)

55

variation, especially in the shape, size and colour of the

fruit. This cultivar group is native to Africa and is believed

to have arisen from the wild, weedy and semi-cultivated

but poorly domesticated S. anguivi (Lester & Niakan

1986).

Vouchers: U2: Kyembogo D.F.I., fl. & fr. Oct. 1990, Bukenya 109.

U3: Abwangati Tororo, fl. & fr. Aug. 1991, Bukenya 134. U4: nr Kisasa

Bukoto, fl. May 1972; Kivuuvu Mityana, fl. & fr. Aug. 1990, Bukenya

58.

25b. Solanum aethiopieum Shum group

S. zuccagnianum Dunal (1813, 1852).

Subshrub about 0.6 m high; stem glabrous. Leaves

ovate, about 150 x 100 mm, apex acute, base oblique,

attenuate or truncate, margin repand; young leaves on both

surfaces with small, sessile, stellate hairs with 5-8 more

or less equal arms; mature leaves subglabrous; petiole 50-

60 mm long. Inflorescence 3-10-flowered, sessile or

flowers solitary, lateral; flowers hermaphrodite; pedicel ±

7 mm long with scattered stellate hairs. Corolla white,

5-7 mm long x ± 10 mm wide, glabrous; flower buds

with dense stellate hairs. Calyx as long as the corolla tube.

Style with stellate hairs. Fruit globose, 15-35 mm diam.,

green with dark green stripes when young, shiny red when

ripe; fruiting calyx ± 7 mm long; stalk up to 12 mm long.

Seeds 2.8 x 3.0 mm.

In Uganda the Shum group is a popular leaf vegetable

in Buganda region, from where it has been introduced by

migrants from Buganda to a few areas in western and

eastern Uganda. It is native to Africa and is frequently

cultivated in tropical Africa. It is less polymorphic than

its relatives, the Gilo group and S. anguivi.

Vouchers: U2: Nyaruzinga, fr. Oct. 1990. Bukenya 106. U3: Budondo

Jinja, fr. Aug. 1991. Bukenya 127. U4: 2 km E of Budo, fl. & fr. Nov.

1972, Katende 1727.

26. Solanum albicaule Kotschy ex Dunal in A.P.

de Candolle, Prodromus systematis naturalis regni

vegetabilis 13,1: 204 (1852); C.H. Wright: 255 (1906);

Bitter: 101 (1923); Carvalho & Gillet: 237 (1960); Heine:

204 (1963). Type: Sudan, Kordofan, Kotschy 309 (G,

holo.).

The species is sunk under S.forskalii Dunal by D'Arcy

& Rakotozafy (1994).

For synonymy see Heine (1963).

An undershrub about 1 m high. Branches slender,

shortly hairy-tomentose; hairs stellate. Leaves rather small,

ovate, 20-70 x ± 10 mm, subentire, covered with sessile

stellate hairs on both surfaces. Inflorescence 3-6-flowered.

Flowers bluish purple. Calyx about 3 mm long. Corolla

8-13 mm long. Fruit green with white stripes when

young, yellow when ripe, up to 10 mm diam.

S. albicaule has been collected from the dry region of

Karamoja. It extends into NE Africa (Sudan, Somalia,

Eritrea and Egypt) and west and NW Africa (Senegal,

Chad, Mauritania). Elsewhere, it has been recorded for

Arabia, Pakistan and India. It seems to be a species of

arid lands. It has no close relatives in section Oliganthes.

Vouchers: Ul: 5-6 km N of Lothea Bokora Karamoja, fl. May 1970,

Lye 5455; 5 km S of Kantaku Bokora Karamoja, fl. & fr. June 1970,

Katende 442.

27. Solanum eyaneo-purpureum De Wild., Plantae

bequaertianae 1: 425 (1922). Type: Zaire, Kabare, Be-

quaert 5333 (BR).

For synonymy see Whalen (1984) and Jaeger (1985).

Clambering shrub 2-3 m high; young stems, leaves on

both surfaces with stalked stellate hairs; stem, petiole and

midrib armed with short recurved prickles. Leaves ovate,

30-80 x 20-40 mm, entire or sinuate; petiole 5-20 mm

long. Inflorescence 3-10-flowered, racemose. Corolla

purple, up to 10 x 180 mm; pedicel up to 10 mm long.

Fruit red when mature, ± 10 mm diam.; fruit stalk 10-20

mm long.

5. eyaneo-purpureum is common in the shrub forests

in the plains of southwestern Uganda. It also occurs in

Rwanda, Burundi and eastern Zaire and often grows on

termite mounds. It is related to S. taitense Vatke and S.

hastifolium Hochst.

Vouchers: U2: nr Gayaza Rwampara Ankole, fl. & fr. Oct. 1932,

Eggeling 661. U4: 17-18 km SE of Ntusi Mawogola. fl. & fr. Oct. 1969,

Lye 4479.

28. Solanum hastifolium Hochst. ex Dunal in A.P.

de Candolle, Prodromus systematis naturalis regni

vegetabilis 13,1: 284 (1852). Type: Sudan, Kordofan,

Kotschy 393 (MPU, holo.; K!, iso.).

For synonymy see Whalen (1984).

Shrub, often scandent, up to 2 m high; branches, petiole

and midrib armed with small recurved prickles; stems and

both surfaces of leaves covered with stalked stellate hairs.

Leaves ovate, entire or sinuate, base shortly cuneate, about

50-70 x 20-30 mm. Inflorescence 2-6(-10)-flowered,

umbelliform. Corolla violet, up to 10 x 15 mm. Fruit red

when mature, ± 10 mm diam.

S. hastifolium is a deciduous bushland species of NE

Africa, from northern Tanzania through Kenya and eastern

Uganda to Sudan, Ethiopia and Somalia. It is rather

polymorphic and often confused with S. taitense Vatke. It

is also related to S. eyaneo-purpureum De Wild.

Vouchers: Ul: nr Rupa, Karamoja, fl. & fr. Sept. 1958, Wilson 585.

U4: Katuugo Buruli. fl. & fr. Oct. 1970. Katende 716.

29. Solanum taitense Vatke in Linnaea 43: 327

(1882). Type: Kenya, between Ndi and the river Tsavo,

Hildebrandt 2605 (Bf; isotypes possibly at L, M, P).

Slender-stemmed scandent shrub to 2 m tall; stem,

petiole and midrib armed with small recurved prickles.

Leaves lanceolate, entire, repand or lobed, 40-80 x 20-40

mm, on both surfaces with stalked stellate hairs. Inflores-

cence ( 1 )2^f-flowered. Corolla pinkish purple, 8-15 x

56

Bothalia 25,1 (1995)

15-20 mm. Fruit green with white stripes when young,

red when mature, ± 10 mm diam.

S. taitense is closely related to S. hastifolium and the

two are often confused with each other. The former has

a simpler inflorescence than the latter and leaf shapes are

different in the two species. S. taitense is found in bush-

lands of Masaka, Ankole and Karamoja. It also occurs in

Kenya and Tanzania.

Vouchers: Ul: 5 km of Kantaku Bokora Karamoja, fl. & fr. June

1970. Katende 436. U4: 1-2 km E of Kikoma Mawogola, fl. & fr. Oct.

1969, Lye 4435.

30. Solanum usambarense Bitter & Dammer in

Bitter, Feddes Repertorium Beih. 16: 40 (1923). Syn-

types: Holst 380, 8921 , 8925a (K!), 9091a ; Hick 28a, 200,

273, 382\ Volkens 616\ Buchwald 380. All collected in

northern Tanzania and all cited with original description.

Shrub up to 3 m tall, armed heavily with prickles, in-

dumentum pilose. Leaves ovate, up to 200 x 150 mm,

margin lobed, base truncate and unequal and may be

shortly attenuate, covered with stellate hairs. Inflorescence

8- to many-flowered rachis simple or branched, racemose.

Corolla creamy white with a purple tinge on the veins, ±

10 mm diam. Fruit orange-red or red, ± 8 mm diam.

S. usambarense has often been confused with S. an-

guivi (S. indicum ). Although the two are closely related,

S. usambarense is easily distinguished from S. anguivi by

inflorescence and infructescence characters.

S. usambarense occurs in the shrub layer of the forests

on mountains of Rwenzori, Elgon and Kigezi. It also oc-

curs in the mountains of Kenya and Tanzania.

Vouchers: U2: Virunga-Kette Nkanda, fl. & fr. Nov. 1954, Stauffer

899. U3: Buhekeke Bugishu, 11. & fr. July 1926, Maitland 1213 (sheet

2).

Section Torva Nees in Transactions of the Linnean

Society: 51 (1837); D’Arcy: 273 (1972). Type species: S.

torvum Sw.

31. Solanum giganteum Jacc/., Collectanea 4: 125

(1790); Jacq.: 11,1. 382 (1792). Type: collected near the

Cape of Good Hope and cultivated in Vienna; in the ab-

sence of a specimen, the illustration t. 328 (fide Jaeger

1985) is taken as the type.

Shrub or tree up to 8 m high; stem and young leaves

covered with white indumentum of tiny sessile stellate

hairs; mature leaves with sessile stellate hairs on the un-

dersurface, upper surface almost glabrous apart from oc-

casional sessile stellate hairs; stem and petioles with fiat

triangular prickles. Leaves ovate to elliptic, up to 280 x

120 mm. Inflorescence paniculate, terminal or subter-

minal, 20 to > 60-flowered. Flowers violet to purple, 8-10

mm long. Fruit green when young, red when ripe,

globose, ± 10 mm diam.

S. giganteum is fairly common in Uganda. It is a dis-

junct montane species stretching southwards from Ethiopia

to South Africa, and west to Cameroon. It has also been

recorded in S India and Sri Lanka.

Vouchers: U2: Budongo, fl. & fr. Nov. 1932, Harris 161. U3: Mt

Elgon, 11 Apr. 1930, Lubenberg 1664. U4: Entebbe, fl. & fr. Sept. 1922,

Maitland 217 .

32. Solanum kagehense group

This group was brought together by Jaeger (1985) for

the convenience of dealing with material belonging to four

closely related species: S. kagehense Dammer: 187

(1906). Type: Tanzania, Muansa, Fischer 78 (isotype

LE?); S. muansense Dammer: 243 (1912). Type: Tanzania,

Muansa, Stuhlmann 4504 (iso. HBG?); S. sordidescens

Bitter: 260 (1921). Type: Mozambique, Tschumo, in

Matumbi Mts, Basse Hi 3097 (iso. E.A); S. wittei Robyns:

82 (1943). Type: Zaire, Kabasha, De Witte 1142 (holo.

BR?).

Jaeger (1985) did not make new combinations since

his grouping was based on published descriptions. He sug-

gested that a study of the types should be undertaken

before new combinations are made. His approach was

adopted in this study.

Shrubs up to 3 m high. Stem with broad-based prickles

and scattered sessile stellate hairs with ± equal arms.

Leaves elliptic, entire or repand, up to 150 x 70 mm;

upper surface with sparse sessile stellate hairs about 6-

armed, middle arm much longer than rest, lower surface

with sparse or heavy cushion of same type of hairs as on

upper surface. Inflorescence cymose with flowers often

subumbellate on branched peduncle, 10-20-flowered.

Corolla violet, 7-10 x 14 mm. Fruit red when ripe, ± 7

mm diam.

S. kagehense group occurs in all the geographical

regions of Uganda. It is common in the East African

region as a whole, especially in thickets and areas of light

shade between 600-1 500 m. It is related to S. renschii

Vatke. Some past plant collectors have confused it with

S. giganteum.

Vouchers: Ul: Kaabong Karamoja, Sept. 1950, Dawkins 643. U2:

Ruizi R., Nov. 1950, Jarrett 208. U3: Serere Labori, July 1926, Maitland

1349-. Busoga, June 1925, Maitland s.n.. U4: 5 km S of Sembabule, fr.

May 1971. Katende 881.

33. Solanum renschii Vatke in Linnaea 43: 328

(1882). Type: Kenya, Ukamba, Hildebrandt 2735 (K!,

iso.).

For synonymy see Whalen (1984) and Jaeger (1985).

Shrub or subshrub. Stem covered when young with

small sessile stellate hairs with about 8 ± equal arms; stem

and petiole bear broad-based prickles; veins with tiny

prickles. Leaves 50-80 x 30-40 mm, ovate, entire; upper

surface covered with same type of hairs as above; lower

surface covered with a heavy cushion of sessile stellate

hairs, larger than above and with more than 10 ± equal

arms. Inflorescence corymbose, 20-50-flowered. Corolla

pale violet, ±10x13 mm. Fruit ± 8 mm diam., red/black

when ripe.

Bothalia 25,1 (1995)

57

S. renschii is endemic to eastern Africa, and is a vari-

able species. Jaeger (1985) suggested that S. kwebense

N.E. Br., S. munitum Bitter, S. tettense Klotzsch and the

material in his S. kagehense group could be incorporated

in a broad concept of the species. Whalen (1984) lists S.

tettense and S. wittei A. Robyns as synonyms of S.

renschii. S. wittei was included in Jaeger’s S. kagehense

group. If S. tettense and S. renschii are synonymous, then

S. renschii should be sunk under S. tettense , having been

published 21 years later than S. tettense. In Uganda S.

renschii is restricted to the arid region of Karamoja.

Vouchers: Ul: Moroto Karamoja. ft. & fr. Sept. 1956. Bally 10788 ;

foothills of Mt Moroto, fl. & fr. Oct. 1952, Verdcourt 809.

D. Subgenus Potatoe (G. Don) D'Arcy, in Annals

of the Missouri Botanical Garden 59: 272 (1972); D'Arcy:

750 (1973). Type species: S. tuberosum L.

Herbs, rarely woody, often glandular-pubescent and

aromatic, unarmed; hairs simple. Leaves mostly com-

pound or deeply lobed, but simple, entire leaves often

present at certain stage. Inflorescence mostly paniculate;

peduncles once or temately branched, often pendulous;

pedicels mostly articulating near base or above it. Fruit

10-20 mm diam.

The subgenus is represented in Uganda by two sec-

tions: Petota and Jasminosolanum.

Section Petota Dumort. Florula belgica: 39 (1827).

Type species: S. tuberosum L.

34. Solanum tuberosum L., Species plantarum:

185 (1753); D’Arcy: 752 ( 1973). Lectotype: LINN 248.12

labelled ‘S. tuberosum’ (fide Hawkes 1956) (LINN IDC

microf.! 138: II.4).

Herb ± 1 m high; stem with shallow grooves when

dry, and simple hairs; stolons tuberiferous. Leaves com-

pound, interceptedly imparipinate; major leaflets 3-7, al-

ternating with ± 3 pairs of minor leaflets; leaflets elliptic

to oblong-elliptic, ± 50 x 25 mm; simple hairs on both

surfaces; petiole up to 60 mm long. Inflorescence panicu-

late, 1 0-30-flowered, terminal on long peduncle ± 80 mm

long; pedicel 7-15 mm long with simple hairs. Flowers

hermaphrodite. Corolla white, bluish purple or purplish

red, 10-22 x ± 20-30 mm. Anthers 5, occasionally 6.

Calyx ± 7 mm long, with simple hairs on outer surface,

glabrous within. Fruits globose, ± 7 mm diam. 2n = 48.

S. tuberosum, is a mainly temperate crop introduced in

East Africa about 100 years ago by missionaries from

Europe. The main area of commercial cultivation of

potatoes in Uganda is Kigezi, a mountainous area with

cool climate. Potato growing has spread to other highland

areas in the country. Lowland areas also in recent years

have started growing especially lowland cultivars. Potato

blight seems to be a more serious limiting factor to cul-

tivation of potatoes than climate.

Vouchers: U2: Kacwekano D.F.I., fl. Oct. 1990, Bukenya 99-101.

U4: Kamenyamigo D.F.I., fl. Oct. 1990, Bukenya 93: Makerere Univ.,

fl. Oct. 1990, Bukenya 92.

Section Jaminosolanum Bitter ex Seithe in Botani-

sche Jahrbiicher 81: 191 (1962); D’Arcy: 757 (1973).

Type species: S. jasminoides Paxton.

35. Solanum seaforthianum Andrews, The Bota-

nist’s Repository: pi. 504 (1808). Lectotype: the plate cited

(fide Symon 1981).

Woody climber; stem terete, glabrous apart from oc-

casional tiny simple hairs. Leaves compound, imparipin-

nate to deeply pinnatifid with about 7-9 leaflets or lobes;

leaflets up to 50 x 30 mm. elliptic; lower leaflets with ±

winged petiolule up to 5 mm long; upper leaflets (i.e.

lobes) webbed together; lamina glabrous; margin ciliolate

with sparse simple hairs; petiole up to 50 mm long. In-

florescence glabrous, mostly terminal or lateral, paniculate

with up to 30 flowers or more; pedicel up to 7 mm long.

Flowers hermaphrodite. Corolla blue to violet, up to 15

x 20 mm. Fruit spherical, ± 10 mm diam., red when ma-

ture; up to 40 from an inflorescence; fruit stalk 10-14

mm long. Seeds 2x2 mm.

S. seaforthianum is an introduced decorative climber

seemingly naturalized. It is native to central America and

the West Indies, but has spread to many parts of tropical

Africa, where it has been introduced for decorative pur-

poses.

Vouchers: Ul: Mulago hospital, fl. & fr. Apr. 1990, Bukenya 45: fl. &

fr. Aug. 1990, Bukenya 55: Makerere Univ., fl. & fr. Apr. 1990, Bukenya

48.

CONCLUSIONS

Solanum americanum, S. tarderemotum, S. incanum, S.

macrocarpon (wild group), 5. anguivi and S. kagehense

group are the most widespread taxa in Uganda, occurring

in all four geographical regions of the country. On the

other hand, S. scabrum, S. terminale subsp. inconstans, S.

benderianum, S. albicaule, S. renschii and S. seaforthia-

num are the least widespread, occurring in only one

region. The rest of the taxa are either fairly well dis-

tributed (occurring in three regions) or relatively rare (oc-

curring in two regions). This is associated with habitat

preference and/or utilization.

There is tremendous morphological variation within

the S. macrocarpon complex. Bukenya (1993) recognised

four groups belonging to the S. macrocatpon complex in

Uganda. These include S. macrocarpon (wild group — S.

dasyphyllum) ; S. macrocarpon (semi-wild group), S.

fnacrocarpon 'Mukono’ cultivar and S. macrocarpon

‘Nabingo’ cultivar. Sections Solanum and Oliganthes are

also very variable. Experimental work on these taxa is

still necessary to resolve their taxonomy.

At least 25 species in Uganda are useful or economi-

cally relevant. Pharmacological studies are needed to

authenticate the medicinal potency of the various species

used in traditional medicine. More attention should be

paid to crops in terms of research to control pests and

diseases, to increase yield and to produce well-adapted

cultivars.

58

Bothalia 25,1 ( 1995)

ACKNOWLEDGEMENTS

We are deeply indebted to Makerere University, UN-

ESCO, the Linnean Society of London and the British

Council for financial support to the senior author.

We would like to thank Dr R.N. Lester for comments

and suggestions which were very useful. Thanks are ex-

tended to Dr J.M. Edmonds for identifying materials

belonging to section Solatium and for useful discussion.

We are grateful to Dr K. Vollesen and Prof. J. Grau for

assistance during herbarium studies at Kew and Munich

respectively. We are grateful to all individuals who

provided us with material and other forms of assistance

during this study.

REFERENCES

ANDREWS, H.C. 1808. The Botanist's Repository, t. 504. London.

BEAMAN-MBAYA, V. & MUHAMMED, S.I. 1976. Antibiotic action of

Solatium incanum L. Antimicrobial Agents and Chemotherapy 9:

920-924.

BECTOR, N.P., A JIT, S.R & GUPTA. R.H. 1971 . A new approach to the

treatment of some chronic respiratory diseases. Indian Journal of

Medical Research 59: 730-742.

BENTHAM, G, 1861. Flora hongkongensis: a description of the flower-

ing plants and ferns of the island of Hongkong. Lovell, Reeve,

London.

BISTA, M.S. 1983. Biosystematic evaluation of germplasm of eggplants

and related Solanum species with special reference to Africa.

M.Sc. Thesis, University of Birmingham.

BITTER, G. 1912. Solana nova vel minus cognita. Feddes Repertorium

10: 529-565.

BITTER, G. 1913. Solana Africana 1. Botanische Jahrbiicher 49: 560-

569.

BITTER, G. 1917. Solana Africana 2. Botanische Jahrbiicher 54: 416—

506.

BITTER. G. 1919. Die papuasischen Arten von Solanum. Botanische

Jahrbiicher 55: 69-89.

BITTER, G. 1921. Solana Africana 3. Botanische Jahrbiicher 57: 248-

286.

BITTER. G. 1922. Solana nova vel minus cognita. Feddes Repertorium

18: 301-307.

BITTER, G. 1923. Solana Africana 4. Feddes Repertorium Beih. 16:

1-320.

BUKENYA, Z.R. 1980. Studies in taxonomy of Solanum L. in southern

Ghana. M.Sc. Thesis, University of Ghana, Legon.

BUKENYA, Z.R. 1991. Solanum macrocarpon : an underutilised but

potential vegetable in Uganda. Proceedings of the 1 3th AETFAT

Congress. Zomba, Malawi.

BUKENYA, Z.R. 1993. Studies in the taxonomy of the genus Solanum in

Uganda. Ph.D. Thesis, Makerere University.

BUKENYA, Z.R. & HALL, J.B. 1988. Solanum (Solanaceae) in Ghana.

Bothalia 18: 79-88.

BURK1LL, J.H. 1925. Solanum macrocarpon. Kew Bulletin 1925: 333—

341.

CARVALHO, L.A.F. & GILLET, J.B. 1960. In ,/. Agri. Trap. 7: 237.

DALZIEL, J.M. 1937. The useful plants of west tropical Africa. Crown

Agents, London.

DAMMER, C.L.U. 1905. Solanaceae Africanae. Botanische Jahrbiicher

38: 57-60.

DAMMER, C.L.U. 1906. Solanaceae Africanae 1. Botanische Jahr-

biicher 38: 176-195.

DAMMER, C.L.U. 1912. Solanaceae Africanae 2. Botanische Jahr-

biicher AH'. 236-260.

D’ARCY, W.G. 1972. Solanaceae studies 2. Typification of subdivisions

of Solanum. Annals of the Missouri Botanical Garden 59: 262-

278.

D'ARCY, W.G. 1973. Solanaceae. In R.E. Woodson Jr, R.W. Schery &

collaborators, Flora of Panama. Annals of the Missouri Botanical

Garden 60: 573-780.

D’ARCY, W.G. 1974. Solanum and its close relatives in Florida. Annals

of the Missouri Botanical Garden 61: 819-869.

D’ARCY, W.G. & RAKOTOZAFY, A. 1994. Solanacees. In Ph. Moral,

Flore de Madagascar et des Comores.

DE CANDOLLE, A.P 1852. Prodromus systematis naturalis regni

vegetabilis 13,1. Paris.

DE WILDEMAN, E. 1922. Plantae bequaertianae 1: 425. Brussels.

DILLENIUS, J.J. 1732. Hortus Elthamensis II: 366-368.

DUKE, J.A. 1970. Ethnobotanical observations on the Choco Indians.

Economic Botany 24: 344—366.

DUMORTIER, B.C.J. 1827. Florida belgica. Tournay.

DUNAL, M.F. 1813. Histoire naturelle, medicate et economique des

Solanum. Paris, Strasbourg, Montpellier.

DUNAL, M.F 1852. Solanaceae. In A.P. De Candolle, Prodromus sys-

tematis naturalis regni vegetabilis 13,1: 1-690.

EDMONDS, J.M. 1971. Solanum L. In W.I. Stearn, Taxonomy and

nomenclatural notes on Jamaican gamopetalous plants. Journal of

the Arnold Arboretum 52: 634, 635.

EDMONDS, J.M. 1972. A synopsis of the taxonomy of Solanum sect.

Solanum ( Maurella ) in South America. Kew Bulletin 27: 95-114.

EDMONDS, J.M. 1977. Taxonomic studies of Solanum sect. Solanum

{Maurella). Botanical Journal of the Linnean Society 75: 141—

178.

EDMONDS, J.M. 1979a. Nomenclatural notes on some species of

Solanum found in Europe. Botanical Journal of the Linnean

Society IK. 213-233.

EDMONDS, J.M. 1979b. Taxonomic studies of Solanum sect. Solanum

(Maurella). Linnean Society Symposium 7: 529-548.

EDMONDS, J.M. 1984. Solanum L. section Solanum — a name change

for S. villosum Mill.. Journal of the Linnean Society of London 89:

165-170.

EDMONDS, J.M. 1986. Biosystematics of Solanum sarrachoides

Sendtn. and S. pitysalifolium. Botanical Journal of the Linnean

Society 92: 1-38.

FAO 1988. Solanum species. Traditional food plants: a resource book for

promoting the exploitation and consumption of food plants in

arid, semi-arid and subhumid lands of eastern Africa. FAO, Food

and Nutrition Paper 42: 450^-66. Rome.

FEDEROV, A. A. 1969. Chromosome numbers of flowering plants.

Academy of Sciences USSR, Moscow.

FORSSKAL, P. 1775. Flora aegyptiaco-arabica. Moller, Copenhagen.

GBILE, Z.O. 1979. Solanum in Nigeria. Linnean Society Symposium 7:

113-120. Academic Press, London.

GRANT, V. 1971. Plant speciation. Colombia University Press, New

York.

GRAY, J.M. 1 968. Taxonomy of the Morelia section of the genus Solanum

L. within South America. Ph.D. Thesis, University of Birming-

ham, England.

HAWKES, J.G. 1956. Taxonomic studies of the tuber-bearing solanums

1. Solanum tuberosum and the tetraploid species complex.

Proceedings of the Linnean Society of London, session 166: 97-

144.

HEINE, H.H. 1960. Notes on Solanum. Kew Bulletin 14: 245-249.

HEINE, H.H. 1963. Solanaceae. In R.W.J. Keay & F.N. Hepper, Flora of

West tropical Africa , 2nd edn: 325-335. Crown Agents, London.

HEISER, C.B. 1971. Notes on some species of Solanum (sect. Lepto-

stemonum) in Latin America. Baileya 18: 59-65.

HENDERSON, R.J.F. 1974. Solanum nigrum L. (Solanaceae) and related

species in Australia. Contributions from the Queensland Her-

barium 16: 1-78.

HENDERSON, R.J.F. 1977. Notes on Solanum (Solanaceae) in Australia.

Austrobaileya 1: 13-22.

HEPPER, F.N. 1976. The West African herbaria of Isert & Thonning: a

taxonomic revision and an index to the IDC microfiche. Bentham-

Moxon Trust, Kew.

HEPPER, F.N. 1978. Typification and name changes of some Old World

Solanum species. In J.G. Hawkes, Systematic notes on the

Solanaceae. Botanical Journal of the Linnean Society 76: 287-

292.

HEPPER, F.N. 1979. On typifying Linnean names of Solanaceae. Lin-

nean Society Symposium 7: 131-134.

HEPPER, F.N. & JAEGER, PM.L. 1985. The typification of six Linnean

names of Solanum. Kew Bulletin 40: 387-389.

HOOKER, W.J. 1887. Solanum wendlandii. Curtis's Botanical Magazine

1 13: t. 6914.

IRVINE, F.R. 1961. Woody plants of Ghana, with special reference to

their uses. Oxford University Press, London.

JACQUIN, N.J. VON 1781. leones plantarum rariorum 1: 5, t. 41.

Vienna.

JACQUIN, N.J. VON 1790. In Collectanea 4: 1 25.

Bothalia 25,1 (1995)

59

JAEGER, P.M.L. 1985. Systematic studies in the genus Solarium in

Africa. Ph.D. Thesis, University of Birmingham.

KHAN, R. 1979. Solanum melongena and its ancestral forms. Linnean

Society Symposium 7: 629-636. Academic Press, London.

LAMARCK, J.B.A.P.M. DE 1794. Tableau Encyclopedique et

Methodique 2: 2131. Paris.

LESTER, R.N. & HASAN, S.M.Z. 1991. Origin and domestication of the

brinjal eggplant, Solanum melongena from S. incanum , in Africa

and Asia. Solanaceae III: 369-387. The Royal Botanic Gardens,

Kew, Richmond, Surrey, U.K.

LESTER, R.N. & NIAKAN. L. 1986. Origin and domestication of scarlet

eggplant (S. aethiopicum L. from S. anguivi Lam.). In W.G.

D'Arcy, Solanaceae: biology and systematics : 433^156. Colum-

bia University Press, New York.

LIND, E.M. & TALLANTIRE, A.C. 1975. Some common flowering

plants of Uganda. Oxford University Press, Nairobi.

LINNAEUS, C. 1753. Species plantarum, 2 vols. Stockholm.

LINNAEUS, C. 1754. Genera plantarum, 5th edn. Stockholm.

LINNAEUS, C. 17 59. Ainoenitates academicae. Stockholm.

LINNAEUS, C. 1771. Mantissa plantarum altera. Stockholm.

MARTIUS, C.F.P. VON 1846. Solanaceae. Flora brasiliensis. Leipzig.

MILLER, P. 1768. The gardener’s dictionary, 8th edn. London.

NEE, M. 1979. Patterns in biogeography in Solanum section Acan-

thophora. Linnean Society Symposium 7: 569-580. Academic

Press, London.

NEES AB ESENBECK, C.G.D. 1837. Monograph of the East Indian

Solaneae. Transactions of the Linnean Society of London 17:

37-77.

PLUKENET, L. 1696 . Almagestum botanicum. Londini.

RICHARD, A. 1850. Tentamen florae abyssinicae 2: 101. Paris.

ROBYNS, A.W. 1943. Solanaceae. Bulletin duJardin Botanique de I'Etat

Bruxelles 17: 82, 83.

ROE, K.E. 1972. A revision of Solanum section Brevantherum. Britton ia

24: 239-278.

ROE, K.E. 1979. Dispersal and speciation of Solanum section Brevan-

therum. Linnean Society Symposium 7: 563-567. Academic Press,

London.

SCOPOLI, J.A. 1788. Deliciae florae et faunae insubricae. Ticini.

SEITHE, A. 1962. Die Haararten der Gattung Solanum und ihre

taxonomische Verwertung. Botanische Jahrbriicher 8 1 : 26 1-336.

SEME, E.N. 1983. Evaluation of morphological and biochemical charac-

ters of African Gboma eggplant (Solanum macrocarpon L.).

M.Sc. Thesis, University of Birmingham.

SENGENDO, B.K.A. 1982. Studies in the taxonomy of Solanum L. sect.

Oliganthes. B.Sc. Dissertation, Makerere University, Kampala,

Uganda.

STAFLEU. F.S. & COWAN, R.S. 1976. Taxonomic literature Vol. 1:

A-G. Regni vegetabilis 94.

STEBBINS, G.L. 1950. Variation and evolution in plants. Columbia

University Press, New York.

SYMON, D.E. 1981. A revision of the genus Solanum in Australia.

Journal of the Adelaide Botanic Gardens 4: 1-367.

VATKE. G.W. 1882. Plantas in itinere africana ab J.M. Hildebrandt

collectas determinare pergit. Linnaea 43: 305-334.

WALTERS, B. 1965. The role of steroid saponines on antibiotic effect of

S. dulcamara. Planta medica 13: 188-193.

WHALEN, M.D. 1984. Conspectus of species groups in Solanum sub-

genus Leptostemonum. Gentes Herbarum 12: 179-282.

WRIGHT, C.H. 1894. Solanaceae. Kew Bulletin 1894: 126-129.

WRIGHT, C.H. 1897. Solanaceae. Kew Bulletin 1897: 275.

WRIGHT, C.H. 1902. Solanaceae. In H.H. Johnston, Uganda Protec-

torate: an attempt to give sone description of the physical geog-

raphy, botany 1: 362. Hutchinson, London.

WRIGHT. C.H. 1906. Solanaceae. In W.T. Thiselton-Dyer. Flora of

tropical Africa 4: 207-261. Reeve, London.

MARY GUNN LIBRARY

NATIONAL BOTANICAL INSTITUTE

PRIVATE BAG X 101

PRETORIA 0001

REPUBLIC OF SOUTH AFRICA l

Bothalia 25,1: 61-63 (1995)

A new species and a change of status in Ophioglossum (Ophioglossaceae:

Pteridophyta) in Africa

J.E. BURROWS* and T.J. EDWARDS**

Keywords: Africa, new species, Ophioglossum. Pteridophyta

ABSTRACT

A new species, Ophioglossum gracile Pocock ex J.E. Burrows, is described from South Africa. A lectotype is chosen for O.

vulgatum var. kilimandscharicum Hieron. and its status upgraded to a subspecies.

U1TTREKSEL

Ophioglossum gracile Pocock ex J.E. Burrows, 'n nuwe spesie uit Suid-Afrika word beskryf. ‘n Lektotipe word vir O.

vulgatum var. kilimandscharicum Hieron. gekies en die status word opgegradeer tot 'n subspesie.

The species concept in Ophioglossum L. is greatly

misunderstood and the species are consequently poorly

delimited, largely because of the simple morphology of

the genus and the resulting lack of characters upon which

to base species. Recent research by the senior author (Bur-

rows 1992) has resulted in a better understanding of the

African taxa, largely due to extensive SEM studies of the

spores. Consequently it has become necessary to describe

and clarify the two following taxa.

1. Ophioglossum gracile Pocock ex J.E. Burrows ,

sp. nov. ab O. vulgato subsp. africano trophophoris

minoribus (longitudine plerumque < 33 mm) anguste el-

lipticis vel lanceolatis, sporisque muris secundariis

tenuibus in quoque lumen descendentibus differt.

TYPE. — Western Cape, 3318 (Cape Town): Table

Mountain. Woodhead Reservoir, (-CD), 30-6-1947, Ester-

huysen 13915 (BOL, holo.!; GRA!, NBG!, PRE!).

O. vulgatum L. var. gracile Pocock, ined.

O. nudicaule L.f., sensu Roux: 9 (1979).

Rhizome linear, 13-22 x 2-3 mm. Roots horizontal,

fleshy, proliferous. Leaves one or two. Stipe ( 14— )21— 48(—

83) mm long, 50-70% of its length subterranean,

stipe :trophophore length ratio 0.8— 1 .7: 1 . old stipe bases

not persistent. Trophophore variable in shape, narrowly

elliptic to lanceolate, (1 7-) 26-32(^13) x (4— )7— 9(— 1 3)

mm, length: width ratio 2.3-7.5:l, apex acute, apiculate,

base narrowly cuneate to attenuate, trophophore held at

50°-60° from the horizontal, margins Hat. Venation with

secondary veins and blind veinlets laterally and distally

but sparse or lacking proximally, epidermal cells elongate

throughout with wavy to sinuous anticlinal cell walls.

* Buffelskloof Nature Reserve, P.O. Box 710, Lydenburg 1120.

** Dept, of Botany, Natal University, P.O. Box 375, Pietermaritzburg

3200.

MS. received: 1994-04-19.

stomata aligned. Sporophore (46— )76— 1 08(— 1 45) mm

long, with ( 1 3—) 1 8— 22(— 28) pairs of sporangia, sporo-

phore: trophophore length ratio 2.0-5.3:l. Spores 40—47

pin in equatorial diameter, trilete; muri continuous, some-

what uneven, with thin, secondary muri descending into

each lumen; lumina conical, minutely granular. Figures

1C & D; 2A.

Specimens examined

WESTERN CAPE.— 3218 (Clanwilliam): Clanwilliam Dist.,

Boskloof, (-BB), 20-1-1947, Schelpe 1982 (BOL, K); Piquetberg Mtn,

between Mounton’s Vlei and Gruys Kop, (-DA), 7-11-1934, Pillans

1257 (BOL). 3219 (Wuppertal): Cedarberg, Langberg, (-AC), 15-12-

1941, Esterhuysen 7334 (BOL); 16-12-1941, Compton 12764 (NBG).

3318 (Cape Town): Table Mtn. near ‘Slaangully’, (-CD), 4-1908,

Diimmer 1297 (NBG, SAM); Wood Ravine and Slangolie, pipe track,

at the spring, (-CD), 8-1907, collector unknown ex Herb. Marloth

3778 (PRE); Table Mtn. lower reservoir, (-CD), 19-5-1956, Ester-

huysen 25846 (BM. BOL); Table Mtn. Carmichael s.n. (K). 3322

(Oudtshoom): S slopes of Outeniqua Mountains, next to Montagu Pass,

(-CD), 750m, 3-11-1983, Vlok 748 (BOL); next to Outeniqua Pass,

(-CD), 7-2-1985, Vlok 923 (PRE).

Historically this taxon has caused much confusion and

has been given several names: O. nudicaule (Roux 1979),

O. lancifolium (Schelpe & Anthony 1986) and O. vul-

gatum var. ambiguum (Pocock herb, annotation). Pocock

named the taxon O. vulgatum L. var. gracile in an un-

published manuscript on southern African Ophioglossum.

However, she confused the issue by including within the

species collections which are now known to belong to O.

lusoafricanum (Figure 2B) and O. vulgatum subsp.

africanum (Figure 2D). While O. gracile appears to be

most closely related to O. vulgatum subsp. africanum , it

differs in its much smaller and narrower trophophores,

and spores that consistently display primary muri that are

interconnected by smaller and narrower secondary muri

(Figure 2A), a character only seen elsewhere in the unre-

lated Namibian taxon currently included under O.

polyphyllum. Its restriction to the winter rainfall area of

the Cape separates it geographically from the summer

62

Bothalia 25,1 (1995)

FIGURE 1 . — Ophioglossum vulgatum subsp. kilimandscharicum'. A.

Schelpe 7177 , whole plant; B, Schlieben 4987, trophophores.

O. gracile, Esterhuysen 25846 : C, whole plant; D, trophophore

shape variant. All x 0.5.

rainfall O. vulgatum subsp. africanum, and Pocock’s

species should therefore be awarded specific rank.

Within the circumscription of her original taxon,

Pocock included two collections from the same locality

near Grahamstown in the eastern Cape which most

resemble O. vulgatum subsp. africanum. However, since

I (senior author) have not examined the spores from this

population, I cannot be certain of its identity. From the

collection data, it seems that O. gracile is restricted to

sandstones of the Table Mountain Series in ericoid scrub

(fynbos) on moist soils. A collection from the Cedarberg

( Esterhuysen 7334) was made on a 'damp spot at the base

of a shale band’. The type collection was made on ‘shal-

low soil on the banks of a small stream .... in shade of

pines’. Collections have been recorded in January,

February, April, May, June, August, November and

December, indicating that this species may be evergreen

in suitably moist situations. Its distribution falls within the

winter rainfall area which receives most of its rain from

May to September.

Although O. gracile shares the same phytogeological

association as O. bergianum , there is no record of the two

species growing in mixed communities. The third Ophio-

glossum species of the winter rainfall area of the Cape,

O. nudicaule, is confined to shale-derived soils and does

not occur with O. gracile.

Key to O. gracile and two similar species

la Trophophore lanceolate to ovate; trophophore length; width

ratio < 3.3:1;

2a Trophophores narrowly ovate to lanceolate, trophophore

length: width ratio 1. 8-3.5: 1; spores uniformly and

smoothly reticulate

O. vulgatum subsp. africanum Pocock ex J.E. Burrows

2b Trophophores ovate, trophophore length : w idth ratio

< 1.8:1; spores with muri coalescing irregularly to

form flattened areas

O. vulgatum subsp. kilimandscharicum (Hieron.) J.E. Burrows

lb Trophophore elliptic, rarely lanceolate; trophophore length:

width ratio 2. 3-8. 8:1:

3a Trophophore elliptic or lanceolate; spores trilete, with

secondary muri descending into the lumina

O. gracile Pocock ex J.E. Burrows

3b Trophophore elliptic, rarely oblanceolate (never lanceolate);

spores commonly alete, occasionally trilete, without

secondary muri O. lusoafricamtm Welw. ex Prantl

2. Ophioglossum vulgatum L. subsp. kilimandscha-

ricum (Hieron.) J.E. Burrows , stat. nov.

Ophioglossum vulgatum L. var. kilimandscharicum Hieron. in Engler:

89 (1895). Type: Tanzania, Kilimanjaro. Kifinika Volcano, 2 800 m, Oct.

1893, Volkens 1161 (B [destroyed]; BM!, lecto., here designated).

Differs from O. vulgatum subsp. vulgatum and O. vul-

gatum subsp. africanum (Pocock) J.E. Burrows by its

more broadly ovate to elliptic trophophores with a

length: width ratio of 1.8:1, and the spores in which the

muri coalesce irregularly into broad, flattened areas.

Figures 1A & B; 2C.

Although Hieronymus designated no type specimen for

his taxon (Engler 1895), all his new fern taxa from that

expedition appear to have been collected by Volkens in

1 893. Engler also visited Kilimanjaro in the previous year

FIGURE 2. — Spores of Ophioglossum, distal face. A, O. gracile, Vlok

748\ B, O. lusoafricanum. Burrows 523 1 ; C, O. vulgatum subsp.

kilimandscharicum, Lebrun 6016,. D, O. vulgatum subsp.

africanum. Chase 3758. Scale bar: 5 pm.

Bothalia 25,1 (1995)

63

(Gillett 1962) but since Engler mentions Volkens’ collect-

ing locality (Kifinika Volcano) in a subsequent account of

the East African Ophioglossum (Engler 1908), it is likely

that Volkens is the original collector of this taxon. A search

for the type material in Berlin revealed that all pre- World

War II material of Ophioglossum was destroyed (R.J.

Johns pers. comm.). We can therefore assume that, if the

specimen from which Hieronymus described the variety

was in Berlin, it no longer exists.

There is, however, a collection made by Volkens on

that expedition from the Kifinika Volcano, Kilimanjaro in

the British Museum which is almost certainly isotypic.

On the assumption that the Berlin material was destroyed

and since no type was designated by Hieronymus, the

sheet of Volkens 1161 in BM is designated as the lec-

totype.

Hieronymus distinguished his variety on the basis of

the plants having ‘leaves nearly always in pairs, the sterile

lamina obovate or elliptic (max. 30 x 17 mm), obtuse or

slightly acute, spores 36-44 pm broad, areolae 2^1 pm

wide'. The six collections seen that match the type

specimen, differ from typical O. vulgatum. However, oc-

casional trophophores within variable populations of nor-

mal O. vulgatum match subsp. kilimandscharicum quite

well. Conversely, no trophophores on the collections of

subsp. kilimandscharicum resemble those of typical O.

vulgatum.

In addition, spores from the Natal and Transvaal col-

lections reveal a morphological pattern distinct from both

typical O. vulgatum and subsp. africanum (Figure 2D).

The spore muri frequently coalesce into large flattened

areas and the lumina are correspondingly smaller and

more widely separated. A specimen from Zaire ( Lebrun

6016 , Figure 2C) displays similar spore sculpturing and

trophophores which match the Volkens collection. If these

three collections belong to the Kilimanjaro taxon, then the

spore pattern is distinct and constant. The trophophores

are also subtly different from the typical plants in that

they are broader, have a rounder apex and a thinner tex-

ture.

Very little is known of the ecology of this subspecies.

The Kilimanjaro specimens were both collected at 2 800

m but no ecological data were given. The Drakensberg

collection was made in burnt marsh under Miscanthus

(Poaceae). The Transvaal collection was made at 1 645

m in submontane grassland, growing under Helichrysum

(Asteraceae) bushes. As the subspecies is presently

known, it is an afromontane taxon and could be expected

from the Zimbabwe/Malawi gap which currently exists.

Specimens examined

KENYA. — 0035: Kericho Dist., western Mau Forest to Mt Blacket,

Kipkunyu R., (-BA), 13-6-1972, Faden, Faden & Grumbley 72/358 (K).

ZAIRE. — 0226: between Kinolu and (KatoKikombe?), Maniema, (—

AA), 8-1932, Lebrun 6016 (BR).

TANZANIA. — 0337: Kilimanjaro, Bergwiesen, (—A?), 24-3-1934,

Schlieben 4987 [B? (K, photo.!), BM. BR. PRE],

SOUTH AFRICA (EASTERN TRANSVAAL).— 2530 (Lydenburg):

Lydenburg Dist., Buffelskloof Nature Reserve, below Stone Huts, (-BC),

8-1-1991, Burrows 5133 (Herb. J.E.B., K, MICH, NU, PRE, TI).

SOUTH AFRICA (KWAZULU-NATAL).— 2829 (Harrismith):

Cathedral Peak area, Mhlonhlo Valley, (-CC), 1-1-1965, Schelpe 7177

(BOL, NU).

ACKNOWLEDGEMENTS

Bob Johns of Kew is gratefully acknowledged for search-

ing for the subsp. kilimandscharicum type in Berlin.

REFERENCES

BURROWS, J.E. 1992. The taxonomy of the genus Ophioglossum L.

( Ophioglossaceae ) in southern Africa. M.Sc. thesis, University of

Natal, Pietermaritzburg, Natal.

ENGLER, A. 1895. Die Pflanzenwelt Ost-Afrikas und der Nachbar-

gebiete. Dietrich Reimer, Berlin.

ENGLER, A. 1908. Die Pflanzenwelt Afrikas insbesondere seiner tropi-

schen Gebiete. 1, Die Pteridophyten, Gymnospermen und mono-

kotyledonen Angiospennen. Engelmann, Leipzig.

GILLETT, J.B. 1962. The botanical exploration of the area of the FTEA.

In A. Fernandes, Comptes Rendus de la TV Reunion Pleniere de

I' Association pour T Etude Taxonomique de la Flore d'Afrique

Tropicale. Lisboa.

ROUX, I P. 1979. Cape Peninsula ferns. National Botanic Gardens of

South Africa.

SCHELPE, E.A.C.L.E. & ANTHONY, N.C. 1986. Pteridophyta. In O.A.

Leistner, Flora of southern Africa. Botanical Research Institute,

Pretoria.

Bothalia 25,1:65-85 (1995)

The generic delimitation of Lachnaea and Cryptadenia (Thymelaeaceae)

J.B.P. BEYERS* and J.J.A. VAN DER WALT**

Keywords: Cryptadenia , generic delimitation, Lachnaea , Thymelaeaceae

ABSTRACT

The cunrent generic delimitation of Lachnaea L. and Cryptadenia Meisn, was investigated. A multidisciplinary approach

was followed. This included a study of the habit, phyllotaxis, morphology and anatomy of the leaves and flowers, pollen

morphology and karyology.

No major differences were found between the two genera. The range of variation of certain diagnostic characters of these

genera is such that the genera merge into each other. It was concluded that the two genera are congeneric and the inclusion of

Cryptadenia within Lachnaea is proposed.

UITTREKSEL

Die huidige generiese afbakening van Lachnaea L. en Cryptadenia Meisn. is ondersoek. 'n Multidissiplinere benadering is

gevolg. Die groeiwyse, blaarstand, morfologie en anatomie van die loofblare en blomme, stuifmeelmorfologie en kariologie is

bestudeer.

Geen noemenswaardige verskille tussen die twee genera is gevind nie. Die variasiegradient van sekere diagnostiese

kenmerke van hierdie genera is van so 'n aard dat die twee genera oorvleuel. Die insluiting van Cryptadenia binne die

omskrywing van die generiese grense van Lachnaea word voorgestel.

CONTENTS

Introduction 65

Materials and methods 66

Results 67

Habit 67

Morphology of foliage leaves 67

Anatomy of foliage leaves 69

Cuticle 69

Epidermis 70

Stomata 71

Mesophyll 71

Venation and vascular bundles 72

Morphology of flower and fruit 72

Pedicel/floral articulation 72

Hypanthium 72

Calyx 74

Petals 74

Androecium 74

Scales 75

Disc 75

Gynoecium 75

Fruit 76

Vascular anatomy of the flower 76

Pollen morphology 78

Karyology 79

Discussion 79

Habit, phyllotaxis and foliage 79

Morphology of flower and fruit 81

Pedicel/floral articulation 81

Hypanthium 81

Calyx 81

* National Botanical Institute, RO. Box 471, Stellenbosch 7599.

** Department of Botany, University of Stellenbosch, Stellenbosch 7600.

MS. received: 1994-01-24.

Androecium 81

Scales 84

Gynoecium 84

Fruit 84

Pollen morphology 84

Karyology 84

Conclusions 84

Acknowledgements 85

References 85

INTRODUCTION

The genera Lachnaea L. and Cryptadenia Meisn.

belong to the family Thymelaeaceae. Their close affinity

is reflected by their placement in the different classifica-

tion systems of the Thymelaeaceae (Endlicher 1847, sec.

Domke 1934; Meisner 1857; Bentham & Hooker 1880;

Gilg 1894 and Domke 1934). Only Baillon (1880) did not

recognize Cryptadenia as a separate genus but treated it

as a section of Lachnaea. He found that the flowers of

Lachnaea varied from regular to irregular and that inter-

mediate forms made it impossible to split the genus.

The genus Lachnaea , based on two species, was es-

tablished by Linnaeus in 1753. In subsequent publications,

such as Lamarck (1792, 1804), Salisbury (1808) and Sims

(1813, 1814), new species were described, but only in

1 840 was a detailed taxonomic account of the genus pub-

lished by Meisner based on Drege’s collections. Three sec-

tions were established, namely Sphaerocliniwn Meisn.,

Conoclinium Meisn. and Microclinium Meisn., based on

the inflorescence structure. Thirteen species and six in-

fraspecific taxa were recognized (Meisner 1840).

Meisner’s earlier contribution on Lachnaea , with some

alterations, was published in De Candolle’s Prodromus in

66

Bolhalia 25.1 ( 1995)

which 18 species and 17 infraspecific taxa were recog-

nized (Meisner 1857). The most recent taxonomic treat-

ment of the genus was that of Wright (1915) in Flora

capensis. In this treatment no mention was made of the

sections established by Meisner. Wright (1915) recognized

19 species and eight infraspecific taxa, and referred to one

imperfectly known species. In the Addenda to the same

volume, a further two species and two infraspecific taxa

were added (Wright 1925). Since then four new species

have been described, namely three by Compton (1934a

& b, 1953) and one by Fourcade (1934). When Thoday

(1924) published the revision of the genus Passerina , he

noted that the type of Passerina laniflora C.H. Wright

was not a Passerina species but indeed a species of Lach-

naea. Bond (1982) proposed the new combination, Lach-

naea laniflora (C.H. Wright) Bond. L. marlothii Schltr.

(Schlechter 1896) and L. gracilis Meisn. (Meisner 1840)

were both overlooked by Wright. At present 29 species

and eight infraspecific taxa are recognized (Van Wyk

1993).

The genus Cryptadenia , comprising five species, was

established by Meisner in 1840 and was also based on

Drege’s collections (Meisner 1840). According to Meisner

(1840) the main characters by which Cryptadenia can be

distinguished from the closely related genus Lachnaea,

are the glands which are positioned much lower down in

the hypanthium just above the ovary, and by the con-

spicuous li laments which are adnate to the hypanthium.

In De Candolle’s Prodromus the same five species were

recognized by Meisner ( 1 857). The most recent taxonomic

treatment of Cryptadenia was that of Wright (1915) in

Flora capensis , in which four of the species in Meisner’s

treatment were recognised, whereas one was reduced to

synonymy. A new species was also described. The

criterion used by Wright (1915) to delimit the two genera

was the position of the flowers: ‘Flowers axillary, solitary'

in Cryptadenia opposed to ‘Flowers terminal, capitate,

rarely solitary’ in Lachnaea. At present Cryptadenia is

considered to comprise five species (Van Wyk 1993).

Both Lachnaea and Cryptadenia are endemic to the

Cape Province. Lachnaea is found mainly at high altitudes

from Clanwilliam in the north to the Cape Peninsula in

the south and eastwards along the coastal areas as far as

Uitenhage. Further inland it occurs on the mountain ran-

ges from Ceres through Worcester to Ladismith (Bond &

Goldblatt 1984). The centre of diversity is in the south-

western area. Cryptadenia occurs on sandy fiats and lower

mountain slopes from Piketberg in the west to the Cape

Peninsula, and along the coastal areas to Bredasdorp and

inland to Ceres (Bond & Goldblatt 1984).

Lachnaea and its closest related genus, Cryptadenia ,

are both characterized by the absence of petals and by a

hypanthium having scales below the insertion of the

stamens. Beyers & Van der Walt (1994), contrary to

Wright (1915), found that the two genera could not be

distinguished on the inflorescence structure. With the large

number of herbarium specimens now available, compared

to the small range of material on which earlier studies

were based, a sound delimitation of the two genera is

possible. The aim of the present investigation was to com-

pare the two genera using a multidisciplinary approach,

with the view to establishing whether they are congeneric

or not. Besides the morphological study accompanied by

field work, anatomical, palynological and karyological in-

vestigations were undertaken to assess the taxonomic

value of the evidence derived from these sources.

MATERIALS AND METHODS

Eighteen taxa were selected, 14 from Lachnaea and

four from Cryptadenia. The aim in selecting the taxa was

to have as broad a representation as possible of all the

taxa in the two genera. The criteria used for selecting the

Lachnaea taxa were as follows: a, taxa representative of

the three sections instituted by Meisner (1840), taking into

account the variation in each section; b, all taxa with

solitary flowers; c, the position of the scales in the floral

tube; d, the stigma types; e, the degree of asymmetry of

the calyx lobes.

Wright’s revision of Lachnaea and Cryptadenia in

Flora capensis (1915 and 1925) was followed. Type

specimens and quoted specimens were examined to verify

the species concepts for the taxa used in this study. In-

fraspecific variation has been included in each taxon

studied and has not been treated individually.

Four of the five species of Cryptadenia currently

recognized were studied. Cryptadenia breviflora Meisn.

was excluded as it is an intermediate taxon between Cryp-

tadenia grandiflora (L. f.) Meisn. and Cryptadenia

uniflora Meisn., which Levyns (1950) considered to be a

hybrid between these two species.

The following species were studied: Lachnaea aurea

Eckl. & Zeyh., L. axillaris Meisn., L. burchellii Meisn.,

L. buxifolia Lam., L. capitata (L.) Meisn., L. densiflora

Meisn., L. diosmoides Meisn., L. ericoides Meisn., L.

eriocephala L., L. filamentosa (Thunb.) Meisn., L. ner-

vosa Meisn., L. penicillata Meisn., L. ruscifolia Compton,

Cryptadenia filicaulis Meisn., C. grandiflora , C. laxa C.H.

Wright and C. uniflora (author citations according to Van

Wyk 1993).

For the anatomical study of the foliage leaves, both

material preserved in FAA and herbarium material was

used. The herbarium material was soaked in 50% ethanol

for 24 hours, thereafter treated as preserved material. The

preserved material was dehydrated and embedded in wax

according to the tertiary butyl method (Johansen 1940).

Transverse sections 12 pm thick were made through the

middle of the leaf blade with the aid of a rotary microtome

and stained with Alcian Green Safranin (AGS) (Joel

1983). Scanning electron micrographs of the leaf surfaces

were obtained by mounting portions from the same region

of the leaf blade of herbarium material on stubs which

were coated with gold and studied with a Joel JSM-35

scanning electron microscope. Epidermal cell patterns of

the leaf blade were studied under a light microscope after

the adaxial and abaxial epidermal layers were peeled off

and mounted in water.

The anatomical study of the flowers was done on

material preserved in FAA. The preserved material was

dehydrated and embedded in wax according to the tertiary

butyl method (Johansen 1940). For the anatomy, transverse

Bothalia 25, 1 (1995)

67

TABLE 1. — Voucher specimens of Lachnaea species for the leaf

anatomy, floral anatomy, pollen morphology and karyology

each genus. The terminology used to describe the pollen

morphology is that of Walker & Doyle (1975).

For chromosome studies inflorescences were fixed in

Camoy’s fixative in the field. After 24 hours, the fixative

was replaced with 85% ethanol for 15 minutes and then

finally replaced with 70% ethanol. Flower buds were

stained in Snow’s fluid (1963). The anthers were removed

and squashed in 45% acetic acid.

Voucher specimens for the anatomy of leaves and

flowers and for pollen morphology and karyology are

given in Tables 1 and 2.

RESULTS

Habit

I n Lachnaea the species are sparsely to densely

branched, erect shrubs or erect to sprawling shrublets

0.15-2.00 m high. There is considerable variation in ap-

pearance and size among, and sometimes within species.

The Cryptadenia species are erect or sprawling, moderate-

ly to densely branched shrublets 0.12-0.80 m high. In

both genera the species are either basally single-stemmed,

multi-stemmed, or single-stemmed and multi-stemmed

within a single species. The branches are leafy, usually

with an adpressed pubescence, later becoming naked and

glabrous, and usually marked with the leaf scars of the

fallen leaves.

Morphology of foliage leaves

In Lachnaea the leaves are alternate or decussate, but

in L. burchellii they are opposite or subopposite and in L.

diosmoides they are alternate to subopposite. The leaves

are adpressed, ascending, patent or occasionally reflexed,

and imbricate to scattered. The leaves in Cryptadenia are

decussate, adpressed, ascending or sub-patent, and imbri-

cate or scattered. In both genera the orientation and posi-

tion of the leaves on the axis may vary inter- and

intraspecifically.

sections were used almost exclusively and longitudinal

sections were resorted to only to verify the points of at-

tachment of the floral structures. Serial sections were cut

12 pm in thickness. AGS staining (Joel 1983) as well as

Crystal Violet and Erythrosin staining (Jackson 1926)

were used. The AGS staining technique was found to give

better results.

Pollen was taken from herbarium specimens and from

material collected in the wild. For both light microscopy

(LM) and scanning electron microscopy (SEM), pollen

was acetolyzed according to Erdtman (1960). Acetolyzed

samples were prepared for both SEM and LM according

to the method of Marais (1990). At least 20 pollen grains

of each taxon were measured. Transverse sections ± 15

pm thick were made with the aid of a freeze-microtome

to study the structure of the pollen wall of two taxa in

TABLE 2. — Voucher specimens of Cryptadenia species for the leaf

anatomy, floral anatomy, pollen morphology and karyology

Species Voucher specimens

68

Bothalia 26.1 ( 1995)

The leaves in both genera are always entire and usually

sessile except in L. ruscifolia where they may be sub-

petiolate. The shape of the leaves in Lachnaea can be

linear-elliptic to elliptic, oblanceolate, obovate or ovate

(Figures 1, 2). Again, the variability is not only among

the species but may be within a particular species (Figure

2). The leaves range in size from 2-34 x 0.6-9.0 mm.

Within some species, as in L. filamentosa (Figure 2), the

variation in leaf size may be considerable, namely from

5-34 x 1.8-12.0 mm. In Cryptadenia the shape of the

leaves can be lanceolate, narrowly elliptic, narrowly ovate

or obovate. In C. uniflora the leaves are always narrowly

elliptic but in C. grandiflora they are narrowly elliptic,

narrowly obovate, obovate or oblanceolate. As in Lach-

naea the shape and size of the leaves vary inter- and in-

traspecifically (Figure 3). The range in leaf size is

2.9-12.8 x 0.6-3.2 mm. The leaves in both genera are

coriaceous but in L. densiflora they are occasionally semi-

succulent.

In Lachnaea the abaxial side of the leaves may be Hat,

obtusely convex to convex, or faintly keeled or keeled in

the distal half. Adaxially they may be flat, obtusely con-

cave or concave. Both surfaces of the leaves are usually

smooth. However, the abaxial surface in L. ruscifolia and

L. nervosa has several longitudinal ribs, whereas in L.

diosmoides there is only a single midrib. The abaxial sur-

face is usually green and shiny, whereas the adaxial sur-

face is usually pale green, dull and white-punctulate but

in some species the leaves are concolorous. The con-

FIGURE I . — Lachnaea. Variation of leaf shape and size, abaxial view.

A, L. eriocephala , Beyers 54; B & C, L. ericoides , Beyers 172 ;

D, L.funicaulis, Beyers 125; E, L. diosmoides, Beyers 181 ; F,

L. capitata, Beyers 128. G & H, L. burchellii: G, Oliver 9251 ;

H, Spreeth 155. 1-L, L. densiflora'. I & J, Oliver 9129; K. Taylor

12023; L, Beyers 171 . M-O, L. aurea, Beyers 135; P & Q. L.

nervosa , De Kock 152; R, L. ruscifolia, Beyers 193; S, L.

penicillata, Beyers 173. T-V, L. axillaris : T, McDonald 1395;

U & V, Beyers 197. Scale bars: A-R, 3 mm; S-V, 1 mm.

FIGURE 2. — Lachnaea buxifolia and L. filamentosa. Variation in

leaf shape and size, abaxial view. A-I, L. buxifolia'. A-C,

McDonald 1683; D. Marais s.n.; E-G, Beyers 118; H & I.

Beyers 122. J-R, L. filamentosa; J & K, Low 1246; L & M,

Beyers 109; N & (X Beyers 118; P-R, Taylor 12001. Scale

bar: 6 mm.

colorous leaves are usually green but in L. buxifolia , and

sometimes in L. filamentosa , they are glaucous. In Cryp-

tadenia the abaxial side of the leaves is convex, faintly

FIGURE 3. — Cryptadenia. Variation in leaf shape and size, abaxial

view. A-C, C.filicaulis; A, McDonald 1988; B & C, Oliver 9803.

D, C. uniflora, Beyers 136; E & F, C. laxa, 1. Oliver 4. G-K, C.

grandiflora; G, Beyers 136; H, VI ok2179; I K, Beyers 129. Scale

bar: 3 mm.

Bothalia 25.1 (1995)

69

c

200J-im

D

200,urn

FIGURE 4. — Lachnaea and Cryptadenia. Leaf transections, adaxial face above. A, L. eriocephala, Beyers 54; B, L. densiflora, Beyers 115; C, L.

biaifolia , Beyers 122; D, C. grandiflora , Beyers 151; E, C. laxa , 1. Oliver 4. c, large cells of inner layer of biseriate epidermis; s, stoma. Scale

bars: A, B, E, 100 |im; C, D. 200 |im.

keeled or keeled, smooth or transversely rugulose towards

the margins with a shiny, green surface, whereas adaxially

they are concave with a smooth, dull, pale green, white-

punctulate surface.

In Lachnaea the leaves are glabrous or pubescent at

the margins but the marginal pubescence is often

deciduous, whereas in Cryptadenia the leaves are always

glabrous.

Anatomy of foliage leaves

In transverse sections the leaves of most Lachnaea

species and all Cryptadenia species appear concave

adaxially and convex abaxially (Figure 4A, D, E), but in

L. densiflora both surfaces are flat or the adaxial one may

be slightly convex (Figure 4B); in L. bnxifolia (Figure

4C) and L. filamentosa both surfaces are flat.

Cuticle

In Lachnaea the cuticle is well developed and of even

thickness on both sides, or thicker on the abaxial side,

whereas in Cryptadenia the well-developed cuticle is al-

ways thicker abaxially. Under the SEM the adaxial and

abaxial cuticles of the Lachnaea species are unsculptured

(Figure 5A, D), undulate (Figure 5B, E) or minutely papillate

(Figure 5C, F). In Cryptadenia the cuticle is usually minutely

papillate adaxially, as in C. grandiflora (Figure 6A), but in

C. filicaulis it is unsculptured (Figure 6B). The relief of the

70

Bothalia 25,1 (1995)

abaxial cuticle is either minutely papillate (Figure 6C) or

undulate (Figure 6D). In C. grandiflora , however, both

types of cuticular sculpturing occur abaxially.

Waxes in the form of small Hakes occur commonly on

the adaxial leaf surfaces of Cryptadenia species and most

Lachnaeci species (Figure 7 A, C). In both genera, when

waxes are present on the abaxial surface, they are sparse

and flakelike (Figure 7B, D). Cutter (1978) is of the

opinion that it is this wax deposit that gives the 'bloom'

to glaucous leaves.

Trichomes, when present, are non-glandular and

uniseriate in both genera.

Epidermis

In both genera the adaxial epidermis is composed of

small cells with thin anticlinal and inner tangential walls

and slightly thicker outer tangential cell walls. In L.

eriocephala the outer tangential cell walls are the thickest

(Figure 4A). In Lachnaeci and Cryptadenia the adaxial

epidermis is uniseriate as in L. eriocephala (Figure 4A)

FIGURE 5. — Lachnaeci. SEM micrographs of cuticular relief of leaf. A-C, adaxial surface: A, L. funicaulis, Beyers 125 , smooth; B, L. buxifolia,

Beyers 122 , undulate; C, L. ericoides, Beyers 172 , minutely papillate. D-F, abaxial surface: D, L. ericoides, Beyers 172 , smooth: E, L.

diosmoides , Beyers 181. undulate; F. L. funicaulis. Beyers 125. minutely papillate, s. stoma. Scale bars: A-F, 20 pm.

Bothalia 25,1 (1995)

71

FIGURE 6. — Cryptcidenia. SEM micrographs of cuticular relief of leaf. A, B, adaxial surface: A, C. grandiflora, Beyers 129 , minutely papillate;

B, C. filicaulis, Boucher 5071, smooth. C, D, abaxial surface: C, C. grandiflora, Beyers 129, minutely papillate; D, C.filicaulis, Boucher

5071, undulate, s, stoma. Scale bars: A-D, 20 jam.

and C. grandiflora (Figure 8D. E), incompletely uniseriate

(occasional cells with periclinal walls) as in L. ericoides

(Figure 8 A) and C. uniflora (Figure 8C), incompletely

biseriate (occasional cells without periclinal walls) as in

L. densiflora (Figure 4B) and C. laxa (Figure 4E) or

biseriate as in L. buxifolia (Figure 4C). The abaxial epider-

mis is usually biseriate in both genera. However, in L.

axillaris and L. ericoides (Figure 8A) it is incompletely

biseriate and in C. grandiflora it may be uniseriate (Figure

8D) or incompletely biseriate (Figure 8E).

When the epidermis is biseriate, the cells of the inner

layer are usually larger than those of the outer layer. The

epidermis contains mucilage, except in C. grandiflora

where it was not always observed. The mucilage may be

present in the cells of the inner and outer epidermal layers,

or in the cells of the inner layer only.

Tannin occurs in the epidermis of L. axillaris , L. bur-

chellii, L. diosmoides, L. ericoides, L. funicaulis, L.

penicillata, C. grandiflora and C. uniflora.

Druse crystals were observed in the epidermis of L.

buxifolia, L. filamentosa and L. ruscifolia, and sometimes

in the adaxial epidermis of C. grandiflora.

Stomata

The stomata in both genera are anomocytic and sunken

except in L. funicaulis and L. penicillata where they are

on the same level as the outer walls of the surrounding

epidermal cells. In Lachnaea the leaves are epistomatic

or amphistomatic but in Cryptadenia they are only episto-

matic. When the leaves are amphistomatic. there are

usually more stomata adaxially.

Mesophyll

In both genera the cells of the mesophyll are

chlorenchymatous and palisade-like (elongated perpen-

dicular to the axis) with the exception of C. grandiflora

where they are either palisade-like (Figure 8E) or short

and more or less round (Figure 8D). As a result of the

size of the intercellular spaces, the leaves can be classified

as dorsiventral or isobilateral. When dorsiventral, the

smaller intercellular spaces occur abaxially and the larger

ones adaxially as in L. axillaris, L. diosmoides (Figure

8B), L. ericoides, L. eriocephala, L. funicaulis, L. nervosa,

L. penicillata, C. filicaulis, C. laxa and C. uniflora (Figure

8C). The isobilateral arrangement occurs in L. aurea, L.

burchellii, L. buxifolia, L. capitata, L. densiflora (Figure

4B), L. filamentosa and L. ruscifolia. In C. grandiflora

72

Bothalia 25,1 (1995)

however, a dorsiventral or an almost isobilateral arrange-

ment is found.

Tannin occurs in the parenchymatous cells surrounding

the median vascular bundle in L. penicillata and some-

times in the mesophyll in C. grandiflora.

Druse crystals are present in the mesophyll of L. aurea,

L. axillaris, L. burchellii, L. buxifolia, L. capitata, L. den-

siflora, L. filamentosa, L. nervosa and the four Cryptadenia

species, and in the vascular tissue of L filamentosa.

Venation and vascular bundles

The venation in both genera is reticulate and the vas-

cular bundles are collateral. As Metcalfe & Chalk (1950)

stated, no internal phloem occurs in the midrib.

In both genera, extraxylary fibres occur abaxially in

association with the vascular bundles. A prominent median

fibre strand (Figure 8) is present in all the species except

in L. buxifolia and L. filamentosa. In addition to the

prominent median fibre strand, equally prominent fibre

strands occur in association with the larger lateral vascular

bundles in L. nervosa, L. ruscifolia and C. grandiflora

(Figure 8E). In L. ruscifolia the median and larger lateral

strands extend to the abaxial epidermis.

Extraxylary fibres not associated with the vascular

bundles, occur along the lateral margins immediately

below the epidermis in L. aurea (Figure 8F).

Morphology of flower and fruit

Pedicel/ floral articulation

The flowers in Lachnaea are either sessile or shortly

pedicellate with pedicels up to 2.5 mm long. In Cryp-

tadenia the llowers are always sessile. In L. eriocephala,

L. funicaulis and L. penicillata the sericeous to villous

pedicels elongate in the fruiting stage and remain on the

plant long after the fruit has been shed. These hairy

pedicels are responsible for the woolly appearance of the

old inflorescences. The llowers of Lachnaea and Cryp-

tadenia are at their bases surrounded by a ring of

trichomes, which occurs at the floral articulation of the

sessile flowers or on the pedicels.

Hypanthium

In Lachnaea the basal portion of the hypanthium is

ellipsoid to widely ellipsoid, ovoid to widely ovoid, or

obovoid, whereas the portion above the plane of cir-

cumscision is funnel-shaped, cylindrical, obconical, cup-

shaped or cup-shaped with a sigmoid curve at the base

(Figure 9). In Cryptadenia the basal portion of the hypan-

FIGURIi 7.- Lachnaea and Cryptadenia. SEM micrographs of flaky surface wax on leal'. A, B, L. aurea, Beyers 135: A, adaxial surface; B, abaxial

surface. C, D, C. laxa. Bolus 7875: C, adaxial surface; D, abaxial surface. Scale bars: A-D, 5 pm.

Bolhaliii 25. 1 (l‘W5)

73

FIGURE 8. — Lachnaea and Cryptadenia. Leaf transections, adaxial face above. A— E, median vascular bundle: A, L. ericoides, Beyers 172\ B, L.

diosmoides , Beyers 181 ; C, C. uniflora , Beyers 136\ D, C. grandiflora, Beyers 129\ E, C. grandiflora , Beyers 151 . F, leaf margin of L. aurea ,

Beyers 54, showing extraxylary fibres (ex) along lateral margin, p. palisade-like chlorenchyma; me, median extraxylary fibre strand; le, lateral

extraxylary fibre strand. Scale bars; A. D, E, 100 pm; B, C. F, 50 pm.

thium is ovoid or ellipsoid, whereas the upper portion

above the plane of circumscision is funnel-shaped or cup-

shaped (Figure 9). The position of the plane of circum-

scision, expressed as a percentage of the length of the

base to the total length of the hypanthium. varies from

20-80% in both genera. The upper portion of the hypan-

thium abscises soon after seedset at the plane of cir-

cumscision. The length of the hypanthium ranges from

1.5-17.5 mm in Lachnaea and from 2.2-11.0 mm in

Cryptadenia.

The outer surface of the base of the hypanthium is

pubescent or glabrous in Lachnaea and pubescent in

Cryptadenia, whereas the inner surface is usually glabrous

in both genera, except in L. bwdfolia and C. uniflora

where it is pubescent. The outer and inner surfaces of the

upper portion of the hypanthium in both genera are usual-

ly pubescent, but in L. axillaris the outer surface is

glabrous.

In both genera the shape of the trichomes and type of

pubescence on the outer surface of the hypanthium often

changes around the plane of circumscision. In L. burchellii

the trichomes on the basal portion are clavate, whereas

those on the upper portion are longer and acicular (Figure

10). In C. laxa the trichomes of the basal portion are ob-

tuse, whereas those of the upper portion are acicular. The

pubescence on the outer surface of the hypanthium is

denser in the vicinity of the scales in both Lachnaea and

Cryptadenia. The upper portion of the hypanthium and

the calyx are concolorous.

74

Bothnlia 25,1 (1995)

Calyx

The calyx in both genera comprises four sepals which

are fused to the rim of the hypanthium. In Lachnaea the

sepals vary from being almost equal to distinctly unequal,

but with a gradual transition from one form to the other

(Figure 1 1 ). In the distinctly zygomorphic forms the anterior

lobe is usually the longest. The sepals are narrowly to widely

elliptic, ovate to widely ovate, or obovate to widely obovate.

The size of the anterior sepal ranges from 1 .3-22.0 x 0.8-7.5

mm, that of the two lateral sepals from 1. 3-7.5 x 0.6-4.5

mm, and that of the posterior sepal from 1. 2-5.5 x 0.7-3.5

mm. Because the flowers of L axillaris and L. ruscifolia are

solitary, the outer sepals represent the posterior and anterior

sepals and the two inner sepals represent the two lateral

sepals. The abaxial surface of the sepals is pubescent, except

in L. axillaris where it is glabrous. The adaxial surface of

the sepals is usually pubescent, but in L. aurea and L. nervosa

it is glabrous and in L. axillaris and L. funicaulis it is partly

glabrous and partly pubescent. The calyx is white, cream,

pink, pale blue, white or cream tinged with pink or blue, or

yellow.

The sepals in Cryptadenia are almost equal (Figure

12), naiTowly to widely elliptic, ovate or orbicular, varying

in size from 1.6-10.7 x 1.2-5. 8 mm with the apices acute,

rounded or obtuse. The abaxial surface is pubescent

whereas the adaxial surface is either glabrous or pubes-

FIGURE 9. — Lachnaea and Cryptadenia. Hypanlhia, showing plane of

circumscision, lower dotted line; and scales, upper dashed line.

A, C. uniflora, Beyers 136', B, C. grandiflora, Beyers 129; C, L.

densiflora, Oliver 9129; D, L. capitata, Beyers 138; E, C.

filicaulis. Thompson 3789; F, L. axillaris, Bolus 8596; G, L.

ericoides, Beyers 172; H, L. hurchellii, Oliver 9241; I, L.

funicaulis, Beyers 125; J, L. penicillata, Beyers 173; K, L. rus-

cifolia, Vlok 2084; L, C. laxa, Bolus 7875; M, L. nervosa, De

Kock 152; N, L. diosmoides, Compton 7580; O, L. aurea, Beyers

135; P, L. filamentosa, Beyers 110; Q, L. buxifolia, Beyers 118;

R, L. eriocephala, Beyers 54. Scale bar: 2 mm.

FIGURE 10. — Lachnaea burcliellii, Oliver 9241. SEM micrograph of

trichome structure on outer surface of hypanthium around cir-

cumscision zone, b, basal portion of hypanthium; c, plane of cir-

cumscision; u, upper portion of hypanthium. Scale bar: 200 pm.

cent. The calyx is white, cream, white tinged with pink,

pink or mauve-pink.

The aestivation of the sepals is decussate-imbricate in

Lachnaea and Cryptadenia.

Petals

Petals are absent in all taxa.

Androecium

Eight stamens in two whorls of four are present in all

laxa. In Lachnaea the outer whorl, the antisepalous

stamens, are episepalous, inserted on the basal portion of

the sepals, or they are inserted on the rim of the hypan-

thium (Figure 11). The stamens of the inner whorl, the

antipetalous stamens, are inserted on or very close to the

rim of the hypanthium or a third of the way down the

throat of the upper portion of the hypanthium as in L.

capitata (Figure 1 IF).

In Cryptadenia the outer whorl, the antisepalous stamens,

are episepalous, fused to the basal portion of the sepals as

in C. grandiflora and C. laxa, or inserted in the upper throat

of the hypanthium as in C. uniflora and C. filicaulis (Figure

12). The inner whorl, the antipetalous stamens, are inserted

at the rim of the hypanthium in C. laxa, a quarter of the

way down in the upper portion of the hypanthium in C.

uniflora and a third of the way down in C. filicaulis and C.

grandiflora (Figure 12). The stamens are fully exserted in

Lachnaea (Figure 1 1 ). In Cryptadenia the antisepalous

stamens of all the taxa and the antipetalous stamens of C.

laxa are exserted. In the other Cryptadenia species only the

anthers of the antipetalous stamens are exserted or semi-ex-

serted (Figure 12).

The anthers are basifixed and introrse. In Lachnaea

they are either spherical or oblate-spherical, whereas in

Cryptadenia they are ellipsoid or ovoid.

Bolhalia 25.1 (1005)

75

FIGURE 1 1 . — Lachnaea. Flowers. A,

L. buxifolia, Beyers 722; B, L.

filamentosa, Kruger 977', C, L.

aurea, Gillett 4343: D, L. ner-

vosa, De Kock 152; E, L.funi-

caulis. Marsh 758; F, L. capi-

tata, Beyers 138; G, L. den-

siflora, Boucher 1767; H, L.

ericoides , McDonald 2027; I,

L. penicillata, Oliver & Felling-

ham 9145. D-I: flowers split

longitudinally with gynoecia

removed; floral scales (arrow

in I) shown in each flower

below stamens. Scale bars: A-

C, 4 mm; D-I, 2 mm.

Scales

Eight scales are inserted on the upper portion of the

hypanthium below the attachment of the antipetalous

stamens and alternating with the stamens. The position of

the scales varies in different species of both genera. In

most Laclmaea species the scales are inserted at the mouth

of the hypanthium but in some species they are inserted

two thirds or halfway down the upper portion of the

hypanthium (Figure 11). In Cryptadenia they are inserted

either at the mouth of the hypanthium or from midway

to three quarters of the way down the upper portion of

the hypanthium (Figure 12).

The shape of the scales varies in the different taxa.

They can be linear, clavate, subrotund, capitate, ovoid or

obovoid in Lachnaea, whereas in Cryptadenia they are

subrotund, narrowly oblong, oblong or ellipsoid. In both

genera the scales are either translucent-white or yellow.

They are usually glabrous, except in L. buxifolia where

they have long stiff trichomes at their apices. In all the

Lachnaea species and in C. laxa the scales are basally

attached to the hypanthium. In C. fdicaulis the scales are

basally and partly abaxially attached to the hypanthium.

and in C. grandiflora and C. uniflora they are abaxially

attached to the hypanthium.

Disc

A disc is absent in both genera.

Gynoecium

In Lachnaea the ovary is sessile, oblong, narrowly el-

lipsoid, ellipsoid, ovoid or obovoid, 0.5^F4 x 0.2-2. 5

mm, completely glabrous or glabrous with a tuft of hairs

at the apex. In L. filamentosa the ovary is usually glabrous

with a tuft of hairs at the apex but occasionally it may

also be adpressed hairy and tufted at the apex.

In Cryptadenia the ovary is sessile, ellipsoid, 0.7-1. 8

x 0.4-0.7 mm. usually glabrous, but in C. filicaulis it is

FIGURE 12. — Cryptadenia. Flowers split longitudinally with gynoecia

removed. A, C. laxa, 1. Oliver 4; B, C. uniflora, Beyers 197; C,

C. grandiflora, Beyers 151; D, C. fdicaulis, Thompson 3789. sc,

scale. Scale bars: A-D, 2 mm.

76

Bothalia 25,1 (1995)

FIGURE 13. — Lachnaea and Cryptadenia. Gynoecia. A, L. funi-

caidis , Marsh 758: B, L. ericoides, Beyers 172 ; C, L. rus-

cifolia , Goldblatt & Manning 8870: D. L. nervosa, De Kock

152: E, C. laxa, 1. Oliver 4\ F, C. uniflora, Beyers 197: G, C.

grandiflora, Leighton BOL21 142: H, C.filicaulis, Thompson

3789. Scale bars: A-E, H. 1 mm; F, G. 2 mm.

Vascular anatomy of the flower

The basic vascular pattern is the same in both genera.

The eight traces supplying the calyx, depart from the vas-

cular tissue of the pedicel or the floral axis, where they

are arranged in a continuous cylinder (Figure 14A) or in

a ring of bundles (Figure 15 A) in Lachnaea and in a con-

tinuous cylinder in Cryptadenia (Figure 16A). The eight

sepal traces extend upwards in the hypanthium. The lateral

bundles of the sepals arise commissurally after the depar-

ture of the antipetalous stamen traces (Figure 17A, C, G,

H, I) or at the same point of departure of the antipetalous

stamen traces (Figure 17B, D, E, F, J) at various levels

on the hypanthium. The vascular supply of the sepals con-

sists of a midrib bundle and two lateral bundles.

The stamens are single-trace organs. The antipetalous

stamen traces are fused to the commissural sepal traces,

and the antisepalous stamen traces to the sepal midrib

traces at their point of origin from the stele. The stamen

traces separate from the sepal traces in two whorls at dif-

ferent levels in the hypanthium. The antipetalous stamen

traces diverge from the commissural sepal bundles before

either glabrous or pubescent towards the apex on the side

where the style is inserted.

In both genera the ovary is unilocular, with a solitary

anatropous ovule laterally attached near the top of the

ovary; the style is laterally attached, linear or linear-ob-

conical. The style is either completely glabrous or pubes-

cent in the distal third to two thirds, or completely

pubescent in Lachnaea , whereas in Cryptademk it is

usually glabrous but in C. laxa the distal half is pubescent.

The stigma in Lachnaea is either brushlike or capitate

and papillate, or capitate and elongate-papillate (Figure

13). In Cryptadenia it is either conical or capitate and

elongate-papillate, or brushlike (Figure 13).

Fruit

The fruit in both genera is an achene which is enclosed

in the persistent base of the hypanthium. The pericarp is

dry and thin, whereas the seed coat is thick and crus-

taceous.

FIGURE 14. — Lachnaea capitata, Beyers 128. Floral anatomy. A, tran-

section through floral axis. B, C, transactions of flower: B,

vasculature of hypanthium, separation of antipetalous stamen

trace (P ST) from commissural sepal bundle (SC); C, splitting of

commissural sepal bundles into sepal lateral bundles (SL). SM,

sepal midrib bundle. Scale bar: 500 pm.

Bothalia 25, 1 (1995)

77

FIGURE 15. — Lachnaea diosmoides,

Beyers 151. Transactions of

flower. A, pedicel showing ring

of vascular bundles; B, base of

hypanthium showing departure

of whorl of eight bundles; C,

hypanthium at base of ovary; D,

lower portion of ovary; E, mid-

dle of ovary; F, top of ovary; G,

base of style, o, ovary; ov,

ovule; dci, dorsal carpellary of

fertile carpel; cc, commissural

carpellary; dci, dorsal carpel-

lary of sterile carpel. Scale bar:

500 pm.

the antisepalous stamen traces diverge from the sepal

midrib bundles (Figure 14). The antisepalous stamen

traces leave the sepal midrib bundles at or just below the

rim of the hypanthium, except in C.filicaulis (Figure 17H)

where they depart a quarter of the way down the upper

portion of the hypanthium. The antipetalous stamen traces

leave the commissural sepal bundles at various levels on

the upper portion of the hypanthium. In L. buxifolia (Fig-

ure 17F), L. diosmoides (Figure 17B), L. eriocephala (Fig-

ure 17D), L. ruscifolia (Figure 17E) and C. laxa (Figure

17J) these stamen traces depart near the rim of the tube,

in L. axillaris (Figure 17C), C.filicaulis (Figure 17H), C.

grandiflora (Figure 171) and C. uniflora (Figure 17G) they

depart midway and in L. capitata (Figure 17A) they depart

three quarters of the way down the tube. The stamens

become free from the hypanthium a short distance above

the divergence of their traces as in L. buxifolia (Figure

17F) and C. fdicaulis (Figure 17H), or are adnate to the

hypanthium for some distance before becoming free as in

L. capitata (Figure 17 A) and C. uniflora (Figure 17G). In

L. buxifolia (Figure 17F), L. diosmoides (Figure 17B), L.

eriocephala (Figure 17D), C. grandiflora (Figure 171) and

C. laxa (Figure 17J) the antisepalous stamens are adnate

to the basal portion of the sepals. In Cryptadenia, with

the exception of C. Ictxa , the hypanthium. above the in-

sertion of the scales, is further augmented by small median

sepal bundles whose stelar extensions have been aborted

(Figures 17G, H, I; 18 A). Branches of these median sepal

bundles may anastomose with the midrib bundles and the

commissural sepal bundles. A rich vascular plexus exists

in the hypanthium of C. laxa at the level of insertion of

the scales (Figure 17J).

The eight scales arise as emergences from the adaxial

wall of the hypanthium (Figure 19) at different levels on

the upper portion of the hypanthium. They emerge a short

distance below the divergence of the antipetalous stamen

traces from the commissural sepal bundles, as in L.

capitata (Figure 17A) and in C. grandiflora (Figure 171),

or just above the divergence of the antipetalous stamen

traces and the lateral sepal traces from the commissural

sepal bundles, as in L. buxifolia (Figure 17F) and in C.

laxa (Figure 17J). No vascular tissue is present in or at

the base of the scales, even in those taxa in which the

scales are inserted in close juxtaposition to the sepal lateral

bundles and the sepal midrib bundles. In Lachnaea the

scales are usually non-glandular (Figure 20 A) but in L.

axillaris (Figure 20B) they are composed of compact thin-

walled cells rich in cytoplasm with large nuclei, thus ap-

pearing glandular (Esau 1965). Similarly in Cryptadenia

the scales are either non-glandular or appear glandular.

No secretion was observed from the scales in either genus.

The ovary is bicarpellate. In Lachnaea the dorsal car-

pellary bundle of the fertile carpel is always present,

whereas the dorsal carpellary bundle of the sterile carpel

is present in L. capitata and L. diosmoides (Figure 15),

but absent in L. eriocephala (Figure 21) and L. axillaris.

In Cryptadenia the dorsal carpellary bundle of the fertile

caipel is usually present, but absent in C. fdicaulis (Figure

16) whereas the dorsal carpellary bundle of the sterile car-

pel is always present (Figure 16).

i i

FIGURE 16. — Cryptadenia filicaulis , Oliver 9803. A, transection

through floral axis. B-D, transections of flower: B. base of flower

showing departure of whorl of eight bundles; C, lower portion of

ovary showing abortion of dorsal carpellary of fertile carpel; D.

base of style showing commissural carpellaries and dorsal car-

pellary of sterile carpel entering style, cc, commissural carpel-

lary; dc2, dorsal carpellary of sterile carpel. Scale bar: 500 pm.

78

Bothalia 25, 1 (1995)

FIGURE 17. — Lachmea and Cryp-

tadenia. Vasculature of hypan-

thium, stamens and sepals. A, L.

capitata, Beyers 128 ; B. L dios-

moides, Beyers 15 1; C, L. axil-

laris, Beyers 197; D, L.

eriocepluila, Beyers 54;E,L rus-

cifolia , Vlok 2084 ; F, L biocifolia,

Beyers 118; G, C. uniflora ,

Beyers 136; H. C. filicaulis ,

Oliver 9803; I, C. grandiflora ,

Beyers 196; J, C. laxa, I. Oliver

4. Scale bar: 1 mm.

The commissural carpellary bundles (Heinig 1951) are

closely associated with the dorsal bundle of the sterile

caipel (Figures 15, 16), or when the latter is absent they

are in the associated position (Figure 21).

The vasculature of the style is variable. It may consist

of: the commissural carpellary bundles as in L. diosmoides

(Figure 15), L. axillaris and L. capitata ; the two ventral

bundles of the sterile carpel arising from the split com-

missural carpellary bundles as in L. eriocephala (Figure

21); the dorsal bundle of the fertile carpel, the dorsal

bundle of the sterile carpel and the commissural carpellary

bundles as in C. grandiflora', the dorsal bundle of the

sterile carpel and the commissural carpellary bundle as in

C. filicaulis (Figure 16) and C. uniflora ; or the dorsal

bundle of the fertile carpel and the dorsal bundle of the

sterile carpel as in C. laxa.

Pollen morphology

Pollen grains of the Lachnaea and Cryptadenia species

examined, have the same basic morphology. The grains

are monads, globose and polyforate (Figure 22) with a

diameter of 22-52 pm in Lachnaea and 40-50 pm in

Cryptadenia (Table 3).

The pollen is tectate and supra-ornate (Figure 22).

Erdtman (1971) describes the sculpture pattern as

crotonoid. The sexine is thicker than the nexine. The

sexine is attached to the nexine by means of columellae,

which unite into a tectum above. The tectum is reticulate

with supratectal triangular projections. These supratectal

structures are trihedral to shallowly trihedral with the basal

sides straight as in L. buxifolia (Figure 23A), L. ruscifolia

(Figure 23B) and C. uniflora (Figure 23E) or emarginate

as in L. aurea (Figure 23C), L. eriocephala (Figure 23D)

and C. laxa (Figuie 231'). A vestigial spinule occurs at

FIGURE 18. — Crypnnhnia Jilieaiilis. Oliver 9803. Transection of

hypanthium of flowei . A. at level where scales ( Sc) separate from

hypanthium; B, where antisepalous stamens (S ST) separate from

tube. P ST, antipetalous stamens; SM, sepal midrib bundle; SC,

commissural sepal bundle; SL, sepal lateral bundle; MS, median

sepal bundle. Scale bar: 500 pm.

Bothalia 25,1 (1995)

79

FIGURE 19. — Lachnaea and Cryptadenia. Floral anatomy of hypanthia. Vasculature and scales (Sc) arising fromadaxial wall. A, L. capitata, Beyers 128\

B, C. grandiflora , Beyers 196. SM, sepal midrib bundle; SC. commissural sepal bundle; Sc, scale; MS, median sepal bundle. Scale bars: A. B, 100

pm.

the base of each of the three main ridges in some Lach-

naea species (Figure 23 A, B) but was absent in all the

Cryptadenia species (Table 4). The surface of the lateral

sides of the supratectal trihedral projections is striate in

all the species (Figure 23).

Cryptadenia (Table 5). The meiotic chromosome be-

haviour was normal.

DISCUSSION

Karyology

Habit, phyllotaxis and foliage

The haploid chromosome number of n = 9 was ob-

served in six species of Lachnaea and in two species of

The habit, phyllotaxis and macromorphological char-

acters of the foliage leaves are rather variable among the

!)%

'dip''.' A"!

vfY m '>T4;,'r

■ ju-u-v

- ;sV ■ sc

■ i-ri V. ' ■;■■■■ a

1 -rii V

V'. Y 'v'd.

- a y ( v.ty

s • vUr.

I :

k

FIGURE 20. — Lachnaea. Longitu-

dinal section of scale. A, L.

capitata , Beyers 128\ B, L.

axillaris , Beyers 127. Sc,

scale. Scale bars: A, B, 50 pm.

80

Bothalia 25,1 ( 1995)

TABLE 3. — Size range of pollen grains (mean followed by range in

parentheses) in Lachnaea and Cryptadenia

species and sometimes even within individual species of

Lachnaea and Cryptadenia. The variation range within

Cryptadenia is smaller than in Lachnaea but one must

take into account that Cryptadenia is a much smaller

group of species. With regard to the habit, phyllotaxis and

TABLE 4. — Comparison of sculpturing of supratectal projections of

tectate pollen in Lachnaea and Cryptadenia

TABLE 5. — Chromosome numbers of Lachnaea and Cryptadenia species

studied

macromorphological characters of the foliage leaves no

major differences were found which could be used to

demarcate the two genera.

Based on the anatomy of the leaves, two leaf groups

can be distinguished within Lachnaea-. 1, those which are

amphistomatic and either isobilateral or almost centric; 2,

those which are epistomatic and either isobilateral or dor-

siventral. The leaves of Cryptadenia are epistomatic and

either almost isobilateral or dorsiventral. The anatomy of

the latter resembles that of the second group in Lachnaea.

Similar cuticular patterns, namely unsculptured, minutely

papillate or undulate, occur in both genera. These charac-

ters confirm the close relationship between Cryptadenia

and Lachnaea.

FIGURE 21. — Lachnaea eriocephala , Beyers 54. Transections of

Bower: A, base of ovary; B, upper portion of ovary; C, top of

ovary; D, base of style, o, ovary; dc,. dorsal carpellary of fertile

carpel; cc,. commissural carpellary of fertile carpel; cc2, com-

missural carpellary of sterile carpel. Scale bar: 500 pm.

Bothalia 25,1 (1995)

81

FIGURE 22. — Lachnaea and Cryptadenia. SEM micrographs of pollen morphology. A, L. aurea, Beyers 135, pollen grain; B, L. buxifolia, Beyers

122, transection of pollen grain wall, n, nexine; c, columella; t, tectum; s, supratectal structure. Scale bars: A, 5 pm: B, 1 pm.

Morphology of flower and fruit

Pedicel/floral articulation

The study of the floral morphology revealed the strong

similarity between Lachnaea and Cryptadenia. Sessile

flowers are found in both genera. The ring of trichomes

at the base of the flower, occurring at the floral articula-

tion, corresponds to that found by Heads (1990) in Kel-

leria Endl. This structure, according to Heads (1990) is

reminiscent of the calycular pappus of many members of

the Asteraceae and the peltate hairs of Elaeagnaceae, and

could be regarded either as a compressed pubescent

'pedicel' or as the ‘calyx’ of a sessile flower. The pedicels

of pedicellate flowers of Lachnaea species are always

pubescent. The trichomes at the floral articulation do not

differ from those on the pedicel and it is therefore more

acceptable to regard the ring of trichomes at the base of

the sessile flowers as a compressed pubescent ‘pedicel'.

Hypanthium

The nature of the hypanthium of the flower of the

Thymelaeaceae has been interpreted as being appendicular

(foliar) and representing a calyx tube by Meisner (1857),

Wright (1915), Peterson (1959) and Dyer (1975), or as

being receptacular by Eichler (1878), Baillon (1880), Gilg

(1894), Wettstein (1935), Rendle (1938) and Bunninger

(1972). According to Lawrence (1951) the hypanthium is

more commonly and inaccurately termed the calyx tube

or floral tube. Heads (1990), Heinig (1951) and Saunders

(1939) regard the floral tube of the Thymelaeaceae as a

hypanthium. The vascular anatomy undertaken in this

study has shown that the floral tube in both genera can

be regarded as a hypanthium.

The position of the plane of circumscision expressed

as a percentage of the total length of the hypanthium

varies in both genera between 20% and 80%. The shape.

length and pubescence of the hypanthium of Cryptadenia

falls well within the variation range found in Lachnaea.

Calyx

The sepals in Lachnaea vary from being almost equal

to distinctly unequal. The transition from one form to the

other is gradual. The sepals of Cryptadenia are almost

equal which was one of the reasons why Baillon (1880)

incorporated Cryptadenia as a section in Lachnaea.

The aestivation of the sepals is the same in both genera

and a basic floral vascular pattern is common to both

genera.

Androecium

Eight stamens in two whorls of four each are present

in both genera. The position of insertion on the floral tube

shows a varying degree of adnation within both Lachnaea

and Cryptadenia. The stamens are always exserted in

Lachnaea , but in Cryptadenia they are either exserted or

only the antisepalous stamens are fully exserted.

Scales

In the different genera of the Thymelaeaceae organs

comparable to the scales in Lachnaea and Cryptadenia

have been interpreted by various authors as petaloid

glands, scales, mere outgrowths of the perigynium,

aborted stamens, squamellae which should be regarded

either as new structures of uncertain origin or as modified

parts of the androecium, stipules or petals which might

be reduced or greatly modified in form (Heinig 1951).

In Lachnaea various views regarding the nature of

these scales have been expressed, namely that they should

be regarded as staminodes, scales, glandular or scale-like

receptacular effigurations, or petaloid scales homologous

82

Bothalia 25,1 ( 1995)

FIGURE 23. — Lachnaea and Cryptadenia. SEM micrographs of pollen grains. A, L. buxifolia , Beyers 122: B, L. ruscifolia , Vlok 2084: C, L. aurea ,

Beyers 135: D, L. eriocephala, Beyers 54: E, C. uniflora , Haynes 477: F, C. laxa, Bolus 7875. s, supratectal structure; t, tectum; v, vestigial

spinule. Scale bars: A-F, 2 pm.

to petals. In those species of Lachnaea where these struc-

tures are narrowly straplike or filiform, Domke (1934)

regarded them to be in their original form and for that

reason Meisner (1840), according to Domke (1934), mis-

takenly considered them to be staminodes. These struc-

tures, in their original position according to Domke

(1934), are inserted at the base of the calyx lobes and

slightly higher than the insertion of the antipetalous

stamens but through zygomorphy these structures have

positioned themselves in a single whorl at the same height

as, or sometimes below, the filament insertion. This situa-

tion is not met in other genera such as Gnidia and

Struthiola , where the petal-like structures are inserted

above the insertion of the filaments at the mouth of the

tube. From the vascular anatomy of the flowers one would

rather believe the converse, namely that the scales were

orginally positioned below the antipetalous stamen whorl

and through zygomorphy have been slightly displaced to

above the antisepalous stamen insertion, as in L. filamen-

tosa. Heinig (1951) considered these structures to be

stipules of the calyx lobes and found no reason to consider

them to be vertical extensions of the disc as Gilg (1894)

Bothalia 25, 1 (1995)

83

did. Taxonomists have not been eager to accept the inter-

pretation of the petaloid scales as stipules on the grounds

that stipulate sepals would not occur in a group in which

the foliage leaves are exstipulate. Heinig (1951) supported

her interpretation on the grounds that the absence of

stipules on the foliage leaves and the presence of stipular

appendages in the lloral organs should be considered

another aspect of the progressive reduction from a more

primitive state. Heads (1990) rejects Heinig’s interpreta-

tion of the petaloid scales being stipules on the grounds

that she used the concepts of ‘leaf’ and ‘stipule’ of Eames

& MacDaniels (1948) which were in turn derived from

Van Tieghem’s (1871 in Heads 1990) morphology which

he regards as being fundamentally Hawed. Heads (1990)

refers to the comparable organs in Kelleria as gland scales.

In Cryptcidenia , as in Lachnaea , different views regard-

ing the scales have been expressed. They have been

referred to as glands, scales, glandular or scale-like recep-

tacular effigurations, or petaloid scales. Although Domke

(1934) referred to them as ‘glands’, he regarded them as

petaloid scales homologous to those of Lachnaea. Accord-

ing to Domke (1934) the ‘effigurations’ in L. axillaris , L.

diosmoides and L. ericoides, because of their position and

shape, lead directly to those in Cryptadenia where these

structures are nearly completely attached abaxially to the

hypanthium.

Because of their position and anatomy, we regard these

structures in both Lachnaea and Cryptadenia , as scales

which are mere emergences of the adaxial wall of the

hypanthium.

The position of the scales in both genera is usually

below the insertion of the antipetalous stamens. In the

more zygomorphic forms in Lachnaea and in one species

of Cryptadenia , where they are in close juxtaposition to

the stamens, they emerge between the insertion of the an-

tipetalous and antisepalous stamens. The position of the

scales in the upper portion of the hypanthium is variable

in both genera. Those of Lachnaea are inserted at the

mouth of the hypanthium to midway down the upper por-

tion of the hypanthium, whereas those in Cryptadenia are

inserted at the mouth of the hypanthium to three quarters

of the way down the upper portion of the hypanthium.

On the grounds of the position of the scales, a generic

character used by Meisner (1840, 1857), no cutoff point

can be made to separate the two genera as the displace-

ment of the scales is gradual. In both genera the scales

may be non-glandular or glandular. Another generic char-

acter which Domke (1934) used, was the attachment of

the scales to the hypanthium. According to him the scales

in Lachnaea are basally attached, whereas those in Ctyp-

tadenia are abaxially attached. In L. axillaris the cup-

shaped and sigmoid basal part of the upper portion of the

hypanthium gives the impression that the scales are basal-

ly attached. If however, the sigmoid curve is removed by

extending the hypanthium upwards, the scales would in

fact be abaxially attached and would correspond to the

situation in the flower of C. frlicaulis.

Gynoecium

The ovary in both genera is sessile, pseudomono-

merous (consisting of one expanded or fertile carpel and

one contracted or sterile carpel) and unilocular with a

solitary anatropous ovule laterally attached near the top

of the ovary. The style is always laterally attached to the

ovary. The stigma is either brush-like or capitate in both

genera but may also be conical in Cryptadenia.

Zygomorphy is expressed in both genera through the

lateral placement of the style and the single locule.

Fruit

In both genera the fruit is an achene which is enclosed

within the persistent base of the hypanthium. According

to Domke (1934) the achene is, without exception, char-

acteristic of the Thymelaeoideae and thus at the generic

level the fruit is not of taxonomic importance.

Pollen morphology

The pollen grains of Lachnaea and Cryptadenia have

the same basic morphology. Based on the sculpture of the

supratectal structures two pollen grain types can be dis-

tinguished within Lachnaea. namely those with vestigial

spinules at the base of the three main ridges and those

without spinules. In Cryptadenia the supratectal structures

are devoid of spinules at the base of the three main ridges

and are thus similar to the latter pollen grain type of Lach-

naea. Furthermore, in both genera, the lateral sides of the

trihedral structures are striate and the basal sides are either

straight or emarginate. The pollen grain size, following

Radford et al. (1974), is medium to large in Cryptadenia

and small to large in Lachnaea. This feature, therefore,

also overlaps in the two genera.

Karyology

In most Thymelaeaceae the haploid chromosome num-

ber (n = 9) and the somatic chromosome number (2n =

18) have been observed (Neviing & Woodbury 1966 in

Ornduff 1968. The following references are given in

Moore 1973: Sharma & Sarkar 1967, 1968, Bjorkquist et

al. 1969, Delay 1969, Gupta & Gillett 1969, Majovsky et

al. 1970. Bhat et al. in Love 1974. In Moore 1977: Kuz-

manov 1973, Loeve & Kjellqvist 1974. Majovsky et al.

1974. Heads 1990). Polyploidy has been recorded in

Daphne (Heads 1990), Daphnopsis (Neviing & Woodbury

1966 in Ornduff 1968), Edgeworthia Meisn. (Heads

1990), Pimelea Banks & Sol. (Heads 1990), and Wik-

stroemia Endl. (Gupta & Gillett 1969 in Moore 1973;

Heads 1990).

The chromosome number of Lasiosiphon eriocephalus

Decne., according to Subramanyan & Kamble in Love

1967, is n = 8. The genus Lasiosiphon Fresen. was incor-

porated in the genus Gnidia for southern Africa (Dyer

1975). Although the above species is not indigenous to

South Africa, it was the only chromosome number pre-

viously available for a genus of the Thymelaeaceae repre-

sented in South Africa.

The chromosome counts of n = 9 here reported for the

species of both Lachnaea and Cryptadenia , are the first

records for these two genera. The basic chromosome num-

ber of x = 9 was recorded for most genera of the

84

Bothalia 25,1 (1995)

TABLE 6. — Comparison of characters in Lachnaea and Cryptadenia

Thymelaeaceae and can therefore not be used to demar-

cate genera.

CONCLUSIONS

The Thymelaeaceae is a family in which it is difficult

to find sound characters for generic delimitation. Accord-

ing to Peterson (1959), only a few characters can be used

and they are of superficial significance. While revising

the thymelaeaceous genera of the African continent, Peter-

son (1959) experienced difficulties with the classification.

According to him these difficulties were not so much con-

nected to the species concept but rather to the delimitation

of genera which showed morphologically continuous

characters which merged them into each other.

According to Baillon (1880), the only other worker

who has discussed generic characters in the Thyme-

laeaceae, the following characters are generally of generic

value:

* the number of floral parts,

* the point of insertion of the stamens and the dimen-

sion of their filaments which render them exserted

or enclosed,

* presence or absence of the scales in the throat of the

hypanthium,

* presence or absence of a disc at the base of the

gynoecium,

* whether the base of the hypanthium is shed or persists

growing around the ripe fruit, and

* the arrangement of the inflorescence.

Bothalia 25,1 (1995)

85

In evaluating the results obtained from the present mul-

tidisciplinary study, consideration was given to those char-

acters which Baillon (1880) advocated as being of value

at generic level as well as those used by Heads ( 1990) to

illustrate the major differences between Kelleria Endl. and

Dmpetes Lam. The results of this multidisciplinary ap-

proach, as summarized in Table 6, illustrate the great

similarity between Lachnaea and Cryptadenia. These also

show that several characters within Lachnaea exhibit a

variation range which includes the variation in Cryp-

tadenia, and that in a few other characters the variation

range is extended in Cryptadenia. No distinguishing char-

acters of generic value were found to support the present

demarcation of Lachnaea and Cryptadenia. Furthermore

Beyers & Van der Walt ( 1994) concluded that the inflores-

cence morphology revealed no distinct differences be-

tween Lachnaea and Cryptadenia.

With the view of establishing natural genera, and on

the basis of the findings of this multidisciplinary study,

we support the viewpoint of Baillon (1880) that Cryp-

tadenia should be included within Lachnaea. A detailed

taxonomic treatment of Lachnaea sensu lato should now

be undertaken.

ACKNOWLEDGEMENTS

Mr E.G.H. Oliver is thanked for constructive discus-

sions throughout this research project.

REFERENCES

BAILLON, H. 1 880. The natural history of plants 6. Reeve, London.

BENTHAM, G. & HOOKER. J.D. 1880. Genera plantation 3. Reeve,

London.

BEYERS. J.B.P. & VAN DER WALT. .1.1. A. 1994. Inflorescence mor-

phology of Lachnaea and Cryptadenia (Thymelaeaceae).

Bothalia 24: 195-202.

BOND, R 1982. A new combination in Lachnaea L. (Thymelaeaceae).

Journal of South African Botany 48: 527.

BOND. P. & GOLDBLATT. P 1 984. Plants of the Cape Flora. A descrip-

tive catalogue. Journal of South African Botany Suppl. Vol. 13.

BUNNTNGER, L. 1972. Untersuchungen iiber die morphologische Natur

des Hypanthiums bei Myrtales und Thymelaeales-Familien. II.

Myrtaceae. III. Vergleich mit den Thymelaeaceae. Beitraege zur

Biologie der Pflanzen 48: 79-156.

COMPTON. R.H. 1934a. Lachnaea elegans. Journal of Botany. British

and Foreign 72: 20.

COMPTON, R.H. 1934b. Lachnaea naviculae folia. Journal of Botany,

British and Foreign 72: 21.

COMPTON, R.H. 1953. Lachnaea ruscifolia. Journal of South African

Botany 19: 132.

CUTTER, E.G. 1978. Plant anatomy. Part 1: Cells and tissues, 2nd edn.

William Clowes, London.

DOMKE, W. 1934. Untersuchungen iiber die systematische und

geographische Gliederung der Thymelaeaceen nebst einer

Neubeschreibung ihrer Gattungen. Bibliotheca Botanica 27: 1-

151.

DYER, R.A. 1975. The genera of southern African flowering plants, Vol.

1. Department of Agricultural Technical Services, Pretoria.

EAMES, A.I. & MACDANIELS, L.H. 1948. An introduction to plant

anatomy, 2nd edn. McGraw Hill, New York.

EICHLER. A.W. 1878. Bliithendiagramme II. Leipzig.

ERDTMAN, G. 1960. The acetolysis method. A revised description.

Svens k botanisk tidskrift 54: 561-564.

ERDTMAN, G. 1971. Pollen morphology and plant taxonomy. Hafner,

New York.

ESAU, K. 1965. Plant anatomy, 2nd edn. Wiley, New York.

FOURCADE, H.G. 1934. Contributions to the flora of the Knysna and

neighbouring divisions. Transactions of the Royal Society of

South Africa 21 : 75-102.

GILG, E. 1894. Thymelaeaceae. Die Natiirliehen Pflanzenfamilien 3, 6a:

216-245. Leipzig.

HEADS, M.J. 1990. A revision of the genera Kelleria and Drapetes

(Thymelaeaceae). Australian Systematic Botany 3: 595-652.

HEINIG, K.M. 1951. Studies in the floral morphology of the

Thymelaeaceae. American Journal of Botany 38: 1 13-132.

IACKSON, G. 1926. Crystal violet and erythrosin in plant anatomy. Stain

Technology 1: 33, 34.

IOEL, D.M. 1983. A.G.S. (Alcian Green Safranin) — a simple differential

staining of plant material for light microscopy. Proceedings of the

Royal Microscopical Society 18,3: 149-151.

IOHANSEN, D.A. 1940. Plant microtechnique. McGraw-Hill, New

York.

LAMARK. I.B.A.P.M. DE 1792. Encyclopedic methodique ( Diction -

naire encyclopedique de botanique) 3. Paris.

LAMARK, J.B.A.P.M. DE 1804. Encyclopedic methodique ( Diction -

naire encyclopedique de botanique) 5. Paris.

LAWRENCE, G.H.M. 1951. Taxonomy of vascular plants. Macmillan,

New York.

LEVYNS. M.R. 1950. Thymelaeaceae. Flora of the Cape Peninsula.

Juta, Cape Town.

LINNAEUS, C. 1753. Species plantarum, edn 1. Stockholm.

LOVE, A. 1967. IOPB chromosome number reports XII. Taxon 16: 341,

349.

LOVE, A. 1974. IOPB chromosome number reports XLIH. Taxon 23:

193.

MARAIS, E.M. 1990. Maak van stuifmeelpreparate vir ondersoek met

lig- en skandeerelektronmikroskoop (SEM). Department of

Botany, University of Stellenbosch. Unpublished.

MEISNER, C.F. 1840. Synopsis Thymelaearum, Polygonearum et

Begoniarum Africae Australis, imprimus a cl. J.J.Drege lectarum.

Linnaea 14: 385-516.

MEISNER. C.F. 1857. Thymelaeaceae. In A. de Candolle, Prodromus

systematis naturalis regni vegetabilis XTV: 573-580. Victoris

Masson, Paris.

METCALFE, C.R. & CHALK. L. 1950. Anatomy of the dicotyledons 1 &

2. Clarendon Press, Oxford.

MOORE, R.I. 1973. Index to plant chromosome numbers 1967-1971.

Regnum vegetabile 90: 282.

MOORE, R.J. 1977. Index to plant chromosome numbers 1973/74. Reg-

num vegetabile 96: 201.

ORNDUFF, R. 1968. Index to plant chromosome numbers for 1966.

Regnum vegetabile 55: 77.

PETERSON, B. 1959. Some interesting species of Gnidia. Botaniska

Notiser 112,4: 465-480.

RADFORD, A.E., DICKISON, W.C., MASSEY. I.R. & BELL, C.R.

1974. Vascular plant systematics. Harper & Row, New York.

RENDLE, A. B. 1938. The classification of flowering plants, II.

Cambridge University Press, Cambridge.

SALISBLfRY. R.A. 1808. Lachnaea glauca. In W. Hooker, Paradisus

Londinensis 6,1. London.

SAUNDERS. E.R. 1939. Floral motphology II. Heffer, Cambridge.

SCHLECHTER, F.R.R. 1896. Decades plantarum novarunr Austro-

Africanarum. Journal of Botany. British and Foreign. London 34:

500-504.

SIMS, J. 1813. Lachnaea purpurea. Curtis's Botanical Magazine 39: t.

1594.

SIMS, J. 1814. Lachnaea buxifolici (a) virens. Curtis’s Botanical

Magazine 39: t. 1657.

SNOW, R. 1963. Alcoholic hydrochloric acid-carmine as a stain for

chromosomes in squash preparations. Stain Technology 38: 9-13.

VAN WYK. C M. 1993. Thymelaeaceae. 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: 515-519.

THODAY, D. 1924. A revision of Passerina. Kew Bulletin 1924: 146—

168.

WALKER, W.J. & DOYLE, LA. 1975. The basis of Angiosperm

phylogeny: palynology. Annals of the Missouri Botanical Garden

62: 664-723.

WETTSTEIN, R. 1935. Handbuch der systematischen Botanik. Leipzig

and Vienna.

WRIGHT, C.H. 1915. Thymelaeaceae. Flora capensis 5,2 part 1. Reeve,

London.

WRIGHT. C.H. 1925. Thymelaeaceae. Flora capensis 5,2 part 4. Reeve,

London.

Bothalia 25,1: 87-95(1995)

Studies in the Ericoideae (Ericaceae). XVI. Six new species of Erica from the

Western Cape, South Africa.

E.G.H. OLIVER* and I.M. OLIVER*

Keywords: Erica, taxonomy, new species. South Africa, Western Cape

ABSTRACT

Six new species of Erica L. from the mountains of the Western Cape are described: E. alnea E.G.H. Oliv., E. hexensis

E.G.H. Oliv., E. hispiduloides E.G.H. Oliv. and E. tarantulae E.G.H Oliv. from the inland areas centred on the Hex River

Mountains; E. hottentotica E.G.H. Oliv. and E. magistrati E.G.H. Oliv. from the Hottentots Holland Mountains between

Stellenbosch and Somerset West.

UITTREKSEL

Ses nuwe spesies van Erica L. van die berge van die Wes-Kaap word beskryf: E. alnea E.G.H. Oliv., E. hexensis E.G.H.

Oliv., E. hispiduloides E.G.H. Oliv. en E. tarantulae E.G.H. Oliv. van die binnelandse streke gesentreer op the Hexrivierberge;

E. hottentotica E.G.H. Oliv. en E. magistral E.G.H. Oliv. van die Hottentots-Hollandberge tussen Stellenbosch en Somerset-

Wes.

INTRODUCTION

During our investigations of ‘incertae’ material housed

in herbaria, a number of new taxa have been encountered.

Five small-flowered species that would currently be

placed in the section Arscice and one in the section Eury-

storna, according to Flora capensis (Guthrie & Bolus

1905), are dealt with in this paper. Most of the material

has resulted from the remarkable collecting done by T.R

Stokoe and then Miss Elsie Esterhuysen on the high

mountains of the Western Cape during the period from

the 1930’s to the 1980’s.

1. Erica hispiduloides E.G.H. Oliv., sp. nov. (§ Ar-

sace), E. hispidulae L. pervulgatae, E. maestae Bolus et

E. glandulipilae Compton affinis sed ab E. hispidula pilis

mollibus in ramulis foliis pedicellis et sepalis. ab E. glan-

dulipila pilis longis eglandulosis vel glandulosis in

ramulis, glandibus sessilibus in marginibus sepalorum, ab

E. maesta bracteis bracteolisque subaequalibus saepe esul-

catis et glande apicali distinguitur. Figura 1.

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

Div., Hex River Mtns. Milner Ridge Peak, cliffs on S side,

1 525-1 678 m, (-AD), 10-11-1943, Esterhuysen 9379

(BOL, holo.; PRE, STE).

Shrub erect, densely branched, 0. 3-1.2 m tall.

Branches puberulous and with very long soft gland-tipped,

rarely eglandular hairs and shorter gland-tipped hairs in-

termixed, with no infrafoliar ridges, bark splitting ir-

regularly. Leaves 3-nate, 2. 0-3.0 x 0.6 mm, elliptic-

* Stellenbosch Herbarium, National Botanical Institute, P.O. Box 471,

Stellenbosch 7599.

MS. received: 1994-05-04.

oblong, erect to spreading, subopen-backed with revolute

rounded edges, puberulous, edged with a few very long

stout gland-tipped hairs and one apically; petiole ± 0.5

mm long, ciliate. Flowers 3-nate at ends of short lateral

branchlets, erect to spreading; pedicel 1-2 mm long,

puberulous; bract submedian, 0.5-1. 0 x 0.3-0.5 mm, nar-

rowly elliptic, sulcate or occasionally esulcate,

puberulous, with large apical gland; bracteoles opposite,

median, similar to bract but slightly smaller. Calyx 4-lobed

almost to base, 0.7- 1.5 mm long, cyathiform, green to

pink, lobes ovate, acute, sparsely puberulous, edged with

large glands. Corolla 4-lobed. 1.5 x 2.0-2. 5 mm,

cyathiform-urceolate, glabrous, pink, occasionally creamy

yellow, lobes erect, broadly rounded, about quarter length

of corolla. Stamens 8 included; filaments ± 0.5 mm long,

oblong, slightly broadening to base with sigmoid bend

below anther, glabrous; anther erect, muticous. thecae

oblong-elliptic, 0.4 x 0.8 mm, glabrous; pore half length

of theca; pollen in tetrads. Ovaiy 4-locular, 4-lobed, 0.5

x 0.9 mm, obovoid and emarginate, sparsely pilose, with

no basal nectary glands, ovules 15-20 per locule, spread-

ing to pendulous on a subapical placenta; style 0.3- 1.0

mm long, glabrous; stigma broadly cyathiform. Capsule

depressed globose, 1.0 x 1.2 mm, glabrous, yellowish

brown, septa free from central axis; seeds ellipsoid. ± 0.4

mm long, elongate reticulate, golden brown. Figure 1 .

This species is distinguished by the short soft hairs on

the stems, leaves, pedicel and sepals with long soft bend-

ing hairs, which may be glandular and eglandular, inter-

mixed with them on the branches, by sessile glands edging

the sepals and by the small subequal bract and bracteoles

which have a terminal stalkless gland and are often devoid

of a sulcus.

It is closely related to the very common and wide-

spread E. hispidula L. which has a shiny rather than dull

Bothalia 25,1 (1995)

FIGURE 1 . — Erica hispiduloides. A. flowering branches, x 1 ; B. stem; C, leaf; D, flower; E, sepal; F, anther, side, front & back views; G, gynoecium;

G', ovary cut longitudinally: H, fruiting gynoecium; I, seed, B, x 6, C-I, x 25. All drawn from type, Esterhuysen 9379.

appearance on close examination, stiffer, hispid hairs

which are sparser than the dense puberulous hairs of the

new species, particularly on the branches. E. hispidula has

lanceolate glabrous cartilaginous sepals and the corolla is

more urceolate with spreading lobes. It can also be dis-

tinguished from another species, E. glandulipila Compton,

which has long gland-tipped hairs on the sepals, anther

appendages and a peltate stigma. The other close relative,

E. maesta Bolus, differs in having no sessile glands on

the sepals, a glabrous ovary and plumose hairs on

branches and leaves.

E. hispiduloides occurs at high altitudes on the moun-

tains around the Worcester basin with an outlier to the

north in the southern Cedarberg and an extension

eastwards along the Klein and Great Swartberg as far as

Blesberg (Figure 2). The species could well be more fre-

quent, but may have been overlooked due to its close su-

perficial resemblence to E. hispidula which is probably

the commonest species of Erica in the Western Cape.

The habitat of the species is distinctive, being on moist

rock ledges on cliffs or very steep gullies on the south

FIGURE 2. — The known distribution

of Erica hispiduloides.

Bothalia 25,1 (1995)

89

FIGURE 3. — Erica hexensis. A, flowering branch, x 1; B, stem; C, leaf, adaxial view; C' , abaxial view; D, flower; E, anther, side, front & back

view; F, gynoecium; F', ovary, cut longitudinally. B-F', x 25. All drawn from Esterhuysen 8164.

side of mountains between 1 500 and 2100 m where it

forms dense woody shrubs up to 1 .2 m tall. On the ledges

facing south on Jonaskop, the plants were not common

and displayed two colour forms in the flowers on different

plants — creamy yellow with a tinge of pink or pure bright

pink. The latter were the taller shrubs, otherwise no dif-

ferences could be found between the two forms. Flowers

from September to December.

Specimens examined

WESTERN CAPE.— 3219 (Wuppertal): Cedarberg, Uitkyk Peak,

1 495 m, (-AC), 12-10-1975, Esterhuysen 84008 (BOL, PRE). 3319

(Worcester): Hex River Mtns. Milner Ridge Peak. 1 520-1 680 m. (-AD),

10-11-1943, Esterhuysen 9379 (BOL, PRE. STE); Milner Peak. 1 830

m, (-AD), 11-11-1960. Esterhuysen 28610 (BOL, K. PRE); kloof be-

tween Milner Ridge Peak & Shale Peaks, 1 680 m, (-AD), 2-01-1961,

Esterhuysen 28711 (BOL); Milner Peak, 1 680 m, (-AD), 11-10-1980,

Esterhuysen 35525 (BOL); Du Toits Kloof, Molenaarsberg, 1 520 m,

(-CA), 5-10-1947, Esterhuysen 14095 (BOL); Slanghoek Mtns, Wit-

teberg, 1 830 m, (-CA), 28-10-1979. Esterhuysen 35269 (BOL, K);

Brandwacht Peak, 1 830 m, (-CB), 26-11-1944, Esterhuysen 11022

(BOL); Chavonnesberg, 1 620 m, (-CB), 3-10-1948, Esterhuysen 14581

(BOL); Hex River Mtns. Horseshoe Peak, 1 680 m, (-CB), 1-11-1953,

Esterhuysen 22205 (BOL); Fonteintjiesberg. Meiring’s Plateau. 1 680 m,

(-CB), 20-10-1963, Esterhuysen 30412 (BOL); Robertson Dist., Das-

sieshoek Peak. 1 370 m. (-DB), 3-09-1961, Esterhuysen 29122 (BOL);

Jonaskop, 1 616 m, (-DC), 04-1983, Oliver 7971 (STE); Oliver 7972

(STE); ibid., 8-04-1994, Oliver & Oliver 10450 (BM. K. PRE, STE);

top of Baviaanskloof in Boschjesveld Mtns, (-DC), 02-1940, Stokoe 7488

(BOL, PRE) & SAM 55109A (SAM). 3321 (Ladismith): Klein Swartberg,

Toverkop. 1 980 m, (-AC), 16-12-1956, Esterhuysen 26760 (BOL. K);

Klein Swartberg W of Seven Weeks Poort Mtn, ledges on S side of

Ridge Peak, 2 100 m, (-AD), 27-12-1928, Andreae 1231 (BOL, PRE,

STE); Klein Swartberg, main ridge W of Hoeko Peak, 2 060 m, (-AD),

4-02-1992, Oliver 10015 (STE). 3322 (Oudtshoom); Swartberg, Blesberg,

1 830 m, (-BC), 17-10-1955, Esterhuysen 24919 (BOL, K, PRE).

2. Erica hexensis E.G.H. Oliv., sp. nov., (§ Arsaee),

E. hispidulae L. et E. glandulipilae Compton affinis sed

pilis longis apicibus perlonge furcatis, glandibus sessilibus

paucis in ramulis, aliter eglandulosa, antheris muticis,

ovario papillato-venucoso differt. Figura 3.

TYPE. — Western Cape, 3319 (Worcester): Ceres Dist.,

Hex River Mtns, shale band from Witels Kloof up to Buf-

felshoek Peak, SW aspect, 1 525-1 830 m, (-AD). 8-10-

1956, Esterhuysen 26366 (BOL, holo.; K, NBG, STE).

Shrub erect, compact up to 1 m tall. Branches erect

with numerous soft small lateral branchlets, puberulous,

some hairs up to 0.2 mm long and with long stouter long-

forked hairs up to 0.5 mm long and occasional sessile red

glands intermixed, older branches with more forked hairs,

with no infrafoliar ridges, bark grey, splitting irregularly

when older. Leaves 3-nate, 2.0 x 0.5 mm, oblong-elliptic,

open-backed, sparsely long villous mainly towards mar-

gins, abaxially shortly hispid; petiole ± 0.5 mm long, spar-

sely long ciliate. Flowers 3-nate at ends of very short

lateral branchlets, mostly facing upwards; pedicel ± 1.4

mm long, glabrous or occasionally with a few short hairs;

bract and bracteoles median 0.4 and 0.3 mm long respec-

tively, oblong not sulcate, glabrous sparsely ciliate with

long hairs. Calyx 4-lobed, fused at base only, lobes 0.8

mm long, lanceolate, acute, sulcate in upper half, half as

long as corolla, ciliate with long thin hairs and w ith some

scattered on abaxial side. Corolla 4-lobed. 1.5 x 1.4 mm,

cyathiform, glabrous, dusky red to pink, lobes obtuse,

crenate-fimbriate, erect. Stamens 8 included; filaments 0.7

mm long, narrowly linear from a slightly broader base

with a subsigmoid bend below anther, glabrous; anther

just manifest, 0.8 x 0.4 mm, ovate, obtuse, erect, dorsally

attached near base, muticous, very shortly strigose on

adaxial edge; pore one third to half length of theca; pollen

in tetrads. Ovary 4-locular, 4-lobed, 0.6 x 0.6 mm, broadly

ellipsoid and slightly emarginate, finely papillate, with no

90

Bothalia 25,1 (1995)

nectary glands at base, ± 8 ovules per locule, subpen-

dulous, placenta bulbous in upper half; style 0.5-1. 0 mm

long, slightly to much exserted, glabrous; stigma cyathi-

form, red. Fruit unknown. Figure 3.

E. hexensis is another of the small-flowered species in

the section Arsace that bears a resemblance to the very

common E. hispidula. From this it may be distinguished

by the long soft hairs with long-forked tips on the

branches, lack of glands except a few sessile ones on the

branches, distinct white median line down the sepals,

cyathiform corolla and papillate-verrucose ovary.

From the more eastern E. glandulipila it differs in the

lack of glands and anther appendages, the ovary surface

and the cyathiform stigma, the latter having a subpeltate-

capitate stigma. The type of hairs on the branches and the

ovary indumentum serve to distinguish it further from E.

hispiduloides.

As the name suggests the distibution of this species is

centred on the high peaks collectively known as the Hex

River Mountains in the Worcester/Ceres Districts (Figure

4). It grows on southwest- to southeast-facing slopes

which are cool and moist. Flowers from September to

November.

Specimens examined

WESTERN CAPE. — 3319 (Worcester): Worcester Dist., Hex River

Mtns, shale band near Buffels Dome, 1 700 m, (-AD), 10. 11.1943, Ester-

huysen 9370 (BOL, PRE); Worcester Dist., Hex River Mtns, Milner

Ridge Peak, 1 650 m, (-AD), 10-11-1943, Esterhuysen 9371 (BOL);

Ceres Dist., Hex River Mtns, shale band from Witels Kloof up to Buf-

felshoek Peak, 1 525-1 830 m, (-AD), 8-10-1956, Esterhuysen 26366

(BOL, K, NBG. PRE, STE); Worcester Dist., Hex River Mtns, slopes

below Buffelshoek Peak at head of Sentinel Kloof, 1 670 m, (-AD),

19-09-1965, Esterhuysen 31201 (BOL); Worcester Dist., Mostertshoek,

S face, 300 m, (-AD), 12-09-1976, Mostert s.n. (BOL); Mostertshoek,

left side of Donkerkloof, 1 060 m, (-AD), 17-09-1977, Mostert s.n.

(BOL); Worcester Dist., Chavonnesberg, cliffs below summit, 1 525 m,

(-CB), 3-10-1948, Esterhuysen 14578 (BOL); ibid., 1 220 m, 4-10-1942,

Esterhuysen 8164 (BOL, K, NBG, MO, PRE, STE) & 8165 , 8166 , 8167

(NBG); Worcester Dist., Fonteintjiesberg. I 525 m, (-CB), 20-10-1963,

Esterhuysen 30409 ( BOI , ).

FIGURE 4. — The known distribution of Erica hexensis.

3. Erica alnea E.G.H. Oliv., sp. nov., (§ Arsace),

E. cupuliflorae Dulfer affinis sed corolla late ellipsoidea,

antheris muticis thecis adaxialiter setiferis, ovario glabro

et stigmate late cyathiformi differt. Figura 5.

TYPE. — Western Cape, 3319 (Worcester): Ceres Dist.,

Hex River Mtns, shale band from Wit Els Kloof up to

Buffelshoek Peak, SW aspect, steep banks above stream,

1 220 m, (-AD), 8-10-1956, Esterhuysen 26362 (BOL,

holo.; PRE, STE).

Shrub erect, to 600 mm tall, much branched, dense,

lax and spreading in shady places. Branches hispidulous

with no infrafoliar ridges, bark grey, flaking irregularly.

Leaves 3-nate, appressed to spreading, reflexed in shaded

plants, 2.0-2. 5 x 0.6 mm, linear-elliptic, flat adaxially

with acute margins, convex and sulcate abaxially, glabrous

and shortly ciliolate; petiole ± 0.5 mm long, ciliolate,

otherwise glabrous. Flowers 3 to 6-nate at ends of

numerous short lateral branchlets; pedicel ± 2 mm long,

glabrous or with occasional short hairs at base; bract

median to remote, ± 0.5 mm long, oblong-lanceolate,

slightly sulcate subapically, ciliolate; bracteoles opposite,

just above bract and similar to bract. Calyx 4-lobed, fused

in lower quarter, glabrous, lobes broadly ovate, subacute,

glabrous, ciliolate, pink. Corolla 4-lobed, broadly ellip-

soid, 2.0 x 1 .5 mm, glabrous, pink, lobes broadly rounded,

erect to incurved, one third the length of corolla. Stamens

8 included; filaments ± 0.5 mm long, oblong, almost

straight, glabrous; anthers erect, muticous, glabrous,

thecae ellipsoid, 0.8 x 0.3 mm, with a few stiff hairs ±

0.15 mm long on adaxial edges; pore two thirds the length

of theca; pollen in tetrads. Ovary 4-locular, 8-lobed,

globose and emarginate, 0.5 x 0.7 mm, glabrous, no nec-

taries basally, ovules 20-26 per locule pendent from large

complete placenta; style 0.5- 1.0 mm long, glabrous, red;

stigma cyathiform, manifest to well exserted, red. Capsule

globose, 1.4 x 1.0 mm, septa free from central axis; seeds

0.2 x 0.1 mm, subovoid-ellipsoid, irregularly angled, the

angles often sharp, shallowly reticulate, yellow to yel-

lowish brown. Figure 5.

E. alnea is most closely related to E. cupuliflora on

account of the bulbous joined base of the calyx but differs

in having a broadly ellipsoid corolla, muticous anthers

placed on short broad filaments and with short stout setae

on the adaxial edges of the thecae, a glabrous ovary and

far exserted cyathiform stigma. Dulfer’s (1963) species

has an open cyathiform flower, scabrid anthers with long

awns, a sparsely pubescent ovary and subcapitate-

cyathiform stigma which is included. He likened his

species to E. micrandra Bolus but that species has remark-

able small anther thecae with relatively large narrow

crests, and also to E. monantha Compton which has aris-

tate leaves and belongs to the E. longipedunculata Lodd.

complex. The species could also be mistaken for E. tenuis

Salisb. or E. leptopus Benth. which have the same general

facies but simple stigmas.

All of the species mentioned above occur in the Ceres

District. The close ally, E. cupuliflora, occurs on sandy

flats and lower slopes, whereas the new species is con-

fined to damp slopes, streamsides and ledges at higher

altitudes from the Worcester/Ceres area with an outlier in

the central Cedarberg to the north (Figure 6). The type

Bolhalia 25.1 (1 995)

FIGURE 5. — Erica alnea. A, flowering branch, x 1; B, stem; C, leaf; D, flower; D' , old fruiting flower; E, anther, side, front & back views; F,

gynoecium; F', ovary cut longitudinally; G, fruit; H, seed. B-FI, x 25, with D', x 12.5. All drawn from type, Esterhuysen 26362.

collection comes from the Witels area, named after the

white alder [witels], Platylophus trifoliatus (L.f.) D. Don

(Cunoniaceae), hence the name of the new species. It was

collected at the same time as the type of E. hexensis (q.v.).

Flowers from May to October depending on the locality.

Specimens examined

WESTERN CAPE. — 3219 (Wuppertal); Cedarberg. Sneeuberg area,

Cedarhout Kloof. 910 m, (-AC), 2-09-1963, Esterhuysen 30316 (BOL,

K). 3319 (Worcester): Roodezand Pass, Tulbagh side, (-AA), 1-08-1938,

Stokoe 6836 (BOL); Hex River Mtns, shale band from Witels Kloof up

to Buffelshoek Peak, 1 220 m. (-AD), 8-10-1956, Esterhuysen 26362

(BOL, PRE, STE); Red Kloof at Worcester end of Du Toits Kloof, (-CA),

7-09-1958, Esterhuysen 27890 (BOL); Sanddrift Kloof, 910 m, (-BC),

29-05-1949, Esterhuysen 15402 (BOL, MO, NBG. PRE. STE); Prospect

Peak, 1 070 m, (-BC). 2-10-1949. Esterhuysen 15914 (BOL); Outer

Sanddrift Peaks, 1 220 m. (-BC), 28-10-1962, Esterhuysen 29752 (BOL.

PRE); Audensberg. (-CB), 6-10-1940, Esterhuysen 3343 (BOL); Fon-

teintjiesberg, (-CB), 08/09-1925, Stokoe in BOL 18399 (BOL); Stet-

tynsberg, (-CD), 07-1949, Stokoe in SAM 62334 (BOL, SAM);

Louwshoek Mtn, (-CD), 11-1946. Stokoe in SAM 62341 (BOL, PRE.

SAM).

4. Erica tarantulae E.G.H. Oliv., sp. nov., in genere

distinctissima propter cristas antherarum perasymmetricas

et saepe adscendentes sed in sectione Eurystoma posita,

E. calycinae L. et E. brevicaulis Guthrie & Bolus affinis

sed ab eis pilis paucis simplicibus non dendroideis, stylo

apicale papillato differt. Figura 7.

TYPE. — Western Cape, 3319 (Worcester): Cape, Ceres

Dist., Waaihoek Mtns, peak E of Tarantula Peak, on small

stony plateau, 1 200-1 370 m, (-AD), 7-10-1954, Ester-

huysen 21829 (BOL, holo.; MO, K. PRE, STE).

Shrublets low, semi-sprawling up to 160 mm tall.

Branches trigonous with no distinct infrafoliar ridges.

puberulous when young, soon becoming glabrous, bark

splitting irregularly and with small infrafoliar Oakes.

Leaves 3-nate, erect, adpressed, 1. 5-2.0 x 0.8 mm, nar-

rowly ovate to elliptic, convex and sulcate abaxially with

sharp edges, flat adaxially, puberulous towards base

adaxially and sparsely ciliolate with hairs and a few

stalked, red, gland-tipped hairs, otherwise glabrous;

petiole ± 0.3 mm long, ciliolate, glabrous abaxially,

puberulous adaxially. Flowers 3-6 at ends of main and

short lateral branchlets, mostly pendulous; pedicel ± 0.3

mm long, glabrous: bract approximate. 1.3 x 1.0 mm.

FIGLIRE 6. — The known distribution of Erica alnea. •: and E. hotten-

totica, O.

92

Bothalia 25,1 (1995)

FIGURE 7. — Erica tarantulae. A, branch, x I; B, stem & leaves; C, leaf, adaxial view; D. flower; E, corolla; F, sepal; G, bract; H, bracteole; I,

anther, side, front & back views; J, ovary; J', ovary, cut in half longitudinally; K, style & stigma; L, fruit; M, seed. B-H, x 12.5; I-M, x 25;

L, drawn from Stokoe 7 889; rest drawn from type, Esterhuy sen 21829.

broadly ovate, sulcate in upper half, glabrous, ciliolate

with hairs and stalked red glands, rose pink; bracteoles

approximate, like bract, but 1.2 x 0.8 mm. Calyx 4-partite,

segments 2.0 x 1.3 mm, elliptic to ovate, shortly sulcate

apically, glabrous, ciliolate with hairs and some stalked

red glands, rose pink. Corolla 4-lobed, 4.5 x 2.5 mm,

obconical, glabrous, rose pink, lobes quarter the length of

corolla, broadly obtuse, erect to slightly spreading.

Stamens 8, included to manifest; filaments ± 2 mm long,

linear, slightly bent backwards below anther; anthers dor-

sally attached near base, appendiculate, thecae ± 0.7 mm

long, oblong-falcate, obtuse, roughly papillate, crests very

variable in shape and size even on same anther, large.

FIGURE 8. — The known distribution of Erica tarantulae, •; and E.

magistrate O.

Ileshy. variously serrate, sometimes erect, pore half the

length of theca; pollen in tetrads. Ovary 4-locular and 4-

lobed, 1 .0 x 0.8 mm, broadly ellipsoid and emarginate,

sparsely puberulous apically, with nectary glands basally,

6-8 ovules per locule, pendulous on subglobose placenta

in upper half; style ± 4 mm long, narrow cylindrical, ex-

serted, papillate apically; stigma simple to subpeltate.

Capsule ellipsoid, 1.3 x 0.8 mm, reddish brown with

much darker base, valves splitting to base, septa almost

free from axis; seeds 0.5-0.6 mm long, subtrigonous-el-

lipsoid, yellowish brown, finely and closely reticulate,

reticulations tangentially finely undulate. Figure 7.

This very distinct species is unique in the genus for

the form of the anther appendages which are broad, very

irregularly shaped crests which, in some anthers, may be

absent on one side (Figure 71). They are remarkable in

sometimes being larger above the point of attachment to

the theca and therefore totally ascending.

The superficial facies of the calycine, open-mouthed

flowers with manifest stamens would place the species in

the section Eurystoma of Flora capensis (Guthrie & Bolus

1905) where it shows some resemblance to several species

which occur in the same area: E. calycina, E. brevicaulis

and E. costatisepala H.A. Baker. These latter species have

complex dendroid hairs on many parts of the plant and

have long pedicels. E. tarantulae has a sparse complement

of short fine simple hairs and almost subsessile Bowers.

The style is also unusual in being apically papillate, a

condition found only in the section Platyspora.

E. tarantulae appears to be very restricted in its dis-

tribution, being known only in the Hex River Mountains

complex as two localized collections from Tarantula and

Milner Peaks (Figure 8). It is recorded as growing on

stony ground where it forms low, spreading shrublets.

Flowers from early August to late in October.

Bothalia 25,1 (1995)

93

Specimens examined

WESTERN CAPE. — 3319 (Worcester): Ceres Dist., peak E of Taran-

tula Peak, stony plateau, 1 200-1 370 m, (-AD), 7-10-1953, Esterhuysen

21829 (BOL, K, MO, PRE, STE); Ceres Dist., Hex River Mtns, Ezelsfon-

tein, stony N slopes of Milner Peak, 900 m, (-AD), 24-08-1958, Ester-

huysen 27859 (BOL). Without locality: Stokoe 7889 (BOL).

awns ± 0.4 mm long, sparsely puberulous, pore half length

of theca. Ovary 4-locular, 0.5 x 0.7 mm, oblate to obovate,

slightly 8-lobed, emarginate, with basal nectary glands,

with a few longish hairs apically, ± 4 slightly erect ovules

per locule; style ± 4.0 mm long, glabrous, just exserted;

stigma cyathiform with 4 apical stigmatic papillae. Fruit

unknown. Figure 9.

5. Erica hottentotica E.G.H. Oliv., sp. nov.

(§ Ephebus ), E. parviflorae L. affinis sed sepalis ovatis

petaloideis roseis, corolla puberula, stigmate cyathiformi

differt. Figura 9.

TYPE. — Western Cape, 3418 (Simonstown): Somerset

West area, Hottentots-Holland Mtns, Somerset Sneeuw-

kop, (-BB), Stokoe 6641 (BOL, holo.; K, PRE, STE).

Erect compact shrub 0.5-1. 0 m, rarely up to 2.5 m,

tall. Branches erect, leafy with numerous dense lateral

short shoots, softly puberulous, without infrafoliar ridges,

bark flaking irregularly when old. Leaves 4-nate, erect,

imbricate, 0.5-3. 5 mm long, oblong-elliptic, subobtuse,

flattened adaxially with a distinct sharp margin, convex

and sulcate abaxially, with a few scattered short hairs

mainly towards margins abaxially, adaxially hairy; petiole

± 0.5 mm long, sparsely ciliolate. Flowers 1- to 4-nate

tenninal on short lateral branchlets, upright to pendent;

pedicel ± 2.0 mm long, puberulous; bract remote to

median, 0.6 x 0.2 mm. oblong, not sulcate, sparsely

puberulous; bracteoles 2, median, same as bract. Calyx

4-partite; segments 1.4 x 0.8 mm, ovate to lanceolate,

slightly sulcate apically, puberulous, pink. Corolla 4-

lobed, 3.0 x 2.0 mm, cyathiform to urceolate, puberulous,

red, lobes ± 0.5 mm long, rounded erect or slightly

reflexed. Stamens 8, included; filaments ± 1.5 mm long,

filiform, with sigmoid bend just below anther, glabrous;

anther dorsally attached near base, appendiculate, thecae

1.0 x 0.3 mm, oblong with acute apex, sparsely

puberulous laterally and adaxially mainly towards base.

This species is closely related to the common and

widespread E. parviflora but can easily be distinguished

by its broader, ovate, petaloid, pink sepals, finely hairy

corolla and broad cyathiform stigma. The latter species

has linear-lanceolate foliaceous sepals, a coarsely long-

haired corolla and a small capitate stigma. The flowers of

E. hottentotica are larger than those of E. parviflora al-

though the form of the latter in the Hottentots Holland

Mountains has fairly large flowers. E. parviflora is

generally a scrambling plant in marshes, whereas the new

species appears to form an erect sturdy shrub which may

reach a height of 2.5 m (Stokoe 4009). There is also a

resemblance to some forms of the common E. sitiens

Klotzsch which has, however, a glabrous corolla,

foliaceous sepals and a capitate stigma.

This species is confined to the higher reaches of the Hot-

tentots-Holland Mountains above Somerset West, hence the

name (Figure 6). It has been recorded with the following

species, E. magistrati, on Langkloofberg frequenting similar

habitats — moist southeast-facing slopes. The Hottentots-Hol-

land Mountains host the largest number of species of Erica :

175 spp. have been recorded so far (Oliver et at. 1983).

Flowers from September to late November.

Specimens examined

WESTERN CAPE. — 3418 (Simonstown): Somerset West area. Hot-

tentots Holland Mtns, (-BB); Langkloofberg, 1 220 m, (-BB), 30-10-

1943, Esterhuysen 9150 (BOL, STE); The Triplets, 1 220-1 520 m,

(-BB), 31-10-1943, Esterhuysen 9173 (BOL); Klein Valleiberg, 1 070-

1 220 m, (-BB). 11-10-1962, Esterhuysen 29868 (BOL. K, PRE);

FIGURE 9. — Erica hottentotica. A, flowering branch, x 1 ; B, stem; C, leaf; D, flower; E, sepal; F, anther, side, front & back views; G, gynoecium;

G', ovary, cut longitudinally. B-E, x 12.5, F, G & G', x 25. All drawn from Esterhuysen 9150.

94

Bothalia25,l (1995)

FIGURE 10. — Erica magistrati. A, flowering branch, x 1; B, stem; C, leal ; D, flower; E, sepal; F, anther, side, back & front views; G, gynoecium;

G', ovary, cut longitudinally. B-G', x 25. All drawn from Esterhuysen 33921 .

Landdrost Kop, 910-1 070 m, (-BB), 30-09-1979, Esterhuysen 35249

(BOL, K); vicinity of Landdrost Kop & Somerset Sneeukop, (-BB), 11-

1934, Stokoe 4026 (BOL, PRE); Somerset Sneeuwkop, (-BB), 1 1-1938,

Stokoe 6641 (BOL, K, PRE, STE); ibid., Stokoe 6826 (BOL); between

Somerset Sneeuwkop & Landdrost Kop, (-BB), 26-09-1943, Stokoe 8927

(BOL).

6. Erica magistrati E.G.H. Oliv., sp. nov. (§ Arsace),

E. salad Salisb. affinis sed ab ea ramis ramulisque pilis

multis brevissimis et longis glandulosis tectis, foliis pilis

simplicibus, corolla urceolatiore et ovario villoso differt.

Figura 10.

TYPE. — Western Cape, 3418 (Simonstown): Caledon

Div., Hottentots-Holland, Langkloofberg, SE slopes near

water, 910-1 220 in, (-BB), 30-10-1943, Esterhuysen

9145 (BOL, holo.; PRE, STE).

Erect woody shrub 0.6- 1.8 m tall. Branches densely

puberulous, the older with many long spreading stouter

gland-tipped hairs intermixed, without infrafoliar ridges,

bark splitting irregularly with age. Leaves 3-nate, imbri-

cate to spreading, 4.0 x 0.8 mm, oblong-elliptic, elliptic

in section with rounded margins, slightly open-backed,

apex subacute sometimes the younger ending in a long

gland-tipped hair, the younger pubescent with adpressed

hairs, becoming glabrous abaxially, with scattered short

hairs, marginally rarely also with long stouter gland-tipped

hairs; petiole ± 0.8 mm long, ciliolate. Flowers mainly

3-nate, terminal on short lateral branchlets 0.5-10.0 mm

long, erect to slightly spreading; pedicel ± 3 mm long,

softly puberulous; bract median to approximate, lanceo-

late, 2.0 x 0.6 mm, sulcate in upper half, pubeailous,

white; bracteoles opposite, just above the bract, like bract

but smaller. Calyx 4-lobed very slightly fused at base,

lobes 2.0 x 0.6 mm, ovate, acute, sulcate in upper third,

puberulous and ciliolate, while. Corolla 4-lobed, 3.0 x 2.5

mm, urceolate to cyathiform, glabrous or very rarely with

a few short hairs, lobes ± 0.5 mm long, rounded, erect to

slightly reflexed. Stamens 8 included; filaments filiform,

2.5 x 0.5 mm, erect, subsigmoid below anther, glabrous;

anthers erect, placed just above ovary, dorsally attached

near base, appendiculate, thecae 2.5 mm long, narrowly

ovate-oblong, glabrous, crests ± 0.6 mm long, slightly ser-

rate, attached at top of filament, pore half length of theca;

pollen in tetrads. Ovary 4-locular, 8-lobed, broadly

obovate, emarginate, 2.2 x 2.2 mm, villous mainly in

upper half and in lower half along lobe edges, with no

nectary glands at base, 15-20 ovules per locule spreading

from a rounded central placenta; style ± 2.5 mm long,

straight, glabrous, exserted; stigma cyathiform with 4

central stigmatic papillae, glabrous. Fruit unknown. Fig-

ure 10.

This species in the section Arsace is allied to E. salax

Salisb. which has been collected only a few times in the

same area. The latter has no long gland-tipped hairs on

the branches, but has very short dendroid or gland-tipped

marginal hairs on the leaves, a more cyathiform corolla

and glabrous ovary. We have been able to examine

Salisbury’s type (Masson s.n., Herb. Salisb. in Kew) to

verify the correct identity of the few collections which

were tentatively identified as this species.

There are also slight resemblances to E. sphaeroidea

Dulfer and E. argyrea Guthrie & Bolus from the same

area, but these have puberulous corollas and numerous

gland-tipped hairs on various other parts of the plant in

addition.

E. magistrati is confined to the mountains in the Stel-

lenbosch/Somerset-West area (Figure 8) where it is found

mainly on south-facing slopes from 450 m to 1 068 m in

situations receiving extra moisture. Several collections are

Bothalia 25,1 (1995)

95

associated with Landdrostkop (Magistrate’s Head) in the

Hottentots-Holland range, hence the name of the species.

Here there is often a good deposition of moisture from

the dense moving clouds produced by the strong southeast

winds which occur frequently during the summer months.

Flowers from August to December.

Specimens examined

WESTERN CAPE. — 3318 (Cape Town): Stellenbosch Dist., Jonker-

shoek, Langrivier, 760 m, (-DD), 08-1965, Kerfoot 5392 (PRE, STE).

3418 (Simonstown): Hottentots-Holland Mtns, Langkloofberg, 910-1 220

m, (-BB), 30-10-1943, Esterhuvsen 9145 (BOL, PRE. STE); Landdrost

Pinnacle above Lourensford, 460 m, (-BB), 7-09-1975, Esterhuvsen

33921 (BOL, K, PRE, STE); Jonkershoek, beyond Second Waterfall,

1 220-1 520 m, (-?BB), 11-1925, Krige in STE 2130 (BOL); Somerset

Sneeuwkop, Landdrost Kop & environs, (-BB), Stokoe 6826 (BOL);

ibid., 11-1938, Stokoe 6827 (BOL, SAM); Landdrost Kop, (-BB), 26-

09-1943, Stokoe 8933 (BOL, K, PRE). 3419 (Caledon): Jonkershoek,

ridge between Eerste River Kloof and Victoria Peak, 1 070 m, (-AA),

7-09-1979, Esterhuvsen 35242 (BOL, K).

REFERENCES

DULFER, H. 1963. Neue Arten und Varietaten der Gattung Erica L. aus

Siid-Afrika. Annalen des Naturhistorischen Hofinuseums, Wien

66: 19-33.

GUTHRIE. F. & BOLUS, H. 1905. Erica. In W.T. Thiselton-Dyer, Flora

capensis 4: 4-315. Reeve, London.

OLIVER, E.G.H., LINDER, H P. & ROURKE, J.P. 1983. Geographical

distribution of present-day Cape taxa and their phytogeographical

significance. Bothalia 14: 427-440.

■

Bothalia 25,1:97-109(1995)

Notes on African plants

VARIOUS AUTHORS

ASPHODELACEAE/ALOACEAE

TYPIFICATION OF ALOE SPECIES DESCRIBED BY B.H. GROENEWALD

In the mid- to late 1930’s, B.H. Groenewald (1935,

1936a-e, 1937a-c, 1938a-c, 1939, 1940) published a

number of descriptions of new species and one new

variety of Aloe L. Some of these were not accompanied

by any indication of specimens examined, whereas others

had specimen citations of the form ‘Herbarium No. ***’.

Localities and flowering times were given only in the dis-

cussion, and the name of the collector of cited specimens

was uniformly omitted, as was any indication of where

the ‘type’ might be found. It should be noted that only

on or after 1 January 1958 did the indication of a holotype

become compulsory in order to effect the valid publication

of the name of a new taxon of the rank of genus or below

(Greuter et al. 1988), thus about 20 years after Groene-

wald’s major written contributions to alooid taxonomy.

Furthermore, typification of extant yet untypified names

is not compulsory (Greuter et al. 1988, Art. 7.4 ‘.... a

lectotype or, .... a neotype may be designated ...’, but see

Art. 53.1), but in a taxonomically difficult genus such as

Aloe it is essential in order to establish the correct ap-

plication of the plethora of available names. In this paper

we present the results of our attempts to formally establish

types for the Aloe species published by Groenewald. Un-

fortunately very little is known about Groenewald in

general and his taxonomic and cytogenetic research ac-

tivities in particular, and he has been omitted from Gunn

& Codd (1981). Coincidentally another southern African

student of alooid taxonomy and cytogenetics, at about the

same time, Dr Franz Sebastian Muller (1945), has also

not been taken up in Gunn & Codd’s (1981) benchmark

biographical publication, the reason probably being that

neither of them collected a sufficient number of specimens

to warrant their inclusion. This situation will be rectified

in the second edition of this book which is currently in

preparation. One logical deduction from the way in which

Groenewald cited specimens in his papers is that his

‘types’ were housed in a personal herbarium. However,

evidence presented below indicates that such an herbarium

never existed.

An extensive and detailed search for a personal her-

barium belonging to Groenewald among his associates

and descendants failed to produce any results. This search

included the PRE and BLFU herbaria, the former as being

the most likely institute to have inherited any private her-

barium he made, and the latter as being closest to the

place of publication of his new species and of his book

on Aloe (Groenewald 1941). Not only were no specimens

forthcoming, but no source questioned had any recollec-

tion of there ever having been a Groenewald private her-

barium.

Positive evidence for the non-existence of a Groene-

wald private herbarium is largely indirect, and rests on

four points: 1, for each name published by Groenewald

except four, there is one (and usually only one) specimen

in PRE, collected by Dr Frederick Ziervogel van der

Merwe from the precise locality at the appropriate time

indicated by Groenewald in the protologue, some of which

are marked ‘TIPE’ in Van der Merwe’s handwriting. As

indicated below, most of these specimens are cited by

Reynolds (1950) as ‘Types’; 2, about half of these

specimens have collector’s numbers. Where a collector’s

number has been assigned it corresponds with the number

(if any) given by Groenewald; 3, Groenewald and Van

der Merwe worked closely together; all the illustrations

published with Groenewald’s papers are initialled by Van

der Merwe, who is acknowledged first as having ‘en-

couraged me in this work and greatly assisted’ in

Groenewald’s (1941) book on aloes, and who is often

credited with having found the plants Groenewald

described. The ± 2 500 specimens that Van der Merwe

collected are all housed in PRE, and not in a private her-

barium (Gunn & Codd 1981); 4, Gunn & Codd (1981)

assert positively that Van der Merwe gave Groenewald

‘several new species of Aloe ... for description’. This is

supported by Dr R.A. Dyer (pers. comm.), who supplied

much background information. The crux of this informa-

tion is that, for whatever reason, Van der Merwe, as a

medical inspector of schools, did not want to be seen to

be describing new species of Aloe. He therefore passed

his specimens on to Groenewald, who published the new

species under his own name. Van der Merwe did, how-

ever, after his retirement describe a number of new species

in the bulbous genera Scilla L. and Resnova Van der

Merwe. All his Scilla species have since been transferred

to Ledebouria Roth and Resnova is currently included in

the synonymy of Drimiopsis Lindl. (Reid 1993).

Among Reynolds’s notes used in the preparation of his

book on South African aloes (Reynolds 1950) is the state-

ment ‘A. nubigena v.d. Merwe or Groenewald m/s’, in-

dicating that these two worked so closely together that

there is some doubt as to who contributed which part to

any publication signed by either of them. It seems from

correspondence preserved in these notes that Van der

Merwe was on good terms with both Reynolds and

Groenewald. A letter in Van der Merwe’s handwriting

records the gift of some plants of ‘A. labiaflava Groene-

wald’ (= A. greatheadii Schonland var. davyana (Schon-

land) Glen & D.S. Hardy) to Reynolds shortly after the

name was published by Groenewald. In his notes on A.

branddraaiensis, Reynolds records ‘Mr Groenewald has

this species in hand, these notes are merely for my own

records, and from the plant Dr Van der Merwe gave me’.

98

Bothalia 25,1 (1995)

A note under A. vryheidensis indicates that the flow of

information was not only from Van der Merwe to

Reynolds: ‘I gave Dr Van der Merwe a complete set of

photographs, and informed him that there were thousands

of the species growing This species was published in

April 1937, but Reynolds’s first note on it is dated July

1936. However, it is noteworthy that Reynolds (1950: 67)

did not separately discuss Groenewald’s contribution to

alooid taxonomy in the historical section of his book, but

mentioned him under ‘1935-1940. Dr F.Z. van der

Merwe’ only.

There is no evidence of any direct correspondence be-

tween Reynolds and Groenewald in this archival album.

As there are very few complete letters in it, this does not

preclude their having been such a correspondence. The

only mention of any specimens is in a note on A.

brcmddraaiensis , where Reynolds records having made a

specimen from his plant for PRE, with permission from

Groenewald and Van der Merwe. Although some Van der

Merwe registers are housed at PRE, all of these deal ex-

clusively with Scilla , and none contains an entry with a

number as low as those discussed below. Tolken (1971)

gives no indication of the whereabouts of any register of

either Groenewald or EZ. van der Merwe.

In our opinion it is therefore reasonable to accept that

Groenewald never had a private herbarium and that the

original material of the Aloe species which he described

is deposited in PRE. The F.Z. van der Merwe collections

in PRE bearing the same collection number as that cited

in the protologue are considered to be holotypes, if only

one number was cited. In cases where more than one

specimen is quoted in the protologue or where the F.Z.

van der Merwe specimen from the type locality does not

bear a collecting number, lectotypes are chosen. Neotypes

were selected where no types are cited in the protologue.

Groenewald described new species of Aloe in one of

two journals only. The names affected are listed alphabeti-

cally by journal title:

Tydskrif vir Wetenskap en Kuns

A. decurvidens Groenew. (1937a): specimen cited

‘Herbarium No. 107’. A Van der Merwe 107 specimen

could not be found in PRE. Full citation: Neotype:

Transvaal, Sabie River, Skukuza, F.Z. van der Merwe s.n.

in PRE 24092 (PRE!); here designated. The specimen oc-

cupies two sheets; both are needed to confirm the identity

of this plant. This name is considered to be a synonym

of A. zebrina Baker.

A. dolomitica Groenew. (1938a): specimen cited ‘Her-

barium No. 235’. Full citation: Holotype: Transvaal,

Strydpoort Mountains, F.Z. van der Merwe 235 (PRE!).

The specimen occupies two sheets; both are needed to

confirm the identity of this plant. This name is considered

to be a synonym of A. vryheidensis Groenew.

A. graciliflora Groenew. (1936a): specimen cited ‘Her-

barium No. 77’ . A Van der Merwe 77 specimen could not

be found in PRE. Full citation: Neotype: Transvaal,

Dullstroom, F.Z. van der Merwe s.n. in PRE 24089

(PRE!); here designated. The specimen occupies two

sheets, both of which are needed to confirm the identity

of this plant. This name is considered to be a synonym

of A. greatheadii var. davyana.

A. hlangapies Groenew. (1936b): specimens cited

‘Herbarium No. 102, 103’. Full citation: Lectotype:

Transvaal, Piet Retief Dist., Langgewacht, F.Z. van der

Merwe 102 (PRE!); here designated. This name is con-

sidered to be a synonym of A. ecklonis Salm-Dyck. The

specimen F.Z. van der Merwe 103 is also in PRE, and

both these Van der Merwe specimens are of equal quality.

No. 102 is marked ‘TIPE’ (Eng. TYPE) in F.Z. van der

Merwe’s handwriting, and 103 is marked ‘tweede tipe’

(Eng. second type — a meaningless designation) in the

same hand.

A. labiaflava Groenew. (1936c): no specimen cited.

Neotype: Transvaal, Gemsbokspruit, F.Z. van der Merwe

100 (PRE!); here designated. This sheet is marked ‘TIPE’

(Eng. TYPE) in F.Z. van der Merwe’s handwriting. This

name is considered to be a synonym of A. greatheadii

var. davyana.

A. lusitanica Groenew. (1937b): specimen cited ‘Her-

barium No. 79’. Full citation: Neotype: Mozambique,

Maputo, (Lourenyo Marques), F.Z. van der Merwe s.n. in

PRE 24087 (PRE!). The specimen occupies two sheets,

both of which are needed to identify the plant. This name

has long been shown to be synonymous with A. komatien-

sis Reynolds (1950), which is itself considered to be a

synonym of A. parvibracteata Schonland.

A. nubigena Groenew. (1936d): specimen cited ‘Her-

barium No. 133’. Full citation: Holotype: Transvaal, Gras-

kop, F.Z. van der Merwe 133 (PRE!). This is a good

species.

A. recurvifolia Groenew. (1935): no specimen cited.

Full citation: Neotype: Transvaal, Kappiekoppie near

Graskop, F.Z. van der Merwe 73 (PRE!). Designated by

Reynolds (1950). Van Druten (1956) showed conclusively

that this name is a synonym of A. alooides (Bolus) Druten.

In the protologue Groenewald’s initials are incorrectly

given as ‘B.A.’

A. thompsoniae Groenew. (1936e): no specimen cited,

but this is a good species. Dealt with elsewhere (Glen &

Smith 1995).

A. vryheidensis Groenew. (1937c): specimen cited

‘Herbarium No. 266’. Full citation: Holotype: Natal,

Heine near Vryheid, F.Z. van der Merwe 266 (PRE!). This

is a good species.

The Flowering Plants of South Africa

It is noteworthy that no problems arose as far as

specimen citation is concerned in the papers of Groene-

wald that were published in this botanical journal, which

had a botanist as scientific editor (Dr I.B. Pole Evans for

Vol. 18 and Dr E.P. Phillips for Vol. 20). Types were clear-

ly stated and are cited below as they were included in the

protologues. The following names are therefore listed here

simply for completeness.

Bothalia 25,1 (1995)

99

Aloe angustifolia Groenew.: in The Flowering Plants

of South Africa 18: t. 708 (1938c). Holotype: Transvaal,

Leydsdorp Dist., near Gravelotte, April 1936, F.Z. van der

Merwe in National Herbarium, Pretoria, No. 21,288

(PRE!). This name is invalidated by the earlier use of it

by Haworth (1819: 47) and Salm-Dyck (1849). A. angus-

tifolia Haw. is a synonym of A. africana Mill., and the

same combination as used by Salm-Dyck probably

belongs under Gasteria disticha (L.) Haw. (Steam 1938;

Van Jaarsveld 1992). This species has consequently been

renamed A. vandermerwei, in honour of Dr F.Z. van der

Merwe (Reynolds 1950). A critical taxonomic treatment

of Aloe will show that the species should be included in

the synonymy of A. zebrina.

Aloe branddraaiensis Groenew.: in The Flowering

Plants of South Africa 20: t. 761 (1940). Holotype: Lyden-

burg Dist.; Branddraai, Van der Merwe (probably F.Z. van

der Merwe) in National Herb. 24208 (typus) (PRE!). The

specimen occupies three sheets, all of which are needed

to confirm the identity of this plant. In addition to the

type two further PRE specimens, Van der Merwe 138 and

Reynolds 2490 (!) are cited in the protologue. No

specimen with the number Van der Meme 138 was found

at PRE; however there are three unnumbered specimens

collected by F.Z. van der Merwe in this collection, PRE

22998, 38002, 38003. The specimen PRE 38002 is

marked ‘figured for FPSA by B. Connell July ’39'. This

is a good species.

Aloe davyana Schonland var. subolifera Groenew.: in

The Flowering Plants of South Africa 19: 732 (1939).

Holotype: Pretoria Dist., near Pienaars River, Aug. 1936,

Van der Merwe (probably F.Z. van der Merwe) in Nat.

Herb. 22816 (PRE!). This name is considered to be a

synonym of A. greatheadii var. davyana.

Aloe lutescens Groenew.: in The Flowering Plants of

South Africa 18: t. 707 (1938b). Holotype: Zoutpansberg

Dist.; Chipese, June 1937, Van der Merwe (probably F.Z.

van der Merwe) in National Herbarium, Pretoria, No.

23,301 (= PRE 23005) (PRE!). The type is F.Z. van der

Menve 1377 , and not F.Z. van der Merwe s.n. The

specimen occupies two sheets, both of which are needed

to confirm the identity of this plant. This is a good species.

Leptaloe blyderivierensis Groenew.: in The Flowering

Plants of South Africa 17: t. 651 (1937). Holotype:

Pilgrim's Rest Dist.; on slopes of the Drakensbergen near

the Blyde River, about 30 miles north of Pilgrim's Rest,

Van der Merwe (probably F.Z. van der Merwe) in National

Herbarium, Pretoria , No. 21361 (PRE!). Reynolds (1950)

considered this species to be a synonym of A. minima

Baker, an opinion with which we agree.

ACKNOWLEDGEMENTS

Drs L.E. Codd, G.E. Gibbs Russell and O.A. Leistner

are thanked for discussion and advice on the points raised

in this note. We thank Mrs E. van Hoepen and Prof. H.J.T.

Venter (University of the Orange Free State, Bloemfon-

tein) for help in searching for Groenewald specimens.

REFERENCES

GLEN, H.F. & SMITH, G.F. 1995. Notes on the typification of some

species of Aloe L. Bothalia 25: 37-42.

GREUTER, W. et al. 1988. International Code of Botanical Nomencla-

ture. Regnum Vegetabile 1 1 8.

GROENEWALD, B.H. 1935. 'n Nuwe Aloe -won van Oos-Transvaal [A.

recurvifolia Groenew.]. Tyclskrif vir Wetenskap en Kuns 14: 39-

42.

GROENEWALD, B.H. 1936a. Beskrywing van ’n nuwe makulaat-

aloesoort van Oos-Transvaal [ Aloe graciliflora Groenew.].

Tydskrif vir Wetenskap en Kuns 14: 1 37-1 39.

GROENEWALD, B.H. 1936b. ’n Nuwe aalwyn van Piet Retief [Aloe

hlangapies Groenew.]. Tydskrif vir Wetenskap en Kuns 14: 60-63.

GROENEWALD, B.H. 1936c. ‘n Nuwe Aloe- soort van Pretoria distrik

[A. labiaflava Groenew.]. Tydskrif vir Wetenskap en Kuns 14:

57-59.

GROENEWALD, B.H. 1936d. Beskrywing van ‘n nuwe lepto-aloe van

die Drakensberge [Aloe nubigena Groenew.]. Tydskrif vir

Wetenskap en Kuns 14: 135-137.

GROENEWALD, B.H. 1936e. 'n Nuwe aalwyn van die Wolkberg,

Transvaal [Aloe thompsoniae Groenew.]. Tydskrif vir Wetenskap

en Kuns 14: 64—66.

GROENEWALD. B.H. 1937a. ‘n Nuwe Aloe van die Laeveld [A. decur-

videns Groenew.]. Tydskrif vir Wetenskap en Kuns 15: 126-128.

GROENEWALD, B.H. 1937b. 'n Nuwe Aloe uit Louren$o Marques [A.

lusitanica Groenew.]. Tydskrif vir Wetenskap en Kuns 16: 13-15.

GROENEWALD. B.H. 1937c. ’n Nuwe aalwyn van Natal [Aloe

vryheidensis Groenew.]. Tydskrif vir Wetenskap en Kuns 15: 129—

131.

GROENEWALD. B.H. 1938a. Beskrywing van 'n nuwe Aloe uit Noord-

Transvaal [A. dolomitica Groenew.]. Tydskrif vir Wetenskap en

Kuns 16: 179-181.

GROENEWALD, B.H. 1938b. Aloe lutescens. The Flowering Plants of

South Africa 18: t. 707.

GROENEWALD. B.H. 1938c. Aloe angustifolia. The Flowering Plants

of South Africa 1 8: t. 708.

GROENEWALD, B.H. 1939. Aloe davyana var. sobolifera. The Flower-

ing Plants of South Africa 19: t. 732.

GROENEWALD, B.H. 1940. Aloe branddraaiensis. The Flowering

Plants of South Africa 20: t. 761.

GROENEWALD, B.H. 1941. Die aalwyne van Suid-Afrika, Suidwes-

Afrika, Portugees Oos-Afrika, Swaziland, Basoetoeland, en 'n

spesiale ondersoek na die klassifikasie, chromosome en areale

van die Aloe Maculatae. Nasionale Pers Beperk, Bloemfontein.

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

Africa. Balkema, Cape Town.

HAWORTH, A.H. 1819. Supplementum plaritarum succulentarum. Hard-

ing, London.

MULLER. F.S. 1945. ‘n Chromosoomstudie van 'n aantal spesies van die

genus Aloe Linn. Publikasies van die Universiteit van Pretoria.

Reeks II: Natuurwetenskap No. 8, Pretoria.

REID, C. 1993. Hyacinthaceae (Part B). In T.H. Arnold & B.C. de Wet

(eds), Plants of southern Africa: names and distribution. Memoirs

of the Botanical Survey of South Africa No. 62: 146-155.

REYNOLDS. G.W. 1950. The aloes of South Africa. Aloes of South

Africa Book Fund, Johannesburg.

SALM-DYCK, J.F.M.A.H.I. 1849. Monographic generum Aloes et

Mesembryanthemi, Aloe angustifolia. Fasc. 5, fig. 17 (Sect. 29,

fig. 30). Bonn.

STEARN, W.T. 1938. An annotated Index to Salm-Dyck’s ‘Monographia

Generum Aloes et Mesembryanthemi’. Cactus Journal (Great

Britain) 7: 34-44.

TOLKEN, H.R. 1971. Index herbariorum austro-africanorum. South

African Association of Botanists, Cape Town.

VAN DRUTEN, D. 1956. Aloe alooides (Bolus) Druten comb. nov.

(Liliaceae) Bothalia 6: 544, 545.

VAN JAARSVELD. E.J. 1992. The genus Gasteria, a synoptic review.

Aloe 29: 5-32.

H.F. GLEN*, G.F. SMITH* and D.S. HARDY**

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

** P.O. Box 16201, Pretoria.

MS. received: 1994-03-24.

100

Bothalia 25,1 (1995)

VERBENACEAE

A NEW SPECIES IN THE GENUS CLERODENDRUM

E Clerodendrum louwalbertsii P.P.J. Herman , sp.

nov., C. triphyllo ( Harv. ) H. Pearson affinis sed inflores-

centia thyrsoidea indeterminata, foliis linearibus vel an-

guste ellipticis differt.

TYPE. — Northern Transvaal, 2428 (Nylstroom): 18 km

from Nylstroom on road to Warmbaths near Groot Nyl

turnoff, (-CD), 1985-11-04, Germishuizen 3343 (PRE,

holo.).

Perennial herb with annual stems arising from woody

rootstock, stems unbranched or sparsely branched near

base, 200-5001-800) mm tall, angular, glabrous or pilose.

Leaves temate, sometimes opposite or in whorls of 4, ses-

sile or subsessile, glabrous or pilose, linear to narrowly

elliptic, ( 20—) 25—5 0(— 8 5 ) x (2—) 3—7 (— 15) mm, decreasing

in size upwards, apex acute, base cuneate, margin entire,

semi-succulent, glandular-punctate. Inflorescence indeter-

minate, thyrsoid, bracts similar to leaves but smaller,

hypopodium (peduncles) (5—) 10— 30(— 45) mm long,

mesopodium 5-16 mm long, epipodium (pedicels) 1 .5-4.5

mm long, purplish tinged; prophylls (bracteoles) 2— 8(— 12)

mm long, linear.*

Flowers zygomorphic. Calyx tubular, 5-toothed, tube

2.5-3.0 mm long, teeth 1-3 mm long, purplish tinged,

persistent. Corolla tubular in lower part, with 5 lobes, tube

4 — 6(— 8) mm long, curved; 4 lobes lateral, 5-7 x 3-5 mm,

purplish, fifth lobe forming lower lip, 7-10 mm long,

darker purple marked with white inverted Y-shaped pat-

tern. Stamens 4, purple, long exserted, opposite lower lip

and curved downwards, attached ± 1 mm below mouth

of corolla tube, bases, where attached to corolla tube,

covered with white downwardly directed hairs; filaments

14—21 mm long, anthers 1-2 mm long. Style long ex-

serted, purple, 22-27 mm long, curved, stigma of 2 une-

qual lobes, upper lobe shortest, ± 0.5 mm, lower lobe ±

2 mm long. Ovary 4-lobed in upper part, papillose, 2 mm

long, imperfectly 4-locular, with a single ovule in each

loculus. Fruit 1-4 drupes, ripe fruit not seen. Figure 1.

Clerodendrum louwalbertsii has been included under

C. triphymm since Pearson’s treatment of the genus in

Flora capensis 5,1 (1912). Some voucher specimens

quoted under C. triphyllum belong to C. louwalbertsii,

e.g. Zeyher 1362 (PRE, TCD) and Burke 365 (K). The

thyrsoid inflorescence and linear leaves clearly distinguish

C. louwalbertsii from C. triphyllum which has axillary

cymes in the axils of the lower leaves and leaves which

range from oblanceolate to obovate to elliptic (Figure 2).

C. louwalbertsii occurs in Northern Transvaal, the east-

ern part of the North-West, Gauteng (PWV), Eastern

Transvaal and Swaziland, and one specimen was collected

at the northern border of Kwa Zulu-Natal (Figure 3). It

is found in valleys, kloofs, mountains and hillsides, kop-

* Terminology of inflorescence according to Weberling (1989).

pies, grassland and open woodland on sandy, stony or

rocky soil, dolomite or granite.

The specific epithet ‘ louwalbertsii ’ was chosen in

honour of Dr Louw Alberts, the well-known South African

scientist and Christian.

To enable other herbarium workers to distinguish be-

tween the two species, the specimens of Clerodendrum

triphyllum examined are listed below. The distribution of

the two species is shown in Figure 3.

2. Clerodendrum triphyllum (Harv.) H. Pearson,

in Flora capensis 5,1: 220, 221 (1901); Thomas: 80

(1936).

Cyclonema triphyllum Harv.: 17, 18, t. 27 (1859).

leones: Harv.: t. 27 (1859); Phillips: t. 19 (1921).

C. triphyllum occurs in Northern Transvaal, the eastern

part of the North-West, Gauteng (PWV), Eastern Trans-

vaal, Orange Free State, Swaziland, Kwa Zulu-Natal, Le-

sotho and Eastern Cape. It is a typical grassland species.

Specimens examined

Acocks 1918 (2) PRE. Anderson A41 (2) PRE.

Balkwill & Cadman 3546 ( 1 ) J. Balkwill, Cadman & McCollum 3004

(1) J. Barnard & Mogg 913 (1) PRE. Barrett 306 (1) PRE. Beeton 1!4

(2) PRE. Behr 746 (2) PRE. Bezuidenliout 106 (2) PRE. Bolus 12236

(1) GRA, PRE. Botha & Ubbink 1750 (2) PRE. Botha & Van Wvk 1069

(2) PRE. Braun 1260 (2) PRE. Bredenkamp 141 (2) PRE; 236 (2) PRE.

Breyer 17898 (1) PRE. Brown & Shapiro 209 (2) PRE; 504 (1) PRE.

Buitendag 426 (1) PRE. Burgoyne 1067 (2) PRE. Burke 365 (1) K. Bunt

Davy 2307, 7098. 7290 (2) PRE.

Clarke 1434 (2) PRE. Coetzee 227, 343. 533 (2) PRE. Collins 12188

(2) PRE; 27120 (1) PRE. Comins 886 (1 ) PRE. Compton 27098 (2) PRE;

31232 (1) PRE. Culverwell 399 (2) PRE; 1423 (1) PRE.

Davidse 5981 (2) PRE. Davidson 2145 (1) J. De Souza 638 (2) PRE.

Dieterlen 515 (2) PRE. Dlamini s.n. PRE31357 (2) PRE. Dohse 45 (1)

NU. Du Plessis 1012, 1083 (1) PRE. Du Toit 39, 90 (2) PRE.

Edwards 1089 (2) PRE.

Flanagan 1931 (2) PRE. Fourie 1/2/30 (2) PRE.

Galpin 509M, 9060 (1) PRE; 12364, 14568 (2) PRE. Germishuizen

3343 (1) PRE; 3964 (2) PRE. Germishuizen & Retief 578 (1) PRE. Ger-

rard & M’Ken 1251 (2) TCD. Gilfillan 76 (Herb. Galpin 6166) (2) PRE.

Glen 1840 (1) J; 2479 (1) J, PRE.

Hanekom 1723 (1) PRE. Hards 104 (1) PRE. HBG 26891 (2) PRE;

27051 (2) PRE. Herman 151, 1357-1370 (1) PRE; 1332. 1375-1388 (2)

PRE. Holden 14156 (2) PRE. Holt 139 ( 1) NH, PRE. Huntley 1124 (2)

PRE.

lmmelman 83 (1) PRE.

Jacobsen 1863 (2) PRE; 2780 (1) PRE. Jacobsz 145. 1853, 2002 (2)

PRE. Janse 3166 (1) PRE. Jenkins 6929, PRE 39926 (1) PRE; 9829 (2)

PRE. Joffe 139 (2) PRE. Junod 17449 (2) PRE.

Kok 284 (2) PRE. Kroon 10264 (2) PRE.

Lambinon & Reekmans 82/28 (1) PRE. Lang, Tvl. Mus. 30314 (2)

PRE; Tvl. Mus. 31067 (1) PRE. Leendertz 376a (1) GRA. Le Roux 14

(2) PRE. Liebenberg 2507, 8838, 8943a (1) PRE; 8518 (2) PRE. L'Ons

60/44 (2) PRE. Louw 101 (2) PRE. Lucas 374 (1) J.

Maguire 1264, 1265, 8624 (1) J. Marloth 3823 (1) PRE; 3855 (2)

PRE. Mauve s.n. (1) NU. McCollum 561 (2) PRE. McMuriry 5235 (1)

J. Medley Wood 3381 (2) NH. Miller 3878, 8511 (2) PRE. Mitchley 86

(2) PRE. Mogg 10127, 14656, 15,336, s.n. PRE 39943 (2) PRE; 31555

Bothalia 25. 1 ( 1995)

101

FIGURE 1. — Holotype of Cleroden-

drum louwalbertsii , Germis-

huizen 3343.

(1) J, PRE. Moss 8637 , 10826. 16223, 16764. 17820 (1) .1 Moss &

Oetley s.n. (1) J. Muller 2014 (2) PRE. Murray 11 (2) PRE.

Nation 35 (1) PRE. Nel 290 (1 ) PRE. Netshiungani 665 (2) PRE.

Obermeyer 895 (2) PRE.

Phillips 18, 3113, 3276 (2) PRE. Pienaar 748 { 1) PRE. Pole Evans

H13132 (2) PRE. Potgieter 21837 (2) PRE. Prosser P1062 (2) PRE.

Pupils of Convent 97 (2) PRE.

P. Raal & G. Raal 911 (2) PRE. Radetnacher 7306, 7478 (2) PRE.

Reid 436 (1) PRE. Repton 160 (1) PRE; 422, 3431 (2) PRE; 754 (1)

NH, PRE. Relief 134 (1) PRE. Rogers 2930, 22062 (1) PRE; 12412,

19026. 21580 (2) PRE.

Scheepers 694, 1434 (2) PRE. Schijff 471 (2) PRE. Sekhaolelo 156

(1 ) PRE. Sides 1944 (2) PRE. Smith 1267, 1278 (1) PRE. Smuts & Gillett

3055, 3326, 4129 (1) PRE. Stirton 5739 (2) PRE. Story 1404 (2) PRE.

Sutton 351 (2) PRE.

Theron 705, 1405(1 ) PRE. Tltode A 1338, A 1631 (2) PRE. Thompson

8011 (2) PRE. Thorncroft 11291 (2) PRE. Tiedt & Young 2159a (1) PRE.

Turner 222 (2) PRE.

Ubbink & Van Wyk 514 (2) PRE.

Van Dam 20755, 27099. 27121 (1) PRE. Van der Merwe s.n.

PRE39945 (2) PRE. Van Rooyen 2027 ( 1) PRE. Van Vuuren 369 (1)

PRE. Van W\k 6875 (2) PRE. Venter 7202 (1) PRE. Verdoom 141 ( 1)

PRE; 496 (2) PRE.

Walker s.n. PRE39923 (2) PRE. Wassetfall & Van Niekerk 41 (1)

PRE. Werdennann & Oberdieck 1279 (1) PRE.

Young A202 (1) PRE.

Zeyher 1362 (1) PRE, TCD; 1363 (2) TCD (type of Cyclonema

triphyllum Harv.).

102

Bothalia 25,1 (1995)

FIGURE 2. — Diagram showing the differences in inflorescences of A,

Clerodendrum louwalbertsii and B. C. triphyllum.

ACKNOWLEDGEMENTS

FIGURE 3. — Distribution of Clerodendrum louwalbertsii , □; and C.

triphyllum, ▲.

the loan of material; and to the former Department of

Nature Conservation of Bophuthatswana for permission

to collect material in the Pilanesberg National Park.

REFERENCES

HARVEY, W.H. 1859. Cyclonema triphylla (Verbenaceae). Thesaurus

Capensis I: 17, 18.

PEARSON, H.H.W. 1912. Verbenaceae. In W.T. Thiselton-Dyer, Flora

capensis 5,1: 218-225. Reeve, London.

PHILLIPS, E.P. 1921. Clerodendron triphyllum. The Flowering Plants of

South Africa 1: t. 19.

THOMAS, B. 1936. Die Gattung Clerodendrum in Afrika. Botanische

Jahrbiicher 68: 1-106.

WEBERLING, F. 1989. Morphology of flowers and inflorescences.

Cambridge University Press, Great Britain.

P.P.J. HERMAN

My thanks go to Dr H.F. Glen for the Latin translation;

Mrs A. Romanowski for the photograph; Ms G. Condy

for the line drawings; to the curators of K and TCD for MS. received: 1993-06-08.

GERANIACEAE

LECTOTYPIFICATION OF PELARGONIUM MULTIBRACTEATUM

In the description of Pelargonium multibracteatum in

Richard’s Tentamen florae abyssinicae 1: 119 (1847), two

Schimper collections are referred to in the heading of the

protologue and a locality for a Schimper collection is

given as near Axum in the list of material seen. A further

three collections from various localities made by Dillon

are cited in the list of material seen. These collections can

all be considered to be syntypes.

Up to now no lectotype has been designated for Pelar-

gonium multibracteatum in accordance with Art. 9.2 of

the ICBN (Greuter et al. 1994). In all Kokwaro’s papers

on the Geraniaceae of northeast Africa (1969, 1971a,

1971b) in which he treated this species, only the syntypes

were cited without selecting a lectotype, possibly because

he considered P. multibracteatum to be a subspecies of P.

alchemilloides (L.) L’ Herit. and therefore thought the

selection of a lectotype of minor importance. Van der Walt

& Vorster (1988: 91) cited all the syntypes seen by them,

again declining to select a lectotype. However, for the

treatment of Pelargonium for the Flora of Ethiopia by the

senior author, it has become necessary to select a lec-

totype.

The first question to be addressed is: who is the validat-

ing author of the name? Knuth (1912: 433), Dyer (1940:

t. 794), and Richard (1847: 119) cited Hochstetter as the

author; Engler (1895: 225), Cufodontis (1956: 351), and

Kokwaro (1969: 530; 1971a: 666) cited Hochstetter ex A.

Richard; Kokwaro (1971b: 21) cited A. Richard; and Van

Bothalia 25, 1 (1995)

103

FIGURE 4. — Pelargonium multibrac-

teatum: lectotype sheet (Dillon s.n.

in P). Scale bar: A, 100 mm: B. 10

mm.

der Walt & Vorster (1988: 91) cited Hochstetter in A.

Richard. According to Friis (1993: 185) practically none

of Hochstetter’s new names were accompanied by

descriptions. The name Pelargonium multibracteatum first

appeared in print on a herbarium label for Schimper 51,

distributed by Hochstetter on 26 October 1837; and again

on a label for Schimper 1489, distributed on 23 November

1842. It also appeared in Hochstetter’s (1841) list of

Schimper collections. None of these publications satisfies

the conditions for valid publication (ICBN Art. 32;

Greuter et al. 1994), and the first description of the species

was only published in 1847, by Richard. There is no

evidence that Hochstetter contributed towards this descrip-

tion, and therefore Richard is the sole validating author.

The second question is: which specimens did Richard

use to compile his description? Following the description,

four collections are cited, namely, Dillon 316 (inter Adoua

et Ocbasa), Dillon 205 (Taccaze), Dillon s.n. (Chire), and

Schimper 1489 (Axoum). However, as authority for the

name, Hochstetter’s label for Schimper 57 is also cited.

We here accept that Richard did indeed see Schimper 51.

It is unlikely that he saw a detached label only, and we

surmise that he would have been unable to link the name

with a species unless he saw an actual specimen.

According to Stafleu & Cowan (1983: 764), the

original set of specimens on which Richard’s Tentamen is

based, is lodged in R They also state that the Tentamen

104

Bothalia 25,1 ( 1995)

is based on material collected by R. Quartin Dillon and

Antoine Petit, but P. multibracteatum is clearly also based

on Schimper 51 and 1489. In P there are indeed specimens

of all the collections cited by Richard.

For lectotypification purposes, the three Dillon collec-

tions are not particularly suitable, being relatively poor

specimens with few duplicates in other herbaria. Of

Schimper 51 there are six sheets in P with numerous dupli-

cates in other herbaria, and of Schimper 1489 there are

two sheets in P with several duplicates in other herbaria.

However, the only really good sheets of the Schimper

collections bear stamps ‘Herb. A. Bunge’, ‘Herbier E.

Drake’, and ‘Ancien Herbier E. Cosson’. Strictly these

are not part of the original material and were probably

not seen by Richard, being incorporated into P in 1904

and 1913 (Stafleu & Cowan (1976: 407, 552, 678)). Of

the remainder of the sheets in P, the best is Dillon s.n.

which, although not well annotated, is presumed to be

from Chire. In the absence of contradictory evidence, this

sheet is accepted as being one of those studied by Richard,

and is here designated as lectotype (Figure 4):

ETHIOPIA. — ‘in provinciae Chire’, Dillon s.n. [P,

lecto., here selected; MEL (sub MEL 94226), P, W, iso-

lecto.l.

Other original material seen: ETHIOPIA. — ‘in con-

valle fluvii Taccaze’, Dillon 205 [P, MEL (sub MEL

94227)]; ‘Crescit inter Adoua et Ocbasa’, Dillon 316 (P);

‘montis Scholoda’, Schimper 52 [BM (2 sheets), CGE, G,

K (2 sheets), L (3 sheets), OXF (2 sheets), P (6 sheets),

S (2 sheets), and W (4 sheets)]; ‘prope Axoum’, Schimper

1489 [BM, CGE, G (3 sheets), K, L, MO, P (2 sheets),

TUB, W],

REFERENCES

CUFODONTIS, G. 1956. Enumeratio plantarum aethiopiae sper-

matophyta (Sequentia). Bulletin du Jardin botanique de L’Etat,

Bruxelles 26, Supplement: 350, 351.

DYER, R.A. 1940. Pelargonium multibracteatum. The Flowering Plants

of South Africa 20: t. 794.

ENGLER, A. 1895. Die Pflanzenwelt Ost-Afrikas und der Nachbar-

gebiete. Reimer, Berlin.

FRIIS, I. 1993. C.F. Hochstetter's scientific names for G.W. Schimper’s

early collections of African plants. Fragmenta Floristica et

Geobotanica, Supplementum 2, part 1: 183-201.

GREUTER. W. et al. 1 994. International code of botanical nomenclature.

Koeltz Scientific Books, Konigstein.

HOCHSTETTER, C.F. 1841. Erste Lieferung der vom Reiseverein aus-

gegebenen durch Wilhelm Schimper gesammelten Pflanzen.

Flora (Jena) 24, Intelligenzblatt 1: 17-32.

KNUTH, R. 1912. Pelargonium. Das Pflanzenreich 4, 129, 53: 316-545.

KOKWARO, J O. 1969. Notes on east African Geraniaceae. Kew Bulletin

23: 527-530.

KOKWARO, J.O. 1971a. The family Geraniaceae in northeast tropical

Africa. Webbia 25: 623-669.

KOKWARO, J.O. 1971b. Pelargonium. Flora of tropical East Africa,

Geraniaceae: 14—23. Crown Agents, London.

RICHARD, A. 1847. Tentamen florae abyssinicae. Bertrand, Paris.

STAFLEU, F.A. & COWAN, R.S. 1976. Taxonomic literature edn 2, 1:

407, 552, 678. Bohn, Scheltema, & Holkema, Utrecht.

STAFLEU, F.A. & COWAN, R.S. 1983. Taxonomic literature edn 2, 4:

764. Bohn, Scheltema, & Holkema, Utrecht.

VAN DER WALT, J.J.A. & VORSTER, P.J. 1988. Pelargoniums of

southern Africa 3: 90-92. National Botanic Gardens of South

Africa, Cape Town.

P. VORSTER* and lb. FRIIS**

Botany Department, University of Stellenbosch, Private Bag XI, 7602

Matieland.

Botanical Museum & Library, University of Copenhagen, Gothersgade

130, DK-1123 Copenhagen K, Denmark.

MS. received: 1994-06-03.

ROSACEAE

A NEW SPECIES OF CLIFFORTIA FROM THE SWARTBERG

Cliffortia nivenioides Fellingham, sp. nov., C.

aculeatae Weim. sectionis Multinerviae DC. affinis sed

foliis brevioribus, lateraliter complanatis, versus sulcatis

teretibus, structura interna dissimilibus.

TYPE. — Western Cape, 3322 (Oudtshoom): Eastern

Swartberg, plateau north of Blesberg, 1 900 m, 29 Novem-

ber 1991, Fellingham & Vlok 1588 (STE, holo.; K, PRE,

MO, iso.).

A compact small shrub to 600 mm tall with long

emerging procumbent branches spreading over 750 mm,

monoecious but apparently of one or the other sex at any

given time, female Dowers totally hidden in axils of upper

leaves. Branches reddish brown, glabrous, older parts with

persistent vaginas of fallen leaves with or without inter-

nodes in between, axils of some containing either short

shoots or long shoots; internodes about 1 2 mm long.

Leaves unifoliolate, unifacial, bilaterally flattened, in

transverse section elliptic and anatomically centric, closely

arranged on the short shoot in a flat slightly elongated

fan; vagina 4—5 mm long, dorsally whitish and with three

obscure parallel veins close together, laterally green;

stipules, 3^4 mm long, subulate, glabrous, green, proximal

edge straight, thick and flat, distal edge curved, mem-

branous and whitish tinged with purple; lamina sessile,

glabrous, glaucous green, red-tipped, smooth, drying sul-

cate, 3.0-33.0 x 1. 0-2.0 mm, linear, straight to slightly

falcate, base with two colourless cushions at joint to

vagina, dorsally with a central nerve the length of leaf

and two shorter parallel lateral nerves, ventrally from base

upwards channelled for a third to half of its length forming

a partial sheath to leaf above, apically tapering to an acute,

somewhat dorsiventrally Battened point. Male flowers:

bracteoles lanceolate-acuminate, 0.5-0.6 mm long, green,

glabrous; pedicel and receptacle 1.0 mm long, glabrous;

sepals 3, 7. 0-8.0 x 2. 0-2. 5 mm, linear-lanceolate,

leathery, the young lime green with occasional irregular

longitudinal maroon lines, turning pink then maroon and

eventually brown; stamens ( 1 3—) 1 6(— 20), filaments 6.5

mm long, filiform, maroon, glabrous; anthers 1.2 mm

long, pink to maroon. Female flowers: bracteoles 0.4-0.5

Bothalia 25,1 (1995)

105

mm long, similar to male bracteoles; pedicel absent or up mm, longitudinally faintly ridged, glabrous; style solitary,

to 0.4 mm long, thick and fleshy; sepals 3, 2.5-3.0 x 0.3-0.5 mm long; stigma short, broad, flat, maroon, mar-

1.0-1.2 mm, lanceolate, dorsally concave, persisting in gins long fimbriate. Fruit 4.5-5. 5 x 1. 9-2.0 mm, ellipsoid,

fruit; ovarx narrowly elliptic in outline. 2.0-2. 5 x 0.9- 1.0 apex truncate, base narrowed, resembling a pedicel, lon-

FIGURE 5. — A-H, Cliffortia nivenioides: A, B, fruit; C, male flower; D, female flower; E, long shoot with short shoots, three lower short shoots

showing initial lengthening; F, long shoot with female flowers (above) and developing fruit below; G, male bud; H, male flower in axil of

basal leaf of developing long shoot. I-K, Cliffortia aculeata. I, J, fruit; K, male bud. a, pedicel; b, bracteole; c, calyx lobe; d, basal part of

fruit; e, style. A— H, Viviers & Vlok470 (BM, BOL, K, MO, P, PRE, STE); I— K, Fellingham & Vlok 1590 (K, PRE, STE). Scale bars: 1 mm.

106

Bothalia 25,1 (1995)

FIGURE 6. — Known distribution of

Cliffortia nivenioides , ♦; and

C. aculeata, A.

gitudinally faintly ridged, green with a tinge of red (Figure

5A-H).

Diagnostic features

C. nivenioides is similar to C. aculeata of the section

Multiner\>iae DC., but differs in the more compact, upright

growth form (elongated and trailing in C. aculeata)', the

shorter, laterally flattened, wider leaves (terete and some-

what sulcate in C. aculeata)', the shorter wider calyx lobes;

the somewhat different fruit structure; and the different

habitat (Figure 5A-H versus 5I-K).

Distribution and habitat

C. nivenioides is known only from a very small area

on the plateau north of the Blesberg in the eastern

Swartberg, in grassy mountain lynbos on open, sunny,

marshy flats. C. aculeata occurs nearby, but on a shady

vertical cliff face in the spray of a waterfall. It also grows

on the Waboomsberg on the eastern side of the Swartberg

Pass, again in a shady, moist habitat on a north-facing

cliff face (Figure 6).

Discussion

Superficially C. nivenioides is very similar to C.

aculeata in its distribution, growth form, the colouring of

stems and leaves, and the arrangement of the leaves in

the form of fans. Closer examination, however, reveals

significant differences mainly in the leaf anatomy (Figure

7A & B). In C. nivenioides the leaf is centric with the

palisade continuous except where it is replaced by the

dorsal strand of colourless sclerenchymatous tissue sup-

porting the large middle vein with its double phloem ele-

ments and phloem cap, but lacking a xylem cap. In C.

aculeata the leaf is near centric with the diminished

adaxial surface in the form of a V-shaped sulcus lined

with epidermis lacking stomata and supported by a rec-

tangular strand of tissue comprising large, compactly ar-

ranged, thick-walled, colourless cells. At first this

colourless strand appeared to be a ‘window’ but it became

clear that it is similar to the abaxial ‘vein.’ No comparable

adaxial structure was found in C. nivenioides.

Apart from being shorter and wider, the leaves of C.

nivenioides are also bilaterally flattened throughout the

whole length and channelled, but that, near the base only.

The leaves of C. aculeata, by contrast, are longer, more

FIGURE 7. — Leaf anatomy. A, Cliffor-

tia nivenioides', B, C. aculeata.

a, colourless sclerenchymatous

tissue; b, phloem; c, xylem; d,

xylem cap; e, thick-walled

colourless cells; f, stomata; g,

extraxylary fibre cap; h,

palisade. Scale bar: 10 pm.

Bothalia 25,1 (1995)

107

slender, subterete, canaliculate throughout their length,

and aculeate.

In C. nivenioides the calyx lobes in both male and

female flowers are shorter and wider (note: the scale bar

for Figure 5G is twice the length of that for 5K) and the

fruit shorter and more tapered to the ends than in C.

aculeata (Figure 5B: structure ‘d’ versus Figure 51: struc-

ture 'd').

The habitat of C. nivenioides is sunny, open and Hat,

whereas that of C. aculeata is shady, sheltered and verti-

cal. One collection of C. aculeata ( Fellingham & Vlok

1593) however, was from a level, sunny and dry site, no

more than 4 m away from the vertical stream bank from

which a typical specimen (Fellingham & Vlok 1592) was

collected. The specimen from the sunny position had

shorter leaves which were arranged in more compact fans

than those normally found in this species, but rather like

the leaves of C. nivenioides. It was, in fact, difficult to

recognize the plant immediately as C. aculeata , and

momentary excitement was caused at the thought of this

being a possible new find of C. nivenioides. On closer

examination, however, there was no doubt that it was C.

aculeata.

C. nivenioides is known from only one very discreet

area north of the Blesberg, very close to its related species

C. aculeata. The latter species was originally discovered

on the Swartberg Pass and later on the Waboomsberg. To

date no specimens of C. nivenioides have been collected

from these two localities.

The fan-like arrangement of the leaves in C. niven-

ioides resembles the arrangement characteristic of the

genus Nivenia Vent. (Iridaceae), prompting the derivation

of the specific epithet.

Specimens examined

C. nivenioides sp. nov.

WESTERN CAPE. — 3322 (Oudtshoom): eastern Swartberg, plateau

north of Blesberg, next to track; in moist, broad but relatively dry seepage

area. 1 880 m, 30-04-1980, (-BC), Bond 1754 (PRE, STE); Swartberg,

head of Tierkloof, northwest of Blesberg, marshy flats, 1 900 m, 06-01-

1975, (-BC), Thompson 2275 (PRE, STE); Swartberg Mountains, next

to track near Blesberg, in wet humic sandy soil on north-facing slope,

1 900 m, 18-12-1985, (-BC), Vlok 1326 (PRE, STE); eastern Swartberg,

plateau north of Blesberg, between E-W running track and the track up

Blesberg, open mountain fynbos on peaty soil in marshy area on gentle

N slope, 1 900 m, 29-1 1-1991, (-BC), Fellingham & Vlok 1588 (K. MO.

PRE, STE); Swartberg Mountains, on plateau just north of Blesberg, next

to forestry track, in deep peaty sandy soil in seepage on gentle north-

facing slope in grassy mountain fynbos, 1 900 m, 30-01-1990, (-BC),

Viviers & Vlok 470 (BM, BOL, K, MO, P. PRE, S, STE).

C. aculeata Weim. in Botaniska Notiser4: 410 (1946).

Type: Swartberg Pass growing in wet places near stream,

00-12-1945, Stokoe 9250 (LD, holo.; NBG, iso. !).

WESTERN CAPE. — 3322 (Oudtshoom): upper northern slopes of

Waboomsberg, next to perennial stream in deep moist sand, 1 600 m,

26-12-1984, (-AC), Vlok 879 (PRE, STE); Swartberg, Waboomsberg,

about 15 km along road to Die Hel at foot of mountain, mountain fynbos

on edge of stream, in shade, few plants hanging down from vertical

streambank, 1 860 m, 30- 11-1 99 1 ,(— AC), Fellingham and Vlok 1592

(PRE, STE); Swartberg, Waboomsberg, about 15 km along road to Die

Hel, at foot of mountain, in mountain fynbos in dry streambed in full

sun, 1 860 m. 30-11-1991, (-AC), Fellingham & Vlok 1593 (PRE, STE);

Swartberg Pass, damp places, 00-12-1942, (-AC), Stokoe SAM 58304

(SAM); Swartberg Pass, 1 600 m, 00-01-1944, (-AC), Stokoe SAM 58303

(PRE, SAM); 8 miles west of top of Swartberg Pass, 1 600 m, 00-10-

1951,(-AC) Stokoe SAM 67049 (PRE, SAM); Swartberg Pass, 12-12-

1942, (-AC), Stokoe 8606 (PRE); Swartberg Mountains, mid northern

slopes of Blesberg, near forestry track, in deep moist loamy soil next to

perennial stream on north-facing slopes in grassy disturbed mountain

fynbos, 1 400 m, 1 5- 12- 1986,(-BC ), Vlok 7777 (STE); eastern Swartberg,

Blesberg area, top of Tierkloof, vertical north-facing seep, 29-11-1991,

(-BC), Fellingham & Vlok 1590 (K, PRE, STE).

ACKNOWLEDGEMENTS

I wish to thank the following persons for their assis-

tance: Mrs J.B.P. Beyers, Dr E.M. Marais, Mr E.G.H.

Oliver, Mrs I.M. Oliver, and Mr J.H.J. Vlok.

REFERENCES

WEIMARCK, H. 1946. Further notes on the genus Cliffortia. Botaniska

NotiserA'. 410.

A. C. FELLINGHAM*

*Stellenbosch Herbarium, National Botanical Institute, P.O. Box 471,

Stellenbosch 7599.

MS. received: 1994-04-18.

ASTERACEAE

A NEW SPECIES OF TR1CHOGYNE FROM NAMAQUALAND

Trichogyne lerouxiae Beyers , sp. nov. T. verticillatae

(L. f.) Less, in facie maxime similis sed floribus tribus

femineis in quoque capitulo, bracteis circum floribus her-

maphrodite rotundatis ad obtusis, pappo pauco vel nullo dif-

fert; et T. polycnemoidi (Fenzl) Anderb. affinis sed bracteis

involucralibus dissimiliter coloratis et numero florum

femineorum in quoque capitulo facile distinguenda.

TYPE. — Northern Cape, 3017 (Hondeklip Bay):

Namaqualand, 15 km west of Soebatsfontein on road to

Riethuis, (-AB), 14-08-1982, Le Roux 2918 (STE, holo.;

PRE, iso.). Figure 8.

Annual viscid erect herb, 40-150 mm tall. Stems few

to several from the crown, simple or branching near base,

I OS

Bothalia25,l (1995)

FIGURE 8. — The known distribution of Trichogyne lerouxiae.

leafy, with thin cobweb-like indumentum, becoming

glabrous. Leaves : primary leaves 6.5-10 x 0.5-1 mm,

linear-filiform, usually twisted, apex mucronulate, margins

involute, abaxial surface with thin cobweb-like indumen-

tum sometimes glabrescent towards apex, adaxial surface

white tomentose; secondary leaves similar but much

shorter. Heads campanulate, 2.2-2. 7 x 0.9-1. 5 mm, 3 to

few in small leafy clusters in axils of primary leaves,

which much exceed them. Involucral bracts 10-11, the

outer 2-3 sterile, elliptic to obovate, apex acute or

rounded; following 3 subtending female flowers, readily

deciduous, elliptic, obovate or trullate, adaxially concave,

abaxially keeled, apex acute or rounded; inner 5 surround-

ing hermaphrodite flowers, persistent, of which outer 2

trullate, adaxially concave, abaxially convex, apex

rounded and innermost 3 elliptic, ovate or obovate, flat-

fish, apex rounded or obtuse; all bracts translucent pale

golden brown, abaxial surface pilose but 3 innermost

bracts surrounding hermaphrodite flowers glabrescent.

Flowers 4—6; 3 female, corolla filiform, style divided, 2-

(3) hermaphrodite, corolla cylindric below, campanulate

above, style undivided. Achenes 0. 8-1.0 mm long,

obovoid with minute clavate hairs, ovaries of her-

maphrodite flowers aborted. Pappus absent in female

flowers, scanty or absent in hermaphrodite flowers,

slender setaceous with shaft naked below, shortly plumose

towards tip. Figures 9 & 10.

Anderberg (1991) has reinstated the genus Trichogyne

Less, which Hilliard (1981) had regarded as a subgenus

of Ifloga Cass. This new species has been placed in the

genus Trichogyne as its outer female florets are subtended

by bracts, its functionally male disc florets have undivided

styles, and its pappus bristles are apically plumose and

basally without patent cilia, thus corresponding to the

characters defined by Anderberg (1991). T. lerouxiae

resembles T. verticillata in general facies, but differs in

having 3 [not (0)— 1— 2] female flowers in a head, the

rounded to obtuse (not acuminate) bracts which surround

the hermaphrodite flowers and the scanty or absent (not

present) pappus. Its affinities also lie with T. polyc-

nemoides but it can be readily distinguished by the dif-

ferently coloured involucral bracts (translucent straw-

coloured versus reddish central patch with whitish tip) and

the number of female flowers (3 versus 5-10) in a head.

T. lerouxiae is only known from two localities in the

sandveld of Namaqualand growing in sandy clay soil at

altitudes ranging from 150-250 m. According to Hilliard

(1981), T. verticillata is confined to the southwestern Cape,

with its western distribution as far north as Lambert’s Bay

occurring on coastal dunes and low-lying sandy flats at al-

titudes not above 250 m. T. polycnemoides on the other hand

occurs in Namaqualand, and is confined to altitudes between

600 and 900 m.

Specimens examined

NORTHERN CAPE.— 3017 (Hondeklip Bay): Namaqualand, 15 km

W of Soebatsfontein on road to Riethuis, 150 m, (-AB), 14-08-1982, Le

Roux 2918 (PRE, STE); Namaqualand, Kookfontein 466, 250 m, (-BA),

03-09-1986, Le Roux & Lloyd 458 (STE).

!|l!!ijl!!!|lll!|!ll!jlli!|i

0 1 2

CM

FIGURE 9. — Trichogyne lerouxiae: holotype, illustrating habit, Le Roux

2918 (STE).

FIGURE 10. — Trichogyne lerouxiae. A-D, involucral bracts, abaxial view: A, sterile bract; B, bract subtending female flower; C & D, persistent

bracts surrounding hermaphrodite flowers. E, head with deciduous bracts removed to reveal two female flowers. F-H, hermaphrodite flower:

F. with pappus; G, stamens; H, style. All drawn from the type, Le Roux 2918 (STE). Scale bar: 1 mm.

It gives me great pleasure to name this new species

after Ms Annelise le Roux who has for many years been

involved with the study and conservation of the flora of

Namaqualand and is the co-editor of the forthcoming

Flora of Namaqualand for which this new species is re-

quired.

REFERENCES

ANDERBERG, A. A. 1991. Taxonomy and phylogeny of the tribe

Gnaphalieae (Asteraceae). Opera Botanica 104: 5-195.

HILLIARD, O.M. 1981. A revision of Ifloga in southern Africa, with

comments on the northern hemisphere species. Botanical Journal

of the Linnean Society 82: 293-312.

I B P. BEYERS*

* Stellenbosch Herbarium, National Botanical Institute, P.O. Box 471,

Stellenbosch 7599.

MS. received: 1994-05-13.

Bothalia 25, 1 : 111-119(1995)

Preliminary ethnobotanical studies of the Rwenzori Mountain forest area in

Bundibugyo District, Uganda

H. ORYEM-ORIGA*, E.K.Z. KAKUDIDI*, A.B. KATENDE* and Z.R. BUKENYA*

Keywords: Bakonjo, Baamba. Bundibugyo, ethnobotany, plant uses, inventory, Rwenzori Mountain, transects, Uganda

ABSTRACT

Ethnobotanical studies of the Rwenzori Mountain forest area in Bundibugyo District in Uganda were carried out between

May and December 1991, and covered the northern part of the Rwenzori Mountain slopes occupied by the Bakonjo people.

The presence of a major footpath through the forest with numerous utility trails radiating from it showed that some forest

resources are being sought by the local population. Plant biodiversity is high, as is indicated by the fact that in a study plot of

only 4 250 irf, a total of 1 15 plant species, 101 genera and 57 families were identified from a collection of 300 plant specimens.

Seventy-seven plant species were found to be of some importance to the local communities. Out of the 77 useful plant

species recorded: 22 species were used for medicinal purposes; 16 for firewood; 13 for construction, joinery and furniture; 12

for craftwork; 10 provided edible fruits and vegetables; and 27 were used for a variety of other purposes. These other purposes

include construction of shrines, covering of granary floors, use as toilet paper, carrying luggage, and fodder for goats, sheep and

cattle. Arundinaria alpina K. Schum. (bamboo) is the species that is most extensively harvested from the forest.

UITTREKSEL

Etnobotaniese studies van die woudgebied van die Rwenzoriberg in die Bundibugyo-distrik in Uganda is tussen Mei en

Desember 1991 ondemeem. Die noordelike deel van die hange van die Rwenzoriberg waar die Bakonjo-volk woon, is gedek.

’n Bree voetpad deur die woud met talle dienspaadjies wat daaruit lei, was ’n aanduiding dat bronne in die woud deur die

plaaslike bevolking benut word. Plantbiodiversiteit is hoog, soos blyk daaruit dat in 'n studieperseel van slegs 4 250 m ", ’n totaal

van 1 15 plantspesies, 101 genusse en 57 families in ’n versameling van 300 planteksemplare gei'dentifiseer is.

Sewe-en-sewentig spesies is deur die plaaslike gemeenskappe benut. Van die 77 nuttige plantspesies aangeteken, is 22 vir

geneeskundige doeleindes gebruik, 16 vir vuurmaakhout, 13 vir konstruksie, skrynwerk en meubels, 12 vir handwerk; 10 het

eetbare vrugte en groente opgelewer, en 27 is vir 'n verskeidenheid ander doeleindes gebruik. Voorbeelde hiervan is die

konstruksie van altare, bedekking van graanskuurvloere. benutting as toiletpapier, die dra van bagasie, en voer vir bokke, skape

en beeste. Arundinaria alpina K. Schum. (bamboes) is die spesie wat die meeste uit die woud verwyder word.

INTRODUCTION

In the conservation of forests our governments quite

often have taken no account of the non-timber products

important in national economies. This attitude has also

been observed in Latin America’s humid tropical forests

(Pinedo-Vasquez et al. 1990). Prance et al. (1987) clas-

sified these non-timber products into craft materials,

medicinals, seeds and fruits and other edibles, as well as

sources of latex, construction fibres and poles and many

other goods that cannot easily be categorized.

Studies of the vegetation of Uganda are available

(Snowden 1953; Langdale-Brown 1960; Lind & Tallantire

1975; Hamilton 1984; Howard 1991), but few eth-

nobotanical studies have been carried out. There is also a

general lack of public awareness in Uganda of the values

of wild plants and the need to use them sustainably. This

has led to careless and wanton destruction of many valu-

able plant species with unacceptable consequences

(Karani 1982; Hamilton 1984).

* Botany Department. Makerere University, P.O. Box 7062, Kampala,

Uganda.

MS. received: 1993-06-29.

Sociobotanical studies in Uganda have mainly ad-

dressed the relationship between humans and plants as

defined in terms of cultural use. The main aim has been

to explore human use of certain plants and how such

plants are identified with particular social situations. It is

in this identification that the social or cultural value of a

plant is seen to be institutionalized in people's culture.

For instance, some scholars have focused on the impor-

tance of the banana plant to the Baganda culture, millet

to some aspects of culture of the Iteso and the story of

the barkcloth in Buganda (Musoke 1975; Nyanzi-Makum-

bi 1976; L’Obwolo 1980).

With regard to traditional medicines, there are two dis-

tinct groups of people who deal with illnesses. These are

the ‘medicine man' or the ‘diviner’ who not only issues

medicines but is also socially sanctioned to explain the

cause(s) of illnesses. The other group is made up of her-

balists who literally deal exclusively with herbs and herbal

medicines for specific illnesses but not with their supposed

underlying supernatural causes. Ogwal & Kakudidi (1989

unpublished) have made some preliminary collections of

medicinal plants which have been deposited in the Her-

barium of the Department of Botany, Makerere University.

112

Bothalia 25,1 ( 1995)

FIGURE 1. — Map of Uganda with

study site of the Rwenzori

Mountain forest area in Bun-

dibugyo District, forest

reserve boundary; xxx, Bupom-

poli Ridge (25-9-91).

In Uganda, Anokbongo (1972) undertook a general

pharmacological experimental study of nine local

medicinal plants with a view to confirming or disproving

their pharmacological and medicinal potency. Kokwaro

(1976) in his book entitled Medicinal plants of East Africa

briefly discussed a number of medicinal plants used in

Kenya and Tanzania. Cunningham ( 1 990) reported a mas-

sive trade in herbal medicines among the Zulu people of

South Africa. Other publications on medicinal plants of

Africa include those of Watt & Breyer-Brandwijk (1962)

on Medicinal and poisonous plants of southern and east-